fy ABS TE

OF THE

COUNCIL

AND

OFFICERS

OF THE

ZOOLOGICAL

SOCIETY

OF LONDON.

1908.

COUNCIL.

His Grace Tor Duke or Beprorp, K.G., President.

Grorce A. Boutencer, Esq.,

F.R.S., Vice-President.

Prof. J. Rosr Braprorp, M.D.,

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

Aurrep H. Cocks, Esq., M.A.

THe Rr. Hon. THe Hari oF

Cromer, P.C., G.C.B.

CHARLES DRUMMOND,

Treasurer.

Sir Epwarp Duranp, Br., C.B.

Freperick Gituerr, Esq., Vice-

President.

F. Du Cant GopMman,

IDClly, IOS,

dent.

THe Marguis or HAmMILton,

MP.

Ksq.,

Vice-Presi-

EK. G. B. Meape-Watpo, Esq.

Pror. Epwarp A. Mivycuty,

M.A., Vice-President.

P. Cuatmers MircHEni, Ese

MENS IDSihs IDO), JIBS...

Secretary.

Tue Lorp Newton.

W. R. Ocitvie-Grant, Esq.

ALBERT Pam, Esq.

Tue Hon. N. CHartes Rotu-

SCHILD, M.A.

Davi Seru-Snira, Esq.

OLDFIELD THomas, Ksq.,

E.R.S.

A. Trevor-Bartrye, Esq., M.A.

Henry Woopwarb, Esq.,LL.D.,

E.RAS., Vice-President.

PRINCIPAL OFFICERS.

P. CHatmers Mircuent, M.A., D.Sc., LL.D., F.RS.,

Secretary.

Frank KE. Bepparp, M.A., F.R.S., Prosector.

R. I. Pococn, F.LS., Superintendent of the Gardens.

Henry G. Purmer, M.R.C.S8., Pathologist.

F. H. Warernouss, Librarian.

JoHN Barrow, Accountant.

W. H. Cots, Chief Clerk.

ArrHur THomson, Assistunt Superintendent of the Gardens.

LIST OF CONTENTS.

1908, pp. 1-430. .

January 14, 1908.

Page

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

Menagerie during the month of December 1907 ......... 1

Mr. R. Lydekker, F.R.S., F.Z.S. Exhibition of, and

remarks upon, an abnormally marked Leopard-skin

fromethe Meccan election © wee ueedtt nk Ut doticwe ee aot 1

—

. Description of a Biological Expedition to the Birket-el-

Qurun, Faytim Province of Egypt. By W. A. Cunnine-

TONS CAG. Pht: RE Ze Seeeenc crraeaee ons 2a eacmee sae ies 3

2. The Duke of Bedford’s Zoological Exploration in Hastern

Asia.—VI. List of Mammals from the Shantung

Peninsula, N. China. By Outprimtp THomas, F.R.S.,

BY Ai Soin, eae eel: Saale era dict cabie Ete MOM amet ct cee, SF Oe 5

3. On the Musculature and other Points in the Anatomy of

the Kngystomatid Frog, Breviceps verrucosus. By

Frank E. Bepparp, M.A., F.R.S., Prosector to the

OCIS bY: rene sae sces sss teecenas SO bncidin cc. c.e bape deemed il

4, On the Hermaphroditism of the Amphipod, Orchestia

deshayesti Audouin. By Caries L. Boutencer, B.A.,

EE Z2S ines) College, @amplrrde tierce. -eahin..42 shee 42,

February 4, 1908.

Mr. F. Martin Duncan. Lantern exhibition of, and remarks

upon, the Autochrome Natural Colour Process of the

loreayeloveras) I DjrmuouereS | soadnGoneenomapeoeonSereesacaspsbeboonseocoaces 47

1. The Duke of Bedford’s Zoological Exploration in Hastern

Asia.—VII. List of Mammals from the 'T'su-shima

Islands. By Ouprizup THomas, F.R.S., F.Z.8. ......... 47

2. On the Presence of Gonadial Grooves in a Medusa,

Aurelia aurita. By 'T. Goopry, Student Demonstrator

in Zoology, University of Birmingham. (Plate I.)...... DD

3. The Duke of Bedford’s Zoological Exploration in Kastern

Asia,— VIII. A Collection of Fresh-water Fishes from

Corea. By C. Tare Recan, M.A., F.Z.S8. (Plates IT.

ii BLED Ta Se ee en cs Ag oer ee RTRs ar OI Oem CARRS 59

February 18, 1908.

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

Menagerie during the month of January 1908 ......../... 63

Mr. R. I. Pocock, F.L.8., F.4.8. Exhibition of (on behalf of

Mr. W. Simpson Cross, F.Z.8.), and remarks upon, a

photograph of the rare South-American Wild Dog

COLE OS SITET TIDUS) pM Oi Ree AAR ics titel ce alls ci 64

Mr. R. H. Burne, F.Z.S. Exhibition of, and remarks upon,

preparations of the Olfactory Organs of a Sea-Lamprey

(Petromyzon marinus), Sea- Bream (Pagellus centro-

dontus), and an Albatross (Diomedea exulans) ............ 66

Dr. L. W. Sambon, F.Z.S. Exhibition of a large series of

Internal Parasites from animals recently living in the

Societys (Gardens . {os sg5.0s eee solar cco ae ep erate 66

1. On the Inheritance of Colour in Domestic Pigeons, with

Special Reference to Reversion. By RicHARD STaPpLEs-

Browne, M.A., F.Z.8. (Plates EV.—VII.) ...2........... 67

2. The Duke of Bedford’s Zoological Exploration in Hastern

Asia.—IX. List of Mammals from the Mongolian

Plateau. By OuprreLp Tuomas, F.R.S., F.Z.8. ......... 104

‘3. Descriptions of new Species of Butterflies of the Division

Rhopalocera from Africa and from New Guinea. By

G. T. Bernune-Baker, F.LS., F.Z.8. (Plates VIII. &

TX.)

COC ewe tee eee eee emcee ones esse ner ser eerste eseeeeseeeeereeseeeeere

March 3, 1908.

Mr. R. Trimen, F.R.S., F.L.S., F.Z.8.. On a long spur or

claw-like horny excrescence at the tip of the tail of a

CIOIIESIATE NIN Joke oman ici Bo ApUHE BU nur aR oae a nnnA se, cobmdt oe

The Secretary. Exhibition of, and remarks upon, some

skins of the Coypu, Myocastor coypu ................0..eceee

Mr. F. HE. Beddard, M.A., F.R.S., F.Z.S. Exhibition of,

and remarks upon, the Colon and Rectum of a Badger

Clfelestiivele s) Pac tatetet oc nace an a NS nese ous hme aha da 2

1. On a young Female Kordofan Giraffe. By P. CHAaLmMmErs

MircHet., M.A., D.Sc., LL.D., F.R.S., Secretary to the

ASOCLO DVN Nair ter eta ed ceok SAM uate iehgdls haaraiml Valsad etn etn t a

2. A Comparison of the Neotrapical Species of Corallus,

C’. cookit, with Corallus madagascariensis ; and on some

Points in the Anatomy of Corallus caninus. By Frank

E. Bepparp, M.A., F.R.S., F.Z.8., Prosector to the

NOCIOUY, « Siiu.tisiconued aataseeeounhh oy sa ua hcueG esate ee mae

3. Description of a new Species of Monkey of the Genus

Cercopithecus. By R. I. Pococn, F.L.8., F.Z.S., Super-

intendent of the Gardens. (Plate X.) .....................

March 17, 1908.

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

Menagerie during the month of February, 1908 .........

Mr. G. A. Boulenger, F.R.S., F.Z.8. Exhibition of, and re-

marks upon, a remarkably malformed Plaice (Plewro-

NECLES) DUGLESSE)) ris. 5 os. Ys RRR ee eRe ete Nt ts! tat att

Mr. Walter A. Kidd, M.D., M.R.C.S., F.R.S.E., F.Z.S. An

account of observations regarding the effects of pressure

on the direction of hair in Mammals......................55

Page

110

135

158

161

val

1. The Rudd Exploration of South Africa.—IX. List of

Mammals obtained by Mr. Grant on the Gorongoza

Mountains, Portuguese §8.E. Africa. By OLDFIELD

Tuomas, F.R.S., F.Z.8., and R. C. Wrovucuton, F.ZS8. .

2. Notes upon some Species and Geographical Races of

Serows (Capricornis) and Gorals (Nemorhedus), based

upon Specimens exhibited in the Society’s Gardens. By

R. I. Pocock, F.L.8., Superintendent of the Gardens ...

April 7, 1908.

_ Mr. Thomas Codrington. Exhibition of a collection of stones

taken from the stomach of an Klephant shot in Northern

TBS TOG [GR pn SN re RON ate ha enti ae OV cers NU Ae

Dr. C. W. Andrews, F.R.S., F.Z.8. Exhibition of a restored

model of the skull and mandible of Prozeuglodon atrox

ANTON lal Rite SOON A weeds ke: Cerne RUD MAME A eee DARE: bk iA

The Secretary. Exhibition of a photograph of two young

living examples of the Forest-Pig of Central Africa

(Hylocherus meinertzhagent Thomas)...........c0cceeeeeeees

The Secretary. Notice of the successful acclimatisation of

Clananons sia INGiy /Agalevagl., .:aacccocsacopndnobcooosob aucouosouue

1. A Monograph of the Chiropteran Genera Uroderma, En-

chisthenes, and Artibeus. By Knup ANDERSEN............

2. On certain Points in the Structure of the Cervical Ver-

tebre of the Okapi and the Giraffe. By Sir Ray

IO Nigeaasnises IRC(ORI Bh IMAL acy INAAIS» | oy neeneansnansoanehecss

3. Some Australasian Spiders. By H. R. Hoge, M.A., F.Z.8. ¢

April 28, 1908.

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

Page

164

173

203

Menagerie during the month of March 1908 ............ 345

Myr. C. Tate Regan, M.A., F.Z.S. Exhibition of, and remarks

upon, an Australan Cat-fish (Cnidoglanis megastoma

LESIIOITE SCS): | ERT SOME Pee ACS cA auomaAMC pa ypenGeodD sande 345

Mr. Walter Burton, F.Z.8S. Exhibition of some melanistic

amidgblacls Leopard (skanas oly. gene ener an ner eee rae eee 346

vil

Page

Prof. D’Arcy Wentworth Thompson, C.B., F.Z.8. Abstract

of a paper on the Shapes of Megs .........B..............+5 346

1. A Revision of the Sharks of the Family Orectolobide. By

C. Tate Ruean, M.A., F.Z.8. (Plates XI1—XIII.)...... 347

A Note on the Occurrence of a Species of Phreatothria

(Vejdovsky) in England, and on some Points in its

Structure. By Frank E. Bepparp, M.A., F.RBS.,

F.Z.S., Korresp. Mitgl. d. K. Bohm. Ges. Wiss., &c. ... 365

3. On the Amphipod genus Tvrischizostoma. By H. W.

SPO O Newey (alates Nel Wa NONE) oe sy rac bee ask et aerecs 370

4, On certain Errors with reference to George the Fourth’s

Ciraties) (by HENRY SCHBRREN, HeZiS. G-cenc csc. seaee ase 403

. On the Breeding-Habits of a Cichlid Fish ee nilo-

tica). By Cuaries L. Boutencer, B.A., F.Z.8. ......... 405

6. ‘A Revision of the Oriental Pelobatid Batrachians (Genus

Megalophrys). By G. A. BouLtencer, F.R.S., V.P.Z.S.

(GET EsTe ST NO NUIET NONE Gisele asreotot le oa = intso oats oteineceaiclsle wsloaleui 407

ANTE dalv Ie OMEGA tin ibis al

OF THE

CON hi OU AOuns:

With References to the several Articles contributed by each.

(1908, pp. 1-430.)

ANDERSON, KNup.

A Monograph of the Chiropteran Genera Uroderma,

IEP OMSHORVES , MUNG GAIT bonanoe oops ocddbabondes sousbabcesLeuns

ANDREWS, CHARLES WILLIAM, B.A., D.Sc., F.R.S., F.Z.S.

Exhibition of a restored model of the skull and

mandible of Prozeuglodon atrox And. ...............00...08

BEDDARD, Frank E., M.A., F.R.S., F.Z.8., Prosector to the

Society.

On the Musculature and other Points in the Anatomy

of the Engystomatid Frog, Breviceps verrucosus ............

v Exhibition of, and remarks upon, the Colon and

Rectum of a Badger (Jeles meles)

oOo cho cron BOO ao oro Dae OOOO

A Comparison of the Neotropical Species of Corallus,

C. cookii, with Corallus madagascariensis ; and on some

Points in the Anatomy of Corallus caninus ............+5-

Page

204

203

Bepparp, Frank E. (Continued.)

A Note on the Occurrence of a Species of Phreato-

thrix (Vejdovsky) in England, and on some Points in its

SSGIOUG DUO Reape aret hel os 2 ie rcs Bie ns teas oleate Ree A A ene

Beruune-Baker, G. T., F.L.S., F.Z.S.

Descriptions of new Species of Butterflies of the

Division Rhopalocera from Africa and from New Guinea.

(LLNS SWELL Gs) OS Coe at deeee ee ORC eB

Boutuncer, Cuartes Leorotp, B.A., F.Z.S., King’s College,

Cambridge

On the Hermaphroditism of the Amphipod, Orchestia

MELSON ESI NGO Zrecabisnacue an vbdcunosneddseasnuneeosbosabaees

On the Breeding-Habits of a Cichlid Fish (Tilapia

UMW CRIS eRe Oe Ne AS Re ed RN A ene Nl a

Boutencer, Grorcr AuBert, F.R.S., V.P.Z.S.

Exhibition of, and remarks upon, a remarkably

malformed Plaice (Pleuronectes platessd) ...0.....c00cceeecees

A Revision of the Oriental Pelobatid Batrachians

(Genus Megalophrys). (Plates XXII.-XXYV.) ............

Browne, RicHArpD Srapies-, M.A., F.Z.S.

On the Inheritance of Colour in Domestic Pigeons,

with Special Reference to Reversion. (Plates IV.—

AV ELE) PRAMS Ste SA EL ie) hay 2) DOE LE Eg OR Ran, PRO

Burne, R. H., F.Z.S.

Exhibition of, and remarks upon, preparations of the

Olfactory Organs of a Sea-Lamprey (Petromyzon marinus),

Sea-Bream (Pagellus centrodontus), and an Albatross

(Ovomeded erullaiis Wenge. ie te ee ee

Page

365

110

405

16]

407

Page

Burron, WAurErR, F.Z.S.

Exhibition of some melanistic and black Leopard

SET a Sa et eRe ee ee NR ee, Ba ie ecg cea eee 346

Coprineron, THOMAS.

Exhibition of a collection of stones taken from the

stomach of an Elephant shot in Northern Rhodesia ...... 203

Cross, W. Smupson, F.Z.8. See Pocock, R. I.

Cunnineton, Wiii1aAM Aurrep, B.A., Ph.D., F.Z.8.

Description of a Biological Expedition to the Birket-

el Ourun, Hayumy Provimeeiot Hey pti 2.7.20... .00 sees 3

Duncan, F. Martin.

Lantern exhibition of, and remarks upon, the

Autochrome Natural Colour Process of the brothers

TBR RI aICTA Sioa ast en eee Ur Melee it see ter alae Al ela ai AT

Goopgy, T., Student Demonstrator in Zoology, University

of Birmingham.

On the Presence of Gonadial Grooves in a Medusa,

Aloo, aaron (Weenie ID sect conde ssaese tbabcesosbaceodocnHec

Hoce, Henry Roveuton, M.A., F.Z8.

Storing) AS UTeNIISTE NT SPONMOWETES roc cugoonoss ooo snononsobuevogouod 339

Kipp, Water, M.D., M.R.C.S., F.B.8.E., F.Z.8.

An account of observations regarding the effects of

pressure on the direction of hair in Mammals ............ 164

X11

Lankester, Sir Epwin Ray, K.C.B., M.A., D.Sc., LL.D.,

ID IRIS)

"On certain Points in the Structure of the Cervical

Vertebrze of the Okapi and the Giraffe .................05..

LyprKkker, Ricuarp, B.A., F.R.S., F.Z.S.

Y Exhibition of, and remarks upon, an abnormally

marked Leopard-skin from the Deccan ..................0+5

Mircnent, P. CHaumers, M.A., D.Sc., LL.D., F.RBS.,

Secretary to the Society.

Report on the Additions to the Society’s Menagerie

during the month of December 1907 ..................0.0005

Report on the Additions to the Society’s Menagerie

during the month of J EHaTee wl MEXOKS) sagan aobiocedosnaee ts oods

Exhibition of, and remarks upon, some skins of the

COO, WICCUSEOIP CONFIG sdoodocodsagearsoodosoonsnoasnenesaotes

VOna young Female Kordofan Giraffe ...............-.....

Report on the Additions to the Society's Menagerie

during the month of February 1908 ........................

Exhibition of a photograph of two young living

examples of the Forest-Pig of Central Africa

(Hylocherus meinertzhagent Thomas) .............0.0.0.0.00-

Report on the Additions to the Society's Menagerie

Comat wae imaorain Oe Wleyeeln WYOS .coscc0cvosccovasodosne scone

Pocock, Rucrnap L., F.L.S., F.Z.S8., Superintendent of the

Gardens.

’ Exhibition of (on behalf Mi. W. Simpson Cross, F.Z.8.),

and remarks upon, a photograph of the rare South-

American Wild Dog (Canis jubatus) 2. \cverecese.+-sceenes-

Page

320

161

208

345

Xill

Pocock, Rreinaxp I., F.L.S., F.Z.S., Superintendent of the

Gardens. (Continued).

sé Description of a new Species of Monkey of the Genus

OG REO COINAGUIS, 1 (IEIBIKE D0) cso cteorcatoeausagoos Soauseaot se sAbee: <

Notes upon some Species and Geographical Races of

Serows (Capricorims) and Gorals (Vemorhedus), based

upon Specimens exhibited in the Society’s Gardens

Reean, C. Tare, M.A., F.Z.8., of the British Museum

(Natural History).

The Duke of Bedford’s Zoological Exploration in

Hastern Asia.—VIII. A Collection of Fresh-water Fishes

from Corea. (Plates II. & IIT.)

Exhibition of, and remarks upon, an Australian

Cat-fish (Onidoglanis megastoma Richards. )

A Revisien of the Sharks of the Family Orectolobide.

(GEA Ere SED 9 Te) I IL Nn See avai tore hah oo ME ne Ae

SamBon, L. W., M.D., F.Z.S.

Exhibition of a large series of Internal Parasites from

animals recently living in the Society’s Gardens............

SCHERREN, Henry, F.Z.S.

\ 5 O

On certain Hrrors with reference to George the

Io urcih’ sy Gaiea they sc, 22 ssajsecae Gee ce eee rman, a

Sexton, E. W.

On the Amphipod genus Trischizostoma. (Plates

DOI= KOMBO onal oR RS tee ky

Page

347

403

Page

SraPLes-Browne, R. See Browne, R. Srapes-.

THomas, OLDFIELD, F.R.S., F.Z.S.

“The Duke of Bedford’s Zoological Exploration in

Eastern Asia.—VI. List of Mammals from the Shantung

Peninsula, N. China

The Duke of Bedford’s Zoological Exploration in

Eastern Asia,—VII. List of Mammals from the Tsu-

Slouniayaysl LSI eyaiG ee Raney ph tants UL Nie ice ins eek Re oH. 47

“ The Duke of Bedford’s Zoological Exploration in

Eastern Asia.—IX. List of Mammals from the Mongolian

Plateau 104

ee eken ena omeee sie see4 ine heysvelalesaleialeeleleiselielelataelalvlaleteialslalpleletsieleleleielelataterctercrst ere

Tuomas, OLDFIBLD, F.R.S., F.Z.8., and Wrovcuron, Roperr

CHARLES, F.Z.S.

The Rudd Exploration of South Africa.—IX. List

of Mammals obtained by Mr. Grant on the Gorongoza

Mountains, Portuguese 8.H. Africa ...............2.0.0cc000e. 164

Tompson, Prof. D'Arcy Wentworru, C.B., F.Z.8,

Abstract of a paper on the Shapes of Kegs 346

Trimen, Rouanp, F.R.S., F.L.S., F.Z.S8.

On a long spur or claw-like excrescence at the tip of

the tail of a Domestic Cat 12

FIDO OOD Gono MdDOOCoODDOODOUOODabGanO

WroucutTon, Roperr Cuarnes, F.Z.S., and THOMAS,

OLDFIELD, F.R.S., F.Z.S.

Y The Rudd Exploration of South Africa.—IX. List of

Mammals obtained by Mr. Grant on the Gorongoza

mountains, Portuguese S.K. Africa

LIST OF PLATES.

1908, pp. 1-430.

Plate Page

I. Gonadial Grooves in Awreha aurita .............. 5b

II ie Acanthorhodeus gracilis. 2. Leucogobio strigatus >

CONDO UL LENTES |OCCUION (htm Mel Neh titer et ccek cute ofcberel se soneusus \ 59

.! 1. Ctenogobius bedfordi. 2. Tridentiger coreanus { ~~

"| 3. Acanthogobio longirostris. 4. Liobagrus andersoni. )

IV. ( m1 A :

V.4 Barb=HanballeCrosste yum sceecicn eames ceo maceey:

Vil. 1. Barb-Fantail Cross. 2. Tumbler-Fantail Cross. . 67

WiIUL “Winvealnllermltermimrll ORasS5.0c0000c00ccsdancnogoovobe

Mae) N : : :

op New ee entile=knonwmeyccenvd cera ee eee ee 110

NC New Speciesh ale CZi copii Necis maser nines 158

XI. 1. Chiloscyllium griseum. 2. Orectolobus ornatus e))

NIT. 1. Chiloscyllium plagiosum. 2. Orectolobus tentaculatus. i 347

XIU. 1. Chiloscylhum grisewm. 2. C. indicum ..........

XIV. \ -

DXGV RS el 1S Clit Z OSLOTIUGICCECIIS Cm nae en eee

XV LJ |

XVII. 1-12. Trischizostoma niceense. 13. T. raschii .... | Fe

MOVIUUL, — IPOSOCPROREOMIG TROSG “s scab ot advonboccopoancuonE eu

XIX. 1. Tiischizostoma niceense. 2-11. T. raschii...... |

SOG I SOUFUSSOMUO ROSES gp ocodconababecbooavoonogcs |

XXI. 1-13; 15-18. Trischizostoma raschit. 14. T. niceense )

OM LGR OOS WUNHO Sooo nccaseovesneoddoodaodous a

MOSOUG | WACO VORSTOYOR soocchcooscdesaucounguanonut

XOX Eco alophinys 7 OGUSCOMa ar Petar eee ase saree Lae

XXV. 1. Megalophrys gracilis. 2. M. heteropus. 3. M, |

(WBS “ono nacec RS OR MRE ib iene Jalcuater Si begs ne)

Proc. Zoou. Soc.

LIST OF TEXT-FIGURES.

1908, pp. 1-430.

Page

1. Abnormally marked Leopard-skin from the Deccan .......... 2

2. Anterior cornua of hyoid and part of hyoidean musculature of

BRCUICEDSY A sl peers Hn eat Thy Ur eae, fates ae aha 12

3. Superficial ventral musculature of Breviceps ......... 00.000. 16

4. Superficial dorsal musculature of Breviceps ........... 0.05. 20

5. Ventral abdominal musculature of Breviceps cut so as to display

ditkerent layers. Merve wees he oeehchita See ORRIN, F 23

6. Muscles of the inside of the thigh of Breviceps .............. 25

7. CXsophageal and neighbouring muscles of Breviceps .......... 28

8. Ventral surface of liver of two individuals of Breviceps ...... 30

9. Part of oviduct and stomach of Breviceps (laid open) ........ ol

HOF Intestinaltcanaliof Breviceps ea). snteee ac whenae ce oreo eine © 32

i mWettihymph-heartoneBreviceps. sa eine ama len eaehet ene eae 35

12. Lymph-heart of Breviceps isolated; and view of interior of

SEO Cd io iiceca hh cnc uutaeicn ica bb bach het icup iG iGo. aia heor oaabeactokere Soci 36

jm estesvandukadneys OLManeCveCep Simi t immer eres tat a seen 38

14, Testes of two individuals of Orchestia deshayesii, to show the

positionvor the aval. Exe4ON ATi were Me allel. oy. een eck Mas, 44

lion ManedeWiolit(Canisijubatus) Pron litce ase ers ac a sens ae 64

16. Section of the head of an Albatross (Diomedea exulans) ...... 65

17. Colon of Badger (Meles meles) cut open to show Peyer's

Deube a PRaNe ns 2 SS) cea ae Cd eaten eRe Cette HEART, 129

18. Female Kordotan Giraffe about one week old .............. 15

1OMbendor Kordotan Giratiercaligaemersnt rts Henares i) Peer 152

720) EleadvorelMordotan’ Giratie) caliquenm erty. ere orate: 135

21. Larger lung of Corallus cook cut open to display the entrance

MERSIN) OI WAS WRC, Sooccnnvocosnnoocccvesoanuncuvac 139

1908. é;

22. The larger lung of another example of Corallus cookin .

. Larger lung of Python spilotes

. The liver and certain veins connected with it of Co irae

COOKIE aah ene anes dee MCS Oras oor ae Pir ecors

Posterior end of liver of iw ) andi iduals of Corallus coat

Pancreas and adjacent viscera of Corallus cookii ...

Lungs of Corallus caninus cut open to show interior .....

Coloured side of malformed Plaice, coloured end .

. Blind side of malformed Plaice .......

Skull of Capricornis sumatraensis thar Hodgs..........+...055

. Skull of Capricornis sumatraensis humet Poe.

. Flat skin of Capricornis sumatraensis rodont Poc.

Skull of Capricornis sumatraensis rodont Poe.

. Capricornis sumatraensis jamrachi Poe.

Capricornis sumatraensis robunsont Poe. ..............-+.+-s

. Skull of Capricornis sumatraensis robinsoni Poe.

Nemorhedus goral Hardw.

Nemorhedus raddeanus Heude

. Young Forest-Pigs (Hylocharus meinertzhagent) .........4+5

Right upper and left lower tooth-row of Rhinolophus

trifolidtus . 2:5. - ahs :

Right upper, left laiver. tooth-row, side view of skull and

mandible, and front view of incisors and canines of Artibeus

UOTE. O SEV TSE) QULG Sia ares pes Bed eaten Cay MTOR MAS Te eee

Upper, lower, side view of skull, aud front view of incisors

and canines of Uraderma bilobatum

Upper, lower, side view of skull, and front view of incisors

and canines of Artibeus cinereus cinereus... 2.2 eee

Right upper and left lower tooth-row of Uroderma bilobatum . .

Right upper and left lower tooth-row of Enchisthenes harti....

. Right upper aud left lower tooth-row of Artebeus planirostris

LUE ae 5 Meee eet eee RT I

Front view of lower incisors and canines of Uroderma thomasi . 2

Front view of lower incisors and canines of Artibeus jamaicensis

WEUTOURUS ss Fk Shouse «) «in 2085). ESOT As oe Ie ac HA haawele

. Right upper, left lower tooth-row, and front view of upper

incisors and canines of Enchisthenes harti

| Lateral and upper view of skull of Artidbeus jamaicensis pal-

Manor iit tcc d ao Myson sith Sheteaeleas SO RM AEN oa eo Sith Meat

Right m?, to show strong emargination of hinder margin a

tooth of Artibeus jamaicensis jamaicensis, and right m2, to

show slight emargination of hinder margin tooth of Artibeus

GAMGVCENSUS WELUNGLUS) | -7....1- = 2 Ee eee

Right upper tooth-row of dr Rios nes and A. jamaicensis

JOR ET ODS “S855 cbc A ECG o8 on.4 Bore oe ieee te

Right upper tooth-row of Artzbeus cinereus cinereus

Said

Page

58. Right upper and left lower tooth-row of Artibeus sosenbergr .. 294

56, Right upper tooth-row of Artibeus toltecus ravus and A. cinereus

CUNLED CUSHA Rin sb ate OSS aE EMR UEC CP ben POP uae a aera eA 296

She ars ee ae 92 URC OCT OUI scncpoiaeeko cigs edt: OMA Slo ROA CHARS DOME E 307

Bey roland Ore V2VROUAIS UOUEACUIS GOMES 55 Ga od nee oceashneoedacson 307

59. Sketch of distribution of the races of Artibeus gamaicensis in

Mexico, the West Indies, and Northern S. America........ 319

60. The last three cervical vertebre and the first dorsal vertebra of

ther Gurateyseen tromutherlett siden i alee i yuan 32]

61. The last three cervical vertebree and the first dorsal vertebra of

phe Olapimseenmramb te letuisid ers aera sere oe ale 322

62. The last three cervical vertebre and the first dorsal vertebra of

audomestic Ox, seenuromithe leftisides.j.5..4.408 4000.0. 325

63. The last three cervical vertebree and the first dorsal vertebra of

the @amel, seen from the left side ...7....50....1 40.055. 325

64. The posterior face of the seventh cervical vertebra and the

anterior face of the first dorsal vertebra of the Okapi ...... 326

65. The posterior face of the seventh cervical vertebra and the

anterior face of the first dorsal vertebra of the Giraffe ...... 327

66. View of the articular surfaces which connect dorsal vertebra 1

and dorsal 2 of the Okapi, and of the anterior half of the

articulation between dorsal 2 and dorsal3 ................ 328

67. A similar view of the corresponding vertebree of the Giraffe .. 329

68. The articular facets of the left side on the posterior face of

Cervical vertebra 7 and the anterior face of Dorsal 1 of

Rhinoceros sumatranus ........... rb cUsarenareniaks MERREBL LAURE 2 Spoken 392

69, The articular facets of the left side on the posterior face of

Dorsal vertebra 1 and the anterior face of Dorsal 2 of

UEC RES UM AERCNULSE rn aa e RR Rie eh ne 332

70. Similar drawing, to show the duplicate facets connecting Dorsal

vertebra 2 and Dorsal 3 in the same animal .............. 333

71. Similar view, to show the single facets connecting Dorsal

vertebra 3 and Dorsal 4 in the same animal ........... .. 303

BR EMSSULEN GS OCCALONUAINV al cleans eater nieve ere ae cee 335

Ts ERATION TLDS) TS) Uh UE ie le Aiton dia do bo dns Ao OS bye enone 338

(A Dolomedeshutcone, sp. 0.) ablenraleyen see yeit iar se ental 341

ROD olomedest tipper, Sp. Ne. 5. cane eee ee eve aerate ae ne che 345

76. Ventral pair of setee of segment I. of Phreatothrix canta-

NGS SUSI at RERPCRRE ORS Con, oo oct, 0 60) so 0 Batya Nim ous eon eee 366

77. Anterior end of dorsal and ventral vascular trunks of Phreato-

ALD CH ATIO UO CSU MENS © Y Bic 60d 8st HDG O HOO SHE SH ROTO CEE 367

. Vertebral column of Megalophrys pelodytoites, ventral aspect .. 409

LIST OF NEW GENERIC TERMS

PROPOSED IN THE PRESENT VOLUME (pp. 1-480).

Page Page

Callictita (Lepidopt.) ......... 118 Powellana (Lepidopt.) ......... 114

Hucrossorhinus (Pisces) ...... 307 Thaumaina (Lepidopt.)......... 116

Mambara (Lepidopt.) ........ 119 Upolampes (Lepidopt.) ......... 118°

Acanthogobio

longirostris, 60, 63.

Acanthorhodeus

gracilis, 60, 63.

Acidostoma, 373.

Acomys

selousi, 172.

Acontiostoma, 373.

Acrea

bent, 110.

Allactaga

annulata, 109.

mongelica, 109.

Alytes, 39.

Amaryllis, 373.

Amblystoma

tenebrosum, 146.

Ammoceetes, 47.

Antilocapra

americana, 161.

Antilope

bubalina. 178.

crispa, 196.

goral, 192, 195.

interscapularis, 176.

sumatraensis, 175, 176.

thar, 176.

(Caprina) crispa, 196,

198.

(Nemorhedus)

176.

Apodemus

agrarius, 7, 8, 9, 48.

coree, 8, 9.

geisha, 48, 54.

— sagax, 48, 54.

hokkaidi, 54.

japonicus, 54.

mantchuricus, 8, 9.

minutus, d4.

ningpoensis, 8, 9.

pallidior, 7, 8.

peninsule, 5d.

speciosus, 6, D8.

thar,

INDEX.

Apodemus

speciosus

48.

— peninsule, 48, 53.

ussuricus, D4,

Apteryx, 66.

Apus, 47.

Arctibeus

carpolegus, 267.

jJamaicensis, 267.

perspicillatus, 267.

Ariteus

achradophilus, 267.

Artibeus, 212, 214, 221,

222, 223, 224, 295.

sp., 278, 279.

aztecus, 204, 226, 227,

228, 229, 282, 290,

296, 297, 300; 3

306, 309, 310, 313,

314, 315, 316.

bilobatus, 217, 218.

carpolegus, 228,

265, 267.

cinereus, 204, 226, 227,

229, 230,

290, 291, 292,

294, 295, 296, 297,

5 UR, ail), Gils}

314, 315, 516.

— bogotensis, 204, 228,

229, 291, 292,

295, 313.

— cinereus, 204, 21

229, 290, 291, 29

298, 296, 313.

concolor, 204, 217, 225

, 224, 228, 229,

232, 233, 234,

245, 246,

, 286, 308, :

, oly, BIG,

, 228, 265, 268,

spectosus,

Proc, Zoou, Soc,—1908,

Artibeus

fallax, 2N7, 227, 228;

230, 231, 234, 242,

243, 244,

Femurvillosum, 228,

278, 282, 283.

glaucus, 204, 226, 227,

228; 229 932° 260)

285, 286, 287, 288,

289, 290, 291, 292

294, 296, 310, 312,

314, 315, 316.

grandis, 228, 272,

hercules, 228, 242,

244, 245, 316.

hirsutus, 204, 217, 2

228, 229,

237, 242,

1 26D 28a;

, d12, 314,

316.

insularis, 228,

270.

intermedius,

248, 265, 278,

281.

jamaicensis, 204,

216, 224, 226,

228, 229, 2

275,

243.

26,

231,

232, 233, 234, 235

238, 239, 243,

249, 250,

253, 254, 257,

260, 261, 262,

264, 265,

268, 270, 272,

274, 276, 281,

284, 285, 286

291, 299, 3

312,

317.

— equatorialis,

228, 229, 249,

252, 253, 254,

204,

250,

255,

XX11

Artibeus

257,

Ze 22)

285, 312,

319.

Jamarcensis domiunt-

canus, 249, 250, 252.

— janaicensis, 204,

229, 230, 245,

W2ale

, 296, 257,

, 260, 262,

266, 267,

, 20), 2

278,

285, 286,

317, 318, 319.

— lituratus, 204, 216,

8, 229, 230, 289,

, 241, 247,

» 251, 252; 253)

, 299, 207, 258,

259, 260, 272, 278,

275, 276, 277,

5, 280, 281, 283,

305, 312, ¢

273, 284,

317, 318,

258,

263,

268,

272,

281,

318.

— palmarum, 204, 228,

, 248, 249, 250.

» 208, 254, 255,

7, 258, 259, 260,

, 273, 274, 278,

, 280, 281, 282,

284, 285,

317, 318, 319.

— parvipes, 204,

229, 247, 249,

2, S 255,

258, 259, 260, 261,

262, 264, 266,

272, 275, 312, 319.

— preceps, 204, 228,

229, 249,

264, 265, 270,

274, 288, 284, 285,

312, 317, 318, 319.

— yucatanicus, 204,

228, 229, 247, 249,

250, 252, 253,

255, 257, 258, 259

228;

250,

257,

Ore

260, 261, 263, 264,

265, 266, 272, 273

284, 312, 317, 318.

lineatus, 230.

litwratus, 228, 252, 253,

257, 258, 259, 260, |

272, 284, 319.

macleayt, 228, 261.

nanus, 204, 226, 227

228, 229)

297, 300, 804, 305,

, 260, 270, |

252, 253, |

SD, ||

312, |

254, 260, |

109, |

INDEX.

| Artibeus

asl, |

232, 296, |

307, 308, 309, 310,

313, 314, 315, 316.

palmarum, 228, 275,

276, 278, 280, 281,

283, 319.

parvipes, 228, 2538, 254,

260, 262, 263, 264,

266, 284.

perspiccilalune, 261.

perspicillatus, 227, 228,

230, 231, 234, 235,

253,

264,

272)

280,

236,

260,

239,

261, 263,

265, 268, 270,

274, 278, 279,

281, 283.

pheotis, 204, 226, 227,

228, 229, 232, 300,

243,

303, 304, 305, 306,

309, 310, 318, 314,

315, 316.

planirostris, 204, 205,

212, 216, 217, 226,

PP, Deieh, BBM, BR,

235, 237, 238, 239,

240, 241, 242, 244

245, 246, 247, 251,

252, 253, 254, 260,

261, 265, 268, 271. |

278, 274, 278, 285, |

286, 310, 311, 314,

315, 316, 317.

— fallax, 204, 205,

208, 215, 228, 229,

230, 234, 235, 236,

237, 242, 243, 244,

245, 247, 260, 312,

316.

— grenadensis, 204,

228, 229, 237, 241,

242, 312, 316.

— planirostris, 204,

228, 229, 230, 233,

234, 235, 286, 237,

239, 240, 241, 249, |

243, 250, 260, 274,

287, 311, 316.

— trinitatis, 204, 228,

229, 233, 236, 237,

241, 242, 245, 247,

260, 312, 316.

quadrivittatus, 204,

226, 227, 228, $

232, 293, 296, 302,

303, 807, 308, 3809,

310, 3138, 314, 315,

316.

rosenbergi, 204,

226, 227, 228,

231, 232, 288,

216,

229,

293,

Artibeus

294, 295, 296, 313,

314, 315, 316.

275,

rusbyi, 228, 272,

276.

sulphureus, 267.

toltecus, 204, 226, 227,

231, 232, 290, 294,

296, 297, 299, 300,

302, 308, 304, 305,

306, 307, 308, 310,

313, 314, 315, 316.

— ravus, 204, 224

228, 229, 296, 297,

298, 299, 300,

302, 3038, 304,

309, 315.

— toltecus, 204, 228,

299, 297, 298, 299,

300, 301, 302, 303,

306, 307, 309, 313.

turpis, 204, 227,

228, 229, 232, 296,

297, 300, 305, 307,

208, 309, 310, 318,

314, 315, 316.

watsont, 204, 226, 227,

228, 229, 232, 260,

286, 287, 288, 289,

290, 291, 292, 293,

294, 296, 300, 310,

312, 314, 315, 316.

yucatanicus, 228, 253,

254, 260, 261, 263,

264, 266, 284, 319.

(Dermanura) quadri-

vittatus, 302.

(—) rosenbergi, 293.

(Uroderma) bilo-

batus, 217.

Arvicanthis

dorsalis, 170.

Arvicola

mongolicus, 108.

Astacus, 45,

fluviatilis, 46.

Ateles

varicegatus, 161.

Atilax

vansirt, 167.

Aurelia

aurita, 55, 57, 58.

flavidula, 58.

Austritragus, 174.

Beodon, 206.

Barilius

platypus, 59.

Bdeogale

erassicauda, 168.

Bdeogale

tenwis, 168.

Bitis

nasicornis, 1.

Boa

constrictor, 137, 149.

divinilogua, 186, 137.

dumerili, 135.

madagascariensis, 135, |

Bombinator, 409.

Bos

sp., d24, 325,

taurus, 323,

Brachzlurus, 348.

modestus, 354.

Breviceps, 11-41.

gibbosus, 11.

verrucosus, 11

Bufo, 46.

vulgaris, 34.

Callictita, gen. noy., 118.

cyara, 119, 126.

Calliste

desmaresti, 161.

Candalides

angabunga, 123, 126.

aroa, 122, 126.

cyana, 122, 126.

dinawa, 122, 126.

gloriosa, 121, 126.

grandis, 121.

grandissima, 121, 126.

maria, 120,

newrapacunda, 121, 126.

pratti, 122, 126.

sublutea, 126.

unipunctata, 123, 126.

Janis

jpubatus, OF.

Capra

cylindricornis, 6+.

faleoneri, 194.

sibirica, 345.

Capricornis,

175, 176.

buhalinus, 178.

173,

174, |

sumatraensis, 176, 184, |

190.

— argyrochetes, 175,

185, 190.

—— humei, 178,

179, 190. |

-— jamracht, \75, 176, |

188, 184, 185, 190.

— mitine-edwardsi, 175,

185, 190.

— robinsoni, 175, 185, |

186, 187, 188, 190.

— rodont, 175, 180,

182, 183, 190.

INDEX.

Oapricornis

sumatraensis rubidus,

175.

-—— sumatraensis, 175,

185, 186, 187, 190.

— swettenhami, 175,

186, 187, 190.

— thar, 176, 177, 178, |

179, 181, 190.

thar, 178.

Carassius

auratus, 59,

Cardinalis

pheniceus, 161.

Cariacus

rufus, 161.

Carollina

brevicauda, 274.

Cemas

goral, 195.

Centetes, 1385, 324.

Centorhinus

maximus, 848, 353.

Cepbalophus

monticola, 173.

natalensis, 172.

Ceratohyla

bubalis, 410.

Ceratophrys

montana, 411, 415.

nasuia, 413.

Ceratophryne

montana, 411.

nasuta, 413.

Cercopithecus

alligularis beirensis,

165.

ascanius schnidti, 160

boutourlinii, 160.

diana, 159.

ezré, 108, 159, 160.

hamlyni, 160.

leucampyx, 160.

neglectus, 159, 160.

nigroviridis, 160.

rufoviridis, 165.

Cervicapra

arundinum, 173.

Cervus

sika, D4.

Chersydrus, 149.

Chiloscyllium, 353.

freycinet, 358, 359.

furvumn, 354.

griseum, 348, 358, 560, |

362, 364.

indicum, 358, 361, 362, |

364.

mataianum, 360.

margaritiferum, 362.

modestum, 354.

Chiloseyllium

ocellatum, 358, 359.

phymatodes, 362,

plagiosum, 358, 360,

362, 564.

punctatum, 348, 358,

350, 361.

trispeculare, 358, 359.

tuberculatum, 36%.

Chlorophthalmus

agassizt, 874, 396.

Chrysavra

isosceles, 57.

Citellus

brevicauda, 105.

mongoticus, 105.

Citrinophila

wunipunctata, 111.

Cnidoglanis

megastoma,

Columba

livia, 67, 74, 102.

Corallus

caninus, 135, 154,

156, 157, 158.

coohti, 135, 186,

138, 140, 142,

144, 145, 146,

148, 149, 150,

153, 154, 155,

158.

madagascariensis,

136, 1387, 1388,

143, 144, 146, :

149, 150, 151, 153,

154, 155, 157,

Cordylophora

lacustris, 5.

Cricetomys

adventor, 171.

gambianus, 171.

— cunctator, 171.

viator, 171.

Cricetulus

campbelli, \Q7, 108.

griseus, 10, 107,

108.

— obscurus, 107.

obscurus, 10, 107.

Crocidura sp., 166.

coreeé, 48, D1.

dsi-nezumi, 48.

Crossarchus

fasciatus, 16).

Crossorhinus, 3d4,

barbatus, 395, 356.

dasypogon, 397.

ornatus, 390.

tentaculatus, 3D

Ctenogobius

bed fordi, 62, 63,

ae,

oto.

XXIV

Ctenoplana

korotneffit, 57, 58.

Cyaniris

acesina, 117, 126.

biagi, 117, 126.

drucet, 117, 126.

owgarra, 116, 126.

Dactylethra, 19.

Dasypeltis

scabra, 149.

Dermanura

cinerea, 224, 228, 230,

292, 301.

eva, 228, 265, 267,

268.

Jucundum, 238, 303,

305, 306.

pheotis, 228, 3038.

quadrivittata, 230, 293,

305.

rava, 228, 300, 301.

tolteca, 230, 301, 306.

Deudoryx

eleala, 113.

elealodes, 112, 126.

iturt, 112, 126.

makala, 111, 126.

Diomedea

exulans, 65, 66.

Dipus

jaculus mongolica, 109.

(Jaculus) annulatus,

109.

Discoglossus, 39.

Dolomedes

huttoni,, 340, 341.

imperiosus, 340.

minor, 340, 344.

trippt, 340, 342, 343.

Dymecodon

pilirostris, 51.

Enchisthenes, 221, 225,

3ll.

harti, 204, 215, 2

223, 224, 226.

Engystoma

carolinensis, 13.

Enygrus

carinatus, 137.

Epitola

albomaculata, 126.

mangoensis, 126.

Equus

grevyt, 1.

Erinaceus, 326.

dealbatus, 6.

conicus, 187, 149, 156.

INDEX.

| Hryx

jaculus, 137.

Joh, 137, 156.

turcicus, 137.

Hucrossorhinus, gen.

nov., 348, 357.

dasypogon, 348, 357.

Hunectes

murinus, 136, 149.

noteéus, 136, 137, 142.

Eupagurus, 45.

Hupsychortyx

soning, 161.

| Hyotomys, 49.

Felis

manul, 161.

microtis, 48, 52.

onca, 345.

rufa, 345.

Funisciurus

cepapt, 169.

mutabilis, 169.

Galago

crassicaudatus, 166.

garnetti, 166.

granti, 166.

Galidia

elegans, 167.

galera, 167.

Gebia

major, 46.

Genetta sp., 166.

zambesiana, 166.

Geophagus

brasiliensis, 407.

Georychus

beire, 172.

| Ginglymostoma, 348.

brevicaudatum, 350,

351,

caboverdianus, 350.

cirratum, 350.

concolor, 300, 351, 352.

Serrugineum, 350, 351.

Sulvum, 350.

muelleri, 351.

rippellii, 332.

Girafta

camelopardalis

quorum, 130.

-~ typica, 134.

Guerina, 374, 398.

Guerinella, 398.

antt-

niceensis, 370, 374, 375.

Guerinia, 370, 398.

niceensis, 870, 871,

372, 874, 375.

Hapalemur

griseus, 128.

Helicops

angulatus, 146, 149.

Heloderma, 157.

Hemibarbus

barbus, 60.

Hemipbhractus, 410.

Hemiscyllium, 358.

modestum, 354.

ocellatum, 359, 360.

trispeculare, 359.

variolatum, 349.

Hemitragus

Jemlaicus, 194.

Herpestes

paludinosus, 167.

Heterodon

nasicus, 146.

Hexatihele

hochstetteri,

339.

huttoni, 337.

petreti, 337.

websteri, 337, 338.

Hipposiderus

caffer caffer, 282.

-— centralis, 282.

Holochila

grandis, 121.

Houbara

macqueent, 1.

Hylobates

hainanus, 1.

Hylocheerus

meinertzhagent, 203.

Hypocysta

aroa, 115, 126.

Hypolycena

naara, 112.

Hyrax

mossambicus, 172.

307,

Ichneumon

major, 167.

Inachus, 45.

Tolaus

bilineata, 118.

cottont, 113, 126.

Ixalus

lateralis, 416, 418.

Kemas, J73

henryanus, 200.

todinus, 201.

pinchonianus, 201.

raddeanus, 196, 198.

(Nzemorhedus) xantho-

deiros, 201.

Lafystius, 371.

Lagomys

ogotona, 109, 110,

Lates

niloticus, 4.

Lepidactylis, 371.

Leptobrachium

baluense, 429.

boetigeri, 420.

boulengeri, 425.

cartnense, 427, 429.

fee, 428, 429,

gracile, 421,

hasseltti, 425.

heteropus, 422.

montanum, 425.

monticola, 416, 419.

parvum, 419, 420,

pelodytoides, 428.

Leptomyrina

makala, 115.

Leptoptila

ochroptera, 345.

Lepus

dauuricus, 109.

ladacensis, 109.

ogotona, 109.

swinhoet, 10, 109.

tolat, 10, 109.

Leucogobio

guentheri, 60.

strigatus, 59, 60, 68.

teniatus, 60.

Lilljeborgia, 372.

Liobagrus

andersont, 59, 61, 62,

marginatus, 62.

nigricauda, 61, 62.

reine, 62.

styani, 62.

sugubii, 62.

Lithotragus, 174.

Lonchoglossa, 207.

Lophocheta, 368.

Lutreola

itatsi, 48, 53.

quelpartis, 53.

sibirica, 48, 53.

Lysianassa

magellanica, 371, 399.

Macropus, 128.

Madatzus, 224.

lewisti, 228, 265, 267.

Mambara, gen. nov., 119.

migropunctata, 120,

26.

Mangusta

urinatria, 167.

INDEX.

| Manis, 128.

| Martes

bedfordi, 52.

melampis, 52.

— tsuensis, 48, 52.

| Mazama

rufa, d4.

| Megalophrys

baluensis, 411, 429.

betigeri, 411, 420,

boulengeri, 4M, ADD.

carinensis, 411, 427.

chysti, 413.

fee, 411, 428.

gigas, 418.

gracilis, 411, 421, 430.

hasseltit, 411, 424, 425, |

430.

heteropus, 411,

430.

422,

NDEs, 407, 409, 410, |

Ao 409, 410, 416,

418, 450.

montana, 407, 408, 409,

410, 411, 4138, 415,

420.

-— aceras, 411, 412.

nasuta, 24, 27,407,409, |

410, 413, 415, 430.

parva, 408, 409, 411,

419.

pelodytoides, 409, 411,

24.

423, 4

robusta, 411, 418,

430.

Megophrys

monticola, 411.

Meles

meles, 128, 129.

| Meriones, 108.

psammophilus, 106.

unguiculatus, 106.

Micristodus

punctatus, 358.

Micromys, 7.

geisha hokkaidi, 54.

Micronycteris, 207.

Microtus, 49.

angustus, 108.

arvalis, 108.

gregalis, 108.

raddet, 108.

slowzouz, 108.

tianshanicus, 108.

(Stenocranius) slow-

zowt, 108.

' Missulena

occatoria, 335.

(Hriodon) occatoria,

300.

XXV

Mogera

wogura, 49.

— kanai, 49.

| Mormopterus, 206.

Mungos

ichneumon, 167.

paludinosus, 166, 167,

168.

— rubellus, 166.

Mus

agrarius, 7.

chrysophilus, 71.

confucianus, 6, 7.

— sacer, 6, 7.

coucha, 171.

harti, 9.

huang, 7.

ling, 7

musculus, 10G.

ningpoensis, 9.

norvegicus, Dd.

rattus, 170.

wagneri, 106, 107.

— mongolium, 106.

Mustela

afra, 167.

galera, 167.

Myocastor

coypu, 127.

Myrina

sharpet, 126.

| Myrmecophaga

juhata, dd.

Nemorhedus, 173, 174,

arnouxianus, 201, 202.

bedfordi, 191, 192, 193,

194.

bubalinus, 178.

caudatus, 198, 199,

202.

cinereus, 202.

evanst, 201.

fargesianus, 201.

goral, 190, 191, 192,

193, 202.

griseus, 199, 200, 201,

202.

henryanus, 200, 201.

hodgsoni, 195, 202.

niger, 201.

raddeanus, 196,

198, 199, 202.

sumatrensis, 176, 184.

Nebrius, 350.

concolor, 302.

Nemorhzdus

bubalinus, 178.

sumatrensis, 176, 184.

197,

XXV1

Nemorhedus |

(Antilope) goral, 195. |

Nemotragus, 174.

Nyctinomus, 206.

Ochotona

dawurica, 109.

ogotona, 109.

Odocoileus

americanus savan-

narum, 345.

Odontobutis

potamophilus, 62.

Ogotoma

pallasti, 109, 110.

Okapia

johnstoni, 320-384,

Orchestia

cavimana, 42. |

deshayesti, 42, 43, 44,

gammarellus, 42,43, 45. |

Orcbomenella |

nanus, 396.

Orectolobus, 348, 354.

barbatus, 355.

Japonicus, 355, 356.

ornatus, 385d, 356, 364,

tentaculatus, 355, 357,

304.

Ortalis

rificauda, 345.

Orycteropus

ethiopicus, 1.

Otomys

irroratus, 172.

Pagellus

centrodontus, 66.

Papio

cynocephalus, 165.

Paradisea

apoda, 64.

Paraduba

owgarra, 126.

Paranais

littoralis, 5.

Parascyllium, 348.

collare, 349. |

nuchale, 349.

variolatum, 349. |

Pelagia |

noctiluca, 57.

Pelobates

cultripes, 409,

Pelodytes, 13.

Pelomys

tallax, 172.

Pentila

amenaida, 111.

multiplagata, 111. |

INDEX,

Peltogaster, 46.

Perigonimus, 396.

Petrodromus

tetradactylus, 166.

Petromyzon, 46.

marinus, 65.

Phalangium, 45, 46.

Phocena, 52.

Phreatothrix

cantabrigiensis, 366,

367, 369.

pragensis, 365,

367, 368, 369.

Phyilostoma

Jamaicensis, 261.

lituratus, 228, 272,

. ee.

obscurum, 228, 237,

240, 241.

personatum, 212, 217.

perspicillatum, 2317,

238, 239, 272, 274.

planirostre, 228, 2837,

238, 239, 240, 272.

pusillum, 285, 288.

superciliatum, 228,

241, 272, 275.

Pipa, 15, 19, 26.

Pleuronectes

platessa, 161.

Powellana, gen. nov.,

366,

cottont, 114, 126.

Procayia

brucei, 172.

Propithecus

diademda, 64.

Protocetus, 203.

Prozeuglodon

atrox, 205.

Pseudorasbora

parva, 59.

Pteroderma

perspicillatum, 224,

ope

Putorius

itatst, 58.

nivalis, 213.

sibirica, 53.

Python

molurus, 137.

regtus, 1, 137.

sebeé, 137, 149.

spilotes, 137, 140, 141,

143, 149, 151.

Rana, 11-27, 31-34, 39,

40, 41.

esculenta, 34.

guppyt, 18,

26.

19; 22)

Rana

hasseltii, 425.

liebiqri, 418.

temporaria, 34.

Rhinoceros

sumatranus, 332.

Rhinodon, 352.

pentalineatus, 353.

typicus, 348, 352.

Rhinolophus

cornutus, 49.

trifoliatus, 207.

Rhinophrynus

dorsalis, 29.

Rhogeessa, 206.

Rhynchogale

melleri, 168.

Saccostomus

campestris, 170.

mashone, 170.

Salmo

salar, 46.

Saltator

albicollis, 345.

olinascens, 161.

Sarsia, 5.

Scopelocheirus, 396.

Scotophilus

nigrita, 166.

Scyliorhinus

canicula, 347.

edwardsit, 347.

marmoratum, 347.

Scyllium

Serruginewm, 351.

freycineti, 359.

heptagonum, 364.

malaisianum, 360.

ocellatum, 3859.

ornatum, 362.

plagiosum, 362.

Selenodon, 130.

Silurus

asotus, 61.

bedfordi, 59, 61, 63.

graham, 61.

Simia

satyrus, 161.

Spinax

niger, 373, 396.

Spongilla, 5.

Squalus

appendiculatus, 355.

argus, 300.

barbatus, 350.

caudatus, 362.

cirratus, 350.

cirrosus, 364.

fasciatus, 364.

Squalus

indicus, 362.

lobatus, 355.

longicaudus, 363.

ocellatus, 359.

punctulatus, 350.

punctatus, 350.

tigrinus, 363.

tuberculatus, 362.

Squatina, 348.

Staurotypus

triporcatus, 64.

Stegostoma, 353.

carinatum, 364.

fasciatum, 364.

tigrinum, 348, 363,

364.

Stenoderma, 231.

cinereum, 292.

rufum, 230.

toltecum, 228,

300, 302.

Stylodrilus

vejdovskyt, 369.

Synchismus, 358.

tuberculatus, 362.

297,

Tatera

afra, 170.

draco, 169, 170.

inclusa, 169.

lobengule, 170.

Testudo

elephantopus, 37.

Thaumaina, gen.

16.

uranothauma, 116,

126.

Thrasaétus

harpyia, 64.

Thysonotis

albostrigata, 124, 126.

nov., |

| Uroderma, 205, 209, 222,

INDEX. XXVIl

| Urotragus, 173.

| bedfordi, 191, 192.

| evansi, 200.

Thysonotis

corruscans, 125,

eheiket, 124, 126.

endocia, 124. | goral, 195.

hamilecar, 124. | Urotrichus, 50, 51, 52,

rosselana, 123. | 53.

Tilapia | — adversus, 50.

galilea, 4, 407. talpoides, 50.

nilotica, 4, 405, 406. -— adversus, 50.

Tragelaphus | — centralis, 50.

scriptus, 173. | — hondonis, 50, 51.

Trichodrilus | — pilirostris, 51.

cantabrigiensis, 369. | — talpoides, 50.

Tridentiger |

coreanus, 68. |

Trischizostoma Vampyressa

niceense, 370, 374, pusilla, 288.

375, 885, 392, 394, | Vampyrops, PN, Lai,

395, 396, 397, 398, 257, 288, 294.

400, 401, 402. _lineatus, 230.

raschii, 370, 371, 372, | Vespertilio

eomolamoalOmasa americanus vulgaris,

385, 397, 401, 402. 274, 280.

: ee rans. 52

remipes, 370. murinus superans, D2.

perspicillatus, 274,

| Upolampes, gen. noyv., _ 280.

118. Vipera

striata, 118, 126. russelli, 149.

Viverra

Uranothauma, 119.

civetta, 169.

nems, 167.

224, 225, 230, 257. %

Vulpes, sp., 105.

bilobatwm, 204, 212,

213, 214, 217, 218,

219, 220, 221, 223,

231, 310, 311.

concolor, 231.

convexum, 218,

221.

fallax, 231.

thomasi, 204, 216, 217, |

218; 221 Sie |

validuin, 228, 242,

245.

Waigeum

dinawa, 125, 126.

220, resplendens, 125, 126.

Xenophrys

gigas, 416.

monticola, 408, 416,

418, 419, 420.

| Xenopus, Ld, 19.

THE END.

Printed by Taynor and Francis, Red Lion Court, Fleet Street.

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PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

ZOOLOGICAL SOULETY

OF LONDON,

1908.

Pages 1-126.

Part 1. cONTAINING PAPERS READ IN

JANUARY anv FEBRUARY.

JUNE 1908.

PRINTED FOR THE SOCIETY,

SOLD AT THEIR HOUSH IN HANOVER SQUARE.

LONDON :

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[Price Twelve Shillings. |

LIST OF CONTENTS.

1908, pp. 1-126.

January 14, 1908.

Page

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

December 1907 ...... sis settee ee eae -

Mr. R. Lydekker, F.R.S., F.Z.8. Exhibition of, and notes upon, an abnormally marked

Leopard-skin from the Deccan ........ bisvelejate aye) ainie ete iala:/elete obatelleio euete ciel slapeteiete meets 1

oe Description of a Biological Expedition to the Birket-el-Qurun, Faytim Province of Egypt.

IBya We Az CUNNINGLON) Bo: Aee ten) vel ZS. ces -lepeietensis = Iaieieicieretelabeleyarehelierelarst= apeiene eee 3

2. The Duke of Bedford’s Zoological Exploration in Eastern Asia.—VI. List of Mammals

from the Shantung Peninsula, N. China. By Ouprinnp Tuomas, F.R.S.,F.Z.8....... 5

8. On the Musculature and other Points in the Anatomy of the Engystomatid Frog,

Breviceps verrucosus, By Frank EH. Bepparp, M.A., F.R.S., F.Z.8. ........-.-.000% 11

4, On the Hermaphroditism of the Amphipod, Orchestia deshayesti Audouin. By Cuartns

ii, Bourmnemr, BvA., ¥.Z:8.; King’s College, Camibridge - 2. seis cr. eine ole (erste cheieiaiern te

February 4, 1908.

Mr. F, Martin Duncan. Lantern exhibition of, and remarks upon, the Autochrome Natural

ROC Nora OCR AT ODIO OGG Om ado 66 OK 47

Colour Process of the brothers Lumiére

1. The Duke of Bedford’s Zoological Exploration in Hastern Asia.—VII. List of Mammals

from the T'su-shima Islands. By Otpriz—tp Tnomas, F.R.S., F.Z.8. ......-...00.005 47

Contents continued on page 3 of Wrapper.

PROCEEDINGS

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF TUE

ZOOLOGICAL SOCIETY OF LONDON.

(January to April, 1908.)

January 14, 1908.

Prof. J, Rose Braprorp, F.R.8., Vice-President, in the Chair.

The Secretary read the following report on the additions made to

the Society’s Menagerie during the month of December 1907 :—

The number of registered additions to the Society’s Menagerie

during the month of December was 175, Of these 89 were

acquired by presentation and 4 by purchase, 50 were received on

deposit, 1 by exchange, and 31 were bred in the Gardens. The

number of departures during the same period, by death and

removals, was 174.

Amongst the additions special attention may be directed to :—

One Hainan Gibbon (Hylobates hainanus), 3, from Hainan,

presented by R. Douglas, Esq., on December 6th.

One Grévy’s Zebra (Hquus grevyi), 3, from Abyssinia, received

in exchange on December 9th.

One Hthiopian Aard Vark (Orycteropus athiopicus) from Nai-

vasha, British Hast Africa, presented by the Marquess Gandolfi

Hornyold on December 17th.

One Macqueen’s Bustard (Houbara macqueent), caught in the

Red Sea, presented by Mr. Jos. Vyskocel on December 9th.

Two Royal Pythons (Python regius) and one Nose-horned Puff-

Adder (Bitis nasicornis), from 8S. Nigeria, presented by R. B.

Brooks, Esq., on December 20th.

On behalf of Mr. Lydekker, an abnormally marked Leopard-

skin from the Deccan, India, was exhibited by Mr. Pycraft. The

specimen would shortly be presented to the British Museum by

Proc. Zoou, Soc.—1908, No. I. 1

2 ON AN ABNORMAL LEOPARD SKIN. [Jan. 14,

Mx. F. A. Coleridge, who shot the animal from which it was taken

near Putnam, Cuddapah, in December 1906. The following

Abnormally marked Leopard-skin from the Deccan.

notes on the specimen (of which a photograph is here given) were-

supplied by Mr. Lydekker :—

According to information supplied by the donor, leopards of

the ordinary type abound in the district where the abnormally

1908. | ON AN EXPEDITION TO THE BIRKET-EL-QURUN. 3

marked specimen was killed. In the pattern of the markings

this skin appears to be altogether unique. The most striking

feature is a kind of network of broad yellow lines commencing on

the head in a large, somewhat diamond-shaped area and con-

tinued all down the back in meshes of smaller size. Within each

mesh is a large black-bordered patch, containing a central area.

darker than the big network, upon which is a variable number of

black spots. In many instances the borders of the patches are

incomplete. The other details of the pattern are sufficiently

apparent in the photograph. Beyond referring to the fact that

the markings, so far as they resemble those of any normally

coloured animal, make a slight, although decided, approximation

to the Jaguar-type, I am unable to offer any explanation of the

remarkable style of ornamentation presented by this extremely

handsome skin.

The following papers were read :—

1. Description of a Biological Expedition to the Birket-el-

Qurun, Faytiim Province of Egypt. By W. A.

CunninetTon, B.A., Ph.D., F.Z.8.

[Received January 14, 1908. |

An expedition was undertaken by me in conjunction with

Mr. C. L. Boulenger, to investigate the flora and fauna of the

Birket-el-Qurun, a lake in the Faytm province of Egypt—

the Lake Moeris of the ancients. The expedition left London

in the middle of March 1907, and eight weeks were spent in

collecting on the shores of the lake. Since the railway comes

within some seven miles, there were no difficulties of transport to

contend with, the journeying about the lake being accomplished

in a sailing-boat of moderate size.

The Birket-el-Qurun, or lake of the horns, is still of considerable

dimensions, being about 25 miles long, and having a maximum

breadth of 5 or 6 miles. It is, however, only a remnant of the

historic Lake Moeris, which was many times greater, and was used

as an artificial regulator of the Nile floods by the monarchs of the

Xth dynasty. The lake still communicates with the Nile by

means of the Bahr Yusef, a channel over 200 miles long, which

diverges from the river a short distance north of Assiut. The

inflow and outflow of water was originally controlled by means of

lock-gates, but since these were allowed to fall into disuse, more

and more of the extremely fertile land was reclaimed, until the

lake was reduced to its modern dimensions. It is believed that

the surface of the lake was at one time some 70 feet above the

level of the Mediterranean ; at the present day, it is approximately

140 feet below sea-level.

In addition to the historical. evidence, there are not wanting

4 ON AN EXPEDITION TO THE BIRKET-EL-QURUN. [Jan. 14,

‘signs of a considerable reduction in size, for raised beaches are in

many places very apparent, and the lake has become extremely

shallow, the greatest depth being between four and five fathoms.

Additional evidence is afforded by the nature of the water, which

is sufficiently brackish to be quite unpalatable. The density 1s

slightly above that of fresh-water, and the proportion of soluble

salts about one-fourth that in the ocean.

The almost complete absence of seiche alterations of water-level

is probably due to the shallowness of the lake, as are aiso the

rather striking figures obtained of the water temperature. A

maximum of 94:2° was observed in very shallow water close to

shore about 2.0 p.m., whilst a minimum of 54°8° was obtained as

a surface reading in the early morning. The difference between

the surface temperature and that of the water below may also

be very great, one reading showing the water at three fathoms to

be 12°4° colder than that at the surface, whilst a difference of 8°8°

was detected in water only one fathom below.

The lake would seem to be remarkable more for the quantity of

life which its waters contain, than for the number of different

species inhabiting it. From the greenish water, the tow-net

collects immense swarms of Entomostraca—principally Copepoda

and Cladocera—as well as large numbers of Rotifera. The only

phyto-plankton, however, which is present in the tow-nettings,

consists of two or three species of Diatoms.

The fish collected by the expedition were obtained for the most

part from the native fishermen, who secure large quantities from

the well-stocked waters. The fishing industry is highly organised

and the bulk of the catch is despatched to the Cairo market.

Fifteen species of fish were secured, belonging to seven families.

All of these are well-known Nile forms, but more than half of

them are recorded for the first time from the Birket-el-Qurun.

The so-called Nile perch (Lates niloticus) is stated to attain a large

size in the lake, a specimen which measured 120 cm. and weighed

54 lbs. being considered by the local people as by no means

extreme. Although 15 species are thus known to inhabit the

waters of the lake, two of them occur in such remarkable

abundance as to make the others appear rare. These are Tilapia

galilea and Tilapia nilotica, the latter being the more abundant.

The fish markets very frequently contain nothing but these two

Species, which are caught with casting nets in the swampy

shallows, or with seines in the deeper water.

Tn addition to Copepods and Cladocera from the open lake,

Ostracods are found on the muddy bottom and in the swamps,

while certain Oniscide were collected on the beach. It is rather

singular that no other Crustaceans were obtained: no crabs or

prawns were procured, although they are well-known in the Nile,

neither were any Argulidee observed on the large numbers of fish

examined. In the swampy pools on the lake margin were found

water-beetles of apparently little interest, Hydrachnids of the

genus Hulais, and certain spiders. .

1908. ] ON MAMMALS FROM N. CHINA. 5

From such regions too, and from the lake bottom, come the few

Species of Mollusca obtained. Only a single Lamellibranch was

amongst these. Truly aquatic worms are represented solely. by

the Oligocheete Paranais littoralis, no leeches or Turbellaria being

found, although they might well be expected to. occur, . A

Polyzoan is fairly abundant, growing on the submerged stems of

plants, or on the underside of rocks and boulders. It is a gymno-

lematous form, with a circular lophophore and eight tentacles.

Cordylophora lacustris, a form often associated, with brackish

water, grows luxuriantly on stones in shallow water. A much

more interesting and quite unexpected discovery was that of a

medusa and the hydroid form with which it is associated. The

hydroid was obtained first, being dredged from the bottom in

about a fathom of water: the medusa made its appearance in the

lake in vast quantities at a somewhat later date. It appears to

be a typical Anthomedusan, and has some resemblance to the

marine genus Sarsia. Although the Birket-el-Qurun is now

slightly brackish, it certainly was quite fresh within historic times,

and it is not easy to suggest how such a characteristically marine

organism should have found its way into the lake. It is curious

to note that no fresh-water sponge was obtained by the ex-

pedition, although Spongilla occurs commonly in the Nile.

Algz were collected throughout the lake, being scraped from

the rocks and stones and the submerged stems of plants. Of the

truly aquatic higher plants, Potamogeton interruptus and Chara

vulgaris are both of some interest, as they show slight differences

from the common form.

The expedition was undertaken at the request of Captain

Lyons, F.R.S., Director General of the Egyptian Survey

Department, which has recently paid a good deal of attention to

the study of this lake and the whole Faytm province.

The Duke of Bedford’s Zoological Exploration in Eastern

Asia.—VI. List of Mammals from the Shantung Pen-

insula, N. China. By Otprietp THomas, F.R.S., F.Z.8.

[Received December 2, 1907. |

In March 1907 our President’s collector, Mr. Anderson, landed

at Chefoo on the Shantung Peninsula, in order to commence

work in Northern China. Nothing has been done in this region

since the time of Consul Swinhoe, and good modern specimens of

the species he obtained are required before any further progress

can be made.

As may be gathered from the following notes by Mr. Anderson,

the Peninsula is highly cultivated, and almost treeless, so that it

is naturally poor in Mammalian life. He has nevertheless

obtained good series of the few mammals found there, and these

will be of much use in the further study of Chinese Mammalogy.

6 _ MR. OLDFIELD THOMAS ON (Jan. 14,

Notes on Shantung.

“T arrived in Chefoo on March 12th, 1907, and was obliged

to stay there until the 26th, when I moved into the country eight

miles to the south, to remain until April 10th.

“The Shantung Peninsula consists of a complicated series of

mountains isolated from other ranges by the broad plain of the

Hoang-ho.

“In the region of Chefoo plains and hills alternate, the former

being broad, treeless, and though dry are under cultivation for

wheat and millet; the latter rise to a height of 1000 feet or more,

are steep, rocky, and barren. In general the only trees that occur

are diminutive pines, no taller than a man, which are allowed to

grow in some parts of the hills; but occasionally in some favoured

or sacred spot, a cemetery, or temple land, one finds pines or oaks

of larger size. It was such a place that I found eight miles south

of Chefoo. Here the steep and rocky hills bore a sparse wood of

oak, which flourished under the protection of a Buddhist Temple.

“On April 13th I began work at Ai-san, a mountain about

30 miles west of Chefoo; here I remained till May Ist. Ai-san

is an isolated granite crag, 3200 ft. high. Its foothills, in which

I took up quarters at an altitude of 1200 ft., are of granite and

dry granitic sand, clothed in spots with the usual diminutive

pines, and cut by precipitous canyons through which numerous

streamlets flow from the mountain. Were it not for this

frequency of water the hills would I think be untillable even to

the Chinese, but the valleys, canyon bottoms, and sometimes the

hillsides are cultivated up to 1200 ft.” P. A.

1. ERINACEUS DEALBATUS Swinh.

G6 M4617 “2. W447-" ‘Chetoo:

Originally described from Peking.

“Purchased alive from peasants who had brought them into

Chefoo. Said to be not uncommon, but I failed to find them

myself. Seems to be strictly nocturnal.”—W. P. A.

2. MUS CONFUCIANUS SACER, subsp. n.

3. 1385, 1387, 1388, 1393, 1394, 1397, 1398, 1399, 1401, 1402,

1403, 1405, 1406. ©. 1386, 1389, 1390, 1391, 1395, 1396, 1400,

1407, 1408, 1409, 1410, 1411, 1412. Near Chefoo. 300’.

3. 1424, 1430, 1434, 1440. 9. 1423, 1431, 1432, 1433, 1443,

1445. Ai-san, 30 miles W. of Chefoo. 1200’.

A female in spirit (No. 1404), with 2—2=8 mamme.

A. buffy-grey subspecies of IZ. confucianus; tail long-haired,

white-tipped.

Size about as in true confucianus. Fur soft, not spinous in

specimens killed up to 25th April, and probably never so, as the

members of this group are not known to change seasonally in

this respect, as is the case in Apodemus speciosus. General colour

above greyish-buffy or clay-colour, darkened by longer black hairs

1908. } : MAMMALS FROM N. CHINA. 7

‘on the back, clearer buffy on the sides, a buff line edging the white

of the belly. Under surface pure sharply-defined white throughout.

Ears grey-brown, finely edged with white. Upper surface of

hands and feet pure white, the metapodials not darkened. Tail

long, very well haired, so that the scales are nearly hidden, the

terminal pencil of hairs 5 to 7 mm.in length; its colour brown

proximally above, its under surface and its terminal third

(occasionally half) white all round, though some shorter hairs on

the end of the tail above are also sometimes dark.

Skull as in Fokien specimens referred to J/. confucianus, but

the teeth uniformly larger.

Dimensions of four specimens :—

Head & body. oNanile Hind foot. Kar.

mm. mm. Inm. mm.

ol Ueascee Henne 156 186 dl 21

@ (ley7ive)) coe 144 172 29 20°5

es oil oaae 141 Wa 28 21

OUP ETE ee 130 170 29 20°5

Skull of type—greatest length 38 mm.; basilar length 30;

greatest breadth 17-2; palatilar length 16°5 ; palatal foramina 7:1;

length of upper molar series 6.

Type. Adult male B.M. No. 8.2.8.8. Original number 1398.

‘Collected 30th March, 1907.

This fine series of specimens is remarkably uniform,

there being practically no variation in any important respect.

None of the specimens have any trace of the darker markings

on the metapodials found in true J/. confucianus, nor is there

any material variation in the degree of whiteness of the tail.

Bonhote’s Mus huang and MW. ling are both much more fulvous

in colour, while neither they nor confucianus have the tail so

heavily pencilled as it is in WZ. ¢. sacer.

The belly of these rats would appear to be more yellow in life

than it is in skin, judging from the name given them by

Mr. Anderson.

“‘ Sulphur-bellied Rat.—Common among the rocks in the

temple-woods near Chefoo, rarely met with elsewhere. In the

sacred woods it feeds mostly on acorns, leaving large accumulations

of the shells in cavities beneath the rocks.” —J/, P. A.

3. APODEMUS* AGRARIUS PALLIDIOR, subsp. n.

dg. 1365, 1417, 1419. @. 1420. Near Chefoo. 300’.

6S. 1422, 1425, 1426, 1428, 1429, 1435, 1436, 1437, 1446.

©. 1427, 1439. Ai-san. 30 miles W. of Chefoo. 1200',

g. 1453, 1460. 2. 1448. Near Wei-hai-wei. 300’.

On laying out the whole of the fine series of the A. agrarius

group obtained by Mr. Anderson in the East, there proves to be a

* Kaup, Entwick. Gesch. Nat. Syst. Eur. Thierw. p. 154 (1829).

Type A. agrarius (Mus agrarius Linn.) antedates Micromys Dehne, 1841.

8 MR. OLDFIELD THOMAS ON [Jan. 14,.

certain amount of geographical variation, although this is obscured

by the individual variation in size and the development of the

dorsal stripe already referred to in previous papers of the

present series.

The following synopsis indicates the local races into which the

eastern forms of A. agrarius appear to be divisible :—

A. (Northern.)

a. Colour dark—more or less tawny. Belly hairs washed with

pale tawny. Dorsal streak well developed.

Mantchuria. <A. a. mantchuricus Thos.

6. Colour less rich—approximating to “cinnamon.” Belly hairs

white-tipped. Dorsal streak variable.

Korea and Quelpart. A. a. coree, subsp. n.

B. (Southern.)

ec. Colour comparatively greyish—approaching wood-brown, the

head and fore quarters lighter than the back. Dorsal streak

always present ............ Shantung Peninsula. A. a. pallidior, subsp. n.

d. Colour more uniform and brownish. Dorsal streak usually

Obsolete ..:..:...... Nanking to N.W. Fokien. A. a. ningpoensis Swinh.

The two forms now given new names may be briefly described

as follows :—

APODEMUS AGRARIUS CORE.

Summer pelage more or less spiny. General colour above:

approximating to “cinnamon” of Ridgway, not so ruddy as in

mantchuricus, more so than in pallidior. Belly hairs grey basally,

white terminally, not washed with tawny or buffy. Dorsal streak

variable, but never of the clear well-defined unspeckled black

throughout usually characteristic of true agrarius.

Dimensions of four specimens from the typical locality :—

Head &bedy. Tail. Hind foot. Har.

mm. mm. mm. mm.

(Ciro nares 91 81 21 13°5

Om Clive) ieee 110 9D 20°5 13

DU INR ay, 103 81 21 14

Come ok ais 95 80 21 1355

Skull (of type)—greatest length 27-5 mm.; basilar length 22 ;

length of upper molar series 3°8.

_ Hab. Korea and Quelpart. Type from Min-gyong, 110 miles.

S.E. of Seoul. 1000’.

Type. Adult female. B.M. No. 6.12.6.74. Original number

654. Collected 24th Nov. 1905.

The two series of this form obtained by Mr. Anderson are:

enumerated in the papers on the Korean collections.

APODEMUS AGRARIUS PALLIDIOR.

Size perhaps averaging rather less than in the more northern

forms. Fur not spiny, but no specimens seen dated later than:

May 10. General colour above paler and more greyish-white

than in the allied forms, especially on the head and across the-

1908. | . MAMMALS FROM N. CHINA. 9

fore-quarters; this pallor appears to be due to the hairs being

light, almost whitish subterminally, their extreme tips being of

the usual reddish-brown tone. Belly hairs white-tipped. Dorsal

streak fairly uniform throughout the series examined, very slightly

marked and often obsolete anteriorly, fairly well-defined

posteriorly ; never so strong as in agrarius, manichuricus, and

the more strongly marked specimens of corew, but on the other

hand much more evident than in ningpoensis, which usually has

almost no trace of a stripe.

Dimensions of four specimens :—

Head & body. Tail. Hind foot. Har.

mim. mm. mm. mm.

Gn lye) ila: 93 100 19 13

Guede sce eretaiss 89 82 20 13

Ci Aa 97 83 20 th

Chica Voie 82 77 AO 1g

Skull of type—greatest length 27°5 mm.; basilar length 22:5;

length of upper molar series 4:1.

Hab. Shantung Peninsula. Type from near Chefoo. 300’.

Type. Old male. B.M. No. 8.2.8.29. Original number 1419.

Collected 5th April, 1907.

This subspecies is no doubt most nearly allied to “ Mus ning-

poensis” Swinh.* (JZ. harti Thos.), which I should now consider as

a subspecies of the Apodemus agrarius group. It differs, however,

both by its paler colour and by the uniform presence of a dorsal

stripe, this being but rarely perceptible in the more southern

animal.

“Caught usually in traps set under rocks near water-courses.

At Ai-san their ears were nearly always diseased, and I think

thereby shortened.” —J/, P. A.

4, CRICETULUS TRITON de Wint.

3. 1415, 1416, 1418. 9. 1366. Chefoo, Shantung. 100’.

3. 1454. 2. 1450, 1451. Wei-hai-wei. 300-400’.

These welcome examples, practically topotypes, of the hitherto

little-known (. triton vary unexpectedly in size among themselves,

but none of them equal in tooth-length, and only one very old

specimen in hind-foot-length, the immature type of C. nestor, dis-

covered by Mr. Anderson in Korea. Their tails also show an

unusual amount of variation in length.

A specimen referable to C. tritom has recently been pr esented

to the Museum by Mr. E. B. Howell, who trapped it at Tientsin,

thus carrying its known range to the west of the valley of the

Hoang-ho.

Curiously enough, both Mr. Anderson and Mr. Howell note a

predilection on the part of these animals to make their burrows

in human grave-mounds, on the south side of which they sink a

perpendicular hole.

* P,Z.S. 1870, p. 637; cf. Bonhote, P.Z.S. 1905, 1. p. 397.

10 ON MAMMALS FROM N. CHINA. (Jan. 14,

5, CRICETULUS GRIsEUS M.-Edw.

6. L395, 1360, 1361, 1362, 1363, 1367, 1368) °1372; 1373,

1374, 1376. 9. 1356, 1358, 1359, 1364, 1369, 1370, 1371, 1375,

1377, 1381. Chefoo. Sea-level.

3. 1442. ©. 1438, 1441. Ai-san, 30 miles W. of Chefoo.

1200’.

o. 1452, 1458. Near Wei-hai-wei. 300’.

This fine series is very uniform in general colour and in the

development of the dorsal line, which is as distinct in the young

onesas in the adult. The following are the flesh-measurements

of four of the largest examples :—

Head &body. ‘Tail. Hind foot. Ear.

mm. mm. mm. mm.

ee te 100 27 15:5 15

[EN A 97 28 16 15

Oral My 9] 28 15:5 16

Chola tem 86 29 15:5 16

Mr. Howell also obtained this species at Tientsin, and there

were two examples in the collection from N. Shantung presented

by Mr. Styan and worked out by de Winton. The latter used

for them the name of C. obscurus, but griseus comes first 1n Milne-

Edwards’s book, and even if the two are different, which is most

doubtful, grisews should, for geographical reasons, be the name

for the little hamster of Shantung.

“ Although not common this was almost the only mammal in

the fields about Chefoo. Its burrows were most often in small

banks in the gardens, but sometimes occurred between the rows

in wheat fields.”— 7. P. A.

6. LEPUS SWINHOEL Thos.

Lepus tolat Swinh. P. Z.8. 1870, pp. 449, 450.

6. 1382, 1383, 1384, 1392, 1421.

@. 1357, 1379, 13880, 1414. Chefoo. 300’.

3. 1444 (young). Ai-san, W. of Chefoo. 1200’.

3. 1455-7 (young), 1459. @. 1449. Near Wei-hai-wei. 300’.

These specimens are all of the paler type of coloration described

by Swinhoe as characterising his third ( Q ) specimen, though they

vary among themselves in this respect.

“Common near Chefoo and near Wei-hai-wei, but rare near

Ai-san (where there are wolves). Where common this hare is

conspicuous, for although the body is much the colour of the soil,

the white tail is an advertisement. It is not wild, but will stop

again and again if one whistles sharply. This hare has a voice

of some strength, as wounded ones showed by crying out

plaintively, or sometimes threateningly, as I approached. I found

the flesh very palatable, and many are killed for the Chefoo

market.”—W. P. A.

1908. | ON THE ANATOMY OF A FROG. 11

3. On the Musculature and other Points in the Anatomy of

the Engystomatid Frog, Breviceps verrucosus. By

Frank E. Bepparp, M.A., F.R.8.

[ Received December 11, 1907. ]

(Text-figures 2-13.)

I have examined or dissected three examples of a species of

Breviceps, which I refer to the species ‘“ verrwcosus” on the

following grounds. In the definition of the Frog by Boulenger *

the body is stated to be “ entirely covered with distinctly porous

granular glands.” This was the case with my specimens, which

therefore appear to differ from Breviceps gibbosus. Inasmuch as

our knowledge of this African and ant-eating Batrachian seems

to be confined to its external and osteological characters, I have

thought it worth while to bring before the Society a further

contribution to the knowledge of its structure. The notes upon

which the present communication is founded chiefly relate to

those structures which are known to vary in their characters

among the Anurous Amphibia. Other characters, however, are

not altogether ignored.

§ Pelvis and Coccyx.

It is of course well known 7 that Breviceps is distinguished from

(e. g.) Rana by the widely suvunried transverse processes of the

sacral vertebra and by the fusion of that vertebra with the

ensuing coccyx. Since the latter point at any rate has been

found, “though rarely, to vary among the Anura, it is perhaps

wor th while’ to record here the fact that I found in two specimens

a complete fusion between the sacrum and the coccyx. When

the frog is extended with the dorsal surface uppermost, the ilia

are not visible as they are in Runa to a great extent, and the

ilio-ecoccygeal muscles descend on a plane which only forms a

small angle with the plane of a sagittal section. The strongly

expanded transverse processes of the sacral vertebra show no

connection with the ilia when viewed from above. These bones

are quite invisible if the dorsal surface of the transverse process

has been cleaned and the underlying musculature left alone.

The attachment of the ilia, in fact, is not to the edge of the

broad transverse process as in some other Batrachians, but is

completely ventral leaving the edge entirely free.

Another structure in connection with the pelvis of Breviceps

remains to be described, which I have not noticed, or seen a

description of, in other Frogs. On the dorsal surface of each

sacral transverse process, lying, apparently, freely on that process,

is a flat and somewhat oval plate of cartilage not so long as the

transverse process is in an antero-posterior direction. This plate

* Cat. Batr. Sal. 1882, pp. 176, 177.

+ “Amphibia” in Bronn’s Thierreich, pp. 608 & 640.

12 MR. F. E,. BEDDARD ON THE [Jan. 14,

therefore lies completely on the expanded sacral transverse

process. Anteriorly fibres of the—as I presume—ilio-lumbaris

muscles are inserted upon the cartilaginous plate, and quite

anteriorly it is connected, though feebly, round the edge of the

transverse process, with the expanded end of theilium. Iimagine

that this cartilage belongs to the ilium, and that it is in con-

sequence related to it as the suprascapula is to the scapula. We

have, in fact, in this frog an exaggeration of the grooving which

the anterior end of the ilium of other forms shows at its line of

articulation with the sacral vertebra. It is, moreover, interesting

to observe that we find in this frog a kind of foreshadowing of

the relations which the ila bear to the sacrum in the bigher

Sauropsida—especially birds—where the ilia are not merely

attached to, but cover, the sacral vertebree.

§ Hyoid.

This cartilaginous complex presents some peculiarities of form

in Breviceps. Of these the most salient are shown in the accom-

panying figure (text-fig, 2) illustrating also some of the hyoid

Text-fig. 2.

SEE.

Lf abd. Sth.

Anterior cornua of hyoid and part of hyoidean musculature of Breviceps.

h. Basihyal plate. h!. Anterior cornu of hyoid. !. Lateral process of basihyal..

H.abd. Hyoabdominal muscle. h.g. Hyoglossus. P.. Petrohyoideus. Sh.

Subhyoideus. Sm. Submaxillaris. S¢.4. Two divisions of sternohyoideus.

muscles. It will be there seen that the anterior cornua of the

hyoid are particularly stout and strong which is correlated,

1908. } ANATOMY OF A FROG. 13

perhaps, with the unusually strongly developed subhyoideus

muscle. Again, correlated with these facts of structure is the

mode of origin of the anterior cornua from the body of the hyoid.

As the text-figure referred to shows, the anterior cornu on each

side bifurcates near to its junction with the body of the hyoid,

and shortly afterwards the two branches rejoin, thus forming an

almost triangular foramen. Anteriorly there is no distinction

between the cornu itself and the anterior process of the body of

the hyoid, such asis so marked in Rana. In fact, the connection

of the anterior cornua of the hyoid with the basihyal recalls the

arrangement of the corresponding cartilages in the not nearly

allied Pelodytes, but more so as there is a close approach in the

middle line between the rounded-off ends of the conjoimed

anterior process and anterior cornu of either side; they do

not, however, so nearly meet in Areviceps as is the case with

Pelodytes.

The figures of the hyoid apparatus of other Frogs given by

W. K. Parker* show no types which closely resemble Breviceps

in these particulars. In many forms, however, there is no

anterior process on either side of the basihyal, and in others

there is an approximation between the anterior lateral process of

the basihyal on each side and the corresponding cornu. The

thyrohyals of Lreviceps are also peculiar in certain respects.

Each is ossified as usual and is of the customary hourglass-shape,

being-—that is to say—thinner in the middle than at its two ends.

It is, moreover, rather bent in the middle outwardly. The end

by which it articulates with the body of the hyoid is not affixed

to that cartilage as in many (? most) Frogs. In the latter the

bone is inserted on to the posterior edge of the basihyal cartilage.

In Breviceps it is quite distinctly mserted on to the ventral

surface of the cartilage in front of the posterior edge 7.

§ The Musculature of the Hyoid.

As might be expected from the habits of the Frog, the Hyoid

muscles differ in many respects from those of Rana, &e. I shall

describe under the heading of the abdominal muscles an important

muscle which I term hyoabdeminal, which is a part of the superficial

abdominal sheath inserted on to the hyoid.

The sternohyord is also a peculiar muscle. For it is composed

of two perfectly distinct parts. The anteriorly attached part has

the usual insertion on to the body of the hyoid ventrally between

the (posteriorly) diverging halves of the geniohyoideus of its

side. This part of the sternohyoideus arises almost entirely

from the coracoid dorsally ; ; the second half of the sternohyoideus

is in contact at its origin with the last, and appears also to be

continuous with such fibres of the very feeble rectus abdominis

* Phil. Trans. pt. 1. 1881.

+ Parker’s figure of Hngystoma carolinensis suggests that this is also the case

with that species, which is of the same family.

14 MR. F, E, BEDDARD ON THE [Jan. 14,

sternalis as reach the sternum. Its direction is from the very

first different from that of the anterior portion of the muscle.

It passes downwards at a greater angle with the plane of the

coracoid and to the inside of the hyoabdominal, dorsally to which

it then passes to be inserted opposite to the insertion of the

outer part of the geniohyoid. The hyoabdominal thus passes.

between the two portions of the sternohyoid, and it is able to.

insinuate itself into the narrow space prepared for its reception,

not by a diminution of its fibres or a lessening of its diameter by

becoming converted into a tendon, but by bending over so that

its broad and flat surface comes to be disposed perpendicularly

instead of horizontally.

The geniohyoideus of Breviceps appears to offer no very great.

differences from the same muscle in Rana. But the two inner

halves of the two muscles fuse together some way back over the

subjacent hyoglossus more markedly than in Rana, and in this

the muscle recalls the geniohyoid of the Pelobatidee *.

I could find no trace whatever of an omohyoid muscle.

The hyoglossus completely envelops each thyrohyal. The fibres.

run aggregated into coarse strands over the body of the hyoid. In

the region of the thyrohyal the hyoglossus is overlain by a thin

layer of muscle which appears to be perfectly continuous with the

geniohyoid (see text-fig. 7, A, p. 28), that is of course with the inner

division of that muscle. The fibres, however, are here transversely

arranged to the longitudinal axis of the thyrohyal and are wrapped

round the enormously thick hyoglossus. In view of their direction

these fibres can hardly be referred to the geniohyoid and called

by its name; but they appear to be clearly a differentiation of

the same sheet of muscle. The function of this sheet of muscle

appears to me to be possibly this: the enormously developed

musculature—the hyoglossal—which enwraps each ceratohyal

tends to occlude, particularly during its contraction, the glottis,

and thus to hinder free respiration. ‘This would, however, be

advantageous during swallowing. During relaxation of the

hyoglossal the diameter of that muscle would, perhaps, be still

further reduced by the contraction of the transverse sheet, and

the orifice into the lungs in consequence enlarged.

The petrohyoid muscles are present in the normal number:

and are thick and fleshy leaving no gaps, and indeed overlapping

each other. The posterior division of the muscle seems to be less.

attached to the thyrohyal than is usual among these Batrachians,

The petrohyoideus posterior primus has apparently some con-

nection with the bony thyrohyal bar. When the latter is raised,

the muscle is seen to underlie it (it is of course dorsal to the

bone) and to be posteriorly attached to it. The muscle is, in

fact, anteriorly inserted on to the edge of the body of the hyoid

behind the posterior lateral process, this portion of the body of

the hyoid being, as already explained, overlain by the thyrohyal

* See P. Z. S. 1907, p. 895.

1908. ] ANATOMY OF A FROG. 15:

which does not arise from its edge. When this first section of

the petrohyoideus posterior is raised, it is seen to overlie the

posterior portion of the petrohyoideus anterior, which latter

therefore is a more extensive muscle than is usual among the

Batrachia Salientia.

The petrohyoideus posterior secundus is attached in the present

species as is the petrohyoideus posterior tertius of some other

Frogs. It is inserted, in fact, on to the expanded lower extremity

of the thyrohyal bone. Whether some fibres escape to be con-

nected with the laryngeal apparatus I do not know.

The petrohyoideus posterior tertius is quite different in its.

relations from the corresponding muscle in other Frogs whose-

anatomy is known. In order to see the muscle the secundus has.

to be raised. When this is done, the muscle now under discussion

is seen as a somewhat slender muscle running parallel with the.

other parts of the petrohyoideus posterior. But it has no

relations whatever to the thyrohyal bone. This in itself is a

point of likeness to the Pelobatide. But the resemblance ceases.

with this. For in Breviceps the muscle avoiding altogether the

end of the thyrohyal ends in close juxtaposition to the cesophageal

muscle on the walls of the commencement of the lung (see text-

fig.7,B). Its action on contraction would appear from its position

to be like that of the cesophageal muscle and would dilate the

pulmonary cavity.

§ Superficial Muscles of the Ventral Surface.

The general appearance of these various muscles, after the skin

has been removed, is very different from the corresponding view

of the musculature of Rana. I shall proceed to describe these

several muscles, commencing with the rectus abdominis and

passing forward to the throat. All the muscles now in question

are shown in the accompanying figure (text-fig. 3). As is well

known, the thighs of this frog are included within the area of the

body, the portion of the leg from the knee onwards alone pro-

jecting beyond the contour of the trunk. But when the skin

was reflected and turned back, it was to be observed that the

area lying between the anterior border of the thigh and the

posterior border of the abdomen is not merely covered by skin.

For closely adherent to the skin in this region, and indeed

inserted apon it, is a layer of muscle (to be considered in greater

detail later, p. 26) attached on the other side to the leg which

bridges over the gap. The conditions are not, therefore, very

widely different from those which characterise Xenopus * and

Pipat, where muscles attached to the leg spread into the

abdominal region and thus help to destroy the demarcation

between thigh and abdomen.

The rectus abdominis muscle appears to me to have in pro-

* Beddard, “On the Diaphragm &c. of Xenopus,” P. Z. S. 1895, p. 844, fig. 3.

+ Id. “On... the Anatomy of Pipa,” ibid. p. 888, fig. 4.

16 MR. F. E, BEDDARD ON THE (Jan. 14,

portion to the animal a very great thickness where it arises

posteriorly by the usual two fleshy pillars, one for each half of the

muscle. It is sharply crescentic in outline on each side, a state

of affairs which appears to have been brought about by the

inclusion of the thigh within the body and the consequent and

mutual pressure. This is plainly shown in the figure (text-

fig. 3) and is to be contrasted with the relative form of the same

muscle in Rana. The rectus abdominis of Breviceps also differs

Text-fig. 3.

wee

Pabd.

PR abd.

Sk.

Superficial ventral musculature of Breviceps.

D, Clavicular head of deltoid. P.abd. Pectoralis abdominalis. P.st.ant. Pectoralis

sternalis anterior. P.s¢.post. Pectoralis sternalis posterior. R.abd. Rectus

abdominis. Sk. Skin-muscle of thigh. S.m. Submaxillaris and subhyoideus

hardly distinguishable for some way after their origin. S.ment. Submentalis.

x, 21, Muscles of jaw (not specially studied).

from that muscle in Rana by the fact that there is only one

inscriptio ‘tendinea instead of the four or five of Rana and of

many other Frogs. This one tendinous inscription is placed a

little way behind the origin of the. abdominal portion of the

pectoralis muscle which overlaps it. nowhere. It is hidden

anteriorly by the complete union across the middle line of some

of the fibres of the abdominal pectoral.

1908. ] ANATOMY OF A FROG. 17

The pectoralis muscle comes next in order and consists of the

usual three parts. The portio sternalis anterior is very much

larger than the portio sternalis posterior, and is at its origin in

the middle line nearly three times the diameter of the latter.

It is incompletely divided into two portions. There is no gap

between the two parts of the portio sternalis, nor between the

posterior of these and the portio abdominalis ; nevertheless the

several muscles are not in any way confused at their origins;

they are perfectly distinct. The portio abdominalis is large and

important. It allows no trace to be seen of any scapular portion

of the obliquus externus such as is figured in Ecker’s work upon

the Frog. It arises mainly from the surface of the rectus

abdominis in front of the single tendinous inscription of the latter.

There is also an origin not represented in Rana (or represented

indeed by the posterior part of the portio sternalis of that

Amphibian) from the expanded cartilaginous sternum. The

fibres of the two sides of the body here meet in the middle line of

the sternum. ‘This region of the portio abdominalis is not, how-

ever, separated in any way from the rest of the muscle; its fibres

he side by side with those of the rest of the muscle and there is

no gap anywhere. Indeed, on both sides of the body the sternal

fibres of the portio abdominalis actually overlap the hinder edge

of the portio sternalis posterior, that part of the latter muscle in

fact which arises from the rhomboidal cartilaginous sternum;

for the latter muscle arises more from the edge of the sternum,

while the fibres of the portio abdominalis arise from the ventral

surface of the same cartilage. There being no omosternum in

Breviceps, there is no superficially visible equivalent of the

sternoradialis of Lana. The only thoracic muscle visible in

front of the pectoralis is the clavicular head of the deltoid, which

is Shown in the figure referred to (text-fig. 3, D).

The throat-muscles visible on the superficial view are again

different from those seen in Rana without any further dissection

than the removal of the skin. As in Fana, a large sheet of

muscle occupies the throat which obviously consists, as in that

Frog, of the submaillaris and subhyoidews muscles. The two

halves of each of these are separated along the median line by a

very narrow tendinous raphe. Bui whereas in Rana by far the

greater part of this sheet belongs to the submaxillaris, only a

slender slip posteriorly being referable to .the subhyoideus, the:

precise reverse is the case ih Breviceps. In fact in the Frog

which forms the subject of the present memoir, the diameter

(antero-posterior) of the submaxillaris is 2°5 mm., and of the

subhyoideus is 5‘) mm., the measurements being taken near to

the middle line. This is accounted for of course by the reduced

lower jaw of Sreviceps; but not entirely so, since fibres of the

muscle, which, did they continue in a straight course, would reach

the ramus of the lower jaw, bend posteriorly to form part of the

mass of the subhyoideus. It should be stated in explanation of

the above, that medianly there is no differentiation of the two

Proc. Zoo. Soc.—1908, No. II. 2

18 MR. F. E. BEDDARD ON THE [Jan. 14,

muscles : it is only when they diverge a little to the inside of the

mandible that they can be distinguished. In front of this lies

the submentalis which has quite normal relations. In addition

to these three muscles of the throat which are quite as recognisable

in Rana, Breviceps possesses another small muscle which is not

visible in a corresponding dissection of Rana. This is seen on

one side on the right (text-fig. 3, «), running along the inside of

the jaw as a fairly broad slip of muscle passing out of sight just

behind the submentalis. Between the subhyoideus and the wall

of the skull and the articulation of the mandible, there is a

considerable space left which is not occupied by muscle. It is

filled with a loose tissue which I have not investigated farther.

In it, however, lies a large circular flattened and somewhat

muftin-shaped body which I take to be the thymus gland, on

account of its general (though not minute) agreement in position

with the thymus of Rana.

§ Muscles of Shoulder-girdle*.

The latissimus dorsi is not a large muscle, and it is entirely hidden

for the whole of its course by the muscular origin of the obliqui

externus et internus. And these latter muscles are too thick to

allow of the latissimus dorsi being seen through them; they have

to be dissected away to bring that muscle into view. Not only

is the latissimus dorsi a small muscle relatively speaking, but it

extends for a much shorter way backwards than in Rana, owing

perhaps and partly to the very forward position and the small

size of the suprascapula. The latissimus dorsi does not at all

overlap the infraspinatus.

The cucullaris does not cover the occipital region of the

longissimus dorsi as is the case with Rana guppyi; nor has it so

straight a course from the occiput to the border of the supra-

scapula. Furthermore it skirts the curved dorsal border of the

suprascapula to be inserted into the posterior angle of that

cartilage.

The retrahens scapule belongs in this Anuran to the serratus

(or transverso-scapularis) series, that is to say it arises from

transverse process and not from spinous process. It is necessary

to emphasise this point because I have shown that in Rana

guppyt this muscle does so arise, and we may therefore fairly

speak of it as a rhomboidews. I take this opportunity of con-

firming that fact on ihe results of the dissection of another

individual. But whether Breviceps can be said to possess this

muscle is a matter open to dispute. In Rana (guppyi as well as

esculenta) there are three muscles which have been termed serratz,

but which in Haslam’s edition of Ecker (made use of by myself

in the preparation of the present communication) are described

as transverso-scapularis. Of these muscles two are broad and flat

* The pectoralis as well as the omoabdominal are described under the ventral

musculature on p. 22.

1908. ] ANATOMY OF A FROG, 19

and pass directly upwards, or very nearly so, from the region of

the transverse process of the vertebra to the under surface of the

suprascapula. They really arise from the transverse process. The

third muscle (text-fig. 7, A, t.sc., p. 28) is long and slender, arising

from the tip of a transverse process and inserted lower down on the

scapula. In Breviceps the slender third muscle (transverso-scapule

tertius) is plain; but there are only two of the broad dorsally

running portions, and one of these has been already referred to

as the retrahens scapule. Moreover these muscles seem to me

to arise from the surface of the longissimus, and not from

transverse processes.

Coraco-humeralis—When the posterior part of the portio

sternalis and the portio abdominalis of the pectoralis are cut

through and reflected, two muscles are brought into view without

any further dissection and when the Frog is lying upon its back.

These are of unequal size, the smaller of the two lying nearest to

the coraco-humeral margin. I am disposed to regard this muscle

as corresponding to that which I have termed “ pectoralis minor”

in Pipa and in Rana guppyt. It arises mainly from the coracoid,

but some of its fibres appear to reach the sternum. It is inserted

on the humerus to the posterior side of the insertion of the

pectoralis abdominalis, 2. e., below that muscle as seen in the

position of dissection referred to. The larger muscle I term

therefore coraco-humeralis, which arises from the same bone and

cartilage as the last. Towards its insertion it is a much larger

muscle than the last described, and its insertion is farther down

the humerus, 7. e., nearer to the hand. Both muscles are fleshy

throughout.

$ Muscles of the Dorsal Surface.

In Rana the muscles of the back are covered by, and also in

some cases arise from, the fascia dorsalis which is attached in the

middle line to the spinous processes of the vertebre. In Rana

guppyt, where on account of the size of the frog this fascia is

specially thick, it has not obviously any more relations to the

depressor mandibule than to the latissimus dorsi or the obliquus

externus, all of which arise from it and are in perfect continuity

with it. It cannot be spoken of as the tendon of origin of any

one or indeed of all of these muscles. It is deseribed by Ecker in

his ‘Monograph of the Frog’ in a separate paragraph as something

distinct from the ensuing muscles. It is to be assumed, however,

that in common with many other tendinous structures this fascia

dorsalis is to be referred to a previously existing sheet of muscle

But there is nothing in Rana to connect it definitely with any of

the muscles which arise from it, excepting perhaps the obliquus

externus on account of its larger size. It must be remembered,

however, that in Xenopus (Dactylethra) the latissimus dorsi is of

very large size*, and in shrinking to the dimensions which it

* Maurer, “Die ventrale Rumpfmuskulatur der Anuren Amphibien,” Morph.

JB. 1895; and Beddard, P. Z. 8. 1895, p. 846 (and footnote).

Das

20 MR. F. E. BEDDARD ON THE (Jan. 14,

shows in Rana may have, so to speak, left behind it a tract of

tendon—the fascia dorsalis. The condition of the dorsal muscula-

ture in Breviceps suggests a different explanation. There is no

fascia at all in the middle of the back; only anteriorly between

and upon a portion of the suprascapule is a thin transparent

sheet of ligament to be detached from the underlying structures.

Text-fig. 4.

77e. €. T.

Superficial dorsal musculature of Breviceps.

ec. Cutaneous muscles at end of rectum. Coce.Sac. Coecygeo-sacralis. H. Posterior .

lymph-heart of left side. On right the corresponding heart is represented as

cut open. Inf. Infraspinatus. Z.d. Longissimus dorsi. —m. Extrinsic

muscles of lymph-heart. OdJ.ext. Obliquus externus. Ob/.in¢é. Obliquus

internus. 7. Muscles covering rectum. Se. Suprascapula. Sk. Skin reflected,

showing attachment of extrinsic muscles of lymph-heart.

Posteriorly this is continuous with a sheet of muscle on either side

of the body the fibresof which pass obliquely backwards and ventral-

1908. ] ANATOMY OF A FROG. 21

wards and which arise from the middle line of the back, ¢. e. from

the spinous processes of the vertebree. These fibres are continuous

with some of those upon the ventral surface which I have shown

reasons for believing to be the obliquus externus of Rana.

Immediately beneath them, and equally plainly shown in the

drawing (text-fig. 4), is a set of fibres, also arising from the

spinous processes of the vertebre, which run diagonally to the

former and are in fact disposed at right angles to the longitudinal

axis of the body. These are of course he fibres of whe. obliquus

internus. I imagine that these two muscles together in their

dorsal region represent the fascia dorsalis of Rana. ~In any case,

apart from any question of homologies, it is remarkable to have to

note the extreme muscularity of the back of this small burrowing

toad when compared with Rana. Thissheet of muscle completely

covers the latissimus dorsi, but it only partly covers the infra-

spinatus, the anterior half of which appears, as is shown in the

drawing (text-fig. 4), beyond its anterior margin. In this

region, in fact, it passes into an aponeurosis. A striking feature

of ine flor sal ningerlefanine of this Frog as compared with Rana, is

the absence of a depressor mandibule. Hence the suprascapula

is visible directly the skin is raised. In view of the presumably

feeble action of the jaws in this ant-eating toad, the absence of

this muscle is not surprising. Posteriorly the obliquus internus

ends abruptly at the commencement of the posterior lymph-

heart*. At that point the longissimus dorsi emerges from beneath

its shelter, and is seen to arise from the coccyx to very far back,

in fact within a millimetre of its posterior extremity. This is

quite different to what occurs in Rana, where the greater part of

the coceyx is free from the longissimus dorsi. It may perhaps

be argued from this fact, coupled with the fact that the end of

the coccyx is a long way from the anus, that Breviceps is as com-

pared with “ana a short-tailed frog. It also follows that the

insertion on to the coccyx of the ilio-coces ygeal muscle is hidden

by the longissimus dorsi.

Since only the extreme tip of the coccyx, represented in the

figure to which reference has been made, is free from the attach-

ment of the longissimus dorsi, it is plain that the coccygeo-sacralis

must be either absent or. have rather different relations. In view,

however, of the very large transverse processes of the sacral

vertebre in this Batrachian, the muscle would be hardly likely to

be absent, and indeed I identify it as shown in the drawing (text-

fig. 4, Coce.Sac.). The muscles in question are attached on each

side of the body to the strong transverse process of the sacral

vertebra. Not, however, to the whole of that process. For the

outer part bears the origin of the gluteus. The muscle (the

coceygeo-sacrel is) is pyramidal in form ; it narrows from its wide

origin to the region of the pyriformis, up to which muscle it passes,

and is attached at the extremity of the coceyx between the origins

* Wor the description of which see p. 33.

22 MR. F. E. BEDDARD ON THE [Jan. 14,

of the longissimus dorsi and the pyriformis. This is true of the

superficial layer of the muscle. The deeper fibres are inserted

upon the coccyx still farther forward, underlying the longissimus,

but I have not drawn an exact boundary line between this and

the ilio-coceygeal muscle.

When the region of the ilium is inspected from below, the 2/io-

lumbaris muscle is very plainly seen. It arises quite from the

tip of the ilium, where it overlaps not as might be expected the:

coccygeo-iliacus but the gluteus, or rather a portion of it. Owing

to the abbreviation of the tail the origin of the coceygeo-iliacus is

concealed on this aspect of the body. The ilio-lumbaris (text-fig.

7, A, Zl.1., p. 28) isa strong muscle not broken up into segments as.

in Rana guppyi*, but passing straight forwards to its termination

on the transverse process of the third vertebra. It gives off from

its concealed (7. e. dorsal) surface bundles of muscular fibres to the

transverse processes of the intervening vertebra. This muscle, as.

it appears to me, is in some ways like that of the Pelobatide 7.

For instead of consisting only of detached slips as in Rana running

from transverse process to transverse process, there is also a massive

band of muscle running straight to the most anterior transverse

process to which the muscle is attached. This muscle is not,

however, separate as in the Pelobatide, arising from a lower

(more posterior) part of the ium; it is indistinguishable at its

origin from the anterior end of the ilium from the rest of the

muscle.

§ Abdominal Muscles.

The general aspect of the rectus abdominis as compared with

that of Rana has been already described t, and need not be again

here referred to. While in the Common Frog according to various

authors the rectus abdominis does not extend far laterally, but is.

in those regions replaced by the obliquus eaternus, there is in

Breviceps an absolute continuity between the fibres arising from

the pubic symphysis in the two strong pillars already referred to

and fibres running in a nearly dorso- ventral direction on the sides

of the body (text- -fig. 5). They form obviously one sheet which

may be stripped off. These fibres extend a long way towards the

dorsal median line. More towards the yentral median line, how-

ever, they become attached to the tendinous inscription, and with

that break are continuous with anteriorly running fibres, some of

which end on the sternum. Beyond the tendinous inscription

arises, aS has been already said, the portio abdominalis of the

pectoralis. Beyond this again, 2. e. nearer the shoulder-girdle,

arises a sheet of muscle which is completely hidden by the

pectoralis abdominalis. Nevertheless it lies above (ventral to) an

underlying sheet of muscle, which latter lies on the same plane

as the median region of the rectus abdominis, arising as it does

from it or at least from the aponeurosis covering it. This muscle

* Beddard, P. Z.S. 1907, p. 383, text-fig. 94.

+ Id. ibid. p. 877. tL Supra, p. 15.

1908. | ANATOMY OF A FROG. 23

arising from the surface of the deeper layer is very wide, and

disappears anteriorly beneath the shoulder-girdle, being inserted

dorsally upon the scapula. It would seem to correspond to the

portio omo-abdominalis of the obliquus externus of Rana, but is

clearly much more extensive. The direction of its fibres is on

Text-fig. 5.

om.aba.

—hy.abd.

abd. Wf \) d—Pect. abd.

ObT.

wu.

—P.abd.

Ventral abdominal musculature of Breviceps cut so as to display

different layers.

hy.abd. Hyoabdominal. Obl.inf. Obliquus internus. Om.abd. Omoabdominal

shown below at origin and above near to insertion, the intervening portion

having been cut away. Pect.abd. Origin of pectoralis abdominalis. R.abd.

Rectus abdominis.

the whole obliquely postero-anterior. It arises from near the

edge of the underlying sheet of muscle also just referred to.

24 MR. F. E. BEDDARD ON THE [Jan. 14,

This latter is, as I take it, the obliquus internus. In the middle

line it can be seen to be covered by the very thin sternal portion

of the rectus abdominis. Here its fibres run obliquely forward

from the median line, fairly parallel indeed in direction with the

fibres of the adjacent and covering pectoralis ablominis. From

the inscriptio tendinea the fibres of the obliquus internus run

directly forward parallel to the long axis of the body; more

laterally they run postero-anteriorly but obliquely towards the

median ventral line of the abdomen, Passing round the abdomen

the direction of the fibres is gradually changed, until anteriorly

just behind the shoulder-girdle the fibres of the obliquus internus

run exactly at right angles to the long axis of the animal’s body.

The obliquus internus, therefore, of Breviceps is very different

from that of Hana, where the fibres run obliquely postero-anteriorly

with a main dorso-ventral direction and with but a slight fanning

out from the back towards the ventral surface. In Breviceps the

fanning is much more marked and is in the opposite direction,

a. e. the fibres converge towards a point upon the ventral surface

on each side of the body. This point, or rather area, is formed

by the origin of a very strongly marked muscle, flat and of con-

siderable diameter, from the septum between itself and the

obliquus internus and running forward parallel with the sternum

to be attached to the hyoid. ‘I term this muscle the hy yo-abdom-

inalis, and I regard it as being, like the omo-abdominalis, a

portion of the obliquus externus. Against this view, however, 1s

the fact that it is, in the greater part, covered by the omo-abdo-

minalis. I would further remark that this muscle apparently

has its homologue among the Pelobatide *, where, however, it

is not quite so important as in Breviceps and has a different in-

sertion. I am disposed to regard this peculiar arrangement of the

obliquus internus and its relation to a large hyo-abdominalis and

the large size of the omo-abdominalis, as being connected with

the ant-eating habits of Breviceps. The arrangement of the

muscles in question is such as to produce a powerful pull upon

the hyoid apparatus andtongue. The very slender sternal portion

of the rectus is to be associated with the rudimentary and reduced

state of the cartilaginous sternum. The large omo-abdominalis is

possibly associated with the burrowing habits of the Frog. It

would assist in producing a strong pull upon the shoulder-

girdle.

The abdominal musculature therefore of this Frog agrees with

that of other Anura in the possession of only two layers of muscle.

But the exact homology between the variously metamorphosed

regions in this and other Anura is clearly difficult to settle. The

obliquus internus, as I have termed the inner sheet of muscle,

seems to be comparable not only with the obliquus internus of

Rana but to that muscle plus certain parts of the rectus

abdominis. For the anteriorly directed fibres of the muscle in

* Beddard, “ On Megalophrys nasuta,” P. Z. 8.1907, p. 340 ; id., “On Pelobatide,”

ibid. p. 894.

1908. | ANATOMY OF A FROG. 25

Breviceps ave obviously part of the same muscle, most of whose

fibres have an oblique direction. But elsewhere the rectus ab-

dominis is obviously external to the obliquus internus. It would

seem, in fact, impossible to go further than to declare the two

layers of the abdominal musculature homologous with the two

layers in other Batrachians. An exact homology between indi-

vidual muscles derived from these layers would seem to be

impossible of assertion. It is quite remarkable to note what

great differences in the disposition of the fibres in these two

layers can exist between closely allied Batrachians, and how

plastic these structures prove to be as contrasted with many

others whose functions would also seem to be involved with the

peculiar mode of life of this frog.

$ Muscles of the Thigh.

The muscles visible on the superficial aspect of the thigh are

shown incidentally in the figure (text-fig. 3) representing a

general view of the musculature of Lreviceps, and in a more

Text-fig. 6,

Muscles of the inside of the thigh of Breviceps.

a. Skin-muscle referred to in text. Add.bv. Adductor brevis (or magnus?). Add.D.

Adductor longus. Add.m. Insertion on to knee of one of adductors, perhaps

comparable to the adductor magnus of other Frogs. R.i.maj. Rectus internus

major. R.i.min. Rectus internus minor. V.i. Vastus internus. w. A separate

adductor slip.

elaborate way in the accompanying figure (text-fig. 6), which

represents the thigh-muscles more highly magnified. I take as

26 MR. F. E. BEDDARD ON THE [Jan. 14,

usual the corresponding set of muscles in Rana for comparison *,

in order to set forth those of Breviceps. There is an important

difference at the very beginning of this comparison. When the

skin covering the thigh is removed or reflected, there is removed

or reflected with it a thin sheet of muscle (text-fig. 6, @) which, in

the middle of the thigh, lies superficial to all the other muscles of

the thigh. Its insertion on to the knee is also the most superficial

insertion. ‘There seems to be no doubt that this muscle, which

underlies the skin over a great part of the thigh, actually arises in

part at least from the skin, and is therefore perhaps to be referred

to the series of cutaneous muscles which have been described in

Rana. But there would appear to be no corresponding muscle to

this in Rana guppyi at anyrate. Posteriorly the muscle is not to

be distinguished for a great part of its course from the Rectus

internus minor. Anteriorly it does not spread on to the abdomen.

It seems to have nothing to do with the Rectus abdominis or

adjacent muscles. It is purely a thigh-muscle in its position and

extent. It is necessary to mention this in view of the peculiar

relations of the abdominal muscles to the thigh in Pipa. Although,

as already said, there is a close contiguity posteriorly with one

part of the Rectus internus, its insertion onto the knee seems to

be distinct from that of the said Rectus, which muscle will be dealt

with presently. Apart from this muscle, which is something

superadded, possibly in relation to the inclusion of the thigh within

the contour of the body, the general plan of the femoral muscles

appears to be not unlike that of Rana.

The sartorius courses obliquely over the thigh as in Rana; its

insertion on to the knee is entirely fleshy. It is not a particularly

large muscle, being markedly smaller than the neighbouring

adductors. Nevertheless, its position and relations seem to fix its

correspondence with the sartorius of Rana, &e. When cut across.

the muscle is seen to lie in the hollow between its much more

massive neighbours. These I take to be the adductor longus and

(possibly) the adductor magnus respectively. In the appearance

of the former superficially upon the inside of the knee we have a

character not found in Rana. This muscle, however, is also

inserted on to the inner border of the femur for about the distal

third of that bone, and must be, as I imagine, the equivalent of

the adductor longus of Rana. In front of it lies, as should be the

case if this homology be true, the vastus internus, which is a large

muscle.

Between the insertions on to the knee of the two muscles last

dealt with a portion of the insertion of another muscie is visible, as

clearly shown in the accompanying figure (text-fig. 6, Add.m.). This

also belongs to the adductor series, and may perhaps be regarded

as the adductor magnus, the third adductor described above being

in that case the adductor brevis. The importance of the adductor

muscles in this frog is very striking. In a second specimen, the

* T have figured these muscles in Rana guppyi in P. Z.S. 1907, p. 887, text-fig. 234.

1908. ] ANATOMY OF A FROG. 27

insertion of this adductor magnus (if I am right in so terming it)

did not appear superficially. This variability, it will be observed,

is precisely analogous to that exhibited by the constituents of the

triceps femoris on the outside of the thigh, which will be presently

described. Returning to the muscles visible on the inside of the

thigh, the only ones visible without dissection, in addition to those

already treated of, are the recti interni major et minor. The

connection of the latter with a superficial muscle has already been

dealt with. As in Rana the semitendinosus does not appear

superficially.

On the outside of the thigh the most prevalent muscle is the

equivalent of the triceps femoris of Rana, though its constitution

in Breviceps differs somewhat. There are, however, three distinct

portions which may be termed respectively rectus femoris, vastus

eaternus, and vastus internus. They are, however, all of them

inserted separately, instead of by one tendon as in Rana. More-

over, the most anterior of the three muscles, the rectus femoris

anticus, instead of ending in an aponeurosis, is fleshy and thick

throughout. The two specimens which I have dissected show a

difference in the insertion of the middle of the three divisions of

the triceps femoris. In one this goes as far as the knee; in the

other individual the muscle is inserted on to the thigh up to about

the middle of that bone only. It is therefore not only in the

separateness of the three divisions of the triceps femoris, but also

in their insertion and complete muscularity that Breviceps differs

from Rana. The biceps femoris in Breviceps is a particularly

slender muscle ending in a long tendon, not flattened, which

pushes between the two heads of the gastrocnemius some way

after their origins to be inserted a longish way down the fore

leg. The semimembranosus is of fair size.

§ Esophageo-pulmonary muscle.

This muscle in Breviceps is a very stout muscle obscurely

divided into three or four bundles which have hardly the value

of separate muscles. It has no direct connection whatever with

the muscles of the wall of the abdomen. It is not (that is to say,

obviously) a detached sheet of the obliquus internus, as is the case

with the corresponding muscle in all of the Pelobatide that have

been hitherto examined*. It is in fact similar in many respects.

to its homologue in the Ranide and Bufonide. It arises in them

from the transverse process of the fourth vertebra, and this is also

the origin of the muscle in Breviceps. Its origin lies in front of,

and contiguous with, the insertion of the ilio-lumbaris, and to the

inside of the origin of the transverso-scapularis. The origin and

course of the muscle is shown in the accompanying illustration

(text-fiz. 7). It nearly meets its fellow of the opposite side of

the body in the middle line of the ventral surface of the cesophagus.

* Beddard, “On Anatomy of a Frog of the genus Megalophrys,” P. Z.S. 1907,.

p. 324; and “ On Anatomy of Pelobatide,” zbid. p. 886.

28 MR. F. E. BEDDARD ON THE [Jan. 14,

A large number of its fibres end upon the cesophagus both laterally

and ventrally. In fact the muscle is chiefly an cesophageal muscle,

and has less relation with the respiratory apparatus. It is, how-

ever, connected with the root of the lung where the walls of this

sac, as is Shown in the figure referred to, are non-respiratory and

thicker, and is undoubtedly attached along the dorsal median line

of this region of the lung where it (the lung) forms one cavity with

Text-fig. 7.

M1. coce: i MI

(Esophageal and neighbouring muscles of Breviceps.

A, These muscles in situ without disturbance of adjacent structures, which are,

however, not all included in the figure.

B. A further dissection to show relation of cesophageal muscle to root of lung.

Hg. Hyoglossus muscle. Tl.coce. Tio-coccygeus. J.J. Uio-lumbaris. A. Kidney.

I. lung. es. Hsophagus. ces.m. (Hsophageal muscle. P.h. Petrohyoideus

posterior tertius. ¢.sc. 'Transverso-scapularis.

its fellow. So also of course is the muscle of the opposite side of

the body, and the two muscles can be raised here from the surface

of the cesophagus by pulling up the common cavity of the two

lungs just where it opens into the larynx. The attachment con-

tinues on to the cricoid cartilage of the larynx (“annulus” of

Wilder *) which forms in this Batrachian, as in so many others, a

* Zool. Jahrb., Abth. f. Anat. ix. 1896, pp. 290 &e.

1908. | ANATOMY OF A FROG. 29

complete bar uninterrupted in the dorsal middle line. None of

the fibres of the muscle appear to me to actually run on to the

evicoid bar; but their action must result in moving this bar, seeing

that it 1s intimately and strongly connected by their walls with

the lungs. Any pull on the roots of the two lungs must tend to.

raise the cricoid cartilage dorsally.

§ Abdominal Viscera.

The Liver of this Frog (text-fig. 8) is of large size and has the

unusual character among the Batrachia Salientia that the right lobe

is considerably the larger of the two lobes into which it is divided.

Furthermore, the left lobe can hardly be said to be divided into

two lobes, as is again so frequently the case with Frogs, though an

indentation on its border is an indication of such a subdivision.

The liver is, as a whole, very square-shaped. Anteriorly its.

boundary line is almost straight and is on a level with the

posterior border of the coracoid. The two lobes are nearly in

contact in the middle line and hardly diverge posteriorly, so that

the posterior border of the liver is almost straight. Anteriorly,

however, in the middle line they diverge slightly and form a small

triangular space; this discovers the heart, which is otherwise

quite covered by the liver except for the narrow median ventral

slit between the two liver-lobes, where it is apparent. The apex

of the ventricle is situated a little way in front of the posterior

border of the liver. The smaller left lobe is more triangular in

shape than the right lobe. The gall-bladder is quite invisible on

a superficial view; it lies beneath the inner corner of the right

lobe. As far as I can gather from Dr. Ginther’s account of the

Bufonid Lhinophrynus dorsalis*, the liver of this toad bears

some likeness to that of Breviceps. For he remarks + that the

heart of Rhinophrynus “is surrounded by the liver in a similar

way as in higher animals, asin other Batrachians it is surrounded

by the lungs.” Furthermore, he observes of the liver that it is

divided into a right half and a larger left half. This would seem

to be the exact converse of what I note here in Breviceps. But

elsewhere in the paper Dr. Giinther speaks of the stomach being

situated “quite on the right hand,’ which causes me to doubt

whether right and left may not be used to express the positions as

seen from above during a dissection. Furthermore the liver

extends dorsally to the heart, which thus lies in cavities as it

were excavated in the liver-substance, and it is almost completely

surrounded and hidden by that viscus as in Reptiles and Birds.

The likeness between Lhinophrynus and Breviceps in these features.

of liver construction are remarkable as possibly related to the ant-

eating habit which they have in common, since systematically

they are placed in different families. Part of the stomach is con-

cealed by the left lobe of the liver, and the small intestine passes

* “The Systematic Arrangement of the Tailless Batrachians, &c.,” P.Z. S. 1858,,

p. 339. + Loc. cit. p. 350.

30 MR. F. BE, BEDDARD ON THE [Jan. 14,

directly anteriorly between the lobes of the liver; most of the

coils of the intestine lay entirely above (7. e. quite concealed by)

Text-fig. 8.

ff.

Ventral surface of liver of two individuals of Breviceps.

Ant.abd. Anterior abdominal vein. g.6. Gall-bladder. H. Heart.

L., R. Left and right lobes of liver,

The enormously distended large intestine

the large right lobe.

The peculiar nature of the

was entirely uncovered by the liver.

1908. } ANATOMY OF A FROG. 31

liver and its relations to the heart have been brought about, or at

any rate are accompanied, by an alteration in the normal (7. ¢. that

found in Rana) disposition of the anterior abdominal vein. This

vein in Breviceps gives off no branches to either lobe of the liver

until it reaches and has passed some way beyond the apex of the

heart.

Text-fig. 9.

wae B.

A. Part of oviduct. B. Stomach of Breviceps (laid open).

a. Demarcation between stomach and duodenum. cs. Césophagus.

f. Funnel of oviduct.

In a second specimen which I dissected, a male, and which was

perhaps in consequence rather smaller than the first, measuring

only 33 mm. in total length, the liver showed certain differences.

The relative size of the two lobes was the same, but they were

not by any means so closely approximated in the middle line as

was the case with the larger example. This being so, much more

of the heart was visible on a ventral inspection of the viscus, and

the conditions more approached that to be seen in the Common

Frog. Still, however, there was a considerable difference ; for in

Rana the heart is distinctly ventral of the liver and the lobes

extend far beyond it in the direction of the cloaca. In Breviceps,

32 MR. F. E. BEDDARD ON THE (Jan. 14,

even in this second example to which I am now referring, the

liver only just extends beyond the heart and grasps it firmly, as

it were, with a lobe on either side. And the pericardium comes

into contact and is connected with the absolute ventral edge of

the liver-lobe on either side. I could detect no furrowing of

either lobe of the liver. The gall-bladder in this, the smaller

specimen, instead of being completely hidden, was quite visible

for the greater part of its extent between the ends of the two.

lobes of the liver and extending beyond them even towards the

cloaca.

The Stomach of this Frog (text-fig. 9) seems to be peculiarly

large, and its pyloric projection is sharply marked off from the

ensuing duodenum, not only by a constriction but by the fact that

the walls of the latter are much thinner than those of the pylorus.

The pyloric region of the stomach in fact is quite distinct from

the rest of that organ and forms a projection from it; there is

no gradual passing of the one into the other as in Rana. The

stomach itself was swollen and nearer to the spherical than to the

ovoid in form. It was full of a mass of ants, among which I dis-

tinctly recognised “soldier ants” (with enormous heads) of a

species which I have not attempted to identify. The muscu-

larity of the stomach was very evident, and it thus contrasted

with the gut. The cesophagus, which suddenly expands into this.

large stomach, only just enters the abdominal cavity. On cutting

open the stomach the smallness, relatively speaking, of the

cesophageal aperture into it can be realised.

The Lungs float very freely in the ccelom; it is only at the base

that they are attached by ligaments. The texture is thin with

large alveoli.

Text-fig. 10.

— St.

Intestinal canal of Breviceps.

st. Stomach. Zi. Colon.

The Jntestine (text-fig. 10) does not appear to differ greatly fron»

that of Rana in its proportionate length. I+ differs, however, very

considerably in certain features of itsstructure. The first portion

1908. | ANATOMY OF A FROG. 33

of the small intestine, that immediately following upon the pyloric

constriction, corresponding to the duodenal loop of Rana, is very

short, about 5 mm. long (in the smaller male specimen), and

directed rather forwards and parallel with the stomach as in

Rana. It is of narrow calibre, about that of the pyloric process

of the stomach. This passes into a middle section of the small

intestine, the bore of which is quite twice that of the preceding

part, and which forms therefore a very wide tube. ‘This is no case

of accidental dilatation, for the same structure was apparent in both

examples, and the appearance of the swollen region of the intestine

was quite different from the region in front of and behind it.

Internally its mucous membrance is raised into transverse folds,

which have not the watch-pocket shape of the corresponding folds

in the intestine of Rana, but are thin wavy folds running right

round the lumen. ‘This wider portion of the small intestine passes

into a narrow portion again, which is of greater length and of not

greater calibre than the beginning of the duodenum. This opens

suddenly by a shghtly projecting os into the very wide but not

very short large intestine.

The end of the colon, into the pear-shaped anterior section of

which the ileum opens, bulges equally all round the ileum; that

is to say, there is no unilateral cecum. Shortly after the entrance

into it of the small intestine the colon diminishes in its width and

remains at about the same calibre to the anal apertures. That

is to say, of course, the cloaca is no wider than the antecedent

colon. The cloaca is of considerable length, and is shown in the

figure on p. 20 (text-fig. 4). It occupies the whole of the space

lying between the tip of the coceyx and the rather distant end of

the body. Being, as it is, in this exposed situation and covered

ouly by the skin, it would appear to be rather susceptible to

injury from pressure upon the dorsal surface of the body. ‘It is,

however, protected from such injury, it may be supposed, by a

thick covering of muscle upon the dorsal surface. This is obviously

divided into two longitudinal bands by a furrow, and this arrange-

ment becomes clearer still when the tube is cut across. This

muscle arises, partly at least, from the tip of the Hsoshyle and

corresponds, Ti imagine, to the compressor cloace of Rana*. There

are also two very tiny muscular slips attaching the end of the

cloaca to the skin just above; they are also siatti in the figure

referred to.

§ Posterior Lymph-hearts.

These organs are so extraordinarily developed in Breviceps that

they require a section to themselves for their adequate descrip-

tion. The general topography of their neighbourhood may be

first described, as it differs greatly from that of Rana and is in

relation to the large size of these sacs in Breviceps. As already

mentioned, the thighs of this species are enclosed within the

** Ecker’s ‘ Anatomy of the Frog,’ Haslam’s Translation, p. 348.

Proc. Zoou. Soc.—1908, No. III. 3

34 MR. F. E, BEDDARD ON THE (Jan. 14,

contour of the body which extends for a great distance behind

the coccyx. In an example of Lreviceps measuring 38 mm.

from snout to anus the distance from the tip of the coccyx to the

anus was quite 7 mm., 7.e., more than one-fifth of the total body-

length. Furthermore, the breadth of the thighs adds to the

large space which intervenes between the posterior edge of the

abdominal muscles where they reach the back and the posterior

end of the trunk. Itis in this large space (see text-fig. 4, p. 20),

loosely covered by the skin which does not adhere to the leg until

the knee, that the posterior lymph-hearts lie on either side. They

do not, however, occupy the whole of this considerable tract.

Each, however, is no less than 10 or 11 mm. long in the individual

Breviceps whose total body-length has been mentioned above.

The posterior lymph-heart of Breviceps is therefore between one-

quarter and one-third of its total body-length. With this may be

contrasted the proportions found in Rana, where (in Rf. temporaria

or &. esculenta—very much larger species) the length of the

_ posterior lymph-heart is given in Haslam’s Translation of Ecker’s

‘Frog’ as “about two lines,” z.e. 4 or 5 millimetres*. That is to

say, the posterior lymph-hearts of a frog half or one-third of the

size of Rana esculenta are twice or thrice the bulk of those of that

Rana. This appears to me to bea very remarkable anatomical

fact, and one which argues considerable physiological differences.

The posterior end of each lymph-sac was about 4 or 5 mm. from

the posterior end of the body. This space was ecccupied by a

a lymph-sae corresponding, I presume, to the femoral lymph-sac

of Rana. It lies at any rate on the thigh-muscles. I found this

space on each side filled with a coagulated flocculent mass, pro-

bably lymph. ‘This space bears a relation to its corresponding

lymph-heart similar to that of an auricle to a ventricle. When

the skin of the back is carefully reflected from the middle line,

the lymph-heart is at once exposed. No muscles lie between it

and the integument. It is, however, slightly attached to the

skin here and there by fibres continuous with its own muscular

walls. These fibres arranged in slender bundles spread out in a

fan-shaped fashion over the skin. They form presumably a fixed

point or points to render effective the contractions of the lymph-

heart. It may be also that the fibres thus attached belong really

to the cutaneous muscular system and correspond in particular to

the cutaneus dorsi of Rana; for other cutaneous muscles are

associated with the septa of lymph-sacs. In a general way also

these fibres suggest the “ale cordis” of Arthropods. In any

case the anatomical facts are as has been stated, and are shown

in the accompanying figure (text-fig. 11). These fibres as well

as the lymph-hearts lie dorsally to and unconnected with the

dorsal muscles (which are described on another page 7), although

the anterior end of the lymph-heart overlaps the end of the dorsal

* In a large example of Bufo vulgaris measuring 115 mm., I found a lymph-

heart to be 5 mm.

+ Supra, p. 19.

1908. | ANATOMY OF A FROG. 35

muscles. It is mainly at least by the anterior half or three-

quarters of each lymph-heart that it is fixed to the integument

by these muscular strands, which have to be cut through in order

to free the organ. When it is thus freed it is seen to be of about

the same shape as an acorn (without its cup) and divisible into

two regions, an anterior and posterior, which are however not

very sharply marked off from each other.

Mext ation lia:

o.d.

Left lymph-heart of Breviceps.

a, 6. Extrinsic muscles of heart (H.). G. Tip of coceyx. Obl.int. Obliquus

internus cut off short on both sides. o.d. Oviduct.

The anterior part of each lymph-heart(asis shown in text-fig. 12,

p. 36), which represents one of these structures completely freed

from its attachment to the integument, is of very dense muscular

structure, and in consequence quite smooth. The posterior part

(not half) of the heart, however, is of a basket-work conformation,

strands—varying in breadth, but always broadish—of muscular

fibres crossing each other at right angles. The bands of fibres

which run in a longitudinal direction are ultimately lost in the

regular even and muscular walls of the anterior part of the lymph-

heart. The cross-running bands are also seen in the figure to be

also gradually differentiated fromit. The interspaces between these

bands are considerable, and possibly permit of the free entrance

36 MR. F. E. BEDDARD ON THE - [Jan. 14,

of the lymph into the heart from the lymph-sac already referred

to which lies behind it. When a lymph-heart is cut open the

structure presented is that which is represented in the lower

figure, text-fig. 12. There is the same division into two regions,

the walls of the anterior part being smooth and thick. In the

posterior part the longitudinally-running bands of muscle stand

out from the walls, projecting into the interior of the sac, as is

shown in the figure.

Text-fig. 12.

Upper figure, lymph-heart of Breviceps isolated ; lower figure, view ot

interior of same.

The attachment of each lymph-heart to the integument by

strands of muscles would doubtless increase the efficiency of that

organ as a pump by providing a fixed point for the contraction of

its muscles to pull against. There is, however, another series of

muscles attached to the opposite side of each heart, and acting

in a direction parallel to and in the same plane as, but obviously

1908. | ANATOMY OF A FROG, 37

opposite to, those integumental sheets. The resemblance of the

extrinsic muscles of the heart to the “ale cordis” of the Arthro-

pod heart is thus increased, since there is (or at least may be) a

pull on each side in opposite directions which would clearly dilate

the cavity of the lymph-heart. The contraction of the heart is

probably effected by the intrinsic musculature. This muscle,

when the heart is viewed from the dorsal surface as in text-

fig. 11, is seen to reach the heart as two broad flat bands which

fan out over its dorsal surface. They are seen to dip down

ventrally between the heart and the ilium, and to unite to form

one stout strap-shaped band of muscle. The position of this

muscle is exactly on a level with the tip of the coccyx, and there

is an exact symmetry between the two of opposite sides of the

body. It is inserted on to the symphysis pubis.

In addition to the muscles just mentioned, which are indicated

in text-fig. 4 (p. 20), and shown more in detail in a more enlarged

representation of this region of the body (text-fig. 11), there is

another muscle attached to each lymph-heart more ventrally than

that which has just been described. This completes the mooring

of the lymph-hearts to the adjacent organs of the body. The

muscle now under consideration is single on each side of the body ;

that is to say, there is one of them to each lymph-heart. It is

broad and flat and thin, and shows a metallic glitter on account

of its structure; it arises in the neighbourhood of the edge of

the expanded transverse process of the sacral vertebra, and

thus partly covers over and conceals the coccygeo-sacralis muscle.

The course is backwards, and it reaches the first described skeletal

muscle of the lymph-heart at right angles to that muscle. It

dips under it, and is therefore attached to the lymph-heart rather

ventrally. The muscle is broader than that which runs from the

pubis to the lymph-heart, but could be readily missed owing to

its tenderness and the consequent ease with which it can be

torn *.

§ Organs of Reproduction.

T have had the opportunity of examining both sexes of this frog,

as has already been mentioned in relation to the alimentary system.

In the male (text-fig. 13), the testes have the usual oval form and

are not pigmented. From their considerable size, I gather that the

individual was sexually mature. They lie very close together and

actually indeed in contact, the mesocolon only just being able to

push itself between them, asit were. Nor can they be separated

by any manipulation short of forcibly tearmg them away from

the dorsal mesentery which attaches them and the colon to the

middle dorsal line. The vasa efferentia seem to offer a new

form of these ducts among the Anura. There is only a single

* Theappearance of the lymph-heart is by no means unlike that of the Tortoise as

represented by Fritsch, “Zur Anatomie der Elephant-Schildkréte (Testudo ele-

phantopus),” Prag 1870, from Abh. k. bohm. Ges. Wiss. 1871.

38 MR. F. E. BEDDARD ON THE [Jan. 14,

tube arising from each testis, and each of these meets its fellow

of the opposite testis and forms with it a single duct; this

occurs soon after each has emerged from the testis.

The two ovaries are, like the testes, closely apposed in the

middle line, and as already mentioned each has a very large fat-

body attached to it anteriorly. The eggs were of considerable

size (some of them) and, as I should imagine, mature. But the

ovaries were not extensive as they are in the mature females

of other Batrachia. It may be therefore that the oviducts are

not as complicated in their coiling as they would have been had

the frog lived longer. In this specimen the oviducts were

as is represented in text-fig. 9 (p. 32). Anteriorly the funnel is

spoon-shaped with an elongated aperture on the lower surface.

Text-fig. 13.

Testes and kidneys of Breviceps.

f. Fat-body. p.c. Post-caval vein. ¢. Testes with single vas efferens arising

from anterior end of each. «w. Uvreter.

The proximal section of the oviduct is narrow and straight in its

course. The thick-walled glandular region of the oviduct only

makes two loops, as is shown in the figure referred to. I am

inclined to think that the oviduct is after all mature or very

nearly so, for its walls are as thick as it seems likely they could

become. The glandular part opens into the distal and thin-

walled “uterus” which is flattened and strap-shaped. The two

tubes approach each other in the same straight line behind the

kidneys, and form one tube running of course at right angles to

them towards its opening into the cloaca.

1908. j ANATOMY OF A FROG. 39

§ Fat-Bodies.

Although these organs are known to vary among the Anura,

from individual to individual, from side to side of the body, and

also shrink or become expanded at different times in the life of

the individual *, I think it worth while to describe the appear-

ances seen in the two specimens which I dissected. In both of

them the fat-bodies were large and apparently fully developed.

In the female they were much the larger; but then the female

specimen was considerably larger than the male, a difference

which I am disposed to put down as a sexual character. In the

female the fat-bodies extended forwards a long way and appeared

when the body was opened actually in front of the lungs. In

the male they were much smaller and firmly adherent to the

front margin of each testis. Hach fat-body was divided distally

into five or six finger-like processes of the usual shape.

§ Renal Organs.

The kidneys of Breviceps are like those of Rana in that they

are flat smooth bodies with no division into massive lobes such as

occurs among the Pelobatide. They are represented in text-

fig. 13, which shows also the relations of the testes to them.

The only peculiarity which they show (so far as I have ascer-

tained) is the complete fusion in the middle line of the right and

left viscus in the male. In the female they are very closely

apposed but not fused. This fusion is, however, not complete ;

that is to say, the two organs in the male are not continuous with

each other in the middle line throughout the whole of their

length. It is only for about one-half of their extent that they

are thus fused. Anteriorly the two kidneys are quite distinct

until the emergence of the postcaval vein which bends downwards

just in front of the point where the two kidneys become almost,

if not quite, soldered together. Posteriorly a larger free region

is left.

The common duct thus formed does not, as 1t does in the case

ef Discoglossus and Alytes, enter the ureter direct without passing

through the kidney. In the present species the duct becomes

lost in the substance of the kidney in a way which I did not

follow out more minutely. In any case it did not bend round the

anterior end of the kidney to join the ureter. The two kidneys

are in very close contact below the testes. But the duct showed

no signs of division into a branch of each kidney, but appeared to

enter the middle line, and was at any rate concealed by kidney-

tissue before dividing. As the vasa efferentia referred to were

exceedingly obvious on inspection with a lens, I imagine that

* Boulenger, “ The Tailless Batrachia of Europe,’ Ray Soc. Publication, 1897.

+ I may observe that I found no “ Bidder’s Organ.” a‘

40 MR. F, E. BEDDARD ON THE [Jan. 14,

others have not been missed. At the same time I cannot of

course positively assert that the individual was actually fully

mature, though there was no reason against this view.

§ Resumé*.

As compared with Rana, Breviceps shows the following pecu-

liarities of structure :—

(1) The rectus abdominis has but one tendinous inscription and

its fibres fan out laterally running dorsoventrally, thus

replacing a portion of the obliquus externus.

(2) Part of the obliquus externus is specialised into a strong

muscle ending on the hyoid; on to the base of this is

inserted the fan-shaped obliquus internus.

(3) The sternoradialis is absent.

(4) The subhyoideus muscle is twice the diameter of the sub-

maxillaris.

(5) A special muscle runs from the quadrate cartilage to the

inner side of each ramus of the lower jaw.

(6) The omohyoid is absent.

(7) The sternohyoid is divided into two perfectly distinct

muscles with widely separate insertions into the hyoid.

(8) The obliquus externus is a purely fleshy muscle arising on

either side in the middle line of the back by muscular

fibres. There is thus no fascia dorsalis. The obliquus

internus is similarly fleshy at its origin from the middle

line of the back below the former.

(9) There is no depressor mandibule muscle to be detected.

10) In the thigh the rectus internus minor is connected with a

superficial cutaneous muscle, covering over a portion of

the thigh, which is its chief head of origin. The biceps is

very slender ; the adductors very large.

(11) The posterior lymph-hearts are of enormous size, measuring

more than a quarter of the total length of the body. They

are attached by muscles to the skin dorsally and by two

separate and broad muscles to the transverse process of the

sacral vertebra and the pubis respectively.

(12) The liver is composed of two lobes only which largely cover

the heart.

(13) The testes emit each only one vas efferens; and the two

tubes join before entering the substance of the kidneys

(which are here fused) in the middle line.

(14) The oviducts are very short owing to the fact that they

are but little coiled.

* J do not include in the above reswmé external and osteological characteristics

already well known, nor all minutie of muscular structure.

1908.] ANATOMY OF A FROG. 41

(15) The coccyx is abbreviated and does not extend to the end

of the body. To compensate for this the end section of

the cloaca which naturally extends beyond it is protected

by a pair of strong muscles which are inserted on to the

skin above the anus by short slips.

(16) Upon each sacral transverse process is a detached plate

of cartilage which is related to the adjacent musculature,

and which perhaps corresponds to the supra-sacral portion

of the ilium in Sauropsida.

(17) The hyoid cartilage is marked by the double origin of the

anterior cornua, a foramen therefore occupying the base

of each cornu where it joins the body of the hyoid.

The above list contains a brief epitome of nearly all of the

points in which I have found Sreviceps to differ anatomically

from Rana. Inthe present state of our knowledge of Batrachian

anatomy, it is not possible to use them in order to criticise or

confirm any view which has been held with regard to the sys-

tematic position of Breviceps, except of course to assert that it

is in any case not a near ally of Rana, or of the family

Pelobatide *. Some of its structural features would appear,to be

associated with its ant-eating proclivities; to this category I

would refer the particularly strongly-developed hyoid muscles and

the anterior cornua of the hyoid and the round, globular, some-

what gizzard-like stomach sharply marked off from both cesophagus

and duodenum. The burrowing habits of Breviceps are perhaps to

be associated with some other muscular peculiarities; especially,

as I should imagine, the extent and muscularity of the two

obliqui and of their branches to the shoulder-girdle, and the very

powerful muscles of the anterior part of the thigh, and the

very thick gastrocnemius. Among the remaining characters of

this Frog, those which are particularly noteworthy appear to me

to be the following, viz. :—(1) The enormous size and muscularity

of the posterior lymph-hearts with their special extrinsic muscles ;

(2) the presence of a plate of cartilage overlying the sacral trans-

verse process and representing the supra-sacral portion of the

ilium in higher types; (3) the nearly complete concealment of

the heart ventrally by the liver-lobes ; (4) the existence of only

one vas efferens for both testes.

* Beddard, P. Z. S. 1907, p. 324, & p. 871.

42 MR. C. L. BOULENGER ON (Jan. 14,

4. On the Hermaphroditism of the Amphipod, Orchestia

deshayesi Audouin. By CHarues L. BoutencEr, B.A.,

¥.Z.8., King’s College, Cambridge.

[ Received January 11, 1908. }

(Text-figure 14.)

Otmar Nebeski (1), in 1880, was the first to call attention to

the interesting fact that in the male Orchestia cavimana Heller

(= 0. gammarellus Boeck), the anterior portion of the testis

constantly gives rise to ova instead of spermatozoa.

Della Valle (2), in his Monograph of the Gammarini, partly

confirms this statement but remarks that, at least in the species

which he examined, O. deshayesit Audouin, this phenomenon

was neither as common as Nebeski stated nor had he ever seen

so large a number of ova as figured by that author. He adds

that he has never observed ova in the testes of a fully adult male,

but only in a few individuals which, although of large size, yet

retained juvenile characteristics in the shape of the posterior

gnathopoda.

Geoftrey Smith (3), struck by the discrepancy of the above

results, examined specimens of Orchestia at Naples during

December to March 1905-6. On dissection he found that more

than 50 per cent. of the males belonging to both species, whether

fully developed or not, exhibited ova in their testes. He there-

fore came to the conclusion that Della Valle could not have

examined QO, deshayesii during the winter months, and that in

the summer, when these animals breed, the fully-developed males

lose these ova. As he remarked :—‘“ The males of these species

when breeding is not going on assume a semi-hermaphrodite

condition of a quite indubitable kind which must evidently bear

some relation to the metabolic conditions in the body.”

The particular metabolic condition which calls forth the pro-

duction of these ova is, he thinks, that condition of “adaptive

anabolism ” which he has studied in the infected males of various

animals subjected to ‘“ parasitic castration” and im the middle

males or males of suppressed sexuality in “high and low

dimorphism.”

During a recent stay at Naples I undertook this investigation,

at the suggestion of Mr. Geoftrey Smith, with the object of ascer-

taining what evidence could be obtained in support of his results.

The two commonest sandhoppers at Naples are Orchestia

deshayesti and Orchestia gammarellus; although a_ certain

number of individuals belonging to the latter species were

examined, most of my observations were made on O. deshayesu

as being the more suitable form for an investigation of this kind.

The chief reasons for this choice were :—

(1) O. deshayesii could be obtained in large numbers close to

1908. | HERMAPHRODITISM IN AN AMPHIPOD. 43

the Zoological Station; I was thus able to procure a constant

supply from one locality at different times of the year.

(2) In this species adult males can without difficulty be

distinguished from the young of the same sex by the characters

of the second pair of gnathopoda. In the adult male the pos-

terior margin of the “hand” is produced to form a large spine-

like process, so that the appendage becomes pseudochelate instead

of subchelate as in the female. The young males up to approxi-

mately 9 mm. in length possess gnathopoda of the female type ;

as they increase in size these appendages undergo various modi-

fications, but do not become pseudochelate until maturity (13-15

mim.) is attained.

Barrois (4) was the first to describe and illustrate these stages

in the development of the gnathopoda; a similar series is figured

by Della Valle in his monograph.

Nebeski bas very carefully described the structure of the sexual

organs of Orchestra gammarellus ; I will therefore only briefly

summarise the more salient features in the testis of O. deshayesii,

which agrees in all but details with that organ in the first-

mentioned species.

The male generative organs lie on either side of the thorax

above the gut and are kept in position by a packing of fatty

tissue. The testis itself is of a fusiform shape and extends

approximately from the posterior part of the second thoracic

segment to the anterior part of the fifth. Anteyiorly it is pro-

duced into a longish filament which seems to serve as a kind of

ligament.

The organ is clothed externally by a layer of epithelial cells

with large nuclei; inside is a cavity on the median wall of

which the germ-cells are situated arranged in several layers.

Posteriorly the testis is directly continuous with the narrow

anterior end of the oval vesicula seminalis; in this respect the

generative organs of this species differ from those of Orchestia

gammarellus where, as described by Nebeski, the posterior narrow

prolongation of the testis opens into one side of the vesicula

seminalis a short distance behind its anterior termination. The

vesicula seminalis communicates with a narrow ejaculatory duct

which leads to a small chitinous penis situated at the base of the

seventh thoracic leg.

Whilst at Naples I dissected a number of adult males of this

species at intervals during the months October to March, and

found the testes to be perfectly normal in 135 out of 137

individuals so examined. The two abnormal specimens possessed

a few small ova at the anterior end of the testis, just below the

origin of the ligament-like filament. Both of these were obtained

during the second week of October together with females carrying

ova; breeding was therefore still in progress.

A smaller number (47) of large male Orchestia gammarellus

were also examined by me, but none were found exhibiting any

A4 MR. C. L. BOULENGER ON [Jan. 14,

traces of hermaphroditism. This phenomenon is therefore of

very rare occurrence in the adult males.

My experience with immature specimens was however very

different, for in these I found that the presence of ova was the rule

rather than the exception.

Text-fig. 14.

AN CSNts

les

Testes of two individuals of Orchestia deshayesii, to show the position of

the ova. X 40.

c., cavity of the testis; es., egg-sac; ov., ovum; sp., developing spermatocytes ;

v.s., vesicula seminalis.

In the smallest males which I was able to dissect (approximately

8-10 mm. in length with scarcely differentiated gnathopoda) the

testes had not yet acquired their characteristic fusiform shape,

these organs appearing as long narrow bands extending along each

side of the thorax. Even at this stage ova had already made their

appearance among the germ-cells, and could be easily distinguished

from the spermatocytes by their greater size, oval shape, and by

1908. ] HERMAPHRODITISM IN AN AMPHIPOD. - 4D

the possession of an enlarged nucleus which hardly stained with

carmine or hematoxylin.

Owing to the difficulties of dissection only a few individuals of

this age were examined, but all (8) possessed ova of various sizes.

In older specimens, varying in size from 10-13 mm., these ova

were found to be of greater size and to occur in greater abundance.

Their exact position in the testis varied slightly, but usually the

anterior third only was ovigerous.

The ova at this stage have evidently grown at the expense of

the surrounding germ-cells, which have disappeared, and come to

lie loosé in a cavity surrounded merely by the external epithelium

of the testis. A kind of egg-sac is thus formed separated by a

constriction from the posterior part of the male gland where

sperm-production is proceeding with great activity.

In a few specimens, in addition to the ova at the anterior end,

others were to be found among the developing spermatocytes in

the posterior part of the testis.

The greatest number of ova observed in any testis was 12; this

number falls considerably short of that recorded by Nebeski; that

author figured as many as 29 in a testis of Orchestia gammarellus.

Altogether 198 males were examined at this stage, and of these

only 19 were devoid of ova.

The fate of these ova I have not been able definitively to ascer-

tain, they have never been seen in the vesicule seminales or in

the vasa deferentia, and from their position it is highly improbable

that they ever reach the exterior. In several testes examined by

me the ova were much shrivelled, and it seemed as if they were

being resorbed by the rest of the testis.

As the result of this investigation, it seems that ova are of

normal occurrence in the generative organs of the young males ;

later, when these animals attain maturity and acquire their adult

characters, these ova in most cases disappear and are probably

absorbed by the rest of the organ before the final moult.

These results are therefore much at variance with those obtained

by Smith, and I am at a loss to explain how he arrived at his con-

clusions; I can only suggest that they may be due to his not

having sufficiently separated the adult and the young males during

the course of his observations.

That ova should be developed chiefly in the testes of immature

males of Orchestia need cause no surprise, for similar phenomena

have been described in numerous animals. This type of “ histolo-

gical” hermaphroditism seems particularly common in Vertebrates,

this being probably due to the fact that this group has been more

exhaustively studied than any other. A number of cases have,

however, been recorded among the Invertebrates :—Ova have been

described by Krohn (5) as normally occurring in the testes of

Phalangiwm ; Garnier (6) has found them in Astacus, Ischikawa (7)

in Gebia, and recently so have Smith (3) and Potts (8) in other

Crustacea (Inachus and Hupagurus) as the result of “ parasitic

castration.”

46 ON HERMAPHRODITISM IN AN AMPHIPOD. [Jan. 14,

Among Vertebrates they occur normally at certain periods in

the testes of Petromyzon, Saimo, and numerous amphibians (apart

from the problematical “ organ of Bidder” in Bufo). It is inter-

esting to find that in most of these cases the production of ova is

limited to the developing or regenerating testes, and for this

reason I will enter on a few more details about such forms.

Thus R. Hertwig (9) has recently given an account of his

observations on the sexual organs of young tadpoles. In this

paper he practically confirms the theoretical conclusion, first

arrived at by Pfliiger, that the surplus of females which con-

stantly occurs in broods of tadpoles is due to a more or less

developed tendency to hermaphroditism in the males at this

stage; some of the so-called females proving to be males at a

later stage in their development.

Lubosch (10) has obtained practically the same results from his

studies of the sexual organs of the larval Petromyzon.

To take yet another example. Felix (11), describing the deve-

lopment of the testis in Salmo, calls attention to the “ auftallende

Thatsache das alle von mir untersuchten jungen Mannchen von

Salmo salar in dem vorderen Abschnitt ihrer Genitaldriise

vollstiindig ausgebildete Hier zeigen.”

Tt seems then that the phenomenon described by mein Orchestia

deshayesii is not an unusual one, and finds a parallel in many

other animals; its significance and its relation to the various

theories formulated to explain such cases of hermaphroditism

T cannot discuss here; I will merely refer the reader to the

excellent paper on this subject by Stephan (12), in which will be

found a lengthy discussion of the current theories bearing upon it.

BIBLIOGRAPHY.

(1) Nesusxt, O.—‘ Beitrége zur Kentniss der Amphipoden der

Adria.” Arb. Zool. Inst. Wien, tom. 11., 1880.

(2) Denia Vatie, AA—‘ Gammarini” in ‘ Fauna u. Flora Golf.

Neapel, Monograph 20, 1893.

(3) Smirn, GrorrrEy.—* Rhizocephala.” Tbid., Mon. 29, 1906.

(4) Barrots, Ta.—‘ Notes sur quelques points de la Morphologie

des Orchesties.” Lille, 1887.

(5) Kroun, A.—‘‘ Zur niiheren Kentniss der minnlichen Zeug-

ungsorgane von Phalangiwm.” Arch. f. Nat. 1866.

(6) Garniur, C.—‘‘ Hermaphroditisme histologique dans le tes-

ticule adulte d’Astacus fluviatilis.” C. R. Soc. de Biol.,

1906.

(7) Iscuikawa, C.—‘‘ On the formation of Eges in the Testis

of Gebia major.” Zool. Anz. xiv., 1891.

(8) Ports, F. A.—“ The Modification of the Sexual Characters

of the Hermit Crab caused by the parasite Peltogaster.”

Q. J. M.S. vol. 50, 1906.

(9) Hurrwie, R.—*‘ Weitere Untersuchungen, &c.” Verh. d.

Deutsch. Zool. Ges., 1906.

1908. ] ' ON MAMMALS FROM THE TSU-SHIMA ISLANDS. AT

(10) Lusoscn, W.—‘‘ Ueber die Geschlectsdifferenzirung bei

Ammocetes.” Verh. Anat. Ges. xvil., 1903.

(11) Fetix, W.—‘ Handbuch der Vergleichenden Entwickelungs-

lehre der Wirbelthiere, edited by O. Hertwig, Bd. in,

1906.

(12) SrepHan, Prerre.— De Vhermaphroditisme chez les Ver-

tébrés.” Annales de la Faculté des Sciences de Marseille,

inocu AION Le

P.S.—Since the reading of this paper Dr. Calman has kindly

drawn my attention to a paper by Zograf (Zool. Anz, xxx., 1907),

in which the author records the occurrence of ova in the testes

of Apus.

February 4, 1908.

H.G. THe Dvuxe or Beprorp, K.G., President, in the Chair.

Mr. F. Martin Duncan gave a lantern exhibition of the results

he had obtained with the Autochrome Natural Colour Process of

the brothers Lumiere as applied to zoological subjects.

The following papers were read :—

1. The Duke of Bedford’s Zoological Exploration in Hastern

Asia.—VIT. List of Mammals from the ‘Tsu-shima

Islands. By OLpFieLp THomas, F.R.S., F.Z.S.

[Received December 31, 1907. |

In continuation of his collecting-work in Eastern Asia, Mr.

Malcolm Anderson visited the Tsu-shima Islands, between Japan

and Korea, at the beginning of this year, and made the collection

enumerated below.

The fauna would seem to be comparatively poor, so far as the

number of species 1s concerned, but it is of much interest, owing

to the curious mixture of forms which it contains, some of its

members being absolutely Japanese in relationship, and others

Korean—a mixture the cause of which it is at first sight difficult

to guess.

Putting aside the species which, for one reason or another, are

48 MR. OLDFIELD THOMAS ON MAMMALS | Feb. 4,

indefinite in relationship, we have the following characteristically

Japanese forms :—

Urotrichus talpoides adversus. No Urotrichusrecorded from Korea.

Martes melampus tsuensis. No Marten of this type known

from Korea.

Apodemus speciosus speciosus. Replaced in Korea by the very

different A. s. peninsula.

is geisha sagan. A, geisha common in Japan, absent

from Korea.

(No A. agrarvus. Dominant in Korea, absent from

Japan.)

On the other hand, the Korean forms are :—

Crocidura corec. : Replaced in Japan by C.dsi-nezwmi.

Felis macrotis. No Cat of any sort known in Japan.

Lutreola sibirica. Replaced in Japan by ZL. ttatsi.

The balance even in mere numbers is thus on the side of the

Japanese relationship of Tsu-shima, But further it is to be noted

that three out of the four animals with Japanese affinities belong

to special local subspecies, indicating that they are certainly in-

digenous, having been in the islands long enough to get more or

less modified. On the other hand, all the Korean forms are

identical with their mainland relatives and have perhaps been

accidentally introduced through human agency.

Mr. Anderson’s notes on the physical features of the islands

are as follows :—

“ Notes on T'su-shima.

‘“‘Tsu-shima (literally Opposite Island) consists in reality of two

main islands, the southern being called Shimono, the northern

Kamino-shima. The two are separated by a narrow channel only,

which, I am told, is sometimes dry at low tide. Both islands

are hilly throughout, the highest point being 2100 ft. The hills

are steep and their soil very thin, so, as the valleys are exceedingly

narrow, arable land is scarce indeed, and the population almost

entirely restricted to the fishing-villages on the coasts. The hills

are wooded, especially in Kamino-shima, where pines, oaks,

Cryptomeria, Chamecyparis, and Camelia are the principal trees.

Precipitation is abundant, but, owing to the shortness of the

streams and the stony nature of the soil, which is composed of

fragments of shale, the brooks are usually dry, especially at

their lower ends; it is only during, or just after, a heavy rainfall

that their water reaches the sea.

‘‘¢Tsu-shima is surrounded by a few islets of no importance.

The nearest land is Iki, 32 miles to the south-east, but the

Korean mainland is not more than four or five miles farther,

while some considerable islands of the Korean Archipelago are

less than forty miles from Kamino. Between Shimono-shima and

1908. | FROM THE TSU-SHIMA ISLANDS. 49

Iki the sea reaches no greater depth than 65 fathoms (Japanese

Admiralty chart), while between Kamino-shima and Korea it

falls, in one restricted area near the coast of Kamino, to a depth

of 105 to 118 fathoms.

“I collected in Tsu-shima from Jan. Ist to Feb. 3rd, 1907.

‘No Hares or Squirrels are known to the people of Tsu-shima,

and I could find no traces of Hvotomys or Microtus, or indeed of

any other wild species.” —J/. P. A.

Tn addition to the specimens collected by Mr. Anderson I have

inserted references to a few mammals obtained by Mr. P. A. Holst

in 1891 when travelling on behalf of the late Mr. Henry Seebohm,

by whom they were presented to the British Museum. Among

these are the original examples of the Tsu-shima Marten, described

by me in 1897. Two further new subspecies are now described.

After Tsu-shima Mr. Anderson visited the Iki and Goto

Islands, the resulting collections proving that those groups are

absolutely similar to S.W. Japan in their mammal fauna.

Mr. Anderson’s series amounts to 151 skins, with their skulls,

and is presented as before to the National Museum by His Grace

the Duke of Bedford, K.G.

1. RarNoLopHus corNnutus Temm.

(92.3.20.1-2. P. A. Holst. 1891.)

This is the only Bat as yet recorded from Tsu-shima.

2. MoGERA WOGURA KANAI Thos.

(92.9.2.2. P. A. Holst. 1891.)

Unfortunately Mr. Anderson was not able to obtain further

specimens of the Tsu-shima Mole, which I have provisionally

assigned*, on the single example above recorded, to the smaller

south-western form J/. wogura.

“Not secured or even seen, but natives tell of a black Mole,

and also of a much larger buff-coloured one, which they say they

see in spring and summer.”—J/. P. A,

3. UROTRICHUS TALPOIDES ADVERSUS, subsp. n.

6. 1243, 1244, 1272, 1274, 1275. ©. 1263, 1273, 1279.

Sasuna, N. Island. 100’.

3. 1199, 1204, 1213, 1215, 1221, 1234. ©. 1214, Izuhara,

S. Island. 300’.

General colour very brown, furthest from the slaty black of

the Hondo subspecies. Size less than in the Kiushiu form, the

head and body measurement rarely attamimg 90 mm. Tail com-

paratively long, averaging nearly 34 mm., its hairs also particularly

long. Fur thick and soft—hairs of back about 6:5 mm. in length,

* Supra, p. 463. + Tsu-shima=Opposite Island.

Proc. Zoou. Soc.—1908, No. IV. 4

50 MR. OLDFIELD THOMAS ON MAMMALS [ Feb. 4,

and therefore nearly a millimetre longer than those of the more

northern U. t. hondonis, measured equally on winter specimens.

Dimensions of the type :—

Head and body 89 mm.; tail 34; hind foot 15.

Skull—greatest length 26°5 mm,; basal length 22°3; greatest

breadth 13.

Hab. Tsu-shima Islands—type from Sasuna, N. Island.

Type. Adult male. B.M. No, 8,2.26.1. Original number 1243.

Collected 16th January, 1907.

“ Fairly common in damp wooded ravines in Tsu-shima, where

it was often trapped beneath or beside large stones. Insects are

often among the stomach contents, and here no vegetable matter *

was recognised in their stomachs, but one example was caught in

a trap baited with barley.”—/. P. A.

A renewed examination of the fine series of Urotrichus obtained

by Mr. Anderson in the Japanese Archipelago shows that each

of the three main islands has a form which may be subspecifically

separated from the others, and that Tsu-shima has a fourth.

Their characters are shown in the following synopsis :—

A. Tail comparatively long, averaging about 33 mm.

a. Size larger, head and body 90-101 mm. Colour

Game Joona, — IRGHESIMTUL 53 500 cos oncoo ase sbo spe eGo GEODeS U. t. talpoides Temm.

b. Size smaller, head and body 84-91 mm. Colour

pales brown. alsu-shimlarerseersessesesseeeeeeseeee:

B. Tail comparatively short.

ec. Tail averaging just over 30 mm. (range 28-34).

Colour brown. Shi-koku ........................ U.t. centralis (infra).

d. Tail averaging 27 mm. (range 23-80). Colour

slatyerreya lon donner eceeeeenee erent

U. ¢. adversus (supra).

U. t. hondonis (infra).

The series on which these averages are based are from 15 to

22 in number, and are therefore large enough to eliminate any

material error due to individual variation.

Details of the Shi-koku and Hondo subspecies :—

UROTRICHUS TALPOIDES CENTRALIS Y, subsp. n.

Size averaging rather less than in true ¢alpoides, larger than in

adversus (head and body 90-97 mm., the majority of specimens

92-95). ‘Tail decidedly shorter than in either talpoides or adversus

(average of 22 specimens 30°75 mm., extremes 28-34), Fur

about 6 mm. in length on the back (winter). General colour

deep brown, darker than adversus, lighter than talpoides.

Dimensions of the type :—

Head and body 94 mm.; tail 32; hind foot 16; upper tooth-

row 11:2.

Hab. Island of Shi-koku. Type from Jinrio, Tokushima Ken.

Alt. 500’.

Type. Adult female. B.M. No. 6.1.4.68. Original number

289. Collected 14th February, 1905.

* Of. P.Z.S. 1908, 1. p. 341. eS

+ Central as compared to other forms, both in size, colour, length of tail, and

locality.

1908. } FROM THE TSU-SHIMA ISLANDS, 51

UROTRICHUS TALPOIDES HONDONIS, subsp. n.

Urotrichus talpoides pilirostris Thos. P. Z. 8. 1905, ii. p. 342

(1906), nec Dymecodon pilirostris True, P. U.S. Nat. Mus. 1886,

Tas OE

Size medium, head and body length generally about 90 mm.

Tail shortest in the genus, the average of 17 specimens being

2/ mm., with a range of from 23 to 30. Fur about 5:5 to 6 mm.

in length on the back. General colour “ slate-black ” (grey No. 2),

with a slight tinge of “ mouse-grey,” the brown tone found in

the more western forms absent.

Dimensions of the type :—

Head and body 91 mm.; tail 26; hind foot 14; upper tooth-

row 10:1.

Hab. Hondo. Type from Nakaomi, near Ohito, Izu.

Type. Male. B.M. No. 6.1.4.97. Original number 144. Col-

lected 13th December, 1904.

This form needs a name different from that I applied to it in

1905, for the reasons explained in the footnote*.

4, CRocIDURA CORE Thos.

©, 1242-1247. Sasuna, N. Island.

3g. 1203. Izuhara, S. Island.

This is one of the three T'su-shima species of Korean rather

than Japanese affinity, but must not be considered as of great

importance from a geographical standpoint, as Shrews of this

genus are undoubtedly often carried about on shipboard from

place to place, so that this Shrew may have been accidentally

introduced from Korea.

“Rare. None seen but the three preserved.”—V/, P. A.

* Since I wrote the paper on Japanese Mammals above referred to, the Museum

has received from Mr. K. Kanai, a Japanese who had been taught collecting by

Mr. Anderson, a small series of specimens from Central Southern Hondo. Among

these there are four examples of an Insectivore entirely new to us, allied to Uro-

trichus, but smaller and differmg in various details, and it seemed probable,

therefore, that these represented Mr. True’s Dymecodon pilirostris, which had been

supposed to have been based on a young Uvotrichus.

By Mr. True’s kindness and the great courtesy of the authorities of the United

States National Museum I have been permitted the loan of the type specimen of

D. pilirostris, and a comparison shows at once that the above suggestion is correct,

and that Mr. Kanai’s specimens represent the adult and Mr. True’s type the young

of a species generically quite distinct from Urotrichus. I am therefore now able to

confirm the absolute correctness of Mr. True’s distinction of Dymecodon, on which

I had previously thrown doubt.

The adult dentition of Dymecodon does not differ in number from that of the

young described by Mr. True, and appears to be, so far as I am able to understand

the homologies of the teeth,

1s OR

POS

the permanent p! being present with the milk-teeth, and being included in the

number recorded by Mr. True. The formula of Urotrichus is the same, except that

the lower canine is absent, the statement made by some authors that there are four

52 MR. OLDFIELD THOMAS ON MAMMALS [ Feb. 4,

5. Fexis microris M.-Hdw.

@. 1198, 1278. Sasuna, N. Island. 300’.

(91.10.14.1, 92.1.11.3, and 96.2.28.1. P. A. Holst. 1891.)

This is an essentially Korean member of the Tsu-shima fauna,

no member of the genus Yelis being found in Japan.

“The Wild Cat is common in Tsu-shima, living near the sea-

shore, where it is said to go nightly at low tide to fish. In the

stomach of 1278 I found fish-bones and some feathers of the

pheasant. The natives consider the flesh of the Cat especially

delicate food. Native name: ‘ Yama-niku’ = Mountain-Cat.”—

6. MARTES MELAMPUS TSUENSIS Thos.

3. 1261-1277. Sasuna, Tsu-shima.

(91.10.14.2-3, 92.1.11.1-2. P. A. Holst. 1891.)

When describing this form in 1897* I had not any Japanese

Martens of the bedfordi type to compare it with, that being clearly

the animal to which it is most allied, and not the typical melampus.

Now, however, a comparison of Mr. Anderson’s skins of the two

shows that, in winter pelage, the Tsu-shima Marten is distin-

guishable from that of Southern Hondo by its whitish crown

and the absence of the yellowish tuft at the end of the tail.

“¢ Common in Northern Tsu-shima, and occurring in the South

Tsland. Like the Cat it is most often found near the sea. The

stomachs of the two caught were empty, but the animals were

excessively fat. They are considered a fine food by the peasantry

of the islands, who call them ‘ Wata-boshi’ = Cotton-cap.’—

Wh IP Ale

premolars and only ? incisors being evidently wrong, as a comparison of the milk

and permanent teeth both of Dymecodon and Urotrichus shows that the ante-

penultimate premolar does not change, and is therefore p!, while distinct traces of

the premaxillo-maxillary suture can be seen just behind the third tooth in the upper

jaw. Of the incisors it is probably i, that is missing in the lower jaw.

In the form and relative proportions of the permanent teeth Dymecodon resembles

Urotrichus very closely, the curious alternation in size of the lower teeth, on which

Mr. True based the name of the genus, not existing in the later dentition, and it is

in fact only due to the milk p. being, not unnaturally, smaller than the permanent

P; which coexists with it.

But in the milk stages, which have not previously been compared, the differences

are far greater. For while in Dymecodon the two anterior incisors are subequal,

similar in form, scarcely overtop the teeth posterior to them, and in Mr. True’s

words “resemble the teeth of Phocena,’ those of Urotrichus have already pro-

gressed a long way towards the specialisation found in the adults of both genera,

mi! considerably surpassing mi, and this again being much longer than mi?; and

below again the most anterior tooth (mis, as I suppose) is decidedly longer than any

of the next three teeth.

From this it would appear that Dymecodon is in a more primitive stage of

evolution than Urotrichus.

Mr. Kanai’s specimens of Dymecodon were captured at Shibu, on Mt. Yatsugatake,

Nagano Ken, N.W. of Yokohama, at an altitude ot 5700’.

Mr. Kanai also obtained at Kamisuwa-machi, in the same Ken, an example of

Vespertilio murinus superans Thos., a Bat new to the fauna of Japan.

* Ann. Mag. N. H. (6) xix. p. 161 (1897).

1908. | FROM THE TSU-SHIMA ISLANDS. 53

7. Lurreoua siprrica Pall.

3. 1262, 1269, 1276. Sasuna, N. Island.

These specimens agree absolutely in size, colour, and length of

tail with Vladivostok and other mainland exampies of the group,

and equally differ from the Japanese Putorius itatst.

On the other hand, the Quelpart Minks, which I had supposed

to be summer representatives of sibiricus, prove on a closer ex-

amination to be more allied to 7zéatsi, though separable from it*.

In both islands, therefore, the members of the present genus

exactly contradict the conclusions as to the faunistic relations of

Quelpart and Tsu-shima with Korea and Japan, indicated by

Urotrichus and the Mice.

“The most common of the Carnivora of the Islands; not living

near houses, but in the wooded hills. Sometimes eaten by the

fishermen. Called ‘ Yoto-shi’= Night-thief.”—/. P. A.

8. Mus norvecicus Hrxl.

¢. 1267. 9. 1268. Sasuna, N. Island.

“*Common on the sea-shore, where I found it living among the

rocks. Fishermen told me that it swims in the sea.”—J/. P. 4.

9. APODEMUS SPECIOSUS Temm.

Cre 2528 2 OO SOs 2 close GAs Aaley LA80:

Sasuna, N. Island. 100’.

6+, 1202; 1207, 1210, 1227, 1236, 124). 2. 1201, 1208, 1220

1239. Izuhara, 8. Island. 300’.

These specimens are absolutely speciosus and not peninsule,

thus agreeing with the other Rodents in their Japanese rather

than Korean relationship.

* LUTREOLA QUELPARTIS, sp. 0.

Size as in P. itatsi, the Japanese Mink, markedly smaller than in the Korean

P. sibiricus. General colour, in summer pelage, dark tawny or russet-brown, quite

as in P. itatsi. Face dark “seal-brown ”; under surface, tail, and sides of neck

russet; lips and chin white; hands and feet russet, passing terminally into drab.

Skull with a very high and vaulted brain-case, much more so than that of either

P. sibiricus or itatsi. Upper molar with its inner lobe narrow, the antero-posterior

diameter of this lobe little or not more than that of the outer lobe, and therefore

much reduced as compared with either of the allied forms, in which there is a

broadly expanded inner lobe.

Dimensions of a pair, measured in flesh :—

S$ (Type). Head and body 309 mm.; tail 153; hind foot 55; ear 25.

39 265 be) 3 2 130 ; 3) 46 > 3 22.

& (type). 2.

Skull—condylo-basal length ..................c0.0000--2-.. 5975 50°5

basal lenethy | 23.0: o5.vsseensueuecaeateeene ne cea On 47°3

TTAONG! LTHEEEUHN jocccaccseceososspacesceaensgaccosszccy ADU 22°5

combined height of brain-case and bulle ...... 215 17°8

inner lobe of m!, antero-posterior diameter]... 2°2 2

Hab. Island of Quelpart, S. of Korea.

Type. Adult male. B.M. No. 6.12.6.1. Original number 627. Collected 7th

September, 1905, by M. P. Anderson; presented by the Duke of Bedford, K.G.

(Four specimens examined.)

D4 ON MAMMALS FROM THE TSU-SHIMA ISLANDS. _ [Feb. 4,

Four fairly large specimens measure :—

3 - Head and body 107 mm.; tail 100; hind foot 25-5; ear 16.

3. ”? 104. ” 9 OD 98 ; 25 3 99 15:5.

2 : 29 97 ay) 96 ; ” 25°5 > 99 16.

2 . 9 96 9” 908 93 > or) 24:5 5. OD) 6°5

10. APODEMUS GEISHA SAGAX, subsp. n.

6. 1245, 1246, 1249, 1254, 1255, 1256, 1257, 1258. 2. 1248,

1259, 1260, 1265, 1266. Sasuna, N. Island. 100’.

6. 1206, 1211, 1212, 1217, 1222, 1224, 1225, 1228, 1230,

1231, 1232 .el 23 (4 240n O00, 1205, 12097 SG ae.

1219, 1223, 1226, 1229, 1233, 1238, 1239, Izuhara, S-. Usland:

200'—300’.

General characters as in true geisha, but just as the ears average

shorter in specimens from Hokkaido*, so here they are uniformly

rather longer than in Hondo examples. Of the 25 specimens

from Izuhara, in the 8S. Island of Tsu-shima, no less than 16 are

labelled as having ears 15 mm. in length, three have them 14,

two 14:5, two 15°5, and two 16. In true geisha the ears are

about 14 mm., and in nokkaidi 13 mm.

The Sasuna specimens are not quite so uniform as those from

Izuhara.

Dimensions of four specimens from Izuhara :—

6. Head and body 81 mm.; tail 91; hind foot 19°5; ear 15.

3 (Type) ” 79 3 3 39 100 ; ” 19°53; 2 15.

Q “s Saab et he ete OO a LOR eee Ge

2 COPS ENTS Six y 19°53; ,, 15d.

Skull of type—greatest length 24 mm.; basilar length 18;

length of upper tooth-series 3:5.

Hab. Tsu-shima—type from Izuhara, South Island.

Type. Adult male. B.M. No. 8.2.26.50. Original number

1228. Collected 10th January, 1907.

“The most common of the Tsu-shima mammals. Found in

the forested hills and canyons.”—J/. P. A.

Purely a Japanese type, no Mouse at all allied occurring in

Korea.

11. APoDEMUS MINUTUS Pall.

6. 1235. Izuhara, 8. Island.

Head and body 54mm.; tail 53; hind foot 145; ear 9.

The single specimen appears to be more or less intermediate

between the continental subspecies wsswricus and the Japanese

japonicus, but no very definite opinion can be expressed about it

without further material.

[12. Cervus sixa Temm.

“Said to have been very abundant before the introduction of

good firearms; now rare and not seen by me or my assistant.” —

M.P.A.|

* M. g. hokkaidi, P. Z. 8. 1905, ii. p. 350.

ZS), Ue Alea.

E. Wilson, Cambridge.

GONADIAL GROOVES IN AURELIA AURITA.

ne Geneve as

1908. ] ON GONADIAL GROOVES IN A MEDUSA. ay)

2. On the Presence of Gonadial Grooves ina Medusa, Aurelia

aurita*. By T. Goopxy, ;Student Demonstrator in

Zoology, University of Birmingham.

{Received December 23, 1907. ]

(Plate I.t)

While recently examining specimens of the Scyphozoon

Aurelia aurita in the ordinary course of practical work in the

Zoological Laboratory, the structures mentioned in the title of

this paper attracted my attention. It was not clear that they

formed any part of the ordinary radial canal-system, and, so far, I

have been unable to find any account of them in the different

text-books and journals which I have consulted in the expectation

of obtaining information as to their nature. This being the case,

it seemed desirable to investigate the nature and relations of the

structures in question and to give a brief account of the results.

The gonadial grooves were found in all the preserved specimens

examined in the laboratory, both male and female, and they were

also found in living examples which came under my notice during

the recent summer vacation.

In a view of the sub-umbrella surface (Pl. I. fig. 1) the four

grooves were easily visible to the naked eye, and presented the

appearance of four radially arranged canals. Each groove (g.g.)

lies in an inter-radial axis, and is better seen when the prolonged

mouth-angles have been removed. It has about the same dia-

meter as the origin of an ordinary per-radial or ad-radial canal, and

extends from the central gastric cavity into a gastric pouch (q.p.),

where it terminates in a somewhat funnel-like expansion at about

the centre of the pouch and directly dorsad to the external opening

of the corresponding sub-genital pit (s.p.). In its course the

groove extends along the ventral wall or floor of the passage from

the gastric cavity to the pouch and along the floor of the pouch

itself.

Bristles could readily be passed along the groove, and by means

of a fine pipette a coloured liquid was injected through the

groove into the gastric pouch. When the latter became filled

and more of the liquid was injected, the excess began to flow back

into the gastric cavity through the main opening into the pouch

(PI. I. fig. 2, d.e.p.) which is dorsad to, and considerably wider than,

the groove in question.

At first sight the grooves appeared to be the beginnings of

inter-radial canals ; but the fact that each groove does not extend

beyond the centre of each gastric pouch, and also that at this

point it widens out and abruptly terminates, negatives this idea.

In order to determine the true nature of these structures,

portions of the Medusa, including complete gastric pouches and

* Communicated by Professor Bripex, F.R.S., F.Z.S.

iY For explanation of the Plate, see p. 58.

56 MR. T. GOODEY ON GONADIAL [ Feb. 4,

their connections with the central gastric cavity, were removed.

These were then stained with borax carmine for forty-eight

hours, dehydrated, and embedded in paraftin-wax. Sections were

cut at right angles to the inter-radii, passing completely through

the pouches from the dorsal to the ventral surfaces. In the above

preparation there was a considerable amount of contraction due

to the jelly-like consistency of the material, and, for this reason,

many of the parts in several of the sections were displaced from

their natural relations.

From an examination of the most satisfactory complete sections,

however, the nature and relations of the grooves could be easily

made out. It was at once evident that they were not closed

canals similar to the normal radial canals, and that their resem-

blance to canals in a superficial or surface view was misleading.

Each groove is, in fact, formed by a folding of the endodermal

epithelium lining the floor of the pouch and of its passage of

communication with the gastric cavity. This folding is shown in

Pl. I. fig. 3, where the epithelium is seen to be raised into two

parallel ridges (e), one on each side of a median vertical line,

which form the somewhat folded lateral walls of the groove

g-9-):

In the floor of each pouch, almost completely encircling its

outer, lateral, and inner walls, is situated the gonad (figs. 1, 2, &

4, g.), in the form of a characteristic incomplete ring of sex-

cells, the discontinuity occurring at the point where the groove

enters the pouch.

From Pl. I. fig. 4 it will be seen that the endodermal epithelium

(e, e', e’, e’) not only lines the inner surface of the gastric pouch

and invests the gonad above, but also extends downwards in the

median line so as to form the boundaries of the gonadial groove

(g-g.). An epithelial stratum (e*, e’) also invests the ventral or

oral surface of the gonad, and is continuous with that lining the

floor of the gastric pouch. Thus the gonad is completely

ensheathed by the endoderm from which the sex-cells are derived.

As this section passes through the point of discontinuity of the

gonad-ring, it will be noticed that the two halves of the ring lie

one on either side of a median line along which the gonadial

groove passes into the gastric pouch.

The function of these problematic grooves is by no means so

easy to determine as their structure and relations. That they

have anything to do with the conveyance of food-material. from

the gastric cavity to the gastric pouches is highly improbable,

inasmuch as these cavities are already in free and open communi-

cation with one another. A more feasible suggestion is that they

function as channels for the outward conveyance of the ripe

sex-cells when liberated from the gonads. Their position

in the gastric pouches and their somewhat expanded origins in

close relations with the encircling gonads, seem to point to this

conclusion. The liberated sex-cells would fall on to the floor of a

gastric pouch, and the gonadial groove would seem to constitute

1908.) GROOVES IN A MEDUSA. 57

an obvious and natural channel for their transit to the exterior.

The groove is not closed dorsally, it is true, but the epithelial

ridges bounding the groove seem capable of approximating

sufficiently closely that it is by no means difficult to imagine the

groove as practically a closed canal—a functional gonoduct, in

short.

It must be admitted that I have not been able to detect the

presence of sex-cells in the gonadial grooves, but this may be due to

the fact that the gonads were not mature, or the sex-cells ripe for

extrusion. It remains for further investigation to ascertain on

living specimens under suitable conditions whether these views as

to the nature and function of the gonadial grooves are correct.

A careful examination of Pelagia noctiluca and Chrysaora

isosceles, both of which belong to the Pelagide, failed to reveal the

existence in either species of any structures comparable to the

gonadial grooves of Aurelia aurita.

In conclusion, it may be suggested that the observations

recorded above possess certain features of more general interest.

In discussing the evolution of the celom Sir Ray Lankester *

remarks :—“‘ We may suppose the first celom to have originated

by the closing or shutting off of that portion of the general

archenteron of Enterocela in which the gonads develop, as in

Aurelia or as in Ctenophora” ; and, further, “the most important

developments of the celom are in connection with the establish-

ment of an exit for the generative products through the body-

wall to the outer world.” If, therefore, my observations and

inferences as to the nature and function of the gonadial grooves

be correct, it is obvious that in this organism we have an

extremely interesting and primitive condition.

The gastric pouches are special portions of the archenteron,

from the walls of which the gonads have their origin ; they are,

in fact, primitive gonoceles, although not yet completely shut off

from the general archenteric cavity as is the case in so many

Celomata. On the other hand, the gonadial grooves may be

regarded in the light of incipient c¢elomoducts or gonoducts

which in like manner are still but imperfectly constricted off from

the archenteron. Consequently the gonadial grooves and gastric

pouches of Aurelia seem to represent a very primitive stage in

the evolution of both gonocceles and gonoducts.

It has generally been held that, with one exception, the

Coelenterata have no specially differentiated genital ducts, the

sex-cells finding their way to the exterior either directly by the

external dehiscence of the gonads, or indirectly by internal

dehiscence inte archenteric canals or spaces and thence outwards

through the mouth. The single exception referred to is in the

case of Ctenoplana korotneffii, the aberrant Ctenophor discovered

by Willey tin 1896 in the Eastern Archipelago of British New

Guinea, in which, however, only male gonads were found.

* A Treatise on Zoology, part ii. 1900, p. 9.

+ Quart. Journ. Micros. Science, vol. 39, n. s, 1896-97.

58 ON GONADIAL GROOVES IN A MEDUSA. [ Feb. 4,

According to the discoverer, the two pairs of gonads apparently

had their origin from endodermal cell proliferations of the walls

of genital diverticula of the general archenteric canal-system, as in

so many other Ceelenterata. Special genital ducts are present, but

variable in number, and they are described as being continuous

with the tunica propria investing the gonads. The external

apertures of the ducts are situated on the dorso-lateral surfaces

of the body, below several of the ctenophoral bands.

In the present state of our knowledge of the genital ducts of

Ctenoplana korotnefii, and especially in the absence of any

information as to their mode of origin, any attempt to discuss the

question of their morphological character would be of little value.

It may therefore be concluded, at any rate provisionally, that,

while both Awrelia and Otenoplana stand alone among living

Ccelenterata in possessing genital ducts, it is nevertheless only in

the Scyphozoon that these structures can at present be regarded

as coelomic or archenteric derivations, and, in fact, are a very

primitive form of ccelomoduct or gonoduct.

IT am indebted to Professor Bridge for the kindness and

consideration which he has shown to me in these my first attempts

at independent investigation, and also to Mr. F. W. Crispe for the

material help given in the preparation of the sections.

EXPLANATION OF PLATE I.

Gonadial grooves in Aurelia aurita.

1. Surface view of sub-umbrella aspect of Aurelia aurita, the prolonged mouth-.

angles having been removed. The gonadial grooves are shown, and also

their relations to the. surrounding structures.

Fig. 2. Semi-diagrammatic, enlarged three times. The gastric pouch has been cut

through along the inter-radius, and the drawing represents a view taken

in the vertical plane. The space between the dotted line and the body-

wall represents the course of a gonadial groove from the gastric cavty to

its opening into the gastric pouch.

Fig. 3. (X 20) Section passing through the main passage to a gastric pouch,

showing the gonadial groove lying in a median position in the ventral wall,

bounded on either side by an endodermal epithelial fold.

Fig. 4. (X 20) Section passing through a gastric pouch, showing a gonadial groove

in the floor, and the loops of the gonad ring bounding it on every side.

REFERENCE LETTERS.

b.w., body-wall; d.e.p., dorsal entrance to a gastric pouch ; e. (fig. 3), epithelial

folds; e, el, e2, e, epithelium lining inside of pouch and covering gonad above; e*,

e5, epithelium of ventral surface of the gonad and of the floor of a gastric pouch ;

g. gonad ; g.g., gonadial groove ; g.p., gastric pouch ; m.p., main passage from the

gastric cavity to a gastric pouch ; s.p., sub-genital pit.

[March 9th, 1908.—Since writing this paper my attention has

been directed by Mr. E. T. Browne of University College, London,

to a paper published by L. Agassiz in ‘Contributions to the

Natural History of the United States,’ vols. in. & iv. 1860-62,

dealing with Aurelia flavidula, which is now considered to be the

same as Aurelia awrita.

In his account L. Agassiz explains that the gastric pouches

WGEtOevanehar (S/ORSVGLIUES) tS

‘SNLVOIGLS OLdODO0NHT “2 ‘STTIOVGD SNHGOHHOHINVOV T

‘dwt weurmeyy 4seyy. TUL NE TIE REE YNL 7D)

IU lal SOG! S 4 c_

I Aas. tS) OS, Pi, Mt,

G.M.Woodward del.et lith.

West, Newman imp.

1. CTENOGOBIUS BEDFORDI. 2. TRIDENTIGER COREANUS.

3. ACANTHOGOBIO LONGIROSTRIS. 4. LIOBAGRUS ANDERSONI

1908. | ON FRESH-WATER FISHES FROM COREA. 59

retain a connection with the central gastric cavity by means of

channels along which the sex-cells pass on their way to the

exterior. These channels, I take it, are the main passages of

connection between the gastric pouches and the central gastric

cavity referred to in the above paper.

L. Agassiz makes no reference, either written or by figure, to

the ridges of epithelium which form the boundaries of the

“ gonadial grooves” in the above account.

It should be noted that it is within the limits of these grooves

that it is suggested the sex-cells pass on their way to the exterior

at the period of extrusion, and that it is only these parts which

form the primitive gonoduct by a pinching-off from the arch-

enteric cavity. |

3. The Duke of Bedford’s Zoological Exploration in Eastern

Asia.—VIII. A Collection of Fresh-water Fishes from

Corea. By C. Tarr Ruaan, M.A., F.Z.S.

[Received January 14, 1908. |

(Plates II. & III. *)

A small series of fresh-water fishes collected in Corea by Mr. M.

P. Anderson and presented to the British Museum by His Grace

the Duke of Bedford, K.G., is of considerable interest, as of the

eleven species represented, no less than seven are new to science,

some of these extending the range of genera previously known

from China or Japan. The specimens are from two localities,

viz., Chong-ju, Chung-Chong province, and Kimhoa, 65 miles

north-east of Seoul ; from the latter examples of only two species

(Silurus bedfordi and Liobagrus andersoni) were obtained.

CYPRINID&.

1. CaRasstus AURATUS L.

2. PSEUDORASBORA PARVA Schleg.

3. BaRriLius PLATyPUuS Schleg.

4, LEUCOGOBIO STRIGATUS, Sp. n. (Plate 1I. fig. 2.)

Depth of body equal to or a little more than the length of

head, which is 4 in the length of the fish. Snout a little longer

than eye, the diameter of which is 4 in the length of head;

interorbital width 22 to 3 in the length of head. Mouth oblique;

barbels small. Dorsal 10, with 7 branched rays; origin equi-

distant from tip of snout and base of caudal and above that of the

ventrals. Anal 9, with 6 branched rays. Pectoral not reaching

the ventrals. Caudal forked. 36 to 38 scales in a longitudinal

* For explanation of the Plates, see p. 63.

60 MR. C. TATE REGAN ON | Feb. 4,

series, 4 or 5 in a transverse series from origin of dorsal to lateral

line, 3 or 34 between lateral line and base of ventral fin. Dark

longitudinal stripes along the series of scales, broader and stronger

above the lateral line, fading out below; fins pale.

Three specimens from Chong-ju, the largest 75 mm. in total

length.

The genus Leucogobio has hitherto included one species from

Corea, two from Western China and four from Japan. Of these

the Chinese Z. teniatus Gthr., 1896, and the Japanese L. guenthert

Ishikawa, 1901, are very similar to Z. strigatus, but differ from

it in coloration and in having more scales in a transverse series.

5, ACANTHOGOBIO LONGIROSTRIS, sp. n. (Plate III. fig. 3.)

Depth of body 42 in the length, length of head 32. Snout

long, decurved, 13 as long as eye, the diameter of which is 3$ in

the length of head and equal to the width of the flat interorbital

area. Mouth small, inferior, horseshoe-shaped ; barbel shorter

than the eye. Dorsal II 7; origin equidistant from tip of snout

and base of caudal. Anal 8, with 6 branched rays. Pectoral

nearly reaching the ventrals, which are inserted below the

anterior rays of the dorsal. 43 scales in a longitudinal series,

52 ina transverse series from origin of dorsal to lateral line,

34 between lateral line and base of ventral. Silvery, back darker ;

blackish spots on the back and on the sides above the lateral line ;

small dark spots on dorsal and caudal.

A single specimen, 95 mm. in total length, from Chong-ju.

The pharyngeal teeth appear to be in two series (5.2—2.5);

but this species is not very remote from Hemuibarbus barbus

Schleg., and it seems to me that the genus Acanthogobio Herz.,

1892, is scarcely worth recognition, as it differs from Hemibarbus

only in having the pharyngeal teeth in 2 series instead of 3.

6. ACANTHORHODEUS GRACILIS, sp. n. (Plate II. fig. 1.)

Depth of body 22 to 22 in the length, length of head 43.

Snout a little shorter than eye, the diameter of which is 3 to 33

in the length of head; interorbital width 24 in the length of

head. Mouth small; no barbels. Dorsal I1 13; second spine

nearly 2 the length of head, considerably shorter than the longest

soft rays, which are nearly as long as the head; fin highest

anteriorly ; free edge slightly convex. Anal IT 10; second spine

4 or a little more than 4 the length of head; free edge slightly

concave. Pectoral nearly reaching the ventrals, which extend

nearly to the anal. 34 scales in a longitudinal series, 6 or 7 in a

transverse series from origin of dorsal to lateral line, 5 or 6

between lateral line and base of ventral. Silvery, back olivaceous ;

a bluish lateral stripe on the posterior part of the body; dorsal

with alternating series of pale and dark spots on the rays; anal

with a narrow blackish edge.

Two specimens, 65 and 70 mm. in total length, from Chong-ju.

1908. | FRESH-WATER FISHES FROM COREA, 61

SILURIDA.

7. SILURUS BEDFORDI, sp.n. (Plate II, fig. 3.)

Depth of body 6 in the length, length of head 43 to 5i.

Breadth of head about 13 in its length, diameter of eye 74 to 9,

interocular width 14 to 2, length of snout 34 to 32. Lower jaw

projecting ; vomerine patches of teeth either narrowly separated

or united anteriorly. Four barbels, the mandibulary ones less

than 2 as long as the maxillary ones, which extend nearly to the

posterior end of the pectoral fin ‘(adult) or beyond (young).

Gill-rakers short, 6 to 9 on the lower part of the anterior arch.

Dorsal 5, its distinc: from the end of snout 4 its distance from

the caudal (young) or less (adult). Anal 73-78, continued on to

the caudal. Pectoral spine arora its length a little more than

4 that of the fin or nearly 4 the length of head; outer edge

#5 oughened or finely serrated, inner edge entire. Ventrals with

13 rays. Caudal subtruncate. Adult uniformly greyish (in

spirit); young dark greenish above, yellowish below, with the

sides and anal fin marbled.

Two specimens, 130 and 275 mm. in total length, one from

Kimhoa, the other from Chong-ju.

This species is closely allied to Stluwrus asotus L., from China

and Japan, which differs especially in the longer pectoral spine

with both edges serrated. From S. grahami Regan, from

Yunnan, with which it agrees in the structure of the pectoral

spine, S. bedfordi is distinguished by the broader head, shorter

snout, and shorter pectoral spine (2 the length of ‘head in

S. graham).

8. LioBAGRUS ANDERSON], sp. n. (Plate ITI. fig. 4.)

Depth of body 6 in the length, length of head 4%. Head

nearly as broad as long; interocular width 3 in the length of

head. Jaws equal anteriorly ; premaxillary band of teeth 2 to 24

as long as broad; posterior mandibulary barbel extending to

basal part of peeueuel. Dorsal 15; spine ? the Jengus of head.

Pectoral spine 2 the length of the an, which is 2 the length of

head. Anal 15. Cami rounded or gnbammmenie. Greyish ; fins

more or less dusky, with pale edges.

Three specimens from Kimhoa, 65 miles north-east of Seoul,

the largest 110 mm. in total length.

The genus Liobagrus, established by Hilgendorf in 1878, is now

known to include six species, which may be distinguished by the

following synopsis :—

I. Jaws equal anteriorly or the lower slightly projecting ; depth of body about

6 in the length ; interocular width 2# to 3 in the length ot head.

Length of head 33 to 3 in the length of the fish ; premaxillary

band of teeth 4 times: as long as “broad ; 3 posterior mandibulary

barbel extending to base of pectoral 0 or ushily pee ond; dorsal

spine 2 the length Ofeheaaepere tee. Mgricauda.

62 ON. FRESH-WATER FISHES FROM CORBA. [ Feb. 4,

Length of head 4 in the length of the fish ; premaxillary band of

teeth nearly 3 times as long as broad; ‘posterior mandibulary

barbel extending to middle of pectoral ; dorsal spine a little more

than 3 the length of head ... marginatus.

Length of head 4@ in the length of the fish; "premaxillary ‘band of

teeth 2 to 24 as long as broad; posterior mandibulary barbel

extending to basal nee of pectoral dorsal Per 2 the length of

head .. Ao diaper tei : Renae andersoni.

I. ren er jaw Eins than Ti upper.

A. Interocular width 22 to 3 im the length of head.

Depth of body 6 to 7 in the ‘length ; premaxillary band of teeth

2} to 3.as long as broad; dorsal spine + the length of head;

pectoral spine ‘less than 2 the length of the fin ..... styani.

Depth of body about 6 in the length ; premaxillary band of teeth

22 as long as broad ; dorsal spine 3 \'the length of head ; ecco

spine more than the length of the fin.. hcg , reini.

B. Interocular width 2 to 25 in the iisasdh a Thar,

Depth of body 43 to 5 in the length ; pr emaxillar y band of teeth

twice as long as broad; dor sal spine + the length of head;

pectoral spine less than 3 i the length of the fin ..................... sugubii.

LIOBAGRUS NIGRICAUDA.

Regan, Ann. Mag. Nat. Hist. (7) xiii. 1904, p. 193.

Hab. Yunnan.

LIOBAGRUS MARGINATUS.

Amblyceps marginatus Giinth. Pratt’s “Snows of Thibet,”

Appendix, p. 245, pl. i. fig. A (1892).

Hab. Sze-chuen Province, China.

LIoBAGRUS ANDERSONI, Sp. 0.

Hab. Corea.

LIOBAGRUS STYANI.

Regan, Ann. Mag. Nat. Hist. (8) i. 1908, p. 152.

Hab. South Hupeh, China.

LIOBAGRUS REINI.

Hilgendorf, Sitzungsb. Gesellsch. naturf. Freund. Berlin,

1878, joy Ihe joni. ut Fowler, Proc. U.S. Nat. Mus. xxvi.

1903, p. 909, fig. 2.

Hab. Southern Japan.

LIoBAGRUS SUGUBII.

Regan, Ann. & Mag. Nat. Hist. (8) 1. 1908, p 152.

Hab. L. Biwa, Japan.

GoBIID.

9. ODONTOBUTIS POTAMOPHILUS Gthr.

10. CreNOGOBIUS BEDFORDI, sp. n. (Plate ITT. fig. 1.)

Depth of body 54 to 6 in the length, length of head 3} to 34.

Snout longer than eye, the diameter of ation | is 5 in the leach

of head antl twice the interorbital width. Jaws equal anteriorly ;

1908.] | THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 63

maxillary extending to the vertical from the anterior edge of

eye; head naked; nape covered with small scales. Dorsal VI,

18; second ray of spinous dorsal produced into a filament.

Anal 18. Caudal rounded. Pectoral ? the length of head;

ventrals extending less than 4 the distance from their base to the

anal. 36 to 38 scales in a longitudinal series. Brownish olive ;

fins dusky ; caudal with small dark spots; vertical fins with a

narrow pale edge.

Two specimens, 70 and 75 mm. in total length, and some

smaller ones not included in the description, from Chong-ju.

11. TRIDENTIGER COREANUS, sp. n. (Plate III. fig. 2.)

Depth of body 5 in the length, length of head 32. Snout

longer than eye, the diameter of which is 5 in the length of head

and equal to the interorbital width. Head nearly twice as long

as broad; cheeks not tumid. Jaws equal anteriorly ; maxillary

extending to below anterior part of eye. Dorsal VI, 111; rays

of spinous dorsal produced into filaments. Anal 19. Pectoral 2

the length of head. 42scalesina longitudinal series. Brownish ;

a more or less distinct dark lateral stripe, edges of scales darker ;

fins dusky, the pectoral with a blackish spot margined posteriorly

by a pale crescent on the upper part of its base, the anal with a

dark intramarginal stripe.

A specimen of 70 mm. and several much smaller ones not

included in this description, from Chong-ju.

EXPLANATION OF THE PLATES.

Prats II.

. 1. Acanthorhodeus gracilis.

2. Leucogobio strigatus.

3. Silurus bedfordi.

Puate III.

1. Ctenogobius bedfordi.

2. Tridentiger coreanus.

3. Acanthogobio longirostris.

4. Liobagrus anderson.

February 18, 1908.

Dr. Henry Woopwarb, F.R.S., Vice-President, in the Chair.

The Secretary read the following report on the additions made

to the Society’s Menagerie during the month of January 1908 :—

The number of registered additions to the Society’s Menagerie

during the month of January was 173. Of these 125 were

acquired by presentation and 23 by purchase, 9 were received on

deposit, 6 by &xchange, and 10 were bred in the Gardens.

The number of departures during the same period, by death

and removals, was 234.

64 ON THE SOUTH-AMERICAN WILD DOG. [ Feb. 18,

Among the additions special attention may be directed to :—

One Crowned Sifaka (Propithecus diadema), from Madagascar,

new to the Collection, deposited on Jan. 2nd.

Three Caucasian Ibexes (Capra cylindricorinis), from Lago-

decki, 8.E. Caucasus, presented by Prince Paul Demidoff on

Jan. 16th.

Two Greater Birds of Paradise (Paradisea apoda), either females

or immature males, from New Guinea, presented by Sir William

Ingram, Bt., on Jan. 4th.

One Harpy Eagle CUES aes harpyia), from Venezuela, pur-

chased on Jan. 29nd.

One Three-keeled Terrapin (Stawrotypus triporcatus), from

Guatemala, new to the Collection, received in exchange on

Jan. 8th.

On behalf of Mr. W. Simpson Cross, F.Z.8., Mr. R. I. Pocock

exhibited a photograph of the very rare South-American Wild Dog

Text-fig. 15.

Maned Wolf (Canis jubatus). From life.

sometimes called the Maned Wolf (Canis jubatus), and laid stress

upon the importance of publishing photographs of ‘all animals

1908. | ON OLFACTORY ORGANS OF FISHES AND BIRDS. 65

that were rarely imported alive to Europe, so that a correct idea

of their true form might be conveyed to artists and taxidermists.

In the present instance the value and interest of the photograph

submitted to the Society by Mr. Cross might be judged by com-

paring it with the stuffed specimens of this Dog exhibited in

museums and with the published lithographic figures of the

species.

Mr. R. H. Burne, F.Z.S., exhibited a preparation * of the head

of a Sea-Lamprey (Petromyzon marinus), designed to show the

mechanism by which water is drawn into the olfactory chamber

at each inspiration and expelled at each expiration. The current

of water is produced by the alternate compression and expansion

of the blind bulb-like end of the nasal sac. ‘This sac is separated

from the branchial chamber by the thin compressible cesophagus

only and responds to the alterations in pressure within the

branchial chamber that are due to the alternate expansion and

contraction of its walls during the process of respiration. ‘The

current of water entering the nostril is deflected by valves into

the olfactory chamber and among the leaves of the olfactory

rosette before passing into the nasal sac.

Text-fig. 16.

Section of the head of an Albatross (Diomedea exulans).

L.D., opening of lacrymal duct; M.T., maxillo-turbinal ; N.D., opening of duct

of nasal gland; OL.B., olfactory bulb; OL.E., olfactory eminence.

The action of this mechanism has been observed and recorded

by Bert (Ann. Sci. Nat. sér. 5, t. vil. 1867, p. 372.)

* No. E 85, Physiol. Series, R. College of Surgeons’ Museum.

Proc. Zoou. Soc.—1908, No. V. 5

66 DR. SAMBON ON INTERNAL PARASITES. [ Feb. 18,

Mr. Burne also exhibited a specimen showing an aspirator

contrivance of a somewhat similar character in the Sea- Bream

(Pagellus centrodontus)*, by the action of which a current of water

is drawn into the anterior nostril, passes over and through the

olfactory rosette, and is expelled by the posterior nostril.

The mechanism consists of two accessory membranous sacs that

open into the hinder end of the olfactory chamber and are so

placed that in the resting state with the mouth shut they are

compressed respectively by the process of the premaxilla and by

the dorsal margin of the maxilla. As the mouth opens the sacs

expand by the withdrawal of these bones from contact with

them, but are again compressed when the mouth is shut. Similar

sacs have been observed in many Acanthopterygian Fishes.

Mr. Burne also showed a section of the head of an Albatross

(Diomedea exulans)*t, in which the brain and the olfactory chamber

had been exposed to show the relatively enormous development

of the olfactory organ (text-fig. 16, p. 65). The olfactory bulbs

measured 7 mm. in diameter, and received large nerves from the

nasal sepum and lateral wall of the olfactory chamber. (The rest

of the cerebral hemisphere measured 30 mm. in length.) The

olfactory eminence or pseudo-turbinal was also remarkably deve-

loped and lay for the most part ina special backward prolongation

or recess of the general cavity of the nose. Except for the Ki-wi

(Apteryx), this is apparently the most highly developed organ of

smell recorded among birds. In this specimen the opening of

the duct of the nasal gland upon the edge of the atrial pseudo-

turbinal, close to the border of the nostril, was also shown.

Dr. L. W. Sambon, F.Z.8., exhibited a large series of speci-

mens of internal parasites obtained by him from animals recently

living in the Society’s Gardens. He laid stress on the important

additions to knowledge to be derived from an adequate investi-

gation of such material, and on the practical results to the health

of the animals in the Gardens that might be expected.

* No, E 95, Physiol. Series, R. College of Surgeons’ Museum.

+ No. £ 116 a, Physiol. Series, R. College of Surgeons’ Museum.

IV.

PZ.S 1908,P

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H, Goodchild del. et lith,

TUMBLER—FANTAIL CROSS.

1908. | ON INHERITANCE OF COLOUR IN PIGEONS. 67

The following papers were read :—

1. On the Inheritance of Colour in Domestic Pigeons, with

Special Reference to Reversion. By RicHarp STaPLEs-

Browne, M.A., F.Z.S.

[Received January 20, 1903. }

(Plates IV.—VII.*)

Page

Introduction and brief statement of results ........................ 67

Mhe*Barbubigeom!, ssce-cer coms a eee ee oer eee eee aaa

The Fantail Pigeon REA Oe Be Psa obs asco Mh Na so eee

Types of cross- -bred Barb-Fantails ..... aden

Series of Experiments A. (Matings in the direct line.) serine 76

Series B. (Crossbreds mated to Whites.) ........................ 81

Discussion of aberrant results m Series B.............2.....000e000. 84

Series B (continued) _... i cody bea tue cant waceeeel, AOD

Series C. (Blues mated to Blacks.) Oo aat MACUSER GES Cone ap DAay ais) O

Cross between Barb and Nun .. Ae Aerod tS ssa RRA SR ERR A eRe eg QO)

AheyNimnigice One yee kaa a ontener cee puke te Chet BO

Cross-bred birds Ate Bienen 90

Barb-Fantail Crossbreds mated to ‘Barb- Nun Crossbreds dsfarn) LSB:

Nun-Fantail Crossbreds ..... Tey aD:

Cross between White Tumbler and ‘White Fantail.. pai ee Ay

The White Tumbler Pigeon .................. Bae edandabssy MOL

Types of Cross-bred birds produced — AD EPRGRAD SES RE eR Sr ose Mit )5)

Details of the Matings .. UR Teh aad REC AOD BH Os

General account of the colour of the Thetis RE at Retr ma be 98

x Hs aS % Beaks and Claws ......... 102

. _ 3 Bs Bye-wattles psn eeee a Os

Conclusions AI a CR eee REEL Ee ORT. Saat Tee 104.

INTRODUCTION AND BriEF STATEMENT OF RESULTS.

Since Darwin’s classical experiments on reversion, very little

inquiry has been made into the inheritance of colour in Domestic

Pigeons. But with the rediscovery of Mendel’s work a great

impetus has been given to the study of cross-breeding ; and it was

considered that a repetition of experiments with Pigeons, in the

light of modern knowledge of the science of genetics, would prove

both interesting and instr uctive.

The following account contains the description of certain

experiments with Pigeons, begun in 1901, which, although not

identical with the matings used by Darwin, are yet planned on

the lines adopted by him. Most of the pure-bred varieties used

are also those with which he worked.

It will be remembered that the most striking example of

reversion was obtained by Darwin as follows :—He mated a black

Barb to a white Fantail, and also a black Barb to a red Spot,

which is a white pigeon with the tail and tail-coverts red, having,

in addition, a red spot on the forehead. He then mated ‘together

the mongrel offspring obtained from these two crosses, and from

this was produced a bird identical with Columba livia excepting

* Por explanation of the Plates, see p. 104.

68 MR. R. STAPLES-BROWNE ON [ Feb. 18,

that “the head was tinted with a shade of red, evidently derived

from the Spot, and was of a paler blue than in the rock-pigeon,

as was the stomach.” (v. Animals & Plants under Domestication,

2nd edition, vol. i. p. 209.)

In the experiments here described, as a Spot pigeon was not

readily obtainable, a black and white Nun pigeon was substituted.

When the Barb-Fantail mongrels were mated to the Barb-Nun

mongrels, however, no reversionary types appeared (v. Exps. 1,

34, 37, 88, and 39). It was, however, found that, when the

Barb-Fantail crossbreds were mated together, some birds were

produced having certain blue feathers. Various experiments with

birds produced from the cross between the Barb and Fantail

were carried out at some length, but, owing to the limitations of

space, the Barb-Nun crosses were soon discontinued.

The blue colour, when it appeared, was found chiefly in the tail

and neighbouring parts, as had already been observed by Darwin.

Associated with this the black tail-bar was invariably present.

The wing-coverts and backs of the reversionary types obtained

were generally of a smoky-black colour, thus obscuring the two

wing-bars found in the rock-pigeon.

The reversionary type in pigeons may in some cases be obtained

in the first cross-bred generation (F.1). Such a result Darwin

observed when he crossed a Nun with a red Tumbler.

The fact that in the Barb-Fantail cross the reversionary blue

does not appear until F. 2, is interesting. The F.1 generation

contains all the elements introduced by the parental types ; never-

theless it is not reversionary in colour, but resembles the black

Barb except for the addition of some white. From the fact

therefore that the blue reversionary form can be produced by such

F.1 birds, it is clear that they contain some element which

prevents the appearance of the blue. This element is evidently

the factor for black self-colour; and the experiment shows that

this element is dominant, or, more strictly, epistatic, to the blue.

The black factor must thus be regarded as an element not derived

from the wild pigeon, but added to it by some subsequent

variation. When, by recombination of the various elements, the

F.2 forms are produced, those combinations which contain the

blue in the absence of the black factor exhibit the blue, while

those which contain the black in addition cannot exhibit it.

Tn some other cases of reversion on crossing (e.g., Sweet Peas

and Stocks), the reversion can be proved to be due to the meeting

of complementary factors. -In the case of the Barb-Fantail cross

the evidence is not yet sufficient to show whether the factors

needed to produce the atavistic condition are all present in the

Barb, and their effect merely hidden by the presence of the black

factor, or whether a necessary factor is introduced by the

Fantail; but the fact that no blues came in the F. 2 made from

F 1 (Barb x Fantail) x F.1 (Barb x Nun) distinctly suggests

that some factor of the blue did come from the Fantail.

1908. ] INHERITANCE OF COLOUR IN PIGEONS. 69

By far the greatest number of matings here described belong to

the Barb-Fantail experiment. In all 33 such matings were made.

The descriptions of the pure bred birds and the several types of

cross-bred birds produced are first given. The details of the

various matings are then described. For the sake of clearness

these are not given in the order in which they were made but are

divided into three series.

Series A, comprising Exps. 1-11, deals with matings in the

direct line, together with the testing of extracted whites. The

results are further tabulated in Table I.

Series B (Exps. 12-26) shows the matings of crossbreds, chiefly

blues, to whites, and the subsequent matings of birds derived

from such crosses. Tables IJ. and III. deal with results in this

series.

Series C (Exps. 27-33) deals with the matings of blues and

blacks and the further crossings of offspring produced from such

matings.

The minor characters—irides, beaks, claws, and eye-wattles—are

described at the end of the paper; the details given under the

descriptions of the experiments refer to plumage only.

The general results of the Barb-Fantail experiments may be

briefly summarised as follows :—

The F.1 generation shows a dominance of black to white, and

the further matings show that blue is also dominant to white.

This dominance of the coloured to the non-coloured type is, how-

ever, imperfect, as the majority of birds produced from the mating

of black or blue with white show some white feathers, chiefly on

the rump.

In the F. 2 generation the following types appear :—

Black.

Black, with some white feathers. (Black w. f.)

Blue.

Blue, with some white feathers. (Blue w. f.)

Red.

White, with some coloured feathers.

White.

The blue type may be homozygous or may be dominant to

white. Black was never obtained from the mating together

of two blues. The matings of blues and blacks in Series C

show that blue is a simple recessive to black. The absence of

white feathers in blue birds of the F.2 generation does not

necessarily indicate that they are homozygous, for Exp. 30 shows

such a blue to contain white. Conversely, an F.2 blue with some

white feathers is shown in Exp. 13, when mated with a white, to pro-

duce blues with some white feathers only. The significance of the

presence or absence of white feathers has not been clearly made out.

It was at first thought their presence was indicative of the fact that

the bird was giving off white-bearing gametes. This, however, is not

70 MR, R. STAPLES-BROWNE ON [ Feb. 18,

so inallcases*. From the matings of blues with white feathers, as

in Exps. 11, 24, 25, and 26, whites were produced, and a definite

proportion of homozygous blues was to be expected. With one

exception, however, all the blues produced from these matings

showed some white feathers. We are led to conclude that some

of these are probably homozygous, although the assertion cannot

definitely be made without testing a large quantity of such birds,

of which space did not permit. On the other hand, the matings

of blues with some white feathers to whites in Series B has not

revealed a homozygous bird other than that in F. 2 already

mentioned.

One distinctly abnormal result occurred in Series B from the

mating of blues with white feathers to whites. The proportion of

whites produced was here much higher than the expected equality.

This result, which is discussed later, evidently points to the

existence of a definite complication.

The red birds obtained in F.2 are only briefly mentioned

in the present paper. A further series of experiments, dealing

with them, is now in progress, a full report of which will be

published in another communication,

The majority of reds, produced in the F.2 generation and from

subsequent matings, showed a bluish tail with a very distinct bar,

the under parts also having a bluish tinge. Certain specimens

have, however, been pr oduced in which the bar is absent, and the

amount of blue much reduced. It is possible, therefore, that two

kinds of reds may eventually be demonstrated. In addition, red

has shown itself to be recessive to both black and blue, but

dominant to white >

The extracted whites, which are shown to breed true, need no

comment, with the exception of five produced in the direct line

(Exps. 4, 5, and 8) in which some coloured feathers were present.

These are shown in Exp. 7 to produce whites with and without

coloured feathers in equal numbers. It is probable that this has

* As Crampe and Doncaster have shown in rats, and Hurst in the case of rabbits,

the presence of some white in otherwise self-coloured types, may be an indication of

heterozygosis in respect of a pattern-factor, and the same possibility is to be remem-

bered here.

+ The results and figures so far obtained from the further experiments are briefly

as follows :—

(a) F.2 red X F.2 ned gave 7 red, 1 white with few red feathers, 2 white.

(6) F.2 2 red X F.2 red , red, 3 white.

(ec) F.2 red X F.3 red ‘i 3 red, 1 white with few red feathers, 1 white.

(d) F.2 red X white » 2 black, 8 white.

(e) Black from (@) X white ;, 12 white, 6 blue, 2 black, 1 red.

(f) F.2 ved X black a S red, 4 black.

(g) Red from (f) X red from (f) ,, 7 red.

(h) F.2 red X blue s 6 black, 5 blue.

(¢) Black from (2) X black from (h) (2 pairs) gave 10 black, 7 red, 3 blue.

(7) Blue from (2) X blue from (2) gave 9 blue, 2 red.

In the above no mention is made of white feathers occurring on coloured birds ;

they occurred, however, on a large number, and their distvibution corresponded to

that in similar birds described in the present paper. It will be noticed that the

number of white birds produced was above the expected proportion. Exp. (7) was.

a brother and sister mating.

1908. ] INHERITANCE OF COLOUR IN PIGEONS. 71

been introduced by the Fantail. It is pointed out below that one

of the Fantails in the strain used (Lee 97) showed a_ black

feather. The relation between this bird and Fantail ¢ 23, the

bird used in Exp. 2, is shown in a pedigree of the Fantails.

Although several pairs of pure-bred Fantails were kept, ‘‘ splashed”

birds never appeared in their offspring. An opportunity appears

to have arisen in the crossing, however, by which this latent

character was able to manifest itself.

A further series of experiments was undertaken on the crossing

of two distinct white breeds, Tumblers and Fantails. Here again

blue colour was produced in the F.2 generation. An account of

this cross is included in the present paper.

For testing the various Mendelian ratios the pigeon is not a

thoroughly satisfactory subject to work with, unless the experi-

ments can be carried out on a very extensive scale. To insure

the desired mating it has been found necessary to keep each pair

of birds in a separate aviary. The number of pairs kept is there-

fore limited. Further, the number of offspring produced by a

pair of pigeons is comparatively small, seldom exceeding ten in

any one year. Ina few cases, when a ratio between the numbers

of the various types of offspring produced by a cross was desired,

the same mating has been continued for a second year.

In the following account the ordinary Mendelian terms

are used.

Tue Bars Piczon. (Pigeon polonais.)

In ‘Animals and Plants under Domestication,’ Darwin describes

the points of the Barb thus:—‘“ Beak short, broad, deep; naked

skin round the eyes broad and carunculated ; skin over nostrils

slightly swollen.” It was hoped that these characters might be

traced in the crossbreds, but some ditticulty was experienced in

obtaining accurate measurements, and the attempt was abandoned.

The ‘eye-wattle” or cere, moreover, does not attain to its

maximum development until the bird is in its fourth year. The

Barb is a ‘self-” or whole-coloured pigeon, and is found in black,

red, yellow, dun, and white. Of these black is the commonest.

Blue Barbs are exceedingly rare, and are seldom if ever used by

breeders for crossing with the other colours. No wing~ or tail-

bars, or chequering are found in the usual colours. The eye-

wattle is bright red. The iris is generally white, sometimes

orange, and in white Barbs is black. The beak and claws are

desired by breeders to be white; there is, however, a tendency for

black Barbs to have the beak tipped with black, or even horn-

coloured beaks. This was the case with the specimens used in

these experiments. Four black Barbs were obtained; those used

in Exps. 1 and 34 were bought through a poultry advertising

paper, and their origin is unknown. Barb 2, No. 7, used in

Exp. 2, was obtained from Mr, J. Wilkins of Swindon. This

72 MR. R. STAPLES-BROWNE ON [ Feb. 18,

bird was bred from blacks only for five or six generations. In

the F.2 generation from this bird crossed with a white Fantail

red birds were obtained, and it is quite possible that this colour

figured in the ancestry of the Barb 9, as breeders are in the

habit of crossing blacks with reds in order to obtain white beaks

on the blacks. Barb ¢, No. 100, used in Exp. 27, was obtained

from Mr, Edwards of Exeter. This bird is believed to have been

bred from blacks for four generations, but in the preceding

generation a dun 2 was used. It will be noticed that in Exp. 27,

a dun was produced.

Tue Fanrai Piczon. (Trembleur.)

This variety is well-known on account of the large number of

its tail-feathers; specimens having as many as 42 tail-feathers

having been observed. This character has been noted in the

crosses, and further experiments are still in progress. It is

intended that this should be dealt with later in a separate

communication. By far the largest number of Fantails are white,

and only white birds have been used in these matings. Blue and

silver Fantails, both having wing- and tail-bars, are bred by

fanciers, as well as self-coloured blacks, reds, yellows, and duns.

There are further certain birds possessing well-defined patterns of

which the “saddle-back” is the best known.

It is stated by breeders that deep rich blacks are difficult to

obtain, owing, no doubt, to the frequent crossing with blues.

The irides of the white birds are black, those of the coloured

birds either white or orange. The whites have also white beaks

and claws, and flesh-coloured ceres or eye-wattles. The Fantail

exhibits a curious jerking or twitching movement of the neck ;

this habit has been noticed in certain of the crossbreds, but no

attempt has been made to trace it through the successive

generations. During the experiments a small strain of white

Fantails was kept, the original birds being obtained as

follows :—

From Mr. J. Harrison of Belper, Derbyshire, a judge of pigeons

and breeder of several varieties, 1 9 andl ¢.

From Mr. J. Lee of Ilford, Essex, 4 Q’sand1 ¢.

From the late Mr. J. F. Loversidge of Newark, 1 ¢.

From Mr. W. Stevenson of Beith, N.B., 1 ¢. The three latter

gentlemen are well-known breeders of Fantails.

From the various matings of these birds twelve Fantails used

in the crosses were raised. The appended pedigree, in which the

birds used are numbered and underlined, shows their mutual

relationships. Lee 9 7 was peculiar in the fact that at the

moult following its purchase a black tail-feather made its appear-

ance. At the various successive moults the replacing feather in

that position was always black. Although several white Fantails

were bred in this strain no bird was produced showing any colour

in the plumage. ‘The colours of the irides, beaks, claws, and eye-

-wattles were also observed to breed true.

INHERITANCE OF COLOUR IN PIGEONS.

1908.]

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74 MR. R. STAPLES-BROWNE ON [ Feb. 18,

Typrs or BrrDs PRODUCED IN THE BARB-FANTAIL CROs.

The birds produced in this cross fell naturally into five classes—

black, blue, dun, red, and white. Of these the black, blue, and

white are chiefly dealt with in the present paper.

Coloured birds are divided into two classes; those with some

white feathers in the plumage, and those without white. The

white birds are similarly divided into those having some coloured

feathers, and those with none.

It was possible to see to which type a bird belonged at an

early age.

The descriptions of the types which follow include the more

striking variations noticed in individuals.

1. Blacks.

The blacks produced in these crosses were generally of a rich

deep colour similar in all respects to that found inthe Barb. They

possessed the green and purple iridescence on the neck which is

commonly seen in all dark types of domestic pigeons. There

were, however, certain birds produced, which are specially noted

in Exps. 27 and 32, bred from the mating of black and blue,

which were of a smoky or sooty black colour. In some of these

birds wing-bars of a darker shade of black were observed. The

tail-bar was observed on one specimen only.

2. Blues.

The birds classed as blue in the following experiments were

not identical with Columba livia, as were the birds, produced by

Darwin, referred to in the introduction. Their general colour

was a smoky black with blue tail and black tail-bar. Laying aside

the details of the distribution of white when present, which is

dealt with below, the series of blue birds produced was very

uniform in type. Slight variations occurred in the shade of the

blue colour, and the substitution of chequered feathers (7. e. blue

feathers edged with a variable amount of black) for the smoky

black of the wing-coverts. These points were not very distinctive :

they are, however, noted under the experiments in which they

occurred. The head and upper part of the neck were generally

bluish, slightly darker than in C. livia, but varying to a smoky

black. The lower part of the neck, upper part of the back, wings,

and upper part of the breast smoky black. The wing-bars were

generally obscured, but in some specimens could be distinguished.

The lower part of the breast, lower part of the back and abdomen

were blue, occasionally with slight chequering.

The rump was blue, sometimes of a lighter shade than that in

other parts of the body. The flanks, thighs, vent, upper and

under tail-coverts were blue. The tail-feathers were blue with a

black bar, which was sometimes terminal, but more usually a

1908. | INHERITANCE OF COLOUR IN PIGEONS. 75

short distance from the end of the tail. There was frequently,

but not invariably, a white edging to the outer tail-feathers.

“ Kitiness.”

This term is used by breeders to express a rustiness or bronzing

of the black feathers. It gives to a black feather the appearance

of being edged with a reddish tinge. In certain breeds of pigeons,

notably the Jacobin, this characteristic persists through life in

some individuals. In the majority of cases, however, it is lost

either before or at the first moult. In the cross-bred birds pro-

duced in these experiments kitiness was observed in many of the

young birds. It appeared not only on the blacks but also on the

smoky-black parts of the plumage of birds classed as blues. In

the case of every bird that was allowed to reach maturity the

kitiness disappeared, except in cases where it was present on the

flight-feathers. In this situation it was noticed to persist through

life. There is, at present, no reason to suppose that kitiness

influences inheritance of colour.

3. Blacks and Blues with some white feathers.

It is convenient to describe these two types together. They

differ from the two preceding types only in being more or less

mottled with white. The amount of white varies very greatly.

The black colour has never been observed to be of the sooty shade,

but always deep as in the Barb. The amount of white present

appears to increase with the number of generations from the

original cross. With very few exceptions colour has been in

excess of white in these birds. The white is found in certain

fairly well-defined areas, of which the following is a list arranged

roughly in the order of frequency of occurrence.

1. Rump, vent, and thighs.

2. Certain areas on the head, noticeably the occiput and post-

orbital regions. The neck. The lower part of the

abdomen. Carpal joints. The bastard wings.

3. Primary flight-feathers generally commencing with the most

external, and tertiaries immediately over them. ‘Tail-

feathers generally commencing with those near the middle,

and both upper and upper tail-coverts.

4. Breast, wing-coverts, and scapulars.

Bonhote has shown in the Journ. Linn. Soe. vol. xxix. p. 185,

that the presence or absence of colour tends to make its appear-

ance in mammals and birds in certain definite areas, which he

has named ‘ pecilomeres,’ and of which he gives a list in the

Proceedings of the [Vth International Ornithological Congress.

It will be noticed that the positions of the white in these pigeons

agree closely with Bonhote’s observations.

In the descriptions of the results of matings details of the

various markings are given.

76 MR. R. STAPLES-BROWNE ON [ Feb. 18,

4, Dun.

One bird of this type was obtained in Exp. 27. The head,

neck, back, rump, breast, abdomen and under parts were of a

uniform dark dun colour. The wing-coverts were edged with a

lighter shade. Wing-bars could not be distinguished. The flight-

feathers were lighter than the rest of the plumage. The tail-

feathers showed a bar of a darker shade which was conspicuous

on handling the bird.

5. Reds.

Five red birds were obtained in F. 2 from Experiment 4. The

shade varied slightly in the different individuals, in some the

colour being much darker than in others. In all the rump, tail-

feathers, and both upper and under tail-coverts presented a

bluish tinge. This, however, differed in the various specimens.

In some there was merely a bluish-red appearance on the rump

and tail, in which case the tail-bar appeared to be merely an

aggregation of pigment ; whilst in others the rump and tail were

almost of the same colour as that already described on the “blue”

birds. In these latter the tail-bar was well defined and appeared

to be of a reddish-yellow colour. White feathers were present in

varying amount on all the red birds. The distribution of the

white corresponded with that already described for blacks and

blues.

6. Whites.

The majority of whites produced in these experiments showed

no signs of coloured feathers. A few, however, were raised which

had a few ticks of colour on the neck or rump. These were

either black or red. Full details are given of these birds in the

descriptions of the matings from which they were produced.

DETAILS OF THE SEVERAL MATINGS.

Series A.

F. 1 Generation.

Exp. 1.—White Fantail 9 19 x Black Barb ¢, no number.

Five young birds raised.

Exp, 2.—Black Barb @ 7 x White Fantail ¢ 23. Four

young birds raised.

The colour of the F.1 generation raised from the two above

experiments was practically uniform. The birds were black with

a few white feathers which usually appeared on the rump, vent,

and thighs. In only one instance was there any approach to

mottling, and in this bird the black was greatly in excess of the

white. Birds raised in the two reciprocal experiments were

indistinguishable.

Three matings of the F.1 birds were made, the results of

which are shown in Table I. Thirty-four birds of the F. 2

generation were raised.

INHERITANCE OF COLOUR IN PIGEONS.

1908.]

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FEF, 2 Generation.

As will be seen by the Table no black birds were raised in

Exp. 38. This experiment came to an untimely conclusion early

in the breeding season owing to the accidental death of the ¢.

No other ¢ raised in Exp. 1 was then living. It seems likely,

however, that if the mating could have been continued black

birds would have appeared. Exp. 4 was continued for two years,

and, in addition to the birds included in the table, a young bird

was hatched which died in the nest at the age of ten days. It

was dark in colour, but whether it would eventually have been

black or blue could not at that age be distinguished.

Black birds in the F. 2 generation.—Of the fifteen black birds

raised five showed no trace of white, the other ten having some

white feathers. Birds entirely black were not tested at this point

of the experiment, but in the crosses between Barbs, Fantails,

and Nun pigeons it was shown that extracted blacks in F. 2 bred

true (v. Exps. 40 & 41). Those having white feathers were

noticed to show more white than the birds of the F. 1 generation.

In addition to having white feathers on the same parts of the

body as the preceding generation, these birds frequently showed

white on the head and throat, also on the wings, and more espe-

cially on the flight-feathers. The black, however, always greatly

predominated in the plumage. In one case there were only a

very few white per uberst on one thigh.

Blue birds in the F. 2 2 generation. —In addition to the general

description of the blue birds already given, it may here be stated

that the bird bred in Exp. 5 (No. ‘19) differed from those in

Exp. 3 in having the blue colour much darker. In the young

raised from the subsequent matings of the offspring of this bird,

the dark blue colour was very marked, so that it was possible to

pick them out at sight in an aviary containing many blue birds.

This bird showed two white feathers on the rump which were

afterwards moulted out. Experiment, however, proved that it

did not contain white (v. Exp. 13). Of the blue birds raised in

Exp. 3 two (Nos. 1 & 14) showed no white, while the third

(No. 51) had a few white feathers at the vent. Further matings

proved that two of these birds contained white recessive, while

the other did not (see Exps. 9, 16, & 30). Of the four F. 2 blues,

therefore, two were homozygous to that colour.

Red birds in the F. 2 generation.—A general description of

these birds has already been given. It seems desirable to post-

pone all further details until the investigation is more advanced.

White birds in the F. 2 generation.—Of the ten extracted white

birds six showed no coloured feathers, and four showed a little

colour. From Exp. 4 two birds with coloured feathers were

raised. Of these one (No. 1) showed four or five feathers of a

brownish or reddish tinge on the neck, these were afterwards

moulted out. The other (No. 15) had the head and throat

ticked with some reddish feathers. These birds were not bred

from. In Exp. 5 one white bird (No. 5) was produced which

1908. } INHERITANCE OF COLOUR IN PIGEONS. 79

showed a large patch of black feathers about the size of a five-

shilling piece at the root of the neck. Another (No. 56) raised

in the same experiment had a few feathers on the rump edged

with black. These two birds were mated together in Exp. 7.

Testing the extracted Whites in F. 2.

At the time it was considered desirable to test the extracted

whites only one was available, viz. that bred in Exp. 3. It was,

therefore, mated to a white Fantail.

Exp. 6.—Extracted White 9 53 x White Fantail ¢ 9.

Four young were produced, all white with no coloured feathers.

Exp. 7.— White birds with coloured feathers raised in Exp. 5

described above were mated together.

White with black feathers 2 56 x White with black

patch g 5.

Ten young were raised in this experiment, of which five showed

no coloured feathers; one had one black feather on the rump

which was afterwards moulted out; two had two feathers on the

rump edged with black; and the remaining two had four and five

feathers respectively similarly edged with black.

F, 3 Generation.

A pair of black birds with some white feathers of the F. 2

generation raised in Exp. 5 were mated together in 1905, and the

experiment was repeated in 1907. In the intervening year the

birds were used for mating with blues in Exps. 28 & 31.

Exp. 8.—F. 2 Black w. f. 2 106 x F. 2 Black w. f. ¢ 6.

This mating gave 2 blacks, 6 blacks with some white feathers,

3 blues with some white feathers, 1 white with some coloured

feathers, and 2 whites. Of the blacks with some white feathers,

one bird had only one white feather on the rump, another had

two white feathers on the thighs and three of the under tail-

coverts tipped with white. The remainder had white on the

head, neck, rump, vent, thighs, and under tail-coverts; but one

bird had in addition 12 white flights, and 11 white tertiaries,

while another had some white flights and tertiaries and also 4

white tail-feathers. Of the blues with white feathers, one bird

had white on the rump only, the other two having white on the

head, neck, rump, vent, and thighs, one of them having in addi-

tion three white flights and two white tertiaries. The bird

described as white with some coloured feathers had three feathers

on the rump tipped with black.

Exp. 9.—F. 2 Blue few white feathers 9 51 x F.2 blue ¢ 1.

These birds were raised in Exp. 3.

Nine offspring were reared from this mating, of which three

were blue with no white. The six others all showed white

feathers, four of them having white on the head and throat, vent,

thighs, &c., and some flight-feathers up to the number of ten as

80 MR. R. STAPLES-BROWNE ON | Feb. 18,

well as two or three tail-feathers and some under tail-coverts.

One bird had only two white tail-feathers and some white on the

vent. Another (No. 51 F. 3) had only a very few white feathers

at the vent. This bird, however, when mated to a blue without

white (v. Exp. 10) produced young with white feathers. (The

matings of these several young are shown in Exps. 10, 11, 14,

15, 17, 18, 21, 27, 28, and 31.)

FH. 4 Generation.

Exp. 10.—F. 3 Blue 9 97 x F.3 Blue with very few white

feathers ¢ 51.

These birds were raised in Exp. 9. Four young were reared

from this cross, of which three showed no white feathers, and the

fourth had one white flight-feather, and a large patch of white on

the vent and left thigh. In addition to these, two young birds

were hatched but died at the age of 10 days: on one of these

white quills were seen on the abdomen, but on the other no

white quills were visible. From the above it appears that the 9

bird was homozygous in respect of the blue colour, but that the

¢ contained white. Unfortunately the three F. 3 birds which

showed no white were all Q’s.

Exp. 11.—F. 3 Blue w. f. 9 52 x F.3 blue w. f. ¢ 13.

In this experiment two of the birds with white feathers raised

in Exp. 9 were mated together. Kleven young were raised, of

which 9 were blue with white feathers and 2 were white. Of

the blues with white feathers six showed much white; the

white being in the same situations as in the F. 3 birds but more

extensive. There was much white on the head and neck, the

back and breast were in some cases mottled. The flight-feathers

were mostly white up to the number of 29, and many tail-

feathers up to 18, and several under tail-coverts. The whole

appearance presented was a blue bird mottled with white. The

other three blues, however, had only a few white feathers.

No. 15, F. 4, had only a few white feathers on the head and a

small patch on the abdomen and vent. This bird, how-

ever, was subsequently proved to contain white (see

Exps. 19 and 22).

No. 22, F. 4, had a small patch on the vent and thighs, two

white tail-feathers, and a few under tail-coverts.

No. 76, F. 4, had a small streak of white behind the eyes, and

two white feathers at the vent.

Tt will be noticed that no blues without white were raised in

this experiment as was anticipated. This result may be compared

with those of Exps. 24, 25, and 26, in which birds raised from the

matings of the heterozygotes with whites did not produce the

expected number of blues without white when mated together.

Matings in the direct line have not been carried beyond this

generation.

1908. ] INHERITANCE OF COLOUR IN PIGEONS. 81

Series B.

THE CROSSBREDS MATED TO WHITES.

F.1 xX white.

Exp. 12.—F.1 Black w. f. 2 63 x White Fantail ¢ 32.

The 2 63 was raised in Exp. 1 and is the same bird used in

Exps. 3, 5, and 39. In this experiment only two young were

raised, both black with white feathers. One (46) had only a few

white feathers on the rump, vent, and thighs; the other (47) had

in addition to this a few on the head and neck and a few wing-

and tail-feathers white. Had this experiment been continued

whites would undoubtedly have appeared.

Blues of F.2, F.3, and F.4 mated to whites.

Experiments 13 to 23 (see Table II.) show the results of mating

the blue birds raised in the foregoing experiments to whites.

Some of the whites used were pure Fantails, but the results were

apparently the same when “ extracted” whites were used (Exps.

19, 20, 21, 22).

Exps. 13, 14, and 15 show the result of mating homozygous

blues with whites—24 young were raised in these three experi-

ments, all blue with some white feathers.

Exps. 16-23 show the results of mating blues contaiming white

to whites. In all 41 young were raised, of which 13 were blue

with white feathers and 28 were white. ‘The expected results

from these matings were’blues (with white feathers) and whites

in equal numbers. The irregular numbers are discussed in detail

immediately after the descriptions of the experiments from which

they were obtained.

Details of Hxps. 13—23.

Exp, 13.—White Fantail 9 25 x F.2 Blue (with 2 white

feathers) g 19.

The ¢ has already been described among the blue birds of the

F. 2 generation as being of a darker colour than usual, and having

two white feathers which were subsequently moulted out. Twelve

offspring were reared all blue with some white feathers. The

blue colour varied slightly, in some it was of a darker shade than

in the father, and in all was darker than in birds raised in the

other experiments. The amount of white also varied from a bird

having only a few white feathers on the rump, vent, and thighs,

to one having white on the head, neck, breast, rump, abdomen,

vent, and thighs, as well as several wing-feathers, 10 tail-feathers,

and some under tail-coverts. The average amount of white was

about midway between these two extremes.

Exe. 14._F. 3 Blue 2 98 x White Fantail ¢ 46.

Exp. 15.—F.2 Blue 2 120 x White Fantail 5 26.

The two 9 birds were raised in Exp. 9. No white feathers

were seen. ‘Twelve offspring were raised all blue with white

feathers. The blue colour was uniform throughout. In some

cases the sooty colour of the wing-coverts was to a slight extent

Proc. Zoou. Soc,—1908, No. VI. 6

[ Feb. 18,

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1908. } INHERITANCE OF COLOUR IN PIGEONS. 83

replaced by blue chequered with black. In these cases indication

of black wing-bars were seen. The amount of white varied, but

was in excess of the amount present in the F. 3 blues with white

feathers. The birds were distinctly mottled, and in some cases

the amount of white was in excess of the blue. It appeared on

the head and neck, the rump, abdomen, vent, and thighs—most

of the flight-feathers were white and several of the tail-feathers

and under tail-coverts.

Exe. 16.—F. 2 Blue with few white feathers 2 51 x White

Fantail f 2.

The 2 was raised in Exp. 3. The result of the mating was

4 whites and 2 blues with white feathers. Of these one (No. 13)

had white on the head, rump, vent, and thighs, 6 white flight-

feathers and some other wing-feathers. The other (No. 36) had

white on the head, throat, rump, vent, and under tail-coverts,

16 white flights and some other wing-feathers, and 5 white tail-

feathers. These two birds were mated together in Exp. 26.

Exp. 17.—White Fantail 9 40 x F.3 Blue w. f. ¢ 30.

Exp, 18.—F.3 Blue w. f. 9 14 x White Fantail ¢ 44.

The two blues with white feathers were raised in Kxp. 9.

Fifiteen young were produced from these two experiments, of

which eleven were white and four blue with white feathers. The

amount and distribution of white corresponded roughly to the

description given of similar birds in Exps. 14 and 15.

Exp. 19.—F.4 Blue w. f. 9 15 x F.3 Extracted white ¢ 50.

Exp. 20. F.4 Blue w. f. 2 9 x F.3 as above ¢ 50.

Q 15 was raised in Exp.11. 9 9in Exp.10. The same ¢ 50

was used in the two experiments. This bird was bred from an

F. 2 Red and a white Fantail, which mating is not described in

the present paper. The result of the two experiments was 4

whites and 2 blues with white feathers. These birds were similar

to those bred in Exps. 14, 15, 17, and 18.

Exp. 21.—Extracted white 2 20 x F.3 Blue w. f. ¢ 13.

Exp. 22.—Repetition of Exp. 19.

Exe. 23.—F.4 Blue w. f. 9 9 x White Fantail ¢ 34,

The white 2 20 was raised in Exp. 31 from a blue and a black,

both containing white. F.3 ¢ 13 was raised in Exp. 9 and is the

same bird that is used in Exp. 11. F.4 2 9 is the same bird as

used in Exp, 20. -These three pairs were put up to test further

the proportion of blues with white feathers to whites. The young

birds were killed as soon as they had feathered sufficiently for

their colour to be seen. No details were kept. The blue with

some white feathers in Exp. 22 died in the shell a day or two

before it would have hatched. No hesitation, however, is felt in

recording this bird as blue since it had a black beak. It was

found that, although blue birds with white feathers sometimes

occur with white beaks, no case has been met with in which white

birds had black beaks. In addition to this it was noticed, as the

experimenter became more familiar with the appearance of young

birds in the nest, that birds which were subsequently coloured

84 MR. R, STAPLES-BROWNE ON (Feb. 18,

had a darker down than those which became white. In Exp. 23

only two birds were raised ; eight other eggs were laid, but proved

unfertile. Exps. 21 and 23 produced blues with some white

feathers and whites in equal numbers ; on the other hand, Exp. 22

produced 1 blue and 5 whites. The result of the three exper-

ments being 5 blues to 9 whites. It may further be noted that

when Exp. 22, which is a repetition of Exp. 19, is reckoned with

Exp. 19, we get a result of 2 blues to 7 whites, the exact figures

found in Exp. 18.

Discussion oF ABERRANT RESULTS IN Exes. 16—23.

The expected result from these matings (DR x R) was an

equal number of blues (with white feathers) and whites. The

result obtained, however, was 13 blues to 28 whites. It is there-

fore necessary to analyse the results more closely. The only

matings which gave equality were 21 and 23, these gave 3: 3 and

1:1 respectively. The ratio 2: 4 was obtained in Exps. 16 and

17, and 1:2 in Exp. 20. Seeing that the total number of birds

produced in each family was small, the divergence of these ratios

from the expected equality would not suggest any very marked

irregularity. The totals, however, would point to the need for

repetition of the experiments with greater numbers. But when

we obtain the unlooked for result of 2:7 from Exp. 18 and from

the birds mated together in Exps. 19 and 22, we are forced to

conclude that some definite disturbing factor is present. The

following questions suggest themselves. Do the results obtained

from reciprocal matings differ? Has the fact of the white parent

being a pure Fantail or an “extracted” any bearing on the

matter 2 With regard to the reciprocal matings, only two matings

were made (Exps. 17 and 21) in which the 3 was blue. These

gave ratios of 2:4 and 3:3 respectively. Dividing up our totals

according to the nature of the matings, we arrive at the following

figures:—When the ¢ was white the offspring produced were

8 blues and 21 whites. When the ¢ was blue the offspring pro-

duced were 5 blues and 7 whites. It will be noticed that, although

the whites are in excess in both cases, the divergence is much

more marked when the ¢ was white. The possibility of this

having some influence on the proportions of the offspring cannot

at present be disregarded. The behaviour of the pure Fantail

and the extracted White appears to be identical. Only two

extracteds were used: one of these in Exp. 21 gave a ratio of 3:3,

the other in the mating used in Exps. 19 and 22 gave 2:7, and

the same bird mated to another blue Q in Exp. 20 gave 1: 2.

In connection with the excess of white offspring when the ¢

was white, it is perhaps worth recalling that in certain remarkable

instances recessive forms appear in F.1 when a recessive ¢ is

used. The best ascertained example of this phenomenon occurs

in the Canary. Cinnamon (2. e. pink-eyed) 9 x green (7. e. black-

eyed) ¢ gives F. 1 all black-eyed; but black-eyed 9 x pink-eyed g

may produce some pink-eyed birds, which are said to be always

1908. ] INHERITANCE OF COLOUR IN PIGEONS. 85

females. Professor Whitman also informed Mr. Bateson that in

certain of his crosses between species of Doves white females may

be produced when the father is white, though the reciprocal cross

gives all coloured birds.

Unfortunately no sufficient record of the sexes produced in the

cases of the Barb-Fantail Crossbred birds was made; and in order

to establish a comparison with these other cases, it would be

necessary to show that among the whites here produced there

was an excess of females.

Results of mating together Blue birds with white feathers, raised im

Laps. 13 and 16 respectively. (See Table IIT.)

Exp, 24.—Blue w.f. 9 5x Bluew.f. ¢ 4.

Exp. 25.—Blue w. f. 2? 10 x Blue w.f. ¢ 11.

These four birds were raised in Exp. 13. The results of the

matings were 6 blues with white feathers and 2 whites. No blue

birds without white feathers were raised (cf. Exp. 11). One of

the birds (48), however, had only a few white feathers on the

rump, vent, and thighs; and another (17) had white in the same

position with a few under tail-coverts. The bird having the

maximum development of white (55) showed a few white feathers

on the head, neck, rump, vent, and a little mottling on the breast,

4 white flights, 4 white tail-feathers, and a few under tail-coverts

and other wing-coverts. The amount of white in this bird is not

so much as in the bird showing the maximum development of

white in Exp. 13. In addition to the birds included in the Table

a bird was hatched in Exp. 24 which died in the nest when very

young. It was of a distinct reddish tinge with some white

feathers, having tail and flights “ blackish.” It is probable that

this was “ kitiness ” (v. ante), and the bird would have eventually

been blue with white. Only two birds were raised from Exp. 25.

These were blue with some white feathers. There is no doubt

however that, had the mating been continued, white birds would

have appeared as in Exp. 25.

Tape III.

OFFSPRING.

| | |

| Blue.’ Blue | White.|

74 w. f.

ic}

‘AxGy WOAy ULSIIO

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24. i Blue with some Blue with some |

| ae | 13 | Bae | a / 1h fie | 4

25. | Blue with some Blue with some |

white $10 (te Sue jae | al z

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26. | Blue with some | Blue with some

} waite $36 16 | eaite $18 jie} 4 Bile Was

86 MR, R, STAPLES-BROWNE ON [ Feb. 18,

Exp. 26.—Blue w. f. 9 36 x Blue w.f. ¢ 13. Both these birds

were raised in Exp. 16.

Of the offspring 4 were white. One (69) was the blue form

without any white feathers, This is the only blue bird without

white feathers raised from the mating of two blues with white.

Five were blue with white feathers, and of these one (75) had

only 4 white feathers on the head, a very few on the thighs, and

a few white under tail-coverts. Another (78) had a few on the

rump, vent, and thighs, and some under tail-coverts. The other

three showed more white than those described in Exps. 24 & 25;

one of them haying 12 and another 11 white flights as well as

some white wing-feathers over them, whilst the third had 17 out

of 21 tail-feathers white, in addition to the white on the head,

neck, rump, vent, thighs, and under tail-coverts,

Series C.

BLUES MATED TO BLACKS.

Four different kinds of matings were made of blues and blacks.

Birds which did not contain white and also those in which

white was carried were used. For results of the matings see

Table IV.

Details of Exps. 27—31.

Exp. 27.—F.3 Blue 2 98 x Black Barb ¢ 100.

The 2 had no white feathers, and had previously been shown

not to contain white (v. Exp. 14). The ¢ was received from

Mr. Edwards of Exeter, who said that he believed it to have been

bred from blacks for four generations and before that from a

dun 2. The result of the mating was 8 blacks and 1 dun. No

white feathers were seen on any of the young birds. Very

slight indications of wing-bars were noticeable on some of the

blacks, the ground-colour being rather more sooty or smoky than

in the birds previously described, and so causing the bars to

stand out as a dead black. These bars were much more con-

spicuous in the next generation (v. Exp. 32). In the Dun (wv.

ante) the tail-bar was very obvious, being of a much darker shade

than the ground-colour. This bird was not bred from.

Exp. 28.—F.3 Blue 9 97 x F.3 Black w. f. ¢ 6.

The © 97 had no white feathers, and as already shown in

Exp. 10, when mated to a blue with white feathers gave blues

and blues with white feathers only.

‘The ¢ 6 had white feathers, and has been shown (Exp. 8) to

contain white.

The result gave 9 young, of which 5 were black and 4 blue.

Of the blacks three had some white feathers and two had not,

and of the blues two had white feathers and two had not. One

of the blues with white feathers died when a fortnight old, and

only one white feather could then be seen. The amount of white

in the birds showing this character varied very considerably.

INHERITANCE OF COLOUR IN PIGEONS.

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88 MR. R. S1APLES-BROWNE ON | Feb. 18,

Exp. 29,—F,4 Blue w. f. 9 15 x F. 3 Black d 109.

The 9 15 was raised in Exp, 11, and had previously been shown

to contain white (v. Exps. 19 & 22),

The ¢ 109, raised in Exp. 8, was black with no white. This

bird had not been previously bred from, but seeing that no whites

were produced in this experiment we may conclude that it did not

contain white.

The result of the mating gave 8 young, of which 4 were black

and 4 blue. Of the blacks two (Nos. 50 & 65) showed very few

white feathers at the vent. No white was seen anywhere else on

the plumage. Another black bird (No. 49) was rather light or

sooty in colour, as already described in Hxp. 27, and very slight

indications of a tail-bar were noticed. None of the blue birds

showed any white.

Exp. 30.—F.2 Blue 9 14 x Black w. f. ¢ 46.

Exp, 31._F, 2 Black w. f. 2 106 x F.3 Blue w. f. ¢ 13.

The @ 14 was raised in Exp. 3. The ¢ 46 was raised in

Exp. 12 from F.1 x white. Neither of these birds had been

previously bred from. This experiment, however, showed that

they both contained white.

The 2 106 was raised in Exp. 5, and has been shown in

Exp. 8 to contain white. The ¢ 13 was raised in Exp. 9, and

was shown in Exp. 11 to contain white. The total result of the

two matings was 2 Blacks, 5 Blues, 3 Blues with white feathers,

and 7 Whites. The blues from both matings were lighter in colour

than in the preceding experiments, the sooty colour of the wings

being of a much bluer shade than in the typical blue, and in

some cases being replaced by blue chequered with black. The

wing-bars were very distinct. Where white feathers were present

they were very few in number and were confined to the rump,

vent, and thighs.

Results of mating together Blacks from Hap. 27 and Blues

trom Hup. 30 respectively.

Exe. 32.—Black 9 12 x Black ¢ 1.

These two birds were raised in Exp. 27, and both were without

any white feathers. The plumage of both was somewhat smoky,

and slight traces of wing-bars could be discerned. From this

mating 12 birds were hatched. Of these, however, 4 died under

the age of one week. The colour of the plumage could not be

ascertained. The 8 birds reared consisted of 7 blacks and 1 blue.

Of the blacks, one is of a deep rich plumage showing no traces of

bars. Four are of a sooty-black colour with the wing-bars very

distinct. In one of these a tail-bar is also seen. The remainder

are sooty-black with no, or very slight, indications of bars. it is.

most probable that, if these birds were bred from, the sooty-

coloured specimens with wing-bars would be shown to contain

blue, whilst the deep black with no traces of wing-bars would

prove to be homozygous.

1908. | INHERITANCE OF COLOUR IN PIGEONS. 89

Exp. 33.—Blue chequered black 9 99 x Blue chequered black

with few white feathers ¢ 16.

These two birds raised in Exp. 30 were selected as showing the

lightest type with chequering. The wing-bars were very distinct.

3 16 had a few white feathers at the vent.

The mating produced six birds all of the chequered type with

clearly defined wing-bars. Of these four showed no white, and

the other two had a very few white feathers.

Bars-Nun Cross.

In this cross three matings only were made, and twenty-eight

young produced. The experiment was originally started, as

stated in the introduction, in order to cross the F.1 Barb-Nun

generation with the F.1 Barb-Fantail. Some further matings

were, however, made in order to investigate the inheritance of

the “shell” (v. infra), the results of which, together with some

other crosses in experiments on the same character, are described

in P.Z.S. 1905, vol. ii. p. 550. The Black Barb ¢ used in this

‘mating was obtained through the medium of a Poultry advertising

paper, and no details of its pedigree were obtainable. This bird

had white irides, the beak white tipped with black, the claws

white, the eye-wattles bright red.

Tue Non Picron. (Pigeon Coquille Hollandais.)

A curious structural character presented by this bird is the

“shell,” which is a tuft of reversed feathers standing up at the

back of the head, having an appearance somewhat like that of a

cockle-shell. After this point, attention is paid by breeders to its

colour and markings. It is a white bird with certain coloured

markings forming a very definite pattern. The markings are

found in several colours, of which black, blue, dun, red, and

yellow are the chief. By far the greatest number are black,

and this was the colour of the 2 used in these experiments.

The head, as far back as the “shell,” is black, but the coloured

feathers do not extend into the sheli. The chin and throat are

black, forming what is known as the “bib.” ‘Ten outer flight-

feathers on each wing should be black; in the specimen used,

however, there were only seven black flights in one wing and

eight in the other. There were also two black secondaries and

two black tertiaries, as well as a few black feathers at the carpal

joints. The tail is black, as also are the upper and under tail-

coverts. These markings should be very definite or “clean cut,”

and there should be no black on any other part of the plumage.

The @ used, however, had two black feathers on the back. The

black in the bird used was not a deep rich colour, as is found in

the Barb; and it is stated by breeders that many Black Nuns are

found whose colour is rather smoky. This was noticed in some of

the offspring of the cross. The irides are white, and there

appears to be no difficulty in breeding this character true in this

90 MR. R. STAPLES-BROWNE ON [ Feb. 18,

variety of pigeon. In the Black Nun the cere or eye-wattle is

blackish as are also the beak and claws. No details of the

ancestry of the 9 Nun were obtained. ‘The variety breeds true to

the markings and no self-coloured birds or whites are produced.

CROSS-BRED BrIrRDs.

Although this experiment was taken as far as the F. 2 genera-

tion, no blue birds were produced. The numbers, however, are

small, and it is possible that in further matings they might have

appeared. Blacks were raised in F. 2, but for want of space were

not tested. Three birds also appeared in F.2 which resembled

the Nun in markings, these are alluded to in Table V. as ‘‘ white

with afew blackfeathers.” These birds were not bred from. The

remaining birds were classed in three divisions as, ‘‘ blacks with a

few white feathers,” ‘“ mottled, with black in excess,” and ‘“ mottled,

with white in excess.” This classification, although useful for

purposes of description, is purely arbitrary, and no suggestion is

made that the birds differed gametically. It was very noticeable

that in the mottled birds the markings of the Nun were present,

and in addition black mottling occurred on those parts of the

plumage which are white inthe Nun. Full descriptions of the

cross-bred birds are given in the details of the experiments.

There appears to be no correlation between the presence of a shell

and the Nun markings.

Details of the Matings.

F.1 Generation.

Exp. 34.—Nun 2 (no number) x Black Barb ¢ (no number).

These birds were mated together for two years and twelve

young were produced. All were black with white feathers and in

all the black was in excess of the white. Two distinct types of

birds, however, were produced. One class consisted of black birds

with a few white feathers which appeared usually on the rump,

vent, and thighs, and on the neck or breast sometimes in the

position of the junction of the white of the breast with the black

of the “bib,” as described in the Nun pigeon. Seven of the F.1

birds were of this type. In the other class, of which there were

five, the birds presented a more mottled appearance, black how-

ever being in excess. The head, bib, and tail were always black.

The flight-feathers black with the exception of one or two, the

rump frequently white. The back and wing-coverts black mottled

with some white, the breast and abdomen white mottled with some

black feathers.

F. 2 Generation.

Exp. 35.—F.1 Black with few white feathers 2 8 x F.1 Black

with few white feathers ¢ 54.

Both these birds were raised in Exp. 34 and were of the first

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type described. They were mated together late in the breeding-

season, having both been previously used to cross with Barb-

Fantail F.1 crosses (v. Exps. 88 & 39).

Three young only were produced, of which one was black and

white mottled with the whitein excess. Unfortunately no details

of the markings of this bird were kept. The other two were black

with a few white feathers on the abdomen, and on the rump in

one of them.

Exp. 36.--F. 1 Black mottled with white 9 1 x Black mottled

with white ¢ 3.

These two birds raised in Exp. 34 were of the second type

described.

Thirteen young were produced :—

Black 2.

F. 1 types 8.

Birds resembling Nun 3.

Of the F. 1 types 5 were black with a few white feathers and 3

black mottled with white. Two of these, recorded as belonging to

the first type, died when about a week old in the nest. The others

showed only a few white feathers on the abdomen, rump, ard vent,

and one had three white flight-feathers and a few other white

feathers on the wings.

Tn those recorded as mottled the black was distinctly in excess

in one ease, and in the other two birds the amount of black and

white was about equal. The head, ‘“ bib,” and tail were black in

all cases, and several of the flight-feathers were black. Roughly

speaking, the back and wing-coverts showed more black feathers

than the breast, abdomen, and under parts. In one case (No. 17)

the feathers on the back of the neck and the right wing-coverts

were black edged with white, giving a “ pepper and salt ”

appearance to those parts. This peculiarity has not been noticed

in any other bird. The following are the details of the markings

of the three birds whose plumage somewhat resembled that of the

Nun pigeon :—

No. 7. White. Tail black. 2 black flights. A few black

feathers on the head. 4 black wing-coverts and one black

feather on the breast. The total amount of black was less

than in the Nun.

No. 11. White. Tail black. 7 black flights. A few black

feathers on the head. The wing-coverts were slightly

mottled, the greatest amount of black being in the

scapular region. A few black feathers on the breast and

rump.

No. 14. White. Tail black. 8 black flights. A few black

feathers on the head and at the carpal joint. In this bird

again the amount of black is less than in the Nun.

1908. ] INHERITANCE OF COLOUR IN PIGEONS. 93

Bars-FAantait-Nun Cross.

In this small series of experiments the F.1 Barb-Fantails

were mated to the F.1 Barb-Nuns. No reversionary blues

appeared. The forms produced are scheduled in Table V. Some

were definitely black, and the remainder showed combinations of

black and white in various proportions. The composition of these

latter was not investigated further, but it may be noted that one

bird (No. 70) agreed exactly with the F. 1 Nun-Fantails described

below. As would be expected, no whites appeared.

The further matings show that the extracted blacks in F. 2 breed

true. No blues were produced; and the black colour was rich

and deep in hue, so that no indications of wing- or tail-bars were

discernible.

An extracted black mated in Exp. 42 to an extracted white

from the Barb-Fantail Experiment gave offspring which were

indistinguishable from those produced in F.1 from a black Barb

and a white Fantail.

FE. 1 Barb-Fantails mated to F.1 Barb-Nuns.

Exp. 37.—F.1 Barb-Nun 9 18 x F.1 Barb-Fantail ¢ 68.

Exp. 38.—F. 1 Barb-Nun 92 8 x F.1 Barb-Fantail ¢ 71.

Exp, 39.—F.1 Barb-Fantail 2 63 x F.1 Barb-Nun ¢ 54.

All these birds were black with a few white feathers. The

Barb-Fantails were raised in Exp. 1, the Barb-Nuns in Exp. 34.

The Barb-Fantails 63 & 71 are the same as used in Exp. 3, and the

Barb-Nuns 8 & 54 are those mated together in Exp. 35.

From the three matings twenty young were produced :—

5 Blacks.

7 Black with few white feathers,

7 Mottled.

1 White with a few black feathers in the tail (No. 70).

The notes on the details of the markings of these birds are un-

fortunately very scanty. In the blacks with a few white feathers

the white appeared, when it was noted, on the rump and vent,

In one case there was a white flight, and in another there were

a few white tail-feathers. Of the seven mottled birds three are

recorded as having the white in excess of the black. It was noted

that the tail was black in the mottled birds, one bird, however,

had one white tail-feather. The head was generally black; the

flight-feathers varied, in some cases being all white, and in others

mostly black.

Testing the Extracted Blacks from the three foregoing

Haperiments.

Exp. 40.—Black BF. BN 92 53 x Black BF. BN ¢ 45.

Exp. 41.—Black BF. BN 2 65 x Black BF. BN ¢ 45.

2 53 was raised in Exp. 38, 2 65 in Exp. 37, and g 45 (used

in both matings) in Exp. 39.

Nine young birds were produced, all being black.

94 MR. R. STAPLES-BROWNE ON [Feb. 18,

Extracted Black mated to Extracted White.

Expr. 42.—F. 2 Barb-Fantail White 9 53 x Black BF. BN

3 45.

The extracted White in F. 2 of the Barb-Fantail experiments

was raised in Exp. 3, and has in Exp. 6 been shown to breed true

to white with a White Fantail. The extracted Black BF. BN is

the same used in the two preceding experiments. Seven young

were produced, one of which was black without white; the rest

were black showing white feathers varying in amount from a small

patch on the vent to a distinct mottling. In all cases, however,

the amount of black was in excess of the white.

Nun-FANTAIL CROSSBREDS.

No crosses were made in these experiments between the Nun

and the Fantail. Through the kindness of Miss Thiselton-Dyer,

however, two @ birds were received, the result of such a cross.

For the purpose of comparison with the birds produced in the

Barb-Fantail-Nun experiment a description of them is given.

These birds were white with some black feathers in the tails. One

had a tail consisting of fourteen feathers of which five were black.

The other had eleven tail-feathers of which three were black. A

few of the upper tail-coverts were also black in each bird, but

beyond these the plumage was quite white. The irides were black,

the beak and claws white, and the eye-wattle or cere flesh-colour,

slightly reddish in one. In view of the recessive behaviour of the

white from White Fantail elsewhere, it is perhaps remarkable that

these birds showed so little colour.

Wut1re TUMBLER-W HITE FANTAIL Cross.

This cross was made between two white strains of different

varieties of pigeons. The Tumbler was of a white strain which

is described below. The Fantail used was from the strain

kept and already described. Only four matings were made and

the experiment carried to the F. 3 generation, and only thirty-six

birds were produced. The numbers are scarcely sufficient to give

any quantitative results. It was, however, found that colour

was produced in the F.1 generation, and the reversionary blue

appeared in F. 2 in conjunction with red and white. These birds

are described as Tricolors. Further, in F.3 a bird was obtained

having one blue tail-feather with the terminal black bar.

The appearance of some coloured birds from these matings

suggests the possibility that in the White Tumbler a dominant

white factor, comparable with that known in fowls, may exist.=

Wuire Tumpier Piczon. (Culbutant.)

Nearly every variety of colour, shade, and marking existing in

domestic pigeons is found in the Tumbler. There are a few strains

1908. | INHERITANCE OF COLOUR IN PIGEONS. 95

of whites. The 2 bird used in this cross was obtained from

Mr. G. 8. Fayle of Birr, King’s Co. It was white without any

trace of coloured feathers, having the iris white, or “ pearl.”

It was also “ long-faced” and “ clean legged,” 7. ¢. free from

feathering. Mr. Fayle very kindly gave the following details

of his experience in breeding the strain, which has been in his

possession for over twenty years. The greatest difficulty in

breeding these birds is to obtain a white pigeon having a

white eye, as there is a very great tendency for white birds to

have dark eyes (v. infra). If, however, one or two coloured

feathers are present in the plumage the correct eye-colour is

more easily produced. Breeders are therefore in the habit of

occasionally introducing into their strain a splashed bird in order

to improve the eye-colour. Mr. Fayle introduced two hens

splashed with red about fifteen years ago, and used them for one

breeding-season only. Since then he has never introduced any

bird that to his knowledge was bred from other than white parents.

He believes, however, that there is hardly a strain of white

Tumblers in existence into which splashed birds have not, at some

period, been introduced. It was found that the strain occasionally

produced birds having a few coloured feathers, seldom more than

two or three, which were either black or red. These usually

appeared on the head or neck, and sometimes a secondary wing-

feather might be tipped with colour. ‘These coloured feathers

were frequently not reproduced at the moult. Birds showing these

coloured feathers were never selected for breeding. It was

further noted that there was a greater tendency for birds having

the desired white eye to produce splashed offspring than those

having part or the whole of the eye dark. The eye-wattle is

white as are also the beak and claws.

Typrs oF Cross-BRED BIRDS PRODUCED.

The birds produced from this series of experiments fell naturally

into the three classes of whites, whites with a few coloured

feathers or “splashes,” and “ Tricolors.” If a larger number of

birds had been produced, it is possible that there might have been

some overlapping between the two latter classes.

1. Whites.

Tn every mating of this series of experiments whites have been

produced. The number of whites bred was 17; and of the birds

showing coloured feathers 19. Owing to limitation of space,

extracted whites were not mated together.

2. Whites with a few coloured feathers.

The number of coloured feathers on birds described under this

class was very small, varying from two isolated feathers to a small

patch of about a dozen. In no case was there any approach to

mottling. The coloured feathers were in most cases on the neck

96 MR. R. STAPLES-BROWNE ON _[Feb. 18,

or scapular region, and occasionally on the rump or tail. The

colour generally found was red, but black, bluish black, and blue

were seen. In the few cases in which birds were kept a second

year, it was seen that the coloured feathers in one case did not

reappear after the moult, and there was also one instance of red

feathers being replaced by blackish ones.

3. ‘ Tricolor type.”

In the F.2 and F.3 generations birds were produced which

were red and white with some blue feathers. With one out-

standing exception (F.3; 9) birds of this type were fairly uniform

in marking. The head and upper part of the neck and throat

were white slightly mottled with red. Lower down, especially

on the back and sides of the neck, the amount of red increased

gradually, and at the root of the neck there were no white

feathers. The upper part of the breast was red with a slight

bluish tinge between the red and the white of the lower part of

the breast and abdomen, on which there were no coloured feathers.

Here the line of demarcation between the colour and the white

was very distinct and “clean cut.” The upper part of the back

and scapulars was red, generally of a lighter shade, and resolving

itself into a mere powdering, or the ‘“ strawberry” shade of the

fancier. This was continued slightly on to the proximal wing-

coverts where there was mottling with white. The external

wing-coverts and all the flight-feathers were white. The lower

part of the back below the origin of the wigs was white. The

rump was light blue, the shafts of the feathers being black. This

colour was carried on to the upper tail-coverts. The tail-feathers

were generally white, but in two cases some of the external

feathers showed a slight blue tinge with the shafts black. Very

slight indications of a terminal bar could be made out, especially

when the tail was viewed from the under surface. In most cases

one or both flanks were blue, the colour being carried down onto

the thighs. The rest of the under surface was white.

DETAILS OF THE MATINGS.

Colour.

The details which follow relate to the successive matings and to

the young produced respectively from them. The results are also

given in tabular form in Table VI.

F.1 Generation.

Exp. 43.—White Tumbler 2 9 x White Fantail ¢ 18.

Nine young were produced of which six were white, and three

showed a very few coloured feathers, as follows :—

No. 1. Two red feathers on the neck which were afterwards

moulted out.

1908. ] INHERITANCE OF COLOUR IN PIGEONS. 97

No. 5. A few feathers tinged red on the right scapular region.

This bird changed slightly during the moult. It then had

four blackish tinged feathers on the right scapular region,

the red colour having disappeared.

No. 7. Also had a few feathers tinged red on the scapular

region.

TABLE VI.

emead (OL aes Ge pa

2 1 ase cll OFFSPRING.

: Fee |

Exp. Be ame

| y z Ss | ey | White

No. | = 2 | with few] Tri-

ico) SS | White} coloured calor

gS S| feathers. :

| | !

43.) White Tumbler ...... 9) = || White Fantail ......:.. 18 — 6 3 —

| 1

B.1 White with few) -| 4. F.1 White with few 2 I}

44. coloured fthrs. y >, ae coloured fthrs. ) Be 4 4 2

| le. 2 White with few 2 || B. 2 White with few2,-) 4, ‘||

45. | coloured fthrs. § 13) 44 coloured fthrs. bas Be 4 5 ere

46. pHi) Bricolors sie: 0- 7| 44 || F.2 Tricolor ......... 8| 44 3 — 5

F. 2 Generation.

Exp. 44.F.1 White with few coloured feathers 9 5

x F.1 White with few coloured feathers ¢ 1.

These birds, raised in Exp. 43, are described above. Ten young

were produced, of which four were white, and six showed some

coloured feathers. These latter were divisible into two groups:

one, of which there were four birds produced, was white with a

few coloured feathers as in the F.1 generation; the other, of

which two birds were produced, was of the tricolor type,

previously described. Of the whites with a few coloured feathers,

No. 13 had one feather showing both red and black on the right

scapular, and one black feather on the rump.

No. 14 had two black feathers on the neck and one on the back.

No. 15 had two red feathers on the neck and three on the right

scapulars.

No. 18 had several red feathers on the scapulars.

The details of the two birds recorded as tricolors (Nos. 7 and 8)

coincide very closely with the description of the type already

given.

Proc. Zoou. Soc.—1908, No. VII. if

38 MR. R. STAPLES-BROWNE ON [ Feb. 18,

F. 3 Generation.

Exe. 45.—F.2 White with few coloured feathers 2 13

x F. 2 White with few coloured feathers ¢ 15.

These birds, raised in Exp. 44 and described above, produced

nine offspring when mated together, four of which were white,

and five white with some coloured feathers. Of those in the

latter class four showed a few red feathers on the back of the

neck, and in one instance on the left scapular. No black or blue

feathers were seen on these birds. The fifth (No. 112), however,

had no coloured feathers on the neck, but on the right wing-

coverts was a large patch of feathers tinged red and edged with

black, also one tail-feather blue with a terminal bar, the rest of

the tail and plumage being white.

Exp, 46.—F. 2 Tricolor 9 7 x F.2 Tricolor ¢ 8.

These two birds raised in Exp. 44, and of the type described,

produced eight offspring, three of which were white and five

tricolors. The details of four of the tricolors approximate closely

to the type. In two of them blue was present on the breast,

flanks, and rump. On one it was present on the flanks only, and

in another there were only slight indications of blue on the rump.

In the bird showing most blue there was a slight blue tinge

on some of the tail-feathers. The fifth coloured bird (No. 9),

produced from this mating, showed, in addition to the typical red

on the neck and breast, a large amount of bluish-black colour.

The left scapulars and proximal wing-coverts were light reddish

feathers edged with black, and blue feathers chequered with black

were interspersed among them. The right scapulars were blue

chequered with black. The remainder of the plumage, including

the under parts, was white with the exception of a single blue

feather on the rump.

TRIDES.

The irides of pigeons are chiefly of three colours, white, orange,

and black. On each the blood-vessels of the iris can be seen

very distinctly and, in some cases, give a very well-marked red

appearance most noticeable at the periphery.

The white, or, as it is called by fanciers, the “‘ pearl” eye breeds.

true in many varieties of pigeons, but in some it is apt to throw

orange- and black-eyed birds. The latter is stated to be more

easily “bred out” from a strain than the former. The orange

iris is found in Columba livia, and appears to breed true in

several fancy varieties. The black iris (termed by fanciers

“hazel” or “bull” eye, which is, more correctly speaking, a

brownish-black), seems to breed true invariably. In the nest the

irides of all the young pigeons examined were black, but in birds

in which this was not the eye-colour of the adult, the colour

1908. | INHERITANCE OF COLOUR IN PIGEONS. 99

changed to white or orange usually within two months after

hatching. In the foregoing experiments no matings were made

specially to test the inheritance of the colour of the iris, and this

character has only been studied incidentally. From lack of space

it was found necessary to kill many of the young birds before the

colour of the iris in the adult state could be noted; and in the

earliest experiments no notes were taken of the minor characters

presented.

Some few cases were met with in which the colour of the iris

did not completely change, but a small segment of one iris re-

mained permanently black, or the greater part. of both irides was

black and only a small part showed white or orange. One bird

was bred having one iris black and the other orange, but this

appeared to be a very exceptional case.

The Barb pigeon has generally a white eye, although black

Barbs are sometimes seen with orange eyes. White Barbs have

been seen with white eyes, but nearly always have black. Of the

four black Barbs used, those in Exps. 2, 27, and 34 had white

irides. That used in Exp. | is believed to have had an orange

iris, the notes of this experiment referring to plumage colour only.

The white Fantails had black irides; coloured Fantails, on the

other hand, have either white or orange. The black and white

Nun had white irides. The white Tumbler also had white irides ;

this character has been specially referred to in the description of

the Tumbler.

In the four series of experiments irides were obtained as

follows :—

Barb-Fantail Cross gave white, orange, and black.

Barb-Nun Cross gave white only.

Barb-Fantail-Nun Cross gave white, orange, and black.

Fantail-Tumbler Cross gave black only.

There appears to be a very distinct association between white

plumage and black iris, and, in a lesser degree, between black

plumage and white iris, as the following tables show (p. 100).

Here the birds are grouped irrespective of the generations to

which they belong.

In the case of the blue birds with some white feathers there

appeared to be a general, though not invariable, rule that the

birds showing the greatest amount of white had black irides and

those with fewer white feathers orange irides.

The tables show very conclusively the relation between the

white plumage and black irides. This correlation extends to the

whites with some coloured feathers, and even to the mottled birds

in which white is in excess. It is further seen that no black bird

has a black iris, and the black iris is also exceptional in black

birds with some white feathers. In the Barb-Fantail cross there

appears to be a correlation between the white iris and black

plumage; the figures, however, of the Barb-Fantail-Nun cross,

although small, do not bear this out.

100 MR. R. STAPLES-BROWNE ON [Feb. 18,

The doubtful record of the iris of the Barb used in Exp. 1,

and the absence of notes on the irides in some of the earlier

experiments, makes the tracing of inheritance of this character

very unsatisfactory.

Barb-Fantail Cross.

White. | Orange, Black.

[ Black ........sses sees ses eee ees sees eee eteee cee 15 | 1 0)

| Black with some some white feathers...... 18 6 2

1) DCL aRn ee ee aaa eee A MERC ont Rta ne oe $) 13 2

Blue with some white feathers ............... 10 | 20 20

IWIRIHES Meiwekide nee terse See eae ess. Seen Pe 0) 0) 48

White with some coloured feathers......... | 0) | 0 11

18.10 U9 Manno tadorcosa agdcdeeacdbaeasdy sconce snc a on: | 5 | 0 0)

Dun | 1 10) 0

Barb-Fantail-Nun Cross.

IBY EO g™ Laer aaneuatanpenicee ai anne sone eacaNcadacsmr and: 3 8) 0)

| Black with few white feathers ............... 6 3 1

| Mottled, black im excess. ............-....005. / 2 | 1 0

Mettled, white im excess..-.................005. (0) | (0) 3 |

White with few coloured feathers ......... 0) 0 | il |

A very clear result is, however, obtained in Exp. 2, which was

made later. In this case a Barb with a white iris was mated to

a Fantail with a black iris, and the four young produced had

white irides. Two of these mated together in Exp. 4 gave

13 birds with white irides, and 4 with black. Three other young

were produced, but were killed before the colour of the iris could

be ascertained. This, therefore, gives the simple 3: 1 Mendelian

proportion in F. 2, white being dominant.

The remaining crosses, both of the Barb-Fantail and the Barb-

Fantail-Nun experiments were, unfortunately, all made with birds

descended from the Barb used in Exp. 1.

The results of these experiments are grouped together in the

following tables irrespective of the generations to which the birds

belonged.

1908, | INHERITANCE OF COLOUR IN PIGEONS. 101

Barb-Fantail Cross.

Colour of irides of parents. | White. Orange. | Black.

Orange X Orange ysl ee | 0) 21 8

IB EVelie 6 LBIBNGLS a5 donood on ont aooeae 0 0) 14

| White x Black ttc 7 0 2

Oramcce@ lac kareena erro ere 4, 9 26

| White X Orange .........0..c00.. | 25 6 | 17

White X White 2h all 6 (0) 2 |

Orange X Orange 2 2 0) |

Orange lackieeee selene 0) 3 1 |

| White X Orange ...............:.. 3 8 2 |

In connection with the above tables it may be noted that in

the matings of orange x black irides, the excess of blacks over

oranges in the offspring ‘is in some measure associated with the

excess of white-plumaged birds over blues in Exps. 16-23 where

equality was expected. Further, in the White x Orange ex-

periments, all the offspring from two matings, amounting to

12 birds, had white irides.

The bird already mentioned as having one iris black and the

other orange was raised from 9 with black irides x ¢ with

orange. This bird when mated to a bird with black irides gave

5 young all with black irides.

In the Barb-Nun experiment 21 birds were reared to an age

at which the adult colouring of the iris showed itself. In all

these foregoing birds the irides were white. In the two Nun-

Fantails described the irides were black. The Tumbler-Fantail

cross is exceedingly interesting in respect of the irides. As has

been stated, white irides cannot be kept in a strain of white

Tumblers without the occasional introduction of birds having

some coloured feathers. In the cross between the white-eyed

Tumbler and the black-eyed Fantail, 36 birds were raised. Of

these 35 had black irides, and one had a small part of the left iris

white, the remainder being black. This was a splashed bird in

the F. 2 generation (Exp. 44, No. 14).

The foregoing facts as far as they go suggest the conclusion

that the black iris is correlated with the white plumage, and that

102 MR. R. STAPLES-BROWNE ON [ Feb. 18,

possibly there may also be, although in a much lesser degree, some

correlation between the white iris and the black plumage. The

figures respecting blue birds are not so conclusive, but seeing that

the orange iris 1s the only one found in C. livia, the suggestion

may be hazarded that there is possibly some correlation between

the orange iris and blue plumage. The matter becomes more

complex when we study the blue and black birds in the plumage

of which white feathers occur. It is, however, suggested that the

amount of white present in the plumage may have some influence

on the determination of the colour of the iris. Whether the

association of certain eye-colours with certain types of plumage-

colour arises through gametic coupling or not cannot yet be

positively asserted.

With regard to the question of dominance, it has already been

shown that white is a simple dominant to black in the case where

the record of the irides of the original parents was kept, and a

Mendelian ratio of 3:1 was obtained in F. 2. In the table of

the Barb-Fantail-Nun cross also, a 3:1 ratio was given when

birds: having white irides were mated together. It is further

shown that extracted black irides breed true without exception.

Further experiments are necessary before the relation of

orange to black, and white to orange can be definitely asserted.

It appears probable, however, that black will be eventually found

to be recessive to orange, and that orange may be recessive to

white.

BEAKS AND CLAWS,

In the Barbs used the beaks were either white tipped or tinged

with black, or were horn-colour. The claws also were horn-colour.

In the white Fantails the beaks and claws were white; they were

also white in the white Tumbler, and in the Nun they were

black. In the Barb-Fantail and Barb-Nun-Fantail crosses the

following types of beaks and claws were met with :—(1) Birds

having quite black beaks; in these the claws were usually black,

sometimes white, and sometimes mixed, some of the claws of an

individual being black and others white. (2) Birds having white

beaks with some dark pigment. This might be a white beak

tipped with black, or one mandible might be black and the other

white, or the beak might be’of a general horn-colour. In these

birds the claws were usually mixed, but some individuals were

produced having all the claws black, white, or horn. (3) Birds

having white beaks; these invariably had white claws. [One

bird only is recorded as having a white beak and black claws, it

was black in plumage, and was killed when only just over a fortnight

old. The record is probably erroneous, and had the bird reached

maturity it would have been found to have a white beak tipped

with black.] There is a very marked correlation between the

colour of the beak and claws and that of the plumage. White

beaks and claws have been found on every white-plumaged bird

bred, also on whites with some coloured feathers, and reds. Black

1908. | INHERITANCE OF COLOUR IN PIGEONS. 103

and Blue birds with or without white feathers, on the other hand,

have beaks of types 1 or 2, there being some black present in the

beaks. Five exceptions, however, occurred ; these were blue birds

with a large amount of white in the plumage, having white beaks

and claws. In the Barb-Nun cross the beaks and claws of the

birds in the F. 1 generation were black. In F.2 four birds were

produced having white beaks tipped with black and some white

claws. The remaining birds of the F. 2 generation had beaks and

claws black. In the Tumbler-Fantail cross the beaks were white

with the exception of a bird in F, 2 (No. 14, Exp. 44), which had

the lower mandible tinged blackish. The claws were white

throughout. The general conclusion is that pigment in the beak,

and to some extent in the claws, is correlated with certain types

of plumage. Acting thus it is allelomorphic to white, and is a

simple Mendelian dominant.

EYE-WATTLES OR CERES.

In the Barb the eye-wattle is large. It increases in size with

age, and has been seen measuring one inch in diameter. It

presents the appearance of a series of naked nodules of skin

arranged in two or three concentric rows. In colour it is bright

red. In the three other varieties used in these crosses the wattles

are very small. In colour those of the white Fantail and white

Tumbler are white or pale flesh-coloured, that of the Nun blackish.

In size the wattles of the F.1 generation from a Barb were

intermediate ; some large wattles were observed in the F.2 and

subsequent generations, but these never assumed the proportions

of those of the pure Barb. It was noticed that these large

wattles were always red, but they were present on birds having

black, blue, or white plumage.

Observations on the colour of the wattles of birds in the Barb-

Fantail crosses have been somewhat complicated by the fact that

many birds, which were subsequently seen to have red wattles,

when young showed wattles of a yellowish colour which were

scarcely distinguishable from the flesh-coloured wattles of the

Fantail. The general result, however, appears to be that red is a

simple dominant over flesh-colour. In the F.2 generation one

blackish wattle was obtained, and some others were found in the

further generations. In one experiment birds possessing these

wattles were mated together, with the result that offspring

showing all these kinds of wattles were produced. Occasionally

birds with wattles coloured partly red and partly black, or partly

white and partly black, were produced. Whether these wattles

would have changed their colour later in life cannot be stated.

The colour of the wattles in the Barb-Nun cross is not very

clear. Birds of the F. 1 generation had blackish wattles with, in

one or two cases, a little red at the periphery. In F.2 some birds

with red wattles were obtained, two with white wattles, and some

with mixed wattles. Little reliance, however, can be placed on

104 MR. OLDFIELD THOMAS ON MAMMALS [Feb. 18,

this result as the experiment was very small and the young birds

were killed before they reached maturity. In the Barb-Fantail-

Nun crosses red, yellow, black, and mixed red and black wattles

were obtained. ‘The yellows might possibly have changed to red.

No flesh-coloured wattles were recorded. The two Nun-Fantails,

however, had flesh-coloured wattles. In the Tumbler-Fantail

experiment the wattles were flesh-coloured throughout.

Owing to the various changes occurring in this character during

the life of an individual, it is by no means a satisfactory subject

for experiment.

CoNncCLUSION.

The experiments here recorded have been subsidised by the

Government Grant Committee of the Royal Society. The writer

is indebted to Mr. J. H. Elwell for much kind assistance.

The matings, throughout, have been made in consultation with

Mr. Bateson, who has most kindly supervised the experiments.

He has also read the manuscript for the present report, and made

many very valuable suggestions and alterations. To him the

writer desires to express his sincere thanks.

EXPLANATION OF THE PLATES.

Black Barb-White Fantail experiment.

Prate lV. Fig. 1. Black with some white feathers.

Fig. 2. White with black patch. (Exp. 5.)

Pratt V. Fig. 1. Reversionary blue.

Fig. 2. Reversionary blue, dark type. (Exp. 5.)

Pratt VI. Fig. 1. Blue with some white feathers.

White Tumbler-White Fantail experiment.

Pratr VI. Fig. 2. White with few coloured feathers (black).

Puate VII. Fig. 1. White with few coloured feathers (red).

Fig. 2. Tricolor.

2. The Duke of Bedford’s Zoological Exploration in Eastern

Asia—IX. List of Mammals from the Mongolian

Plateau. By Otprretp THomas, F.R.S., F.Z.S.

[Received January 21, 1908. ]

After making the collection in the Shantung Peninsula re-

ferred to in a previous part of the present series, Mr. Malcolm

Anderson made a trip to the Mongolian Plateau, reaching a point

about 100 miles N.W. of Kalgan, and collected there the series

now enumerated.

The fauna of this region, as was pointed out by Pere David, is

exceedingly poor, and Mr. Anderson was in consequence only able

to get nine species, but these are all of interest, and form a

valuable nucleus for further work in Northern China. Most of

1908. ] FROM THE MONGOLIAN PLATEAU. 105

them are represented by excellent series of the perfectly prepared

skins to which Mr. Anderson has now accustomed us, skins of a

very different character to any on which work in Hastern Asia

has hitherto had to be done.

Mr. Anderson’s notes on the trip are as follows :—

“On the 16th July, 1907, I left Kalgan (Jang-kia-kou), in

North-western Chih-li Province, for the Mongolian Plateau. Some

twelve miles north-west of Kalgan we began the ascent of the

escarpment, and about 18 miles from that city found ourselves at

the summit of the range of mountains which, in this part, borders

the plateau. From here we descended some hundreds of feet to

reach the general level of the tableland. We travelled two days

over the plateau, till, on the evening of the second, we reached

Taboul (Five Hills), at a point 100 miles north-west by north

from Kalgan. Here, at an elevation of approximately 5000 feet,

the country is of rolling hills with only occasional level stretches.

There is not a tree, nor even a bush, in the region, but the hills

bear abundant grass, which makes this district the best pasture-

land in Mongolia. It is, indeed, the district from which the

Chinese Government draws its supply of cavalry-horses. Sheep,

goats, kine, camels, and horses are raised by the natives, who

have no other means of livelihood. |

“Tn general the rainfall is meagre, but it chanced that during

my stay, between July 18th and August 13th, there were almost

daily rains, which often came in the shape of cloud-bursts,

coming up in the south-west and disappearing in the north-east.

Heavy dews fall nightly. I was informed that the winters are

cold with piercing winds, but the snowfall is not great.”’—/. P. A.

1. VULPES sp.

3. 1487.

Too young for determination.

‘“‘ A family of foxes was in the Taboul neighbourhood when I

went there, but they were evidently very shy, for on seeing us

one day they forsook the place and did not return.”—J/. P. A.

2. CrrELLUS MonGoLiIcUS M.-Edw.

3g. 1475, 1477, 1493, 1499, 1500, 1502, 1516.

@. 1479, 1483, 1501, 1508, 1504, 1510, 1533.

These specimens are certainly referable to C. mongolicus,

whether that animal is or is not synonymous with C. brevicauda

Brandt, as has been asserted.

Mr. Campbell also obtained some examples of the species at

Hara-ussu, not far from the present locality.

The Old-World Citelli are very unsatisfactorily known, and

this plentiful series will be of much assistance in working out the

group.

“Very common; diurnal.

“On our trip into Mongolia, both going and returning, I saw

great numbers of these animals along the road. Particularly in

106 MR. OLDFIELD THOMAS ON MAMMALS [ Feb. 18,

one locality, about 70 miles from Kalgan, they have a large

colony, and there the burrows are about the sides of mounds

overgrown with bush-grass. At Taboul I found them living

about the hill-sides, and their burrows not distinguished from

those of other mammals.”—W/, P. A.

3. MERIONES UNGUICULATUS M.-Hdw.

3. 1473, 1478, 1485, 1486, 1489, 1492, 1495, 1506, 1507,

1514, 1539.

@. 1464, 1469, 1470, 1471, 1474, 1496, 1505, 1511, 1512.

A female in spirit, with 2—2=8 mamme.

These specimens quite agree with a typical specimen in the

Museum from “Chinese Mongolia” received from the Paris

Museum. None of them shows any approximation in the

character of the claws to an example of J/. psammophilus

M.-Edw., which was collected in the near neighbourhood of

Kalgan, but which Mr. Anderson did not chance upon.

“This, the most abundant mammal, was literally almost every-

where throughout this part of Mongolia. They are diurnal to

some extent, but may be most frequently seen between sunset

and dark, when they sit spermophile-like before their burrows.

I frequently succeeded in approaching within about eight feet of

a sitting individual, during which manceuvre the animal would

eye me steadily and, finally, with one rapid move, plunge into

his hole, but reappear after a few moments if I remained per-

fectly still, These animals make a curious sound beneath the

earth ; it sounds ver y much like the distant galloping of a horse

on a hard road, and I was much puzzled about it for some days.

How the sound is produced I do not know.”—JV/, P. A.

4. Mus WAGNERI MONGOLIUM, subsp. n.

6. 1472, 1520. @. 1480, 1481, 1484, 1532.

A white-bellied Mouse of the mzsculus group, not so pale as

true wagner.

General colour above rather paler than idgway’s ‘* broccoli-

brown,” the light rings on the hairs below ‘ écru-drab” and

pinkish buff; sides rather paler than back, but not approaching to

clear buffy of true wa gneri. Whole of under surface, hands, and

feet pure white, the hairs white to their bases; line of demareca-

tion on sides very sharply defined. Ears like head, their proectote

little darker. Tail rather short, inconspicuously bicolor, brown

above, dull whitish on sides and melons

Dimensions of the ae SMESELGL in the flesh :—

Head and body 82 mm.; tail 50 (range up to 56); hind foot

15:5; ear 12:5.

Skull—ereatest ee 5min.; basilar ae 17 ; zygomatic

breadth 11: 3; nasals 7-9 Soalealee length 9:5; palatal foramina

4:7; length of upper molar series 3°3.

Type. Old female. B.M. No. 8.3.5.36. Original number 1484.

Collected 28th July, 1907.

1908. | FROM THE MONGOLIAN PLATEAU. LOT.

Accepting Dr. Biichner’s identification of the pale Central-

Asian White-bellied House-Mouse as MW. wagneri, I feel compelled

to give a racial name to its representative in Eastern Mongolia

and China, on account of its much darker colour, which is quite

uniform in all the specimens obtained by Mr. Anderson. Of the

true JZ. wagneri we have for comparison two specimens from the

Prjewalski collections, received from the St. Petersburg Museum.

‘“‘ Found only about the tents of the Mongol village where I

lived. They were very bold, but did not do much damage.”—

Wik Jie, Zale

5. CRICETULUS GRISEUS OBSCURUS M.-Edw.

3. 1513, 1522, 1523, 1529, 1530, 1536, 1537, 1545.

@. 1518, 1519, 1524, 1528, 1538, 1543, 1544.

@ in spirit.

These specimens, all in summer pelage, differ from the Chefoo

series of griseus, which are in winter dress, by being slightly

darker in tone, as compared with the very grey colour of the

latter animals. Whether this difference in colour is solely due

to season remains to be seen when further specimens representing

other seasons are available for examination

In addition, the Mongolian specimens seem to have on the

average rather larger teeth, but the difference is not quite

constant.

Milne-Edwards’s Cricetulus obscurus was described from

Sartchy, some 200 miles S.W. of the present locality, and I

think Mr. Anderson’s specimens may be provisionally referred to

it, and that it might be considered as a subspecies of C. griseus.

6. CRICETULUS CAMPBELLI Thos.*

3S. 1465, 1466, 1468, 1498, 1525, 1527, 1534, 1535, 1546.

@. 1467, 1482, 1497, 1526, 1547, 1548.

Although there are certain discrepancies between the measure-

ments of these specimens, as taken in the flesh by Mr, Anderson,

and those recorded by me from Mr. Campbell’s spirit-specimens,

the agreement in all other respects is too exact, and the localities

are too close to each other, for there to be any genuine racial

difference between the two. Probably variations in the method

of measuring the minute tail, and the contracting effect of

alcohol on the ears would account for such differences as exist.

The beautifully marked Hamsters of this group have hitherto

been exceedingly rare in collections, and the nice series obtained

by Mr. Anderson is of much value.

The following are the flesh-measurements of two old examples :-—

g. Head and body 88 mm.; tail 14; hind foot 12; ear 13.

2 : 7 ” 90 oD ? 11 > oy) ” 12; ”? 14.

* Ann. Mag. N. H. (7) xv. p. 322 (1905). The position of the type locality was

incorrectly given in this description, owing to a confusion between two similar

names. ‘The proper position is about 42° 40’ N., 116° 20’ KE.

108 MR. OLDFIELD THOMAS ON MAMMALS [ Feb. 18,

“‘T did not discover much difference in the habits of these two

Hamsters, but I believe that the long-tailed form (C. griseus)

burrows for itself, while the short-tailed one (C. campbell) is

more inclined, at least in this region, to take possession of holes

of the Red-tailed Rat (eriones). Both species live on the seeds

and leaves of small plants, among which they can often be seen

running about in the lateevening. Both species are abundant.” —

Mi eae

slLe oe ale

7. MIcROTUS ANGUSTUS, Sp. 0.

$ (old). 1517. Mongolian plateau. Alt. 5000’. 4th August,

1907. B.M. No. 8.3.5.63. Type.

A large pale species of the “* Stenocranius” group

Size about as in I/, (Stenocranius) slowzowi Kastch. General

colour pale isabella, resulting from a coarsely lined mixture of

dull buffy and brown. Sides clearer buffy ; a marked buff patch,

probably glandular in nature, just in front of the hips. Hairs of

under surface cream-buff terminally, the grey bases to the hairs

showing through. Sides of iruzzles and tufts at bases of ears

rich buffy; the head otherwise like the body. ars rather short,

not projecting above the (summer) fur, the short hairs clothing

its edges pale buffy. Hands and feet uniformly dull buffy above ;

pollex with a small nail. Tail well clothed, dull buffy, with a

narrow rather darker line along its upper surface. Skull of the

extreme ‘“ Stenocranius” type, very long and narrow, as shown by

the measurements given below. Palatal foramina rather short.

Posterior palatal pits deep, the septum between them very

narrow. Opening of posterior nares narrow, angular. Bulle of

average size.

Teeth of the usual arvalis type, m* with four spaces and a

posterior crescent. M, with six spaces and an anterior trefoil.

Dimensions of the type, measured in the flesh :—

Head and body 112 mm.; tail 32; hind foot 17; ear 105.

Skull—egreatest length 26°8 mm. ; basilar length 24; zygomatic

breadth 12°9; nasals 7 x 3:1; interorbital breadth ae sper ey

breadth 11; palatilar length 13°8; palatal foramina 5:2; length

of upper tooth-series (crowns) 5°7

Hab. and type as above.

This Vole is an extreme member of the narrow-headed group

called Stenocranius by Kastchenko, but I fail to identify the

species with any hitherto described. Radde’s Arvicola mongolicus

isa dark-coloured species (‘‘dorso fusco ”), Microtus raddei and

M. gregalis are smaller, while I. slowzowi and MW. transhanicus, to

which the dimensions most nearly relate it, are from ieee so

widely distant, with other species intervening, that it cannot

possibly be either of them.

“Probably rare. Caught by hand among grass and weeds in the

middle of the afternoon. No other specimens could be found,

and there were no burrows about.” —J/, P. A.

1908. | FROM THE MONGOLIAN PLATEAU, 109

[ALLACTAGA MoNGOLICA Radde,

Dipus jaculus, var. mongolica Radde, Reise §.-O, Sib., Sug.

p. 170 (1862).

Dipus (Jaculus) annulatus M.-EKdw. Rech. Mamm, p. 149

(1868-74).

Q. Hara-Ussu, N.W. of Kalgan, C. W. Campbell.

The specimen obtained by Mr. Campbell at Hara-Ussu nearly

resembles a co-type of A. annulata received from the Paris

Museum, and collected by Pére David on the “ plateaux sablon-

neux de Mongolie,” no doubt not very far from the present

locality. But the animal would seem to be referable to the

earlier-published A. mongolica, described by Radde from the

northern edge of the Mongolian plateau. |

8, Lepus TOLAI Pall.

go. 1462. 9. 1463, 1490.

In 1898 two specimens of this same Hare were obtained by

Mr. C. W. Campbell at Hara-Ussu. All are in summer pelage,

so that it is not easy at present to define their differences from the

Chefoo and Peking Hare, L. swinhoei Thos., which Mr, Anderson

collected in winter and early spring.

“Common in certain localities. The Hare in Mongolia is

rather strangely fond of the proximity of the native encampments.

The people say that they come near the camps for protection from

wolves, and this seems probable, for the Hares have nothing to

fear from the people, while the Mongol dogs, though fierce and

strong enough to make even a wolf think twice, are probably

seldom swift enough to catch a Hare, and, being used merely as

watch-dogs, they are kept close to the tents.” —M/. P. A.

9, OGHOTONA DAUURICA Pall.

Lepus dawuricus & L. ogotona Pall.

3. 1488, 1491, 1508, 1515, 1540.

®. 1476, 1509, 1521, 1531, 1541, 1542,

These specimens agree in all essential characters with the

specimen from Urga, N.W. Mongolia, figured by Biichner as

representing Pallas’s species. With the exception of a faded

dealer’s skin from “ Amurland,’ they are the first examples of

this species that the Museum has received.

There can be no doubt that Pallas’s specific names ogotona and

dawuricus both belong to the same animal, as explained by

Biichner ; and Bonhote’s recent differing from the latter * is due

to his having accepted as correct an old determination of the

Museum specimen No. 45.4.21.5, which was bought from the

dealer Brandt under the name of Lagomys ogotona, and said to

come from ‘“ Asiatic Russia—Kirgisen.”

But this specimen, which is certainly not ogotona (=dauurica),

is the type of Ogotoma pallasii, a name given by Gray to Water-

* P. ZS. 1904, ii. p. 216.

110 MR. G. T. BETHUNE-BAKER ON NEW [ Feb. 18,

house’s Lagomys ogotoma, the description of which was based on

this specimen, while the skull characters of the genus Ogotoma, as

given by Gray, were also drawn up from it. And it has been

again redescribed by Bonhote as Ochotona ogotona, and stated to

be nearly related to O. ladacensis Ginth.

The name O. pallasi and the descriptions above quoted will

therefore have to be reckoned with by future writers about this

difficult group.

“Not common—somewhat diurnal. Frequents the hill-sides

where grows a stiff fragrant weed on which it seems to feed. In

several old fox-burrows I found great masses of cuttings of this

weed, and in that immediate vicinity trapped five of the series.

Their burrows are not clean-cut and vary greatly in diameter.

Where several of these animals live near together their holes

are connected on the surface by a network of little trails. The

presence of these animals is generally betrayed by little piles of

spherical droppings at the mouth of the burrow, indicating also

that the occupants are cleanly in habit.”—V/. P, A.

3. Descriptions of new Species of Butterflies of the Division

Rhopalocera from Africa and from New Guinea. By

G. T. Bernunt-Baxer, F.L.S., F.Z.8.

[Received January 21, 1908. |

(Plates VIII. & IX.*)

ACREA BENI, Sp. n.

3g. Both wings bright orange-red. Primaries with termen

and apical half of costa linear, black, with all the terminal parts

of the veins outlined finely with black, tapering finer basewards ;

a black spot in the cell, another at the end of the cell, beyond

this three subcostal confluent small spots with a larger one shifted

outwards between veins 4 and 5 and a smaller spot below it

between 3 and 4, a spot below the end of the cell, two between

veins 1 and 2, one near the base and one near the termen.

Secondaries with costa narrowly black, with a subterminal scal-

loped line confluent with the costa along the veins, terminal part

of veins slightly marked with black, base irregularly restrictedly

black; a series of three subbasal spots, that in the cell shifted

slightly outwards, a spot at the upper end of the cell; a curved

series of postmedial spots, that near the angle of vein 2 shifted

well inwards with one below it shifted well outwards.— Under

side. Primaries as above, but paler. Secondaries ochreous, with

pink internervular stripes; the spots as above, but smaller, and

the base broken up into five or six spots instead of being all

confluent as above.

@. Both wings dirty brown, with smaller spots and no basal

spot in the fold of the primaries. Secondaries with the black -

* For explanation of the Plates, see p. 126.

JEL VAS Sq IESHOXS) Teil WANN:

10 16

E.C. Knight ad nat.del. West, Newman chromo.

NEW OR LITTLE KNOWN LYCAINIDA.

Je, Avs ESOS), IIDC

West, Newman chromo.

TLE KNOWN LYCA NIDA.

E.C. Knight ad nat.del

NEW OR LI

1908. | SPECIES OF RHOPALOCERAN BUTTERFLIES. 111

termen heavily accentuated. Under side like the male, but dirty

ochreous.

Expanse, ¢ 52, 9 56 mm.

Hab. Angola.

Types in my collection.

PENTILA MULTIPLAGATA, Sp. 0.

Thorax black; abdomen pale tawny.

Both wings tawny, with somewhat scalloped very broad black

outer margins. Primaries with costa narrowly black, with two

blackish spots over the cell, a larger one at the end of the cell,

and three postmedial spots in a curve from vein | to 3, one

between each vein. Secondaries with a spot closing the cell, and

a postmedial series of seven spots following the course of the

black termen, the lower three spots being more basewards than

the upper four.— Under side. Primaries tawny, with apical half of

termen yellowish. Three spots over the cell, one closing it and

another spot between veins | and 2 near the angle; a postmedial

series of eight spots beginning at and above the third costal spot ;

a subterminal series of six spots, the apical three being broad

dashes, the tornal three being largish spots. Secondaries paler

than primaries; a curved series of five basal spots, the one on the

costa and that in the cell very small, the three lower curved ones

large, ending below the angle of vein 3; a postmedial series of

nine large spots, extending around the greater portion of the

wing; a subterminal series of seven large spots.

Expanse 40 mm.

Hab. Makala, Congo Free State.

In the Powell-Cotton collection.

This species belongs to the P. amenaida Hew. group.

CITRINOPHILA UNIPUNCTATA, Sp. 0.

2. Both wings lemon-yellow. Primaries with apical area

broadly black in a fairly even curve internally, except that there

is a slight break at vein 4, tapering down to about vein 2, where

it ends. Secondaries with termen very finely black, frequently

interrupted by the yellow ground.—Under side. Both the wings

paler lemon-yellow, with a single black spot at the end of the cell

in each. Primaries with the black apex showing slightly through,

the costa and termen finely dotted with black all round. Secon-

daries with termen finely dotted with black at the veins.

Expanse 34 mm.

Hab. Makala, Congo Free State: March.

In the Powell-Cotton collection.

DEUDORYX MAKALA, sp. n. (Plate IX. fig. 4.)

3. Both wings brilliant metallic bluish green, of a tone of

colour not uncommon in the Neotropical Thecle, but very unusual

in the Western Hemisphere. Primaries with costa narrowly

black ; apex broadly black, tapering narrowly to about vein 2.

112 MR, G, T. BETHUNE-BAKER ON NEW [ Feb. 18,

Secondaries with costa blackish to vein 7, a small round creamy sex-

spot on the upper margin of the cell; termen with a fine linear

black line with white fringes.—Under side. Both wings white.

Primaries with a very broad olive-brown termen, having a fine

intersecting white line from the tornus to vein 4 or 5, and a second

short fine white intersecting external dash from the tornus to

about vein 2; a very broad, olive-brown, waved tapering post-

medial band immediately beyond the cell right across the wing,

a small pencil of black hairs a third from the base of the inner

margin. Secondaries: apex with a short apical olive-brown dash

and a very broad olive-brown subterminal band from just beyond

the apex to the anal angle, the anal portion largely suffused with

orange-red, a deep black oval spot between veins 2 and 3, with a

smaller black lobe-spot somewhat suffused with very pale bluish-

metallic scales ; sex-spot showing through olive-brown,

Expanse 52-36 mm.

Hab. Makala, Congo Free State: June.

In my collection and in that of Major Powell-Cotton.

This species is nearest Hypolycena naara Hew., but has no

sex-spot in the primaries.

DEUDORYX ITURI, sp,n. (Plate IX. fig. 7.)

¢. Both wings lustrous cobalt-blue. Primaries with costa

narrowly black ; apex very broadly black, tapering gradually to

the tornus, the blue area being gradually rounded off externally.

Secondaries with costa blackish to vein 7; termen linear, black; a

fair-sized creamy sex-mark above the cell, and a small pencil of

black hairs between veins 1 and 2.— Under side, Both wings white,

with apex and termen of primaries broadly grey, with a whitish

bisecting line; postmedial line broadly fawn-yellow, occasionally

nearly straight, but generally with an internal curve more or less

strong; inner margin with a pencil of pale yellow hairs about a

third from the base. Secondaries with termen broadly grey,

with a whitish bisecting line, an oval black spot between veins 2

and 3 in a large pale yellow patch extending more or less to the

lobe, which has a small black spot with a slight metallic-blue

edging above, over which is an angled orange-red line. Post-

medial line broadly yellow, broken at vein 4, with a double angle

between veins 1 and 3. The sex-spot showing through very

slightly. Two tails, one at vein 2, another at 3; at 4 a tooth.

Expanse 33-36 mm.

Hab. Makala, Congo Free State: April.

In my collection and in that of Major Powell-Cotton.

DEUDORYX ELEALODES, sp. n. (Plate IX. fig. 6.)

3. Both wings bright blue, not lustrous. Primaries with the

apex broadly blackish, tapering very rapidly off on the costa and

less rapidly on the termen to the tornus, where it is linear.

Secondaries with costa blackish to vein 7. Tail at vein 2 long

and fine, at vein 3 quite short, The small sex-patch between

al 908. | SPECIES OF RHOPALOCERAN BUTTERFLIES. TRS

veins 1 and 2 similar to that in D. eleala Hew. Lobe slightly

developed. Under side. Both wings dirty whitish all over, without

any grey border to the termen. Primaries with a narrow, very

slightly oblique, pale yellow postmedial line tapering into a point

on the fold; subterminal line obscure, very pale yellow; a sex-

pencil of black hairs on the fold. Secondaries with the two

lines of the primaries continued through. Lobe-spot black, very

small, with a minute spot of metallic-blue scales above; second

spot in a patch of obscure very pale yellow, with a trace of

whitish-blue submetallic scales at its upper edge.

Expanse 26-32 mm.

Hab. Makala, Congo Free State.

In my collection and in that of Major Powell-Cotton.

This species is a near ally of D. eleala Hew., but the blue of

the upper side is quite different—being blue, not green ; the under

side is also different and the markings readily distinguishable.

ToLAus Corront, sp.n. (Plate IX. fig. 14.)

¢. Primaries brilliant metallic lustrous greenish blue, extending

over the cell and two-thirds of the radial area below vein 4 and

the entire lower portion of the wing. Costa broadly black, apex

black to the cell; termen tapering rapidly to vein 2. Secondaries

entirely brilliant greenish blue like the primary, with almost

linear black termen, apex with black very restricted, a large

black shiny sexual patch to over the cell. Fringes snow-white.—

Under side. Both wings white. Primaries with apex broadly grey,

rapidly tapering to vein 1 a, a short interrupted linear line from

vein 6 to 3, with the white of the wing broadly edging it out-

wardly. Secondaries with a linear blackish subterminal line

(interrupted at the veins) from the costa to the inner margin,

angled as usual between veins | and 2; a defined small blood-red

spot between veins 2 and 3; lobe-spot black, quite small, edged

externally with bronzy metallic green and above with a small

blood-red patch, over the lower part of which are superimposed

pale metallic-blue scales; three shortish tails, the third little

more than a tooth.

Expanse 38 mm.

Hab. Makala, Beni, Congo Free State: July.

Type in the Powell-Cotton collection.

ToLAUS BILINEATA, sp. n.

Q@. Primaries brown, with three-quarters of the fold and the

cell and the angle between veins 2 and 4 whitish, suffused more

or less with very pale metallic blue, the outer part having less

suffusion. Secondaries brownish, with the basal two-thirds suf-

fused with very pale metallic blue; a white subterminal stripe

bisecting the broad brown termen, a vermilion anal spot,. and

a black marginal spot with slight metallic-bluish scales.— Under

side. Both wings white. Primaries with a defined brown ex-

curved postmedial stripe, very slightly waved between each vein,

Proc. Zoou. Soc.—1908, No. VIII. 8

114 MR. G. I. BETHUNE-BAKER ON NEW [Feb. 18,

but not broken; a similar, much broader, but less-defined sub-

terminal stripe from about vein 8 or 9; apical half of terminal

area suffused with brownish to the postmediai line, tornal half

suffused nearly to the subterminal line. Secondaries with a

similar brown postmedial stripe as in the primaries, but inter-

rupted above the vermilion patch; subterminal line yellowish

brown to the vermilion patch; a dark suffused shading precedes.

the termen ; a largish patch of bright red (almost vermilion) from

above vein 3 into the lobe, with a black spot below vein 3, edged

above with a dash of brilliant metallic pale blue, a similar dash

extending to the lobe, a small black spot in the lobe ; three tails,.

that at the lobe being the shortest.

Expanse 34 mm.

Hab. Makala, Congo Free State : March.

Type in the Powell-Cotton collection.

The measurement of this species may not be normal; it is in.

oor condition and slightly shrivelled, the left secondary being

much smaller than the right.

PowELLANA, gen. nov.

i) te)

Palpi with second segment scaled smoothly and somewhat

swollen, reaching well above the vertex ; third segment longish,

fine, smoothly scaled; eyes smooth; antenne tapering very

gradually into a very fine club.—WNeuration. Primaries with

vein 2 a third from the angle, 3 from well before the angle,

4 from the lower angle, 5 from above the middle of the disco-

cellulars, 6 from the upper angle, 7, 8, and 9 forked, 8 near apex,,

9 well beyond the middle, 10 and 11 stalked from a short stalk

from the cell. Secondaries with vein 2 from beyond the middle,

3and 4 from the lower angle, 5 from well above the middle,

7 from a third before the angle. Wings ample. Primaries:

costa strongly but evenly arched; termen evenly curved. Secon-

daries subtriangular; termen evenly arched; apex angled, anal

angle somewhat acute.

Type, Powellana cottoni B.-B.

PowWELLANA COTTONI, sp. n. (Plate IX. fig. 13.)

$. Both wings cobalt-blue. Primaries with apex broadly

black, extending slightly into the costa; termen very broadly

blackish, invading the blue in a slight inward curve, tapering

very little at the tornus. Secondaries with costa broadish black,

termen less broadly black, fold grey from vein 1. Fringes

whitish in both wings.—Under side. Both wings with broad

zebra-like stripes alternating sublustrous white and olive-brown

with fine linear subterminal lines; termen finely brown. Primaries

with the final zebra stripe in the postmedial area met at right

angles below the apex by a horizontal stripe from the middle of

the costa, over which is a short similar dash; a double curved

subterminal brown line, parallel with the termen, touches the.

1908. | SPECIES OF RHOPALOCERAN BUTTERFLIES. 115

angle just referred to. Secondaries with a curved subterminal

line as in the primaries, but double only between veins 4 and 6.

Expanse 44 mm.

Hab. Makala, Congo Free State: June.

In the Powell-Cotton collection.

The under side of this species at once separates it from all others.

LePTOMYRINA MAKALA, sp. 0.

Both wings dark brownish grey. Primaries with one or two

ocellated spots at the tornus, the pupil blackish, the iris creamy.

Secondaries with two similar spots at the anal angle and above

them a terminal stripe margined on each side by a fine creamy

line interrupted at the veins, a second fine creamy line nearer the

cell also interrupted.—Under side warm stone-grey. Primaries

with cell closed by a pale brownish dash with white centre, above

and beyond this two small brown costal spots ; postmedial stripe

brown edged with white, broken slightly below vein 4; sub-

terminal line brown, white-edged, curved inversely to the

postmedial line, with a black spot in the tornus. Secondaries

with a pale brownish dash closing the cell with a white centre,

above it below the costal vein and nearer the base a twin spot,

another spot at about the centre of the costa; on the inner

margin a spot below the cell-spot; a very interrupted postmedial

line strongly fractured below vein 4; subterminal line slightly

interrupted and excurved, another indefinite line beyond it, a

black spot between veins 2 and 3 and another at the lobe, both

edged above with orange-yellow. A long brown tail edged with

white.

Expanse 29-31 mm.

Hab. Makala, Congo Free State.

In my collection and in that of Major Powell-Cotton.

Hypocista AROA, sp. n. (Plate IX. fig. 3.)

3. Both wings scoty-brown with white areas. Primaries with

the central two-thirds of inner margin and fold white extending

obliquely to vein 4, above which the white disappears in vapoury

smoky-brown. Secondaries almost sooty-black with extreme base

brown; the antemedial, medial, and postmedial areas pure white,

invading the broad blackish subterminal margin in an acute

angle between veins 4 and 5; an oval deep black subterminal

spot between veins 2 and 4 with two minute white pupils, the

upper one smaller than the lower.— Under side. Primaries similar

to the upper side, but the white area more defined. Secondaries

similar to the upper side, but the white area more restricted and

the black oval spot ringed with yellowish, then finely with black,

and again more broadly with silvery; a similar smaller round

spot at the upper apex ; termen with a broad silvery line.

Expanse 44 mm.

Hab. Avoa River, British New Guinea.

Type in my collection.

116 MR. G. T BETHUNE-BAKER ON NEW | Feb. 18,

THAUMAINA, gen. nov.

Palpi: second segment long, haired, reaching above vertex ;

third segment moderately long, almost naked; antenne short,

reaching only just beyond the middle of the costa. Eyes hairy.—

Neuration. Primaries: vein 2 from about a third before the

lower angle, 3 from well before the angle, 4 from the angle,

5 from about the middle of the discocellulars, 6 and 7 from the

upper angle, the latter ending on the costa just in front of the

apex, 8 stalked with 7 near costa, 9 absent, 10 from the cell,

1] rising rapidly to 12 and lying along it nearly all its length, but

not anastomosing. Secondaries: 2 from well before the angle,

3 and 4 from the angle, 5 from above the middle of the disco-

cellulars, 7 from just in front of the angle.

Type, Thawmaina uranothauma B.-B.

THAUMAINA URANOTHAUMA, sp. n. (Plate 1X. figs. 8 5,9 9.)

$. Both wings blackish. Primaries with the basal two-thirds

brightish blue, the blue occupying the lower half of the cell only.

Secondaries with the under side white, showing through on the

costa and as a spot beyond the cell. Fringes tessellated white

and black.—Under side. Both wings white with ocellated spots.

Primaries with a black costal \.-shaped mark at the base, a black

dash at the centre of the costa, a subterminal irregular row of

brownish-grey spots encircled narrowly with blackish; directly

outside this row is a strongly scalloped pale brownish-grey line.

Secondaries largely filled with confluent blackish spots, an irregular

smallish black patch at the base; three spots across the middle of

the wing, the first and third with white pupils, the second at the

end of the cell more or less greyish ; two black ringed spots with

white pupils on the costa in front of the apex; a subterminal

row of six black confluent spots rising on vein 6, the first four

slightly curved, the fifth and sixth shifted right mwards, not

entirely fractured, but coming just below the three spots across

the middle; termen beyond these spots pale greyish brown, with

small white spots on the termen.

Q@. Like the male, but with the blue of the upper surface

replaced by white and with a small whitish costal patch before

the apex and a white spot below it.

Expanse 27 mm.

Hab. Angabunga River, British New Guinea.

Type in the Tring Museum.

CYANIRIS OWGARRA, sp. n. (Plate VIII. fig. 17.)

3. Both wings pale slightly lustrous sky-blue. Primaries

with a trace of white in the angle of vems 3 and 4; termen

rather narrowly black, tapering slightly towards the tornus.

Secondaries with the costa grey to the cell and vein 6; termen

very narrowly black.— Under side. Both wings greyish white with

pale liver-brown markings. Primaries with cell closed by a

1908. | SPECIES OF RHOPALOCERAN BUTTERFLIES. 117

slightly angled dash; postmedian line spotted, the costal spot

shifted well inwards, all the spots interrupted at the veins,

followed closely by the subterminal line of curved internervular

dashes ; termen very finely linear, preceded by a row of brownish

dots. Secondaries with a basal series of three points; a small

spot just outside this series in the middle of the cell, with another

below it on the inner margin; cell closed by a fine line; a post-

medial irregular series of spots, the costal one the darkest, the

second shifted inwards, third and fourth well outwards, with a

fifth small one below, sixth shifted inwards, seventh slightly

angled outwards; subterminal line fine, scalloped, internervular

small spots lying in the scallops; termen finely linear.

2 with a subovate white patch in the primaries in the radial

area ; apical and terminal areas very broadly black nearly up to

the white patch. Secondaries with the termen narrowly black,

preceded by a row of internervular small black spots. Under side

like the male.

Expanse 30 mm.

Hab. Owgarra and Angabunga River, British New Guinea.

Type in my collection, other specimens in the Tring Museum.

CYANIRIS DRUCEI B.-B. (Plate VIII. fig. 16.)

2. Both wings lustrous silvery blue, the primaries more

metallic than the secondaries. Primaries with the costa very

broadly and uniformly black to half over the cell; apical area and

termen yet more broadly black. Secondaries with costa greyish

brown over the cell and below vein 6, a row of subterminal round

small black spots. Under side like the male, as described by me

in the Ann. & Mag. Nat. Hist. ser. 7, vol. xvii. p. 102 (1906).

This female is in the Tring Museum, from the Angabunga

River, B. N. Guinea.

CYANIRIS ACESINA B.-B. (Plate VIII. fig. 10.)

@. Both wings pale sublustrous blue. Primaries with the

costa broadly black; termen rather broader black. Secondaries

with the costa sooty grey to over vein 7; termen with a row of

internervular black spots. Under side like the male as described,

ep. LOS:

In the Tring Museum from the Angabunga River, B. N. Guinea.

CYANIRIS BIAGI, sp.n. (Plate VIII. fig. 11.)

3g. Both wings pale violet-blue, with very fine linear brown

termen.— Under side. Both wings silvery greyish white. Pri-

maries with the cell closed by a yellowish dash ; postmedial line

yellowish, twice broken, the first spot detached well inwards as

also the bottom spot below vein 3, the middle portion between

veins 6 and 3 straight and confluent, subterminal row of

yellowish scallops; termen with pale yellowish internervular

dashes. Secondaries with three subbasal yellowish dots, the

middle one in the cell shifted outwards, another dot on the inner

118 MR. G. T. BETHUNE-BAKER ON NEW | Feb. 18,

margin nearly below the yellowish dash closing the cell; post-

medial row of spots broken at vein 6, the third, fourth, and fifth

nearly confluent and shifted well outwards, seventh well inwards

and isolated, eighth outwards ; subterminal line scalloped, followed

by a vow of internervular dots.

Expanse 34 mm.

Hab. Biagi, 5000 ft., B. N. Guinea.

Type in my collection.

UPoLAMPES, gen. nov.

‘Palpi roughly scaled; end segment smooth, porrect, not

upturned, second segment not as long as up to the vertex. Hyes

hairy.— Veuwration. Primaries: vein 2 from just beyond the middle

of the cell, 3 from well in front of the angle, 4 from the angle,

5 from above the middle of the discocellulars, 6 from the upper

angle, 7 and 8 absent, 9 from the cell, 10 from the cell, 11 anasto-

mosing very shortly with 12. Secondaries with vein 2 from near

the middle of the cell, 3 from just before the angle, 4 from the

angle, 5 from above the middle, 7 from well before the upper

angle. Primaries with costa strongly arched, termen boldly

curved. Wings broad. Secondaries evenly rounded, of moderate

size.

Type, Upolampes striata B.-B.

UPoOLAMPES STRIATA, Sp. n. (Plate IX. fig. 15.)

Both wings dull purple, with a subdued but strong metallic

lustre ; termen in each wing broadly blackish, rather indefinite.

The under-surface markings show dimly through.— Under side.

Both wings white, with oblique broad brown bands. Primaries

with a very oblique subbasal band; medial band broad, but rather

less oblique ; postmedial band almost spotted, less oblique, again

slightly broken at vein 7, a subterminal row of scalloped spots ;

termen with fine white dashes in the scallops just mentioned.

The postmedial band has a trace of a pale internal band.

Secondaries with the same bands as in the primaries, but the

postmedial one is entirely broken at vein 6 and it has a distinct

whitish internal jagged line. Cell closed by a brown dash.

Expanse 30 mm.

Hab Aroa River, 4000-5000 ft.: June.

Type in the Tring Museum.

CALLICTITA, gen. nov.

Palpi with second segment fringed with long hair reaching

above the vertex, third segment shortish, smoothly scaled ; frons

thickly haired ; eyes densely hairy ; antennz reaching beyond the

middle of costa, ending in a gradually tapered club.—Weuration.

Primaries with vein 2 from a third in front of the lower angle,

3 from well in front of the angle, 4 from the angle, 5 from the

middle of the discocellulars, 6 from the upper angle, 7 and 8

1908. | SPECIES OF RHOPALOCERAN BUTTERFLIES. 119

stalked on a long stalk from just in front of angle, 9 absent,

10 from the cell, 11 rising rapidly to 12 and shortly anastomosing

with it. Secondaries with vein 2 from well before the lower

-angle, 3 and 4 on a short stalk from the angle, 5 from about the

middle of the discocellulars, 7 from just before the upper angle.

Type, Callictita cyara, B.-B.

CALLICTITA CYARA, Sp. n. (Plate VIII. fig. 1.)

g. Primaries very dark brown, with a large subovate black

median patch whose edges are shot with purplish~ blue: this

spot is composed of differently placed scales, as in the African

genus Uranothawma Btl., only in a different position, and I

should expect it to be sexual. Fringes brown. Secondaries

white, slightly creamy, with a very broad blackish costa and

somewhat less broad blackish termen, base very restricted black.

Fringes tessellated black and white-—Under side. Both wings

white with brown markings. Primaries with base brown except

for a white costal dash, a broad outwardly oblique irregular

median band, a broadish postmedial band to vein 2 curved and

slightly tapering ; termen very broadly brown. Secondaries with

base and inner margin brown, the base having, as it were, two

sharply rectangular steps in it, inner margin obscurely marked

with whitish; termen broadly but irregularly brown, with a

black marginal spot between veins 2 and 3 and at the angle

edged above with metallic blue, a short tail at vein 2, An

isolated quadrangular spot in the middle of the costa.

Expanse 30 mm.

Hab. Owgarra ; Angabunga River, 6000 feet.

Type in my collection, other specimens in the Tring Museum.

I have a single specimen from Owgarra; but there is a series

from the Angabunga River, a small tributary of the larger

St. Joseph River.

MAMBARA, gen. nov.

Palpi upturned, second segment thickly scaled, with long hairs,

tapering shorter to the third segment, reaching above the vertex,

third segment smoothly scaled, longish; antenne reaching to

beyond the centre of costa, not long, terminating ina club. Head

and frons thickly haired; eyes hairy. Lees smooth, mid tibie

with a short pencil of hairs on the inner side at the femoral joint.

Wings—Primaries with costa strongly arched, termen slightly

rounded at apex, and tornus almost excavated between veins 3

and 6. Wings broad. Secondaries with termen straight from below

vein 7 to vein 4.—WVeuwration. Primaries with vein 2 from beyond

the middle of the cell, 3 from just before the lower angle, 4 from

the lower angle, 5 from above the middle of the discocellulars,

6 and 7 on a very short stalk from the upper angle in the male;

in the female 6 from the angle, 7 from the cell, 8 stalked from 7

from near the apex, 9 and 10 absent, 11 with a minute bar to 12.

Secondaries with vein 2 from beyond the cell, 3 and 4 from the

120 MR. G. T. BETHUNE-BAKER ON NEW [ Feb. 18,.

angle, 5 from above the middle, 6 and 7 from the upper angle ;.

in the female vein 7 rises from the cell, not from the angle.

Type, Mambara wigropunctata B.-B.

MAMBARA NIGROPUNCTATA, sp.n. (Plate VIIT. fig. 5.)

3S. Both wings white, with restricted black bases. Primaries

with a black spot closing the cell, costa narrowly black ; apex and

termen to vein 1 very broadly black, the black apical area

reaching nearly to the cell and to the middle of the costa, with

a deep indentation between veins 3 and 4. Secondaries with the

black spots of the under side showing through and a black ter--

minal short dash at vein 2.—Under side. Primaries white, with

a basal black spot, a large oblong spot at the end of the cell, a

very broad irregular angled black band from just beyond the

middle of the costa to the termen, the radial and terminal areas.

being black between veins 5 and 2; apical area white with a

small black apical spot. Secondaries creamy white, with a sub-

basal round black spot between vein 8 and the cell, below which

is a three-armed basal mark (reminding one of the Manx Arms);,.

postmedial stripe black, very interrupted, consisting of a costal

apical spot with a broadish dash below it shifted mwards, and

a small black spot below it, a black crescent shifted outwards, a

spot below it in the angle of vein 3, and further in another spot

in the angle of vein 2, below which shifted slightly outwards is a

broad dash to the middle of the inner margin, a black spot below

on this margin, and a black spot on the termen between veins 2.

and 3.

@ like the male; but in the primaries the basal black patch

extends nearly all over the cell, and is confluent with the apical

area, whilst in the secondaries there is a slight subterminal.

brownish dusted indefinite line.

Expanse 30 mm.

Hab. Biagi, 5000 ft., January ; Owgarra.

Types in my collection, other specimens in the Tring Museum..

CANDALIDES MARIA, Sp. n.

$. Both wings uniform dull mauve-blue, with fine black linear

costa and termen, and white fringes.— Under side. Both wings.

pure shining white, with fine linear pale ochreous-brown markings..

Primaries with a twice-broken postmedial line, the middle part.

between veins 2 and 6 shifted outwards, a subterminal line

composed of fine internervular dashes ; termen very fine, linear,

brown; secondaries with four basal points below each other,

a larger point below the angle of vein 2, a fine dash above the

angle of vein 7; the postmedial line broken as in the primaries,

but the middle part shifted further out; subterminal line con-

nected throughout, a very fine brown linear termen, a black spot.

at the angle, and termen finely spotted to vein 6.

Expanse 40 mm.

Hab. Avoa River, October.

Type in my collection.

1908. | SPECIES OF RHOPALOCERAN BUTTERFLIES. 12h

CANDALIDES NEURAPACUNA, sp.n. (Plate IX. fig. 10.)

3. Both wings sooty black, with sublustrous blue areas.

Primaries with the blue area occupying the cell to the Inner

margin and a good half of the radial area, the base of veins 2, 3,

and oe thickened, and the white of the under side showing

through slightly along the fold and the basal half of the imter-

nervular spaces of veins 2 to 4. Secondaries with the blue area

very subdued and suffused with fine black irrorations, occupying

the cell and about half the radial area to vein 2.—Under side.

Both wings silvery white with pale brown markings. Primaries.

with a fine dash closing the cell; a postmedial line of }>-shaped

marks interrupted at the veins, a subterminal line of fine inter-

nervular dashes, a terminal obscure row of fine points. Second-

aries with a basal row of four dots, an antemedial row of three

dots above which is a short erect costal dash. Cell closed by a fine

dash ; a postmedial row of internervular $-shaped marks, a sub-

terminal row of fine similar marks, a terminal row of dark dots.

Expanse 40 mm.

Hab. Angabunga River, 6000 ft.

Type in Tring Museum.

CANDALIDES GRANDISSIMA, nom. nov. (Plate VIII. fig. 15.)

Mr. Grose Smith has already described a species of Holochila

(Rhop. Exot., Lyc. Orient. p. 14, pl. xviii.) under the name grandis ;.

therefore my species C’. grandis requires a new nameas Candalides

and Holochila are synonymous. I propose, therefore, the name

grandissima. In Meek’s collection from the Angabunga River is

a series of males and one female, which latter I now describe.

Both wings brownish black. Primaries with base blackish, the

lower part of the cell, the median part of the fold, and half the

radial area from just above vein 4 white. Secondaries brown

rather than black, with the cell and slightly below it, and half

the radial area above vein 4 to 7 white.

CANDALIDES GLORIOSA, sp. n. (Plate VIII. fig. 3.)

3. Both wings most brilliant metallic lustrous bright cobalt-

blue. Primaries with apex broadly black, tapering rapidly to the-

tornus ; costa black to the costal vein. Secondaries with costa

above vein 6 brownish, termen narrowly black.—Under side..

Both wings spotless silvery white. Secondaries with a trace of

obscure black internervular terminal points.

@. Both wings dark brown. Primaries with half the cell to

the inner margin and half the radial area below vein 4 bright

lustrous pale blue, with the white under side showing through

in the radial area between veins 2 and 4. Secondaries with a

very slight suffusion of blue in the cell and beyord and slightly

below it.—Under side. Both wings uniform spotless shining

cream-colour, with black internervular terminal dashes.

Expanse, ¢ 41, 9 39 mm.

Hab. Angabunga River.

Type, ¢ in my collection, 9 in the Tring Museum.

122 MR. G. 1, BETHUNE-BAKER ON NEW [ Feb. 18,

GANDALIDES AROA, sp. n. (Plate VIII. fig. 7.)

@. Both wings blackish, with pale metallic-blue areas. Pri-

maries with the blue area occupying the lower half of the cell to

the inner margin and half of the radial area to about vein 4.

Secondaries with the blue area confined to the cell to vein 1 a, and

occupying the greater part of the radial area to about vein 6,—

Under side. Uniform creamy spotless whitish. In the secondaries

there are dark short dashes at the end of the veins.

Expanse 36 mm.

Hab. Axvoa River.

Type in my collection.

In shape this species should be a male, as both the wings are

of the normal shape of that sex ; the fore tarsi, however, together

with the very restricted blue areas, show it to be a female.

CANDALIDES PRATTI, sp.n. (Plate VIII. fig. 13.)

3. Both wings bright metallic purplish blue. Primaries with

a linear black costa ; termen narrowly black, increasing somewhat

at the apex. Secondaries with costa white to vein 7, termen

finely black ; fringes white, finely black at the veins.— Under side.

Both wings pure white, spotless except that the secondaries have

a black dot above vein 1.

Expanse 33 min.

Hab. Fak-Fak, Dutch New Guinea.

Type in my collection.

CANDALIDES DINAWA, sp.r. (Plate VIII. fig. 2.)

3. Both wings rich metallic purplish, Primaries with a

broad brown apex rapidly tapering both on the costa and the

termen ; termen narrowly brown below vein 2. Secondaries with

costa pale brown to vein 7; termen narrowly black.— Under side.

Both wings spotless slightly shining white, except that in the

secondaries there is a small black spot above vein 1, and the

termen is distinctly spotted with black at the end of the veins.

Q. Both wings brown. Primaries with a metallic bluish

suffusion along the basal third of the fold and slightly above

vein 1 a, and a good-sized white patch below the end of the cell,

and for about the third of the internervular space between veins 3

and 4, and nearly half the space between 2 and 3. Secondaries

uniform pale brown with scalloped termen. Under side exactly

as the male.

Expanse, ¢ 31, 9 28 mm.

Hab. Dinawa, July and August.

Types in my collection.

CANDALIDES CYANA, sp.n. (Plate VIII. fig. 8.)

$. Both wings bright lustrous pale sky-blue. Primaries

with the costa evenly black to just over the upper margin of the

cell; termen broadly black, tapering slightly to the tornus.

1908. ] SPECIES OF RHOPALOCERAN BUTTERFLIES. 123

Secondaries with costa dark brown to the cell and well over

vein 7, termen broadish black, abdominal fold sooty grey. Fringes

whitish.—Under side. Both wings shining pearly white; the

secondaries with a black dot above vein 1, the end of veins 1 3,

2, 3, and 4 spotted with black.

Expanse 36 mm.

fab. Owgarra.

Type in my collection.

CANDALIDES UNIPUNCTATA, sp. n. (Plate VIII. fig. 14.)

Both wings black, with a large white patch in each. Primaries

with half the cell to the inner margin and about half the radial

area to vein 5 white, with a very shght dusting of fine very

pale greenish-blue metallic scaies at the base and in the cell.

Secondaries with the upper quarter of the cell to the costa and

the upper radial area from vein 4 to near the apex in an increasing

curve white, with a very slight very pale greenish-blue metallic

dusting in the cell. Fringes tessellated white and black.— Under

side. Both wings white, with a broadish indefinite darkly dusted

termen. Primaries with termen rather broader than secondaries,

cell closed by a narrow dark dash. Secondaries with a good-sized

black oval spot closing the cell, and a black dot above vein 1.

Expanse 30 mm.

Hab. Biagi, 5000 ft., and Angabunga River, 6000 ft.

Type in my collection ; another specimen in the Tring Museum.

The specimen in the Tring Museum has the metallic scaling as a

slight iridescence all over the white area of the primaries, and

prominently in the cell of the secondaries.

CANDALIDES ANGABUNGA, sp. n. (Plate IX. fig. 5.)

9. Both wings dull violet-blue, shghtly lustrous. Primaries

with costa broadly black and apex black to the end of the cell;

termen very broadly black, tapering but slightly to the tornus.

Secondaries with costa and termen broadly blackish, the latter

tapering slightly narrower to the tornus.—Under side. Both

wings white, slightly shining. Primaries with a small black spot

beyond the middle of the cell, termen dotted with black at the

veins. Secondaries with a black spot beyond the middle of the

cell, and a black dot above vein 1; termen dotted with black at

the veins.

Expanse 31 mm.

Hab. Angabunga River, 6000 ft.

Type in the Tring Museum.

This is a peculiar species, from the shape of the wings it might

be thought to bea male; but the fore tarsi are fully developed,

and therefore we must regard it as a female.

THYSONOTIS ROSSELANA, Sp. nN.

3S. Both wings dull slightly lustrous violet-blue. Primaries

with costa and termen linear black, and the white of the under

124 MR. G. T. BETHUNE-BAKER ON NEW [ Feb. 18,

side showing slightly through. Secondaries with costa broadly

white, termen narrowly black; tail very fine, white-tipped ; white

of under side showing through; abdominal fold white for the

basal half.— Under side. Primaries white with costa uniformly very

broadly brown extending over nearly half of the cell, termen

moderately narrow brown; a dash of brown from the tornus to

vein 3 separated from the termen by a narrow white line.

Secondaries white, with a broad basal oblique band to the costal

vein ; costa wholly white except the extreme base, which is dusted

with metallic blue; a broad irregular deep black postmedial band

edging the brilliant metallic pale blue terminal area, which is

very broad at the anal angle, tapering in an even curve to the

apex and having a central rew of black spots, those at the angle

itself almost obsolete; termen finely white, preceded by a fine

black line. Fringe blackish.

Expanse 36 mm.

Hab. Rossel Island.

Type in my collection.

Near 7. hamilear G. Smith.

THYSONOTIS EKEIKEI, sp.n. (Plate IX. fig. 1.)

3. Both wings uniform mauve-blue, with termen narrowly

black and costa to primaries linear black, the white of the under

side showing through but slightly.—Under side. Both wings

sooty brown with white areas. Primaries with base, the whole of

the cell, and costa to below vein 6 sooty brown ; termen broadly

black, narrower between veins 3 and 5, the dark termen being

broadest at the tornus; rest of internal area white nearly to the

angle of vein 2. A slight stripe of metallic grey dusting along

the upper margin of the cell and slightly over the margin.

Secondaries with base dark sooty brown to nearly half the cell,

with a short basal metallic-grey curved stripe; terminal third

dark sooty brown, with a terminal row of large deep black velvety

spots tapering smaller to the apex, and edged internally with

metallic-green scallops and externally by a metallic-blue fine

irregular line ; the middle part of the wing white, occupying

the middle third of the inner margin, and expanding outwards

to the costa to near the apex. Without a tail.

Expanse 36 mm.

Hab. Kikeikei, January and February.

Type in my collection.

Near 7’. endocia, H. H. D.

THYSONOTIS ALBOSTRIGATA, Sp. n. (Plate IX. fig. 11.)

3. Both wings brilliant metallic lavender-blue with linear’

black margins. Primaries showing the white of the under side

very slightly through. Secondaries with a broad shining white

irregular median band. ‘Tail fine, tipped with white.—Under

side. Primaries blackish, with the white area restricted to rather

1908. | SPECIES OF RHOPALOCERAN BUTTERFLIES. 125

over half of the radial area up to vein 5, extending just into

the lower part of the end of the cell and to just beyond the

angle of vein 2 to the inner margin; a brilliant metallic pale blue

costal stripe for about half the costa. Secondaries deep black,

with a pure white stripe as on the upper side; base up to this

black, with a brilliant metallic pale blue curved costal stripe ; the

terminal three-fifths black, with a terminal row of large brilliant

metallic pale blue oblong spots tapering smaller to the apex, with

large deep black similar-shaped pupils tapering in lke manner ;

no white terminal fine line.

Expanse 44 mm.

Hab. Fak-Fak, Dutch New Guinea.

Type in my collection.

WAIGEUM DINAWA, sp. n. (Plate IX. fig. 2.)

3. Both wings brown, with white submedian patches and

metallie-blue dusting. Primaries with the white area restricted

to a fair-sized somewhat oblique patch at a quarter from the base

of the inner margin, occupying the middle area of the wing up to

about vein 4, not extending into the cell and for about half the

radial area, ‘surrounded very broadly by brown except at the

centre of the inner margin ; OG brown ground-colour is super-

imposed broadly around the white with brilliant metallic-blue

fine irrorations. _ Secondaries with brown base extremely restricted

and covered with similar brilliant blue fine dusting as in the

primaries, a broad oblique white band; the outer half of the

wing brown, with brilliant blue dusting in a triangular patch

from the base of vein 2 to about vein 4 and halfway to the

termen.— Under side. Primaries black, with the white area

more extended than above, reaching the base below the cell, and

invading the broad blackish terminal area along veins 2, 3, and

4, so as to strongly scallop it; costa with a broad band of brilliant

metallic bluish-green dusting extending into the subapical area

in a short curve, an interrupted terminal bluish-green similar

line, and a similar stripe in the cell. Secondaries with the black

base edged laterally by a broadish metallic-blue, curved, costal

dash, and a fine line on its outer edge; terminal half blackish,

with a subterminal row of large deep black subtriangular spots,

tapering smaller to the apex, and margined entirely by the same

brilliant blue dusting, which dusting tapers rather narrower

towards the apex.

Expanse 42 mm.

Hab. Dinawa, August and September, 4000 feet.

Type in my collection.

This species will stand near 7’. corruscans, G. Smith.

WAIGEUM RESPLENDENS, sp. n. (Plate IX. fig. 12.)

2. Both wings brown, with white areas. Primaries with the

white area from the base below the cell to just beyond the centre

126 ON NEW SPECIES OF RHOPALOCERAN BUTTERFLIES. | Feb. 18,

of the inner margin and extending just over vein 4 for about half

of the radial area; a slight dusting of metallic-blue scales in the

cell, especially along the lower margin. Secondaries with base

brown, most restricted, and with a few metallic-blue scales; a

broad white band across the medial area, more than half of the

posterior part of the wing being uniformly brown.—Under side.

Primaries with the white area occupying a similar area as above,

but more restricted ; the rest of the wing brown, with the costa

very broadly dusted with resplendent metallic greenish-blue fine-

scales, extending round the subapical area ina curve, and recurved

upwards internally so as to margin the apex of the white patch ;

a broadish terminal line of very metallic blue. Secondaries with

black base margined with the same resplendent greenish-blue

scales broadly above, narrowly below; white area as above, the

rest of the wing blackish, edged on its internal edge narrowly

with the greenish-blue scales; a very broad internal band of

resplendent scales tapering slightly towards each end, in which is.

a row of small black spots ; a subterminal broad line uniform in.

width of resplendent metallic greenish blue.

Expanse 42 mm.

Hab. Aru Island, June.

Type in my collection.

EXPLANATION OF THE PLATES.

Prats VIII.

Fig. Fig

1. Callictita cyara. | 10. Cyaniris acesina.

2. Candalides dinawa. ike + biagi.

3. ¥ gloriosa. *12. Epitola albomaculata.

*4. Paraduba owgarra. 13. Candalides pratti.

5. Mambara nigropunctata. 14. os unipunctata.

*6. Hpitola mangoensis. 165. 5 grandissima.

7. Candalides aroa. 16. Cyaniris drucei.

8. % cyan. 17. 55 owgarra.

*9, 5 sublutea. *18. Myrina sharpet.

T have taken this opportunity of figuring some little-known species described by

myself which are not mentioned in the text ; they are marked with a *.

Puate IX.

Fig. | Fig.

1. Thysonotis ekeike?, sp. 0. | 9. Thaumaina uranothauma, sp.n.§ ..

2. Waigeum dinawa, sp. i. | 10. Candalides newrapacuna, sp. 0.

3. Hypocista aroa, sp. n. | 11. Thysonotis albostrigata, sp. 0.

4. Deudoryx makala, sp. n. | 12. Waigeum resplendens, sp. n.

5. Candalides angabunga, sp. 0. | 13. Powellana cottoni, sp. n.

6. Deudoryex elealodes, sp. 0. | 14. Lolaus cottont, sp. n.

ths x ituri, Sp. N. 15. Upolampes striata, sp. n.

Thaumaina uranothauna, sp. n.é .

No. 51.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

January 14th, 1908.

Prof. J. Ros—E Braprorp, M.D., D.Sc., F.R.S., Vice-President,

in the Chair.

The Secretary read a report on the additions that had been

made to the Society’s Menagerie during the month of December

1907.

Mr. W. P. Pycrart, F.Z.8., exhibited, on behalf of Mr. R.

Lyprexker, F.R.S., F.Z.S., an abnormally marked Leopard-skin

from the Deccan, India, which had been presented to the British

Museum of Natural History by Mr. F. A. Coleridi.

Dr. W. A. CunNINGTON gave an account of an expedition con-

ducted last spring, in conjunction with Mr. C. L. Boulenger, to

investigate the flora and fauna of the Birket el Qurun—the lake

Moeris of the ancients. The expedition was undertaken on behalf

of the Egyptian Survey Department, and one important result

was the discovery of a new lacustrine medusa.

Mr. OupFreLD THoMAS read a paper, the sixth of the series, on

mammals obtained in the Shantung Peninsula, N China, by

Mr. M. P. Anderson, for the Duke of Bedford’s exploration of

Eastern Asia. No mammals had come from this region since the

time of Consul Swinhoe, who had visited it in 1866-68. The

present series contained 106 specimens belonging to six species,

of which one was new.

Mr. ¥, E. Bepparp, F.R.S., Prosector to the Society, read a

communication entitled ‘On the Musculature and other Points

in the Anatomy of the Engystomatid Frog, Lreviceps verrucosus.”

* This Abstract is published by the Society at 3 Hanover Square, London,

W., on the Tuesday following the date of Meeting to which it refers. It will

be issued, free of extra charge, to all Fellows who subscribe to the Publications,

along with the ‘ Proceedings’ ; but it may be obtained on the day of publication

at the price of Sixpence, or, if desired, sent post-free for the sum of Six

Shillings per annum, payable in advance,

Mr. C. L. Bounencer, B.A., gave an account of a communi-

cation entitled ‘‘On the Hermaphroditism of the Amphipod

Orchestia deshayesti, Audouin.”

The next Meeting of the Society for Scientific Business will

be held on Tuesday, the 4th February, 1908, at half-past Hight

o'clock P.M., when the following communications will be made :—

1. F. Martin Duncay.—Cinematograph demonstration of re-

sults of Natural Colour Photography with Zoological Subjects.

2. O. THomas.—The Duke of Bedford’s Zoological Exploration

in Eastern Asia— VII. List of Mammals from the Tsu-shima

Islands.

3. T. Goopry.—On the Presence of Gonadial Grooves in Awrelia

aurita.

4. C. Tare Recan.—The Duke of Bedford’s Zoological Explor-

ation in Eastern Asia,—VIIL. A Collection of Freshwater Fishes

from Corea.

Communications intended for the Scientific Meetings of the

ZooLoGicaL Soctery oF Lonpon should be addressed to

P. CHALMERS MITCHELL, Secretary.

3 Hanover Square, Lonpon, W.

January 21st, 1908.

No. 52.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON”

February 4th, 1908.

H.G. Tue Duxe or Beprorp, K.G., President, in the Chair.

Mr. F. Martin Duncan gave a lantern exhibition of the

Lumiére Autochrome Natural Colour Process as applied to

zoological subjects.

Mr. OtpriztD Tuomas, F.R.8., read a paper, the seventh of

the series, on Mammals from the Islands of Tsu-shima, between

Korea and Japan, collected by Mr. Malcolm P. Anderson in

carrying out the Duke of Bedford’s Exploration of Eastern Asia.

So far as its mammals were concerned, the Tsu-shima group was

shown to be predominantly Japanese in character, though three

of its species were distinctly Korean, but these, Mr. Thomas

thought, might possibly have been accidentally introduced from

the mainland. Twelve species were recorded from the Islands,

and, including a few which came from other sources, 151 specimens

were dealt with.

Mr. T. Goopry read a paper “On the Presence of Gonadial

Grooves in Aurelia aurita,” of which the following is an abstract :—

“The ‘ gonadial grooves’ are seen in a view of the sub-umbrella

surface, and lie in the four interradial axes. Each has about the

same diameter as the origin of an ordinary per-radial or ad-radial

canal, and extends from the central gastric cavity into a gastric

pouch, ending there in a slight expansion. The groove is confined

to the ventral wall or floor, and is formed by a folding of the

endodermal epithelium. Itis suggested that the grooves function

* This Abstract is published by the Society at 3 Hanover Square, London

W., on the Tuesday following the date of Meeting to which it refers. It will

be issued, free of extra charge, to all Kellows who subscribe tothe Publications,

along with the ‘Proceedings’; but it may be obtained on the day of publication

at the price of Sixpence, or, if desired, sent post-free for the sum of Six

Shillings per annum, payable in advance,

as channels for the outward conveyance of the ripe sex-cells when

liberated from the gonads. The Ctenophor Ctenoplana is the

only other Celenterate possessing genital ducts, but it is only in

the Seyphozoon Awrelia that the structures can be regarded as

ccelomic or archenteric derivations.”

Mr. C. Tate Reaan, M.A., F.Z.S., read a paper entitled “The

Duke of Bedford’s Zoological Exploration of Eastern Asia.—

VIII. A collection of Freshwater Fishes from Corea.” The col-

lection included examples of eleven species, seven of which were

described as new to science.

The next Meeting of the Society for Scientific Business will be

held on Tuesday, the 18th February, 1908, at half-past Hight

o'clock p.M., when the following communications will be made :—

1. R. Srapres-Browne, M.A., F.Z.8.—On the Inheritance of

Colour in Domestic Pigeons, with Special Reference to Reversion.

2. OupFIELD Tuomas, F.R.S., F.Z.S.—The Duke of Bedford’s

Zoological Exploration in Eastern Asia.—IX. List of Mammals

from the Mongolian Plateau.

3. G. T. Beruune-Baxer, F.LS., F.Z.S.— Descriptions of

new Species of Rhopalocera from Africa and from New Guinea.

Communications intended for the Scientific Meetings of the

ZooLocicaL Society or Lonpon should be addressed to

P. CHALMERS MITCHELL,

Secretary.

3 HANovER Square, Lonpon, W.

February 11th, 1908.

No. 58.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON

February 18th, 1908.

Dr. Henry Woopwarp, F.R.S8., Vice-President,

in the Chair.

The Secrerary read a report on the additions that had been

made to the Society’s Menagerie during the month of January

1908.

Mr. R. I. Pocock, Superintendent of the Gardens, exhibited,

on behalf of Mr. W. Simpson Cross, F.Z.8., a photograph taken

from a living specimen of a very rare South-American Dog (Canis

jubatus), sometimes called the Maned Wolf.

Mr. R. H. Burne, F.Z.8., exhibited and remarked upon a

number of preparations of the olfactory organs of Birds and

Fishes.

Dr. L. W. Samson, F.Z.S., exhibited a large series of specimens of

internal parasites obtained by him from animals recently living in

the Society’s Gardens. He laid stress on the important additions

to knowledge to be derived from an adequate investigation of

such material, and on the practical results to the health of the

animals in the Gardens that might be expected.

Mr. R. SraprEs-Browne, F.Z.8., read a paper ‘‘On the In-

heritance of Colour in Domestic Pigeons, with Special Reference

* This Abstract is published by the Society at 3 Hanover Square, London,

W., on the Tuesday following the date of Meeting to which it refers. It will

be issued, free of extra charge, to all Fellows who subscribe to the Publications,

along with the ‘ Proceedings’ ; butit may be obtained on the day of publication

at the price of Stxpence, or, if desired, sent poust-free for the sum of Si

Shillings per annuin, payable in adyance.

to Reversion,” and exhibited a series of skins illustrating some

experiments on which his communication was based. Crosses

had been made between black Barbs and white Fantails. The

F.1 generation was black with some white feathers. In the F. 2

generation, among other forms, blacks and whites were obtained,

and also some blues. Blues were found to be dominant to whites,

but blacks were dominant, or rather “ epistatic,” to the blues,

which accounts for the fact that the reversionary form does not

appear until the F. 2 generation. When two blues of the F. 2 or

later generations were mated together blacks were never obtained

again. A white in F. 2 mated to a Fantail gave whites only.

A second series of skins illustrated a cross between a white

Tumbler and a white Fantail. Some white birds splashed with

red had figured in the ancestry of the Tumbler, although the bird

itself showed no trace of colour. In the F.1 generation such

splashed kinds occurred, which, when mated together, gave in

F.2 birds which were red and white with some distinct blue

feathers. Possibly the white Tumbler was a dominant white.

Mr. Ouprietp Tuomas, F.R.S., F.Z.8., read a paper on

mammals collected by Mr. M. P. Anderson during a trip to the

Mongolian Plateau, N.W. of Kalgan. Nine species were men-

tioned, of which two were described as new.

The paper formed the eighth of the series on the results

obtained by the Duke of Bedford’s Zoological Exploration in

Eastern Asia.

No properly collected material from the Mongolian plateau had

been previously available to students, and these specimens, repre-

sentatives of its comparatively poor fauna, were therefore of much

interest.

A communication was received from Mr. G. T, Berauns-Baker,

F.L.S., F.Z.S., 12 which the author described as new to science a

number of species of Butterflies of the division Rhopalocera,

from Africa and from New Guinea.

The next Meeting of the Society for Scientific Business will be

held on Tuesday, the 38rd March, 1908, at half-past Hight

o'clock P.M., when the following communications will be made :-—

1. F. E. Bepparp, M.A., F.R.8., Prosector to the Society.—

A Comparison of the Neotropical Species of Corallus, C. cookin

with C. madagascariensis; and on some Points in the Anatomy

of Corallus caninus,

2. P. Caatmers Mrrcweuy, M.A., D.Se., LU.D., F.R.S., Secre-

tary to the Society.—On a Young Female Kordofan Giraffe.

3. R. I. Pocock, F.L.S8., Superintendent of the Gardens,—

Description of a new Species of Monkey of the Genus Cerco-

pithecus.

The following communications have been received :—

1. Dr. Knup Anprersen.—A Monograph of the Chiropteran

Genera Uroderma, Hnchisthenes, and Artibeus.

2. Watrer A. Kipp, Esq., M.D., M.R.C.S., F.Z.8.—Some

Observations on the Effects of Pressure upon the Direction of

Hair in Mammals.

Communications intended for the Scientific Meetings of the

ZOOLOGICAL SociEry oF Lonpon should be addressed to

P. CHALMERS MITCHELL,

Secretary.

3 HANOVER Square, Lonpon, W.

february 25th, 1908,

ConTENTS (continued).

February 4, 1908 (continued).

3 Page

2. On the Presence of Gonadial Grooves in a Medusa, Aurelia aurita. By T. Goopzy,

Student Demonstrator in Zoology, University of Birmingham. (Plate I.) .......... 55

3. The Duke of Bedford’s Zoological Exploration in Eastern Asia—VIII. A Collection of

Fresh-water Fishes from Corea. By C. Tate Reeay, M.A., F.Z.S. (Plates II. & III.) 59

February 18, 1908.

The Secretary. Report on Additions to the Society's Mevagerie during the month of

eI Asya ORetate a cexniolelcversi oye jpishosemeicke afelel ch vicmin ts ee. Siaymarteisetare Ae Sue ciea c shea wore Ga 63

Mr. R. I. Pocock, F.L.S., F.Z.S. Exhibition of, on behalf of Mr. W. Simpson Cross, F.Z.S.,

and remarks upon, a photograph of a rare South American Wild Dog (Canis jubatus).. 64

Mr. R. H. Burne, F.Z.8. Exhibition of, and remarks upon, preparations of the Olfactory

Organs of Fishes and Birds

AUNTS SS CR acon ob CUS a Cre gC aR rane oe cn uaa a ra 65

Dr. L. W. Sambon, F.Z.S. Exhibition of, and remarks upon, a large series of Internal

DREISER ohare ee auaee ote ia alah ave @alcke atiozoner laa ancya) ebenslleron roy srameta so: eae Schema ce ee aoe avian ane oe ee NG

1. On the Inheritance of Colour in Domestic Pigeons, with Special Reference to Reversion,

By Ricuarp Stapies-Browne, M.A., F.Z.8. (Plates [V.-VIL.)........-02000e seen Od

2. The Duke of Bedford’s Zoological Exploration in Eastern Asia.—IX, List of Mammals

from the Mongolian Plateau. By Ouprreip Tuomas, F.RB.S., F.Z.S.............000+ 104

3. ‘Descriptions of new Species of Butterflies of the Division Rhopalocera from Africa and

from New Guinea. By G. T. Bataunr-Baxker, F.LS., F.Z.8. (Plates VIII. & IX.).. 110

LIST-0OF PL MEES:

1908, pp. 1-126.

Plate Page

I. Gonadial Grooves in Aurelia auritd ..++.. ee cece seep eee 55

{ 1. Acanthorhodeus gracilis. 2. Leucogobio strigatus. |

TE 3) Silurus bed fordi. Se. 3 Gel ces teen Nolet ele oe ie ee t 59

lt { 1. Ctenogobius bedfordi. 2. Tridentiger coreanus. |

“(3. Acanthogobio longirostris. 4. Liobagrus andersont...... )

a } BeidoHlanaicMl Oro coc ghossoude op boddocdn oo woos dooK i

: 67

VI. 1. Barb-Fantail Cross. .2. Tumbler-Fantail Cross .......

VII. Tumbler-Fantail Cross .. 1... ce eee eee eee ee ee ees oon

IL.

ee | New or Little-known Lycenide.......-.-.-02++.---000- 110

NOTICE.

The ‘ Proceedings’ for the year are issued in fowr parts, paged consecutivel

d a) Pp » pag yr

so that the complete reference is vow P. Z. 8. 1908, p... . The Distribution

is as follows:—

Papers read in January and February, issued in June.

nm » March and April, BE ak. August.

a » May and June, sy)», October.

ee » November and December,,, ,, April.

‘ Proceedings,’ 1907, pp. 747-1121, were published on June 4th, 1908.

The Abstracts of the papers read at the Scientific Meetings in

January and February are contained in this Part.

—> Ss

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY

OF LONDON.

1908.

Pages 127-430.

Part I]. conTAINING PAPERS READ IN

MARCH anv APRIL.

SEPTEMBER 1908.

PRINTED FOR THE SOCIETY,

SOLD AT THEIR HOUSE IN HANOVER SQUARE.

LONDON:

MESSRS. LONGMANS, GREEN, AND CoO.

PATERNOSTER ROW.

[Price Twelve Shillings. |

LIST OF CONTENTS

1908, pp. 127—4380.

March 3, 1908,

Page

Mr. R. Trimen, F.R.S., F.L.S., F.Z.S. On a long spur or claw-like horny excrescence at

the tipo the tail of a:domestic Cat <iimccse = sels css aes eniem om dee hee eee 127,

The Secretary. Exhibition of some skins of the Coypu, Myocastor coypu ........c0+se+0. 127

Mr. F. E. Beddard, M.A., F.R.S. Exhibition of, and remarks upon, the Colon and Rectum

Ola Bad ger(COleles mieles), cm apisctam ce eee ncn eaee SOMONE AUER OOOUGE dood a: 128

1..On a young Female Kordofan Giraffe. By P. Cuanmurs Mircusty, M.A., D.Se., LL. D.,

HRS; Seeretary: to the Society? @. sce cease eee nt ene eee eee 130

. A Comparison of the Neotropical Species of Corallus, C. cookii, with C. madagas-

cariensis; and on some Points in the Anatomy of Corallus caninus. By Frank BE, ~

Baween MC A., UR. S) Prosector tothe Soriehy<cssc- cm <0. 2s saci <) se eee Sy 18%)

3. Description of a new Species of Monkey of the Genus Cercopithecus. By R. 1. Pocock,

ELS3 Superintendentvor the Gardens:.<CPlate NX.) <.-tactacls cys eerene ace eee 158

March 17, 1908.

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

February 1908 -....... Dei eligig) eee sis stave e olleeane/elaiialsieye ala ere atanese- sicte uate cstelaiane ep ate arcane 161

Mr. G. A. Boulenger, F.R.S., V.P.Z.S. Exhibition of, and remarks upon, a remarkably

malformed. Plaice (Pieuroneetes platessa). sane eae en ee 161

Mr. Walter A. Kidd, M.D., M.R.C.S., F.R.S.E., F.Z.S. An account of observations

regarding the effects of pressure on the direction of hair in Mammals ..............- 164

1. The Rudd Exploration of South Africa.—IX. List of Mammals obtained by Mr. Grant

on the Gorongoza Mountains, Portuguese S.E. Africa. By Oxprrenp Tuomas, E.R.S.,

ZS. and Rh. C2 Wrougnron RUZS: knit once soem ee naya'd oh iaje ae ee ee 164

2. Notes upon some Species and Geographical Races of Serows (Capricornis) and Gorals

(Nemorhedus), based upon Specimens exhibited in the Society’s Gardens. By R. I.

Pocock, F.LS., Superintendent of the Gardens. 2:2. 1224.2... secs ese sco: fee 173

April 7, 1908.

Mr. Thomas Codrington. Exhibition of a collection of stones pee from the stomach of

an Hlephantishot in Northern Rhodesia auc welders mene cto sre clari oe ee 203

. C. W. Andrews, F.R.S., F.Z.S. Exhibition of a restored model of the skull and

mandible tol Prozeuglodon airow VAN sc cmists eae eles mane ea = as cenit ee 203

Contents continued on page 3 of Wrapper.

THE ZOOLOGICAL SOCIETY OF LONDON.

Turs Society was founded in 1826 by Sir Sramrorp Rarrtzs,

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HIS GRACE THH DUKE OF BHDFORD, K.G., President.

Grorce A. Bovtrenenr, Hsa.,

F.R.S., Vice-President.

Pror. Joan Rost Braprorp,

MED Discs bois:

President.

Aurrep H. Cocks, Ese., M.A.

Tur Rr. Hon. Kart oF Cromer,

PC; Gee JB.

CHartes DrummMonn,

Treasurer.

Sir Epwarp Douranp, Br., C.B.

Kse.,

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Pror. EpwaRp ALFRED MINCHIN,

M.A., Vice-President.

| P. Caatmers MircHett, Ese.,

Vice- |

MEAR Di Scs. Des SBeivS.,

Secretary.

Tue Lorp Newton.

W. R. Ocitvie-Grant, Esa.

| ALBERT Pam, Hsa.

Tue Hon. Cuarztes Roruscarip,

M.A.

Davin Seru-Suita, Esa.

OtprieLtp Tomas, Ese., F.R.S.

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M.A.

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3 Hanover Square, London, W.,

May, 1908.

MEETINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON

FOR

SCIENTIFIC BUSINESS.

(AT 3 HANOVER SQUARE, W.)

1908.

orspabye Mia 5... L2rand 26 Torspay, November 38 and 17

er JiGNE a lS fs DecemBEeR 15

The Chair will be taken at half-past Hight o'clock in the Evening

precisely.

1908. | ON THE MAMMARY GLANDS OF THE COYPU. 12

March 3, 1908.

G. A. Boutenesr, Esq., F.R.S., Vice-President,

in the Chair.

The Secretary exhibited for comparison the tail of a young lion

in which a caudal claw is known to occur, and read the following

communication from Mr. R, Trimen, F.R.S., F.L.S,, F.Z.8. :—

“My cat (pale grey with ordinary narrow black stripes much

broken up into short streaks and spots) presents the remarkable

peculiarity of a long spur or claw-like horny excrescence at the

very tip of its tail, This appendage is firmly seated quite at the

extremity of the last vertebra; its base appears to be expanded,

and is covered all round by an elevation of the skin. It projects

posteriorly in the line of the tail, is rather slender, gradually

tapering, almost straight for about two-thirds of its length, and

thence moderately curved downward to its moderately acute tip.

In length it is nearly 7 lines, and more than a third projects

beyond the surrounding fur. The colour of this spine or spur is

dull reddish-brown varied with dull ochry-yellowish, here and

there crossed by some broken, thin, whitish lines.

“The cab in question is a female, small, but rather thick in

body ; the limbs are all rather short and the feet small, but the

tail is noticeably long and broad with long dense fur, I am

informed by the donor that it was born at Witney, near Oxford,

and is now between seven and eight months old, I have

endeavoured, with the kind aid of the donor, to ascertain from the

original possessor of the animal whether any kitten of the same

litter, or the mother, or other known relation, exhibitec the

peculiar appendage or any traces of it; bot without success,

“7 may add that I have found the cat unexpectedly sensitive

to any handling of the caudal claw, however gentle; she first

endeavours to jerk her tail away, then gives a mild vocal remon-

strance, and if the handling is continued employs her paws to

stop it.” a

The Secretary exhibited some skins of the Coypu*, M/yocastor

coypu, kindly lent him for the purpose by Mr. C. Hawkins, and

called attention to the dorso-lateral position of the five pairs of

teats in the female, and remarked that the position of the

mammary glands in the Coypu had been recorded by Sir Richard

Owen, but appeared to have escaped notice in most recent text-

books.

Mr, O. Thomas, F.R.S., F.Z.S., added that the dorso-lateral

position of the mammary glands was not invariably associated

with aquatic habit ; and Mr. E. G. B. Meade-Waldo stated, from

personal observation, that the young of the Coypu swam along-

side the mother and were not carried on her back, as Owen had

supposed,

* See also P. Z.S. 1835, p. 182, for a similar exhibition by Mr. Christy.-

Proc. Zoou. Soc.—1908, No. TX. )

128 MR. F, B, BEDDARD ON THE [ Mar. 3,

My. F. E. Beddard, M.A., F.R.S., exhibited the colon and rectum

of a Badger (MJeles meles), and made the following remarks :—

The specimen of this Carnivore exhibited to-night shows a

remarkably large Peyer’s patch in the colon which extends

through the gr eater part, of that section of the gut and actually

measures eleven inches in length, The accompanying drawing

(text-fig. 17) shows the general aspect of this large agminated

gland and its proportions, as compared with the gut wherein it

lies. At the anterior end of the colon it is of somewhat less

diameter than that which it attains later and retains until its

disappearance about six inches in front of the anus. It ends here

abruptly by a straight margin. Its diameter is about one third

of that of the gut, ‘and is fair ly even throughout except as already

stated at the commencement. More accurate measurements of

this patch and of the gut are as follows: anteriorly the gut is

26 mm. in circumference and the patch is 6 mm. across; more

towards the anus the gut is rather wider, measuring 30 mm.,

and the patch has correspondingly increased in width, being here

of a diameter of 11 mm.

A close examination of the walls of the gut shows that there

is no differentiation of the lining membrane, and therefore the

whole piece cut out of the body “and displayed in the drawing

(text-fig. 17) i is referable to the large intestine. These ‘“ Glandule

agminatee ” are by no means recor rded here for the first time as

occurring beyond the small intestine in a mammal. It would

appear that the first description of these structures as occurring in

the colon is due to Owen*, who found them in the genus Macropus.

Subsequently the late Dr. Dobson f found such Peyer's patches

in a variety of Rodentia and Insectivora and even in the Edentate

Manis. This author quoted my own discovery of the same patches

in the Lemur, Hapalemur griseus, which I subsequently figured {.

Miall and Greenwood record the same glands in their treatise upon

the Elephant; and all of these sources of information, with the

exception of my own observations upon Hapalemur, are to be

found referred to in the monumental work of Oppel §.

It is, however, not only the situation of this Peyer’s patch in

the large intestine that is of interest. Its size also is remarkable ;

but it is by no means unique in this particular. For Oppel

obser ves that “ Beim Rind findet man am Ende des Ileums eine

2-3 M. lange Platte die noch in das Caecum hineinreicht.” There

is also a lars ge Peyer’s patch in the sheep and the pig and in one

or two aie animals belonging to the same order as Meles, viz.,

in the cat and the dog. This large patch in the Badger would

seem to be quite as large as in the other mammals which possess

a similar patch.

P.S. (added June 10th).—In a second specimen (half-grown),

* Article “ Marsupials” in Todd’s Cyclopedia, and in his treatise on Comparative

Anatomy, 1868.

ii Journ. Anat. Phys. xviii. 1884, p. 388.

{ P. Z.S. 1891, p. 453, fig. 2.

§ Lehrbuch der Vergleichenden Mikroscopische Anatomie, Teil ii., Jena 1897.

1908.| — COLON OF THE BADGER. 129

Text-fig. 17,

Colon of Badger (Meles meles) cut open to show Peyer’s patch (about nat, size).

a, 6, commencement and termination of pateh.

130 DR. P. CHALMERS MITCHELL ON [ Mar. 3,

examined on May 5th last, there was no trace of this Peyer's

patch, which may therefore be a character of full maturity. Both

specimens have been sent to the Royal College of Surgeons.

The following papers were read :—

1. On a young Female Kordofan Giraffe. By P. CHALMERS

Mrrcuett, M.A., D.Se., LL.D., F.R.S., Secretary to the

Society.

[ Received February 17, 1908. |

(Text-figures 18-20.)

The female of the pair of Kordofan Giraffes (Giraffa camelo-

pardalis antiquorum) presented to the Society by Col. B. Mahon,

F.Z.8., in 1902, gave birth to a female calf on Sept. 20th, 1907.

There are so many interesting differences in the coloration of

Giraffes, which have been employed by Mr. Lydekker (P. Z.S. 1904,

p- 202) in his valuable attempt to discriminate subspecies, that I

think it is worth while to publish some notes and figures on a

young form, the parentage of which is known, and which seems

to have a good prospect of reaching maturity in the Society’s

Gardens. The drawings from which text-figs. 19 & 20 have been

reproduced were made when the Giraffe was about eight weeks

old, being the work of a very careful artist, Mr. Goodchild, who

had already made similar drawings of the young Nigerian Giraffe,

reproduced with some notes I prepared in the ‘ Proceedings’ of the

Society (P. Z. 8. 1905, p. 244).

A photograph taken by Mr. W. P. Dando, F.Z.8., when the

young Giraffe was a few days old, is reproduced in text-fig. 18.

For the general proportions of the body it 1s interesting to compare

it with the figure of a Giraffe foetus published by Mr. Beddard

(P. Z. 8. 1906, p. 626, text-fig. 107) and with the photograph of the

young female Nigerian Giraffe about a year old published in my

own paper (P. Z. 8. 1905, text-fig. 50). The neck is relatively

longer in the new-born calf than in the fetus, but not so long in

proportion as in the year-old specimen, although in the latter it

had not nearly attained the excessive proportions of the adult.

The long neck of the Giraffe, therefore, is a character that appears

late in ontogeny, as, no doubt, it appeared late in phylogeny.

The frontal horns are well-formed, divergent, large, and covered

at the extremities with very long tufts of hair. Here, however,

ig an instance where phylogeny and ontogeny cannot be regarded as

parallel. It is difficult to suppose that the horn-shaped skin

protuberance was developed in the past history of Giraffes before

the bony core, and yet at birth the core was very much shorter

than the skin protuberance with its tuft of hair, so that the

“horns” could lie almost flat against the head.

1908. ] A YOUNG KORDOFAN GIRAFFE. 131

The colouring and pattern of the skin was well marked in the

new-born Giraffe, the general appearance being a slightly closer

approach to the reticulate type than to the blotched type of its

parents, the edges of the dark fawn-coloured patches being

sharper, and the white reticulum being brighter and better defined.

As in the parents, the front of the face is fawn-colour, the colour

extending more widely than in the Nubian and Nigerian Giraffes,

there being only a rim of white round the eyes; whilst in the other

two forms, and especially in that from Nigeria, the facial

fawn-colour is a relatively narrow band, leaving the sides of the

Text-fig. 18.

Female Kordofan Giraffe about one week old.

face pale. There is a dark spot between the nostrils similar to

that present when the Nigerian Giraffe was younger, but which in

the latter is not now noticeable. The sides of the face have fawn-

coloured patches, larger and more numerous than in the Nigerian

form (text-figs. 19 & 20). The parallel wrinkles on the face, to

which Sir Ray Lankester recently directed attention (P. Z.S. 1907,

p- 115 and text-figs. 42 & 43), are similar to those in the parents.

Sir Ray Lankester informs me that, in examining the Giraffe in

question a few days after birth, he thought there were traces

132 ON A YOUNG KORDOFAN GIRAFFE, [ Mar. 3,

visible of a set of parallel stripes on the forehead between the

horns and the eyes, similar to those figured by him on a coloured

drawing of the fetal Giraffe (P. Z.8. 1907, pl. v.). No doubt

these are ephemeral, for neither Mr. Goodchild nor I could detect

their presence a few weeks later. The interior of the ears shows

dark markings arranged in three distinct pencillings.

Text-fig. 19.

Wit

4 —~ Ww = e “

INS aia. c\\ ae

LAG BO gy ®t G

: SS ve AlN ~ Wy a

= ay \ @ x

Zs ys

Head of Kordofin Giraffe calf.

The coloured patches on the back of the neck (text-fig. 20, B) in

the region just between and below the ears resemble those of the

parents, and differ markedly from those in the corresponding

region in the Nigerian Giraffe. As Mr. Lydekker first pointed

out, and as I was able to confirm from another example, the

Nigerian Giraffe in that region has the broad large series of

blotches extending up almost to the level of the insertion of the

ears, whilst in all other Giraffes a set of much smaller spots

extends considerably lower down.

The white patch which extends round the front of the. throat

Text-fig. 20.

son *

N Sw"

SSS

, Autti!

Ww \ \

say

AWAY!

aft 22 =

Wy) EZ

nN SS=

< 7

yr.

Back view.

Head of Kordofan Giraffe calf.—A. Side view.

134 ON A YOUNG KORDOFAN GIRAFFE. [ Mar. 3,

in the adult, appearing as if a white muffler were tied round the

neck to the roots of the ears and forming a most conspicuous

pattern in both the parents of this young Giraffe, is present, but

1s much less conspicuous.

Along the length of the neck, and especially in front, the large

blotches are much more quadrangular, relatively larger, and more

closely set together than in either the parents or the female

Nigerian Giraffe. Those on the dorso-lateral areas of the neck

are also more numerous and more closely set together than in the

parents or in the Nigerian Giraffe, and are markedly asymmetrical

on the two sides.

On the sides of the body the blotches of the young Kordofan

Giraffe resemble those of its parents in being rather more

numerous and irregular and relatively smaller than in either the

Nubian Giraffe (G. ¢. typica) or the Nigerian form. I cannot quite

follow Mi. Lydekker, however, in the stress he lays on the coiora-

tion of the upper parts of the limbs. He states (Joc. cit. p. 206) that

the Kordofan Giraffe is “‘ easily distinguishable ” from the Nubian

form by the ‘ circumstance that in the fore-limb from just above

the line of the abdomen, and on the hind-limb halfway up the

thigh, the spots suddenly break up intoa series of very small spots

of irr egular size and shape, similar spots occurring on the under-

parts and inner sides of the limbs.” The inner sides of the limbs

and the underparts have small spots as in the parents, and it is

just possible to describe the existence of a sudden change in the

size of the spots on the hind-limb, but hardly so in the case of the

fore-limb; whilst in these respects it is quite impossible to separate

the young Kordofan Girafte from the young female Nigerian form.

I notice, moreover, that in the female Nubian Giraffe, of which

Mr. Lydekker gave a coloured plate (loc. cit. pl. x.), there is a

nearly sudden change in the spots. The lower parts of the legs

are nearly colourless, as in the parents and in the Nubian and

Nigerian forms.

This young Giraffe then, on the whole, conforms with its parents

in the characters selected by Mr. Lydekker as distinctive of the

Kordofan race, but certainly does not furnish additional evidence

for the discrimination. It will be interesting to see if as it grows

it comes to assume these characters more definitely. Whilst at

first sight it resembles the young Nigerian female, a close com-

parison shows that in general tone it is a darker animal with more

numerous, smaller, and more closely-set spots, and that it agrees

with its parents, and does not agree with the Nigerian form, in the

characters separating the Kordofan and Nigerian Giraffes.

1908.] ON THE ANATOMY OF SNAKES OF THE GENUS CORALLUS. 135

2. A Comparison of the Neotropical Species of Corallus,

C. cookii, with C. madagascariensis ; and on some

Points in the Anatomy of Corallus caninus. By FRANK

EK. Bepparp, M.A., F.R.S., Prosector to the Society.

[Received February 7, 1908. |

(Text-figures 21-27.)

e :

CoNTENTS. Page

(1) A Comparison of Corallus cookii with C. madagascariensis ...... 135

(2) Some Notes upon the Anatomy of Corallus caninus .................. 1654

(1) A Comparison OF CORALLUS COOKII AND

C. MADAGASCARIENSIS.

Some little time since* I contributed to the ‘ Proceedings’ of

this Society notes upon the visceral anatomy of the Madagascar

Tree-Boa, Corallus madagascariensis. This particular species is

the only one out of five or six species which inhabits Madagascar.

The rest are all Neotropical in habitat. The distribution of this

genus Corallus is therefore remarkably like that of the allied

genus Boa, of which there are also both Neotropical and Madagascar

species. This very remarkable distribution renders it particularly

desirable to scrutinise carefully the structure of the species of

these two genera, which inhabit two such mutually distant regions

of the world. It is possible that both these instances may prove

to be parallel to the case of Solenodon and Centetes, which were

at one time thought to be more nearly allied than systematists are

now inclined to allow, and I shall give reasons for believing that

the two Species of Corallus are different in structure. Dr. Gadow

observes T that Boa dumerili and Boa madagascariensis, ‘ both

of Macedo cannot be separated from the genus Boa.”

Mr. Boulenger, in the British Museum Catalogue of Serpents {,

places Corallus madagascariensis alone in a special subdivision

of the genus on account of the comparative length of its tail;

but he is unable to find in external characters any features

of sufficient weight to justify its generic separation from the

remaining serpents placed by him and by others in the same

genus. I am able in the present communication to lay some

facts before the Society which bear upon this matter; and, as I

have had the opportunity during the past year of dissecting no

less than seven examples of an American species, Corallus cookii,

I am able to say something about the variations shown in the

anatomy of this snake; the fact that I have been able to check

* “Contributions to the Knowledge of the Vascular System, &e., in Ophidia,”’

P. Z.S. 1906, p. 516.

+ Cambridge Nat. Hist. vol. viii. Ainph. & Rept. p. 602.

~ Catalogue of Snakes. vol. i. p. 99.

136 MR. F. E. BEDDARD ON THE ANATOMY | Mar. 3,

my observations by the dissection of so many examples allows

perhaps of a more reliable comparison of the two species, though

it must be remembered that I only dissected one example of

Corallus madagascariensis.

The seven examples of Corallus cookii which I have dissected

were of different sizes and show among themselves a certain

amount of variability in structure which will be dealt with in

the following pages. For the convenience of reference, in the

course of my remarks I shall refer to them as Specimen A, B, &e.

Their lengths were as follows (the lengths given, it should be

observed, are of the body proper, that is to the cloacal aperture.

I did not trouble to measure the tail) :-—

A. 2 64 inches. B. ¢ 63 inches. C. ¢ 484 inches. D. o

71 inches. EK. ¢ 65 inches. F. 64 inches. G. 53 inches.

Of the last two examples I did not ascertain the sex. I shall

now deal with certain organs and systems seriatim.,

S$ External Characters.

In distinguishing the various species of Corallus the one from

the other in his Catalogue, Mr. Boulenger makes use of a large

number of characters, but does not avail himself of the number

of scales which border the mental groove. This character happens

to be of some use in the determination of the two species with

which I deal in the present communication. I ought to say,

first of all, that a faint doubt hangs over the identification of

specimen A. In this individual I did not ascertain definitely

that it belonged to the species Corallus cookii by counting the

number of scales in the rows upon the body. I do not think,

however, that there is really much danger of a mistake having

been made in its identification; for in every other character

which I examined it agreed broadly with the other individuals

in which I did carefully count these scales. The numbers quite

agreed with the figures given in his Catalogue by Mr. Boulenger.

As to the scales which border the mental groove, I found in

Corallus madagascariensis that the number on each side was

eight. In Corallus cook there are not less than nine and not

more than eleven of these scales on each side, the prevailing

number being ten. There is thus a difference, though doubtless

a slight one, between the two species. The greater length of the

mental groove in Corallus cookii gets perhaps some additional

significance as a character of value from the fact that there is

precisely the same difference, as I have already recorded*,

between the two species of Anaconda, viz., Hunectes murinaus

and #. notwus. In the former there are seven scales on each

side and in the latter oniy five. The same kind of difference

also distinguishes two species of the genus Boa; for in one

individual of Boa divinilogua (a male) the mental groove was

very short indeed and bordered only by two scales on either side,

* P.Z.S. 1906, p. 15.

1908. | OF SNAKES OF THE GENUS CORALLUS. 137

while in another example (a female) by one scale only; whereas

in its close ally, Goa constrictor, I found in one individual three

scales on either side of the mental groove. I may take this

opportunity of mentioning that I found in three examples of the

Boid Enygrus carinatus five or six scales on each side of this

groove. The number of scales bordering the chin-groove,

although apparently useful in discriminating species, does not

help in the division of the Boidz into the two groups Pythoninze

and Boine ; for in Python spilotes I found seven scales on one

side and eight on the other side of the groove, and in P. regius

four on each side. In examples of Python sebe and P. molurus

there were eight of these scales on each side.

As to the anal claws, I have unfortunately no observations

upon these in Corallus madagascariensis ; but in Corallus cookii

they were present and similar in both sexes, being sharp and

curved. Hach was ensheathed in two or occasionally three bract-

like scales, which were situated on the concave side of the claw.

In this latter feature Corallus appears to agree rather with

Hunectes than with Boa; for in the latter genus the claws are

ensheathed (in both sexes of Humnectes noteus) by two scales,

while in both Boa constrictor and B. diviniloqua the female snake

possesses a small sharp claw which does not appear to be ensheathed.

§ Lungs.

Although, as I shall take occasion to point out immediately,

the lungs of Corallus cookii present certain variations in structure

in the individuals of that species, they all agree to differ from

those of Corallus madagascariensis in one important particular.

The latter species on the whole resembles the genus Goa, in that

the bronchus of the larger of the two lungs extends for a long

way into its interior as a shallow gutter; in fact, to a point beyond

the commencement of the liver. I may take this opportunity

of recording the fact that in Hryx conicus*, a representative

of a genus undoubtedly belonging to the Boine as contrasted with

the Pythonine section, the larger lung is also provided with a

very long intrapulmonary bronchus. J have found the same in

the Indian Hrya johni. In an individual of the former snake

I found that the bronchus was continued down to a point corre-

sponding with certainly the end of the first third of the liver,

and that it reached the region of the lung, where it ceased to be

vascular. In this snake I noticed a further peculiarity in this

bronchial gutter: the “ gutter” itself was of course in free com-

munication with .the interior of the lung, as the word used to

describe it implies; but, in addition to this, a series of small

perforations, quite numerous and regularly arranged in pairs, put

the cavity of the gutter into communication with that part of the

lung which was covered over by the semirings of the bronchial

* Tn Cuvier’s ‘ Lecons d’Anatomie Comparée,’ ed. 2, t. vil. 1840, p. 187, the same

is stated of “ Hria turcicus” (= Eryx jaculus).

138 MR. F, E, BEDDARD ON THE ANATOMY [ Mar. 3,

gutter. The exact disposition of the bronchus and its relations

with the lung requires further study in these animals. In all

Pythons, on the other hand, which have been examined from

this point of view, the lungs agree with each other to differ

from those of Boa, Hryx, and Corallus madagascariensis in the

shortness of the intrapulmonary bronchus. This varying dis-

position of the bronchus would appear therefore to be of generic

value at least in some Boide ; it becomes therefore a matter of

some importance to note that in Corallus cookii the conditions

observable in Corallus madagascariensis are not repeated, but

that the intrapulmonary bronchus of the former snake is short.

Thus the two snakes, although belonging, in the opinion of many,

to the same genus, show a point of difference in the structure

of the lungs which in other Boide is at least of generic value.

In Corallus madagascariensis there is not, as it would appear,

any trace of the tracheal lung. In Corallaus cookii, on the other

hand, the extension forward of the soft tissues of the lung

between the separated ends of the semirings of the trachea. is,

for a short distance, invaded by lung-tissue. The cellular

arrangement is quite visible. It is not uncommon among snakes

for the semirings of the bronchus to be continued upon the lung

as a fibrous seam, which seems to mark a shrinkage of the

bronchus itself; this is the case with the lung of Corallus

madagascariensis. It must be borne in mind that this seam is

not a structure differing in any essential from the internal walls

of the lung. It is simply produced by a coincidence in direction

of the walls of a number of lung-alveoli. That there is this

coincidence, and in a longitudinal direction immediately following

upon the end of the intrapulmonary bronchus, would seem to

indicate the gradual obliteration of a formerly more extensive

intrapulmonary bronchus. The shrinkage of this through nar-

rowing and finally obliteration would, in my opinion, leave a

straight lime with pulmonary alveoli on each side, the alveoli

being in consequence symmetrically arranged. This is, at any

rate, a possible explanation of the phenomenon.

Another view might be held which will be explained directly.

While Corallus madagascariensis is characterised by the possession

Explanation of Text-figs. 21 & 22 (opposite).

Text-fig. 21—Larger lung of Corallus cookii cut open to display the entrance

thereinto of the trachea.

a. Forward extension of the lung lying dorsally to the trachea. 67. Aperture

of bronchus of smaller lung. ¢7./. Tracheal lung.

Text-fig. 22.—The larger lung of another example of Corallus cookii, to be

compared with that represented in text-fig. 21.

a, Forward cxcal extension of larger lung. 7.2. Smaller lung, below which is seen

its forward cecal extension projecting to right of trachea. s. “Seam” running

along lung. ¢r./. Tracheal lung.

OF SNAKES OF THE GENUS CORALLUS. 139

1908. |

Text-fig. 22.

Text-fig. 21.

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140 MR. F. E. BEDDARD ON THE ANATOMY [ Mar. 3,

of this seam running down the lungs for some distance after the

intrapulmonary bronchus has ceased as a series of cartilaginous

bars, Corallus cookii has at most a small representative of this

seam, which appears to be in some specimens actually absent.

There is thus in any case a difference between the two species.

It is necessary, however, to describe in greater detail the con-

dition of the lungs in the specimens which I have dissected.

In specimen A. there was no seam at all; in B and D there was

a distinct seam; in EK, F, and G, I could find no seam. Of C,

T have no observation upon the point. It would appear there-

fore that in Corallus cookw the seam is at least beginning to

disappear, while there 1s no evidence that that is the case with

C. madagascariensis, though the latter data being based upon one

specimen are obviously not so secure.

The variations in the lungs of Corallus cookit do not, however,

stop with the presence or absence of the longitudinal seam. The

accompanying figures (text-figs. 21 & 22, p. 139) illustrate the

anterior end of the lung in two individuals of Corallus cookit, and I

have had drawn for comparison with them a corresponding portion

of the lung of Python spilotes, in which snake there is a very con-

spicuous seam continuing the bronchus down the lung (text-fig. 23).

There is, however, in Python no vestige of a tracheal lung and the

intrapulmonary bronchus is quite short. The lungs of Corallus

cookit figured are from the specimens D and E; one shows a seam

and the other does not. Although in the lung which was removed

from specimen E there was certainly not the faintest trace of a

seam to be noted when the lung was examined in a fresh condition,

appearances indicative of such a seam were to be seen in that

lung after it had lain for some little time in a solution of formol.

It occurred to me therefore, on a re-examination of the lung after

it had been thus prepared, that some error must have crept into

my notes. The matter is of obvious systematic importance, as

will be pointed out later on in connection with the variation of

another organ (the gall-bladder), and so a careful study of the

lung was made. I found that the appearance of a seam in this

lung was in truth only an appearance; in fact, a contraction

produced by the formol had manifested itself along the line of

junction of the two lungs, and here it was that the seam appeared.

But on pulling apart the surfaces of the lung the “seam” was

observed to disappear at once and to be therefore not a permanent

structure, but merely a wrinkle produced by the contraction due

to the formol. On the ether hand, no amount of pulling apart

would obliterate the real seam of the second lung figured, that

of specimen D, Here, therefore, was a real seam, a permanent

structure. It is possible, however, that these appearances, due

to contraction, may be an indication of how the seam really arose

in the first place. A contraction along the line of fusion of the

right and left lungs might possibly become permanent in the

course of generations. With regard to the seam, it is noteworthy

and quite apparent in the figure that it did not begin immediately

1908. | OF SNAKES OF THE GENUS CORALLUS. 141

Text-fig. 23.

4 fc0" ate

Larger lung of Python spilotes, for comparison with text-figs. 21 & 22.-

The commencement of the liver is shown to the left of the figure.

Lettering as in text-figs. 21 & 22.

142 MR. F. E. BEDDARD ON THE ANATOMY | Mar. 3,

with the cessation of the intrapulmonary bronchus*. It is first

visible at some little distance beyond this point and extends back

only for an inch or so. In comparing the larger lung in these

two examples of Corallus cookit, another important difference will

be apparent from the figures. The larger individual measured

71 inches, it will be remembered, from snout to cloacal orifice,

the smaller 65 inches. The difference in length between these

serpents is not therefore very large; one might fairly speak of

them as being about the same size. It will, however, be noted

from the accompanying figures, which are drawn to the same

scale, that the lung of one of the examples is very much

larger than that of the other, the difference being much more

pronounced than would perhaps be expected in two snakes so

nearly of the same length. It would, I should think, be imagined

by anyone examining the drawings referred to, that one snake

was half the size of the other. One would hardly expect to find

any difference in the size of the lungs in tie two individuals ;

and yet there is the difference described and figured. As I have

contrasted the lungs not only in these two individuals but in two

others, F and G, measuring respectively 64 and 53 inches, and

which both possess small lungs like those of specimen H, I can

venture to draw the conclusion that the lungs do certainly

vary considerably in capacity within the limits of this single

species. Furthermore, the tract of bronchus which lies within

the lung is by no means equal in the two specimens (D and KE).

In the one with the smaller lung there were, so far as I could

count, 16 bronchial semirings belonging to the intrapulmonary

bronchus, and the length of the intrapulmonary bronchus was

17 mm. In the snake which had the larger lung there were

certainly three or four more semirings, and these rings were

distinctly narrower than in the other specimen, where their

breadth from side to side was clearly greater. The intra-

pulmonary bronchus of the larger specimen measured 31 mm,

In specimens F and G the intrapulmonary bronchus measured

respectively 11 and 10 mm. In specimen B, the only other one

of those in which I identified the presence of a seam at or near

the end of the intrapulmonary bronchus, I have a note that

the intrapulmonary bronchus is at least 25 mm. long. There

is thus a considerable variability also in the intrapulmonary

bronchus ; but it will be observed that the variations in the lung

tend to arrange themselves into two series. In the one the

intrapulmonary bronchus is long and a seam is present; in the

other the bronchus in the same region is short and there is

no seam. These facts seem to point to two stages in the

disappearance of the intrapulmonary bronchus.

§ Arterial System.

The arterial system, so far as I have examined it, also shows

* T find in a newlv born Hunectes noteus a similar gap between the end of the

bronchus and the beginning of the seam.

1908. | OF SNAKES OF THE GENUS CORALLUS, 143

differences between the two species of Corallus with which I am

concerned in the present communication ; and the difference is

rather remarkable in that it is closely paralleled among the true

Pythons, as I shall point out immediately. In Corallus cookii

the aorta gives off comparatively few intercostal branches. Yet

the intervertebral spaces are served by a regularly paired series

of arteries as in the Boide generally. The arrangement of these

arteries is, in fact, like that of the Boine (as contrasted with the

Pythonine) snakes, in that the few branches arising from the

aorta and passing dorsally are connected together by a series of

secondary longitudinal trunks; from these latter arise, at the

requisite regular intervals, the actual branches which bury them-

selves in the intercostal spaces. This is precisely the arrangement

that would have been expected in a snake belonging to the sub-

family Boine of the family Boide according to the facts which I

have been able to record in various genera of that family. It is,

in fact, Corallus madagascariensis which is thus far “ abnormal,”

for, as I have already recorded, in that snake the disposition of

the intercostal arteries is not after the Boine but the Pythonine

plan. ‘This rather unexpected state of affairs to a certain extent

of course throws doubt upon the value of these characters from a

classificatory point of view. I have laid some stress in former

papers upon the value of the intercostal arteries in the classi-

fication of serpents; but it is at least noteworthy that the

difference in structure goes hand in hand with a totally different

range in space. The Old World Corallus madagascariensis agrees

with the Old World Pythons, while the New World Corallus cookii

agrees with the New World Boas. The parallel instance to which

I have referred above is of course Python spilotes; this snake,

while agreeing with Pythons generally in a number of features

of its organisation, has a system of intercostal arteries which is

not like that of the Pythons generally, but agrees with that of

the Bone. Here, again, the difference goes hand in hand with a

difference in habitat ; for Python spilotes belongs to the Australian

region, while all the other Pythons that have been examined

from this point of view are either African or Asiatic in range.

It would be of the greatest interest to study some of the other

members of the genus Python which occur in or near to the

Australian region, and also the other American members of

the genus Corallus. In the meantime I can only point out these

rather anomalous facts. JI may also mention that in those

individuals where I looked for this artery each kidney had only

one renal artery, as in Boide generally.

§ Venous System.

With reference to the venous system of Corallus I have obser-

vations to make only concerning the umbilical and azygos veins.

The wmbilical vein is, so far as I know from my own observa-

tions, small and rudimentary in the Pythons, but well developed

Proc. Zoou. Soc.—1908, No. X. 10

144 MR. F. E. BEDDARD ON THE ANATOMY | Mar. 3,

in Boa and Hunectes. I could not find this vein in Corallus

madagascariensis. In one specimen of Corallus cookii, on the

other hand, there is a small vein arising from the postcaval just

after the commencement of the liver which I regard as probably

representing that vein. It may be that I have missed it in

OCorallus madagascariensis; and yet I rather fancy that it did

not occur in the specimen which I dissected, for it was naturally

carefully sought for, as being of some importance in classification.

Another example happened to allow of a more accurate descrip-

tion of what appears to be the persistent umbilical vein, more

poorly represented, or, it may be, in a poorer condition for obser-

vation in the other snake. The accompanying figure (text-fig. 24)

shows this vein and its connections. Behind the liver a single

epigastric vein runs along the median dorsal line in close contact

with the body-wall, as in all snakes (in which it has been looked

for). Passing forward, this vein divides into two, at first divergent

and afterwards parallel, veins at a point about half an inch or so

in front of the posterior end of the liver. The left-hand of these

two veins continues to bear the characters of the epigastric vein.

It emits branches to the liver, which cross over or rather under

(as the animal is viewed on a dissection) the other vein. These

branches enter the liver and form a part of the hepatic portal

system ; there are altogether five of these, the last entering the

liver quite close to its anterior termination. On the other hand,

the second of the two divisions of the at first single epigastric

vein runs above, but gives off no branches to, the liver. This

can be seen most plainly, and, as I think, is an anatomical fact—

and not merely due to the vessels being invisible owing to their

emptiness of blood. Close to the anterior end of the liver this

vein receives a branch from the parallel epigastric vein, which

runs obliquely backwards from that vein to join the parallel

trunk and at this point bends downwards to reach the liver.

On the surface of that organ it dilates somewhat and narrows

again to be connected with the postcaval vein by at least a

membranous seam in which I did not observe with certainty

any blood. It may be pervious, but I am not quite determined

upon the point. There are clearly reasons for looking upon this

vessel as the persistent umbilical vein. If this be admitted, then

it follows that in this character also the American species of the

genus Corallus shows likenesses to the Boine rather than to the

Pythonine subdivision of the Boidee.

There is therefore in this feature also a difference between the

New World and the Old World representatives of the genus

Corallus, and, moreover, on the whole it may be said that the New

World species resembles more the Boine than the Pythonine section

in this particular. There is, however, by no means a close resem-

blance between Corallus cookit and the Boas in the disposition of

the umbilical vein, as will be seen by a reference to my descriptions

of that vein in Goa and Hunectes. That there should be differ-

ences between individuals of Corallus cookit is not surprising in

1908. ] OF SNAKES OF THE GENUS CORALLUS. 145

view of the fact that the vein is, as it would appear, a vein in

course of disappearance among the Reptilia.

Text-fig. 24.

The liver and certain veins connected with it of Corallus cookii.

a., epigastric vein; w.v., umbilical vein; v.¢.2., postcaval vein.

On the whole the azygos vein appears to be a more extensive

vein among the Boide than among the Colubrine Snakes. I have

146 MR. F. E. BEDDARD ON THE ANATOMY [ Mar. 3,

quoted in other papers communicated to the Society facts from

my dissections relative to the shortness of this vein in both of

those two great divisions of the Ophidia. With regard to the

Colubrine Serpents I have now two other examples to mention

which bear out the view expressed above. In Helicops angulatus

the azygos is smaller than in any snake in which I have seen it.

A stoutish vein on the right side connects the parietes with the

precaval, and nothing further is seen of it after the point at which

it emerges from the parietes ; it is not at all continued down the

body In Heterodon nasicus the azygos is also small, but not

smaller than in some other Colubrines in which I have de-

seribed it. The vein, which is, as usual, on the right side of the

body, is formed by the confluence in regular order of four inter-

costal branches. The last of these emerges from a circular orifice

in the parietes with well-marked edges of very much greater

diameter than the vein which it permits to pass out. In the

case of the other branches no such orifice was visible; they

simply push their way between the various layers of tissue.

In the Boide on the whole the azygos is better developed than

in the non-Boine snakes, but in no Boid which I have had the

opportunity of studying is that vein so continuous for so many

segments as in Corallus cookii. 1 only observed this in one

specimen which happened to be particularly favourable for this

examination. Furthermore, the arrangement was the same on

both sides of the body, and the appearance presented by these

veins was therefore reminiscent of the postcardinals in the

Tailed Amphibians as figured by Hochstetter*, and as I have

myself observed. I believe that there are no embryological

data as to the significance of these veins in the Boide, and I

therefore use the term ‘‘ azygos” as being in the present state of

knowledge a somewhat vague term with nevertheless a definite

meaning of a kind §, but one which implies no homologies with

other vertebrates. As I have no facts about the same veins in

Corallus madagascariensis, and as our knowledge of this vein

in the Ophidia generally is not extensive, I can make no com-

parisons of value except, indeed, to point out that Corallus is, on

the whole, like other Boide in this respect.

S Liver.

Corallus madagascariensis showed a curious peculiarity in the

structure of the liver. The posterior end of one of the lobes of

that organ was enormously prolonged as a thin tail of hepatic

substance. ‘This state of affairs is merely an exaggeration of the

commonly found projection of the one of the two liver-lobes

riorly b 1 the other. In Corall kit there is

posteriorly beyond the other. In Corallus cookii there is some

* Morph. Jahrb., Bd. xii. 1888, p. 119.

i In Amblystoma tenebrosum.

t There is not always a strict homology,.as I believe, even between veins in

different mammals which have been described under this name.

§ T.e., a vein which draws blood from the dorsal thoracic parietes.

1908. | OF SNAKES OF THE GENUS CORALLUS. 147

variability in the form of the posterior termination of the liver

(text-fig. 25), but in no specimen that I have dissected was there

anything at ali approaching the long thin appendage of Corallus

madagascariensis. In one specimen, where this prolongation was

particularly well developed, it only measured ? of an inch in length,

which is far below that of the hepatic process of its presumed

congener. ‘The prolongation, in fact, in Corallus cookii is hardly

an exaggeration of what is found in those other serpents in which

‘the one lobe does extend further backwards than the other.

Text-fig. 25.

Posterior end of liver of two individuals (a & b) of Corallus cookii.

a shows a long tail-like process.

‘Out of the six other examples of Corallus cookii which I examined

three had a more or less well-developed tail to the liver and three

had the barest traces of the same. The two species thus contrast,

it being of course assumed that the one example of Corallus

madagascariensis which I examined represented the normal in

structure.

§ Pancreas and Spleen.

I take these two organs together, because in the Reptilia

(especially in the Ophidia) there is generally a pretty close

connection between them. In Corallus madagascariensis the

spleen is a small rounded body situated at some distance from

the pancreas, which is a solid compact gland abutting on to the

duodenal walls. In Corallus cooki the pancreas is of a different

form, always assuming that I have made no error in the descrip-

tion of C’. madagascariensis. In any case it is certain that in

C. cookii (for I have identified the various structures to be

148 MR. F. E. BEDDARD ON THE ANATOMY [ Mar. 3,.

described in five individuals) the pancreas (text-fig. 26) consists

of a large solid piece abutting upon the duodenal walls, as in the

other species of the genus with which it is here compared; but

in addition to this there is a thin isthmus of pancreatic tissue

connecting the major half of the gland with a round piece of pan-

creatic tissue closely attached to the spleen; the latter is of course.

Text-fig. 26,

Pancreas and adjacent viscera of Corallus cookii.

a. Commencement of coiled region of intestine. . Main mass of pancreas ;

p’., process of the same attached to spleen. spl. Spleen.

to be distinguished from the pancreas by its purplish colour, the:

pancreas being yellowish. In one example of Corallus cookii

(specimen A) the spleen, instead of being a rounded body, is.

thicker at one end and tails off into a long thin process at the

1908. OF SNAKES OF THE GENUS CORALLUS. 149

other. In specimen E the spleen has traces of this peculiar form

in that the rounded spleen closely attached to the nearly detached

piece of the pancreas is reinforced by a splenculus situated three-

quarters of an inch from it, which latter tails off into a thin

process. In the other specimens the spleen is a single rounded

body as I have described it in Corallus madagascariensis. The

pancreas would appear therefore to be pretty constant in its

characters in Corallus cookii. This form of pancreas seems to

be characteristic of the family Boide among the Ophidia. I find

in Hryx conicus a practically identical arrangement; so also in

Python sebe*. I have already described as characteristic of

Hunectes murinus t that the pancreas is divided into two portions,

of which one is apposed to the duodenal wall and the other is

attached to the distantly situated spleen. The duct of the gland

connects these two parts. It is possible that in that snake there

is also an isthmus of pancreatic tissue lymg along the duct; in

any case there is certainly this isthmus in the other serpents

to whose pancreas I have referred above. In Python spilotes

much the same structure of these organs is to be seen; and,

moreover, in that Python, as in Hunectes murinus, the spleen is

reinforced by additional splenculi, seen also rarely in Corallus

cookit. On the other hand, Boa constrictor differs in that the

spleen is close to, but not much in contact with, a solid pancreas

unprovided with any discrete fragment and connecting isthmus.

The organs in Goa constrictor are, in fact, constructed on the

plan of those of the non-Boine snakes, where, however, the

connection between spleen and pancreas is sometimes rather

close. In Vipera russelli the smaller spleen sits like a cap upon

the larger pancreas. In Dasypeltis scabra the two “ glands ” are

in absolute contact, and in Helicops angulatus the spleen is

actually imbedded in the wider distal end of the pancreas,

appearing upon its surface in the form of two patches. Von

Siebold and Stannius, in their well-known and usually (so far as

my own experience goes) accurate and comprehensive text-book of

comparative anatomy t, do not sum up accurately the relations

of the pancreas and the spleen when they write: “ Die Milz ist

getrennt vom Pancreas bei Lryax, Python, Chersydrus, nur wenig

ihm anhiingend bei Boa. Die Verbindung ist inniger bei den

ubrigen Schlangen.” These observers would seem to have missed

the process of the pancreas so common among the Boide. The

fact that it is common in that group, and not, as it would appear,

found elsewhere, has some significance. For among the Lacer-

tilia there is very generally a long and thin process of the

pancreas which comes into actual contact with, or at least gets

* In another example of Python sebe the distal portion of the pancreas was

completely separated from the larger duodenal portion by an actual break in the

connecting isthmus, bridged only by the pancreatic duct.

+ “Contributions to the Anatomy of the Ophidia,” P. Z.S. 1906, p. 25.

~ ‘Handbuch der Zootomie,’ Bd. 11. Amphibia, 1856, p. 187. It is perhaps

remarkable that Cuvier and Milne-Edwards have no observations upon these peint=

in their text-books.

150 MR. F, E. BEDDARD ON THE ANATOMY [ Mar. 3,

very near to, the more distantly situated spleen. As the Boide

are believed, by reason of a considerable number of anatomical

peculiarities, to stand at the base of the Ophidian series, this fact

is obviously not without importance.

§ Relative Position of Viscera.

The relative position of the viscera among snakes and their

proportionate lengths are important as diagnostic characters.

Thus, in the Viperine Serpents the liver follows close upon the

heart, and in the Boide the kidneys have rather a different

position from that in the case of other Ophidia. It is not, there-

fore, without importance to compare the two species of Corallus

in these particulars.

C. madagascariensis. C. cookit.

Total length to vent ..................... 50 inches. 64 inches.

From sy see sis of jaws to auricles

ot heart.. Sages SEES ooo ee 2 18+ inches.

From apex of heart to beginning of

Na Vel ee ce Se oe ten nase aban ee cee INCHES: 4$ inches.

Hen sbhyvom live rseeeeeesee Aone eee Ness 11 inches.

Distance between end of liver and

GIMME VEEGISD soe che ssoneusseg sous saa usgees. | ay HNO 8 inches.

en othvolgkidneys eee 1$ inch. 3x & 25 inches.

Distance of end of posterior kidney

from Vent ...........4...0.e0.-s-000.-..... About 5-6 inches, 10 inches.

It is obvious from the above measurements that there are

differences between the two species, which cannot by any means

be accounted for by the differences in length which they show as

individuals. Thus one kidney, at any rate, of Corallus cookii is

twice the length of that or rather those of Corallus madagas-

cariensis, although the length of the snake is only one-fifth or

one-sixth greater. Furthermore, the distance of the kidneys

from the cloacal orifice in C. cookii is greater, both actually

and proportionately, than in C. madag gascariensis. This Boine

character is therefore more emphasised in the American than in

the Madagascar species. Although the liver of C. madagas-

cariensis 1s slightly longer than that of the American species,

this does not account for the great discrepancy which the two

species show in the distance of the gall-bladder from the end of

the liver. In Corallus cookii the two kidneys hardly overlap.

The posterior kidney begins where the right kidney ends. I find

that here, as in the case with Corallus madagascariensis and other

Boidz, the kidney of each side is furnished with only a single

renal artery. This structural fact can, as I think, be now

regarded as characteristic of the Boide as compared with many

other snakes.

1908. | OF SNAKES OF THE GENUS CORALLUS. 151

In view of the rather slender differences which often exist

between different genera among the Ophidia, it appears to me

that the differences here recorded between Corallus madagas-

cariensis and C’. cookii are worthy of receiving generic value.

But for the present I do not consider the matter of a name

for the Madagascar species, since there is no knowledge of the

structure of the remaining American species of the genus.

The following brief statement embodies the principal ana-

tomical characters of the two species, Corallus madagascariensis

and C’. cook :—

Corallus madagascariensis.

Mental groove bordered by eight scales on each side. Bronchus

continued for a considerable distance into larger lung. ' Intercostal

arteries regularly paired like those of Python*. . No rudiment of

umbilical vein. Liver commences closer to apex of heart, of

considerable length, terminating in a very long thin posterior

prolongation of one lobe. Gall-bladder close to end of liver.

Pancreas with no prolongation to spleen. Kidneys short, nearer

to vent.

Corallus cookii.

Mental groove bordered by ten scales on each side. Bronchus

continued only for a short distance into interior of larger lung.

Intercostal arteries not regularly paired like those of Boa. A

rudiment of or a more fully-developed, umbilical vein. Liver com-

mences further from apex of heart, of less considerable length,

sometimes terminating in a short thin prolongation of one lobe.

Gall-bladder at a considerable distance from end of liver. Pancreas

with a prolongation to spleen. Kidneys longer, further from vent.

For the measurements made above for the purpose of con-

trasting the two species of Corallus, I was compelled to select one

of the larger specimens of Corallus cookii dissected by myself.

For though I have dissected smaller examples of the same length

as the individual of Corallus madagascariensis, the details as to

the position of the viscera were unfortunately imperfect in my

notes. From such details as I have preserved, however, it would

not appear that much has been lost by not being able to make a

more accurate comparison between the two species of snake in

these particulars. Since the examples of Corallus cookii which

I have examined range from a length of 484 inches to 71 inches,

it becomes a matter of interest to note what, if any, alterations

take place in the relative sizes of the various organs and their

position with regard to each other during the process of growth.

There is, furthermore, another rather important variation to be

deduced from these studies, as will be apparent when the facts

* Exc, Python spilotes (see above, p. 143).

152 MR. F, E, BEDDARD ON THE ANATOMY [ Mar. 3,.

have been set forth. These facts are given in the accom-

panying table, which is regrettably imperfect in places.

ae Bona ean) E. ‘S|

(64 inch.)* (63) | (484) (71) | (65) |,(64) | (63) |

From symphysis of jaws to | |

auricle of heart.............| bt | 18 | 14 | 19h | 2 (18 | 17 |

| From apex of heart to liver ... 43 Ae Senos: P A tp |

| Tberaealln Gre WAGE! Sossconovesovopec0| LIL GE Is 8 {| 114 | 102 |104) 8%

| From end of liver to gall-

bladder: sa:csredyecsoenesec sso 8 54 | 5s P 8 6 62)

| | |

| Length of kidneys ............... Sere Gano | 2h | Ging) |e 2 |b

| |

From end of posterior kidney | | |

PORVGMU: Macatee Menten sec ce 10 Nel al 123 | 12 P| 6

The study of the statistics presented in the above table shows,

as it would appear, only one important instance of variation

between individuals. That concerns the distance at which the

gall-bladder is situated from the end of the liver. In specimens

A, E, F, and G, particularly in the first and second of the four,

the gall-bladder is markedly more distant from the end of the

liver than in the remaining examples where I have noted its

position. In view of its much smaller length, specimen C should

perhaps be referred to the same list. It is a fact not without

interest that in specimens H, F, and G the intrapulmonary

bronchus is not continued by a seam down the lung, and that the.

lungs are relatively small. On the other hand, in the only

examples in which I ascertained the presence of a seam, viz., in

B and D, one, at any rate, has a large lung, and the other shows

a short interval between the end of the liver and the gall-bladder.

There is thus, as I venture to maintain, a commencing differentia-

tion of species among these externally quite similar snakes.

There is a distinct tendency to the segregation of the individuals

into two groups: the one with a seam down the larger lung,

which is relatively of larger size than in the other individuals,

and a gall-bladder comparatively far removed from the posterior

end of the liver; the other with relatively smaller lungs without.

any seam, and the gall-bladder not so far removed from the end

of the liver. Apart from these series of variations, which seem

to allow of the sorting into two lots of the individuals considered,

other organs vary in a less regular fashion. In the liver there

* Tt will be recollected that the above measurements are taken from the top of

the snout to the vent. They are set forth in inches.

+ This and all the following measurements are in inches.

+ The thin “tail’’ to one lobe of the liver is comprehended in these measure-

ments, where 1¢ occurs. It does not occur in specimens D, H, G.

1908. | OF SNAKES OF THE GENUS CORALLUS. 152

may or may not bea moderately long and thin posterior prolonga-

tion of one of the lobes; the intrapulmonary bronchus also is:

longer and shorter in some specimens than in others; the spleen

is by no means entirely constant in its form in every individual.

The position of the kidneys varies slightly, as does the amount of

overlap of one by the other. The table showing the relative

position of the ditterent organs of the body indicates that there-

is, as might be expected, a correspondence between the lengths

of the snakes and the distances separating the viscera. The

table, however, does not show satisfactorily where the growth of

the individual takes place, except that it would perhaps be

inferred that it is not in the region following the liver; for in

two individuals measuring respectively 63 and 483 inches the-

distance from the end of the liver to the gall-bladder is precisely

the same. I observed this carefully in the two individuals F

and G. There is a difference of ten or eleven inches in total length

between the snakes, and yet the distances between the end of the

liver, the spleen, and the gall-bladder were exactly the same, as

was conclusively shown by placing the snakes side by side, when

the organs mentioned were seen to correspond exactly in position.

This seems to show that the growth in length takes place both in

front of and behind this particular region of the body. I should

imagine that this region corresponds to the umbilical region of

the new-born snake. In conclusion, it may be pointed out that

the comparative fixity in the relative position of organs in so

many specimens and of different lengths confirms the use of this.

character as of systematic value in snakes,

$ Conclusions.

It will be clear that the characteristics of these two species of

Corallus, even if they be ultimately referred to two distinct,

genera, breaks down my previous attempts to define the sub-

tamilies Boine and Pythonine by anatomical characters, and

renders those characters only generally applicable. Hitherto

these two subfamilies have been distinguished by the existence in

the Pythonine of a supraorbital bone wedged in between the:

prefrontal, frontal, and postfrontal bones; this bone is wanting

in the Boine. I presume that Corallus has been examined in

this respect, and has been found to possess no supraorbital bone.

Otherwise the Boine (and Corallus cookii) agree to differ from

the Pythonine (including Corallus madagascariensis!) in the

mode of distribution of the intercostal arteries, and in the

persistence of a considerable umbilical vein. J am disposed to.

think that in view of facts accumulated since the division of the

Boidee as indicated above, it is not so desirable for the present to

insist upon any such subdivision, which is, after all, by no means

in accord. with geographical range. On the other hand, the.

present communication enables me to substantiate still further

the characteristics of the family Boidee, which differs from all other-

154 MR. F. E. BEDDARD ON THE ANATOMY [ Mar. Dp

families of Ophidia, whose characters are well known, by the

following distinguishing features :—

Vestiges of hind limbs present. Two lungs (except in Ungalia).

Broncht entering lungs at a litile distance from their anterior

extremity. Aorta giving off regularly-paired intercostal arteries

or regularly-paired intercostals derived from fewer branches of

aorta. Renal artery single on each side (rarely two). Umbilical

vein of ‘foetus often persistent in adult to a greater or less extent.

Anterior abdominal vein connected with afferent renals. Azygos

usually long. Postcardinals present in front of kidneys. Pancreas

usually with splenic lobe.

These characters are, for the most part, points of likeness to

the Lacertilia, which are more numerous in the Boide than in

other snakes. ‘They thus argue the basal position of the Boide

in the Ophidian series—a view which is quite generally held.

(2) Some Norges upon THE ANATOMY OF CORALLUS CANINUS.

The comparisons instituted in the above account of the structure

of Corallus cookii are further confirmed by the dissection of an

example of Corallus caninus, which I have been able quite lately

to examine. The specimen of this extremely beautiful species of

Tree-Boa had been, as I understand, identified when alive by

Mr. Boulenger. I identified it myself when dead, before learning

this fact, with the help of that gentleman’s ‘ Catalogue of the

Snakes in the British Museum’*. The catalogue in question,

though allowing of an easy reference of the individual to its

proper species, does not refer to what appears to me to be the

most salient external character of the species when compared

with its immediate ally Corallus cookii; that is, the much greater

size of the scales in the latter species. In Corallus cookii, in fact,

a scale upon the dorsal surface near to the middle line measured

4mm.in length; a scale froma corresponding position in C’. caninus

measured only 2mm. ‘The pitting of a much greater number of

the labial scales than in C. cookii is another character of this

species. I found that in the individual examined by myself the

first lower labial of one side was slightly pitted. This does not

appear to be always the case. The mental groove is longer in

this snake than in ©. cookit. It is bordered by fifteen scales on

each side. Thus there is an exaggeration of the characters of

C’. cookit, and not an approach towards C. madagascariensis. On

the other hand, the short tail of C. caninus (the length to the

cloaca was 34 inches, that of the tail 52 inches) is an approach

to the condition observable in C. madagascariensis. The anal

claws were small, and, on account of their white colour, incon-

spicuous. Hach lay between two scales in front and one behind,

and the area of implantation was rather bare.

I have given reasons for regarding the position of the viscera

* Vol. i. London, 1893, pp. 99 & 102.

1908. ] OF SNAKES OF THE GENUS CORALLUS. lias

within the body of serpents generally as of systematic impor-

tance. The following are the corresponding facts relating to

Corallus caninus :—

Length of body to cloaca, 24 inches.

From snout to heart, 9 inches.

From heart to liver, 33 inches.

Length of liver, 53 inches.

From end of liver to gall-bladder, 14 inches.

These measurements, though incomplete *, indicate a closer

agreement with Corallus cookii than with the Madagascar species,

as will be evident from a comparison with the tables on pp. 150

& 152. At the same time, there is perhaps some approach to the

latter in the comparative closeness of the end of the liver to the

gall-bladder. Moreover, the liver in Corallus caninus has a long

thin ‘ tail,” measuring no less than an inch and a half. This,

again, is a slight approach to Corallus madagascariensis, inasmuch

as the tail of the liver in both species is about one fourth of the

length of the entire liver, while in (. cookii there is evidently a

tendency for this thin posterior prolongation of the liver to

disappear. This, however, is nearly the only point in the internal

anatomy of Corallus caninuws in which I have ascertained a

likeness to the Madagascar species at all more pronounced than

to Corallus cookii. In examining the liver I naturally sought

with care for vestiges of the umbilical vein. These are quite

obvious, and I am even inclined to think that this vein is fairly

well-developed in the present species, though its condition did

not permit of an accurate mapping of the course of the vein.

However, a strongish vein—the further course of which along the

body-wall I am unable to state—dips down from the body-wall to

the liver. Arrived upon the surface of the liver its calibre becomes

increased, as 1s often the case with the umbilical vein in the Boine

snakes ; but, instead of joining the postcaval, it appears to end in

the liver-substance. But its former connection (¢) with the post-

caval is indicated by a furrow deeply marked which connects the

vein with the postcaval, continuing in the same straight line as

the vein would traverse were it to pass directly to the postcaval.

I think, therefore, that this vein is the persistent umbilical and

not merely a branch of the epigastric. I may point out that, if

this opinion is well founded, the present species of Corallus agrees

with its American congener.

As I have already pointed out, the Madagascar and the South

American species of Corallus differ in the relations of the

pancreas to the spleen. I am bound to state that Oorallus

caninus seems to be more like Corallus madagascariensis than

is to Corallus cookii in these matters. That is to say, there is no.

thin process of the pancreas putting the pancreas into actual

contact with the anteriorly lying spleen. On the contrary, the.

* It was desirable to injure the snake as little as possible.

156 MR. F. E. BEDDARD ON THE ANATOMY [ Mar. 3,

rounded pancreas lies upon the gut, and the small spleen lies at

a considerable distance from it and is without any connection

with it. The spleen is rounded, distinctly divided into two lobes,

but with no tail-like process. On the whole, however, it does

not appear to me that these differences have generic value. For

I have pointed out precisely similar differences between the two

species of Anaconda *.

Two important anatomical features remain for consideration,

in both of which Corallus caninus is more like its South American

Text-fig. 27.

Lungs of Corallus caninus cut open to show interior.

try, Trachea. 7. Right lung; the irregular bronchial semirings within this lung

are shown on the right near to the septum between itself and the left lung.

ally. ‘These are the arrangement of the intercostal arteries and

the structure of the lungs. As to the former, Corallus caninus

has the Boine disposition of the intercostal arteries. That is to

say, there are but few of the arteries arising directly from the

* “Contributions to the Anatomy of the Ophidia,” P. Z. S. 1906, p. 25. I may

add that Eryx conicus and EL. johni also differ in this; for in the latter there is no

“tail” to the pancreas. As to the former see above, p. 149.

1908. ] OF SNAKES OF THE GENUS. CORALLUS. 157

aorta ; but these few bifurcate and trifurcate, and from secondary

longitudinal connections arise the actual intercostals. This is

obviously like Corallus cookii.

The lungs of Corallus caninus are illustrated in the accom-

panying figure (text-fig. 27). It is obvious that they are,

generally speaking, more like those of Corallus cookii than they

are like those of the Old- World species Corallus madagascariensis.

This is chiefly seen in the short length of the intrapulmonary

bronchus. There is, however, no tracheal lung. The inter-

annular membrane is readily to be distinguished from the tissues

of the lung by its white colour and thick appearance. It has no

lung-like structure; there is no appearance of alveoli. It is,

therefore, fairly sharply marked off from the lung-tissue with

which it iscontinuous. The bronchus of the larger lung descends

into that lung for a distance of about 14mm. This distance is,

it will be observed, much the same as that found in the case

of Corallus cookvi, In the smaller lung the intrapulmonary

bronchus is only 6 mm. long. In the case of the larger lung I

counted 15 bronchial semi-rings. This, again, is about the

number found in Corallus cookii. It is possible that here, as in

Corallus cookw, there is some variability in the structure of the

lung. But as I have only one example, I can merely point out

that it agrees with one set of specimens of Corallus cookii. The

intrapulmonary bronchus is rather broad; it was 8 or 9 mm.

across in the case of the larger lung a little way down, and the

intervening pulmonary tissue only 6 mm.across. It is important

to notice that no trace of a seam could be detected. The intra-

pulmonary bronchus of this snake does not, as will be apparent

from the figure referred to (text-fig. 27), form a regular gutter

with a series of regular and similar annuli. On the contrary,

‘the annuli are very irregular after the commencent, and are fused

together and sometimes incomplete. They were to be distin-

guished by their very red colour in the example dissected. This

irregularity is much more marked than in Corallus cookit. In

describing above the lung of Hrya* I have drawn attention to

the series of orifices into lung-cells ranged along the sides of the

bronchus and between the annuli. In the present species this

condition is much more marked, and serves to distinguish the

species from Corallus cookii. As will be seen in the figure, these

orifices are numerous and scattered irregularly between various

annuli. The differentiation between lung and intrapulmonary

bronchus is therefore less marked in this species than in many

snakes, and is by no means typically Ophidian. It recalls, in

fact, very distinctly the lungs of such a reptile as Heloderma rt.

There is a final point in the structure of the lungs of this

serpent which demands attention, though it does not bear upon

the main object of the present communication to the Society, viz.,

the relationship of the Neotropical to the Madagascar species of

* Supra, p. 137.

+ See Beddard, P. Z. 8. 1907, p. 61, text-fig. 16.

158 MR. R. I. POCOCK ON A [ Mar. 3,

Corallus. As will be observed in the figure of the lung of Corallus

caninus, the headward running pocket of the lungs occupies a

different position in the two lungs. In the case of the larger

lung it will be seen that the anterior process of the lung in

question is partly roofed by the interannular membrane of the

trachea as well as by the bronchial semirings. On the other

hand, the same anterior process of the shorter lung has no rela-

tion whatever to the interannular tracheal membrane. I do not

know how far this is to be seen in other two-lunged snakes

(Boidee). I hope to look into this matter further, since it has an

obvious bearing upon the degree of degeneration of the smaller

lung, and therefore upon the phylogeny of the group.

The following brief statement of the anatomical characters of

the species will serve for a comparison with the corresponding

definitions of C’. madagascariensis and C’. cookii upon p. 151.

Corallus caninus.

Mental groove bordered by 15 scales on each side. Bronchus

continued only for a short distance into interior of larger lung ;

its rings become irregular and interrupted by orifices into lung-

tissue. Intercostal arteries like those of C. cookii. A rudiment of

umbilical vein persists. Liver with long posterior prolongation of

hepatic substance. Gall-bladder rather near to end of liver.

Pancreas with no prolongation to spleen.

The agreement with Corallus cookvi is obviously much closer

than with C. madagascariensis.

3. Description of anew Species of Monkey of the Genus

Cercopithecus. By R.1I. Pocock, F.L.S8., Superintendent

of the Gardens.*

[Received February 18, 1908.]

(Plate X.7)

The Zoological Society has recently acquired by purchase from

Mr. F. C. Thorpe, of Hull, a young example of a species of Cerco-

pithecus quite different from any form hitherto described. At.

Mr. Thorpe’s special request I dedicate this new species to

Mr. Ezra, of Calcutta.

CERCOPITHECUS EZR& Pocock.

Abstr. P. Z. 8. 1908, p. 10 (March 3).

Skin of upper part of face black ; that of the lips and chin blue,

clothed with white hairs. Head ornamented in front with a

* [The complete account of the new species described in this communication

appears here; but as the name and the preliminary diagnosis were published in the

< Abstract,’ the species is distinguished by the name being underlined.—Eprror. |

+ For description of the Plate, see p. 160.

NEW SPECIES

J-Green delet Chrome

Oe CINCO MMsis CUS ,

1908. ] NEW CERCOPITHECUS MONKEY. 159

conspicuous rusty-brown brow-band like that of C. neglectus,

but melting posteriorly into the general colour of the rest of the

head, and not sharply defined by a jet-black band. The area in

front of the ears slightly tinged with rufous-brown, but the rest of

the head, the cheeks, neck, and dorsal and lateral surfaces of the

body closely speckled greyish-yellow and black. The outer surface

of the thighs and of the upper arms also similarly speckled, but, like

the cheeks, rather more rufous than the body. On the shoulder and

upper arm there is a lighter stripe such as is seen in C. neglectus, and

there is a corresponding stripe on the outer side of the thigh, but

this is not nearly so conspicuous as the thigh-stripe of C. neglectus,

being of a dirty yellow and not white. The lower arms are dark

brown and finely speckled externally, but not jet-black as in

C. neglectus, whitish stained marginally with yellow on the inside.

Hands and feet somewhat rubbed, scantily clothed with white

hair. Tail not black, but covered with speckled hairs, resembling

in tint those of the body, rufous at base. Hairs on the throat

white, but not forming so long a beard as in C. neglectus. Chest

and belly greyish with a decidedly rusty tinge. Belly and inside

of the thighs white, but the hairs beneath the callosities and on

the backs of the thighs not white but tinged distally with rusty

red.

Loc. Unknown, probably Upper Congo.

. This species is evidently related to C. neglectus, as is testified by

the red-brown band, the whiteness of the lips and chin, and the

presence of the pale stripe on the thigh, not .to mention the

corresponding stripe on the upper part of the arm and the

general colour of both the upper and the under side. It might

almost be briefly diagnosed as a species of Cercopithecus dittering

from C. neglectws in lacking the black areas characteristic of the

latter. The two may be contrasted as follows :—

a. The red area of the forehead sharply defined posteriorly by

a jet-black band crossing the summit of the head from side

to side; hairs beneath the callosities white ; tail, with the

exception of a couple of inches at its base, jet-black.

neglectus.

a. The red area on the forehead not sharply defined behind, but

blending with the speckled tint of the top of the head,

which has no black transverse band; hairs beneath the

callosities not white but tinged with rusty yellow; tail not

black, but the same colour as the back ............ ezre.

Up to the present time C. neglectws has occupied in the genus

an isolated position, and opinions as to its affinities have differed.

By Drs. Sclater, Forbes, and Trouessart it was associated with

C'. diana on account of the presence of a beard of longish hairs on

the throat and chin, and of a white stripe on the outside of the

thigh. But in the monograph of the genus Cercopithecus recently

published by this Society (P. Z.8. 1907, pp. 677-746), I put

Proc. Zoot. Soc.—1908, No. XT. lit

160 ON A NEW CERCOPITHECUS MONKEY. | Mar. 3,

forward the opinion that the species must be regarded as an

aberrant type of the Lewcampyx-group (p. 685). The correctness

of this view is shown by the discovery of C’. ezr@, which may be

said to occupy an intermediate position between C. neglectus, on

the one hand, and the Abyssmian Monkey described as C. boutowr-

linii, which I consider to be a subspecies of C. lewcampyx, on the

other.

The only very young example of C. neglectus that I have seen—

namely, one from the Cameroons—resembled the adults in colour.

It is to be regretted that the exact locality of C. ezre is at

present unknown, but since it came to Hull in company with an

example of C. ascanius schmidti, its geographical area is probably

the Upper Congo. The species is so distinct from even its

nearest allies that there can be no difficulty, I think, in its

identification when other specimens come to hand to be named.

In the P. Z. 8S. 1907, p. 739, I described a new species of

Cercopithecus from the Congo under the name nigroviridis. Since

the figure of the head of this species represented on pl. xlil.

fig. 5, and taken from a dried skin, is inaccurate in two particulars,

namely, in showing the chin black and in omitting the narrow

black streak running back from the corner of the eye, I take this

opportunity of figuring the whole animal from an example now

living in the Gardens.

IT also figure the type of that remarkable new species said to

have come from the Ituri Forest and described by myself in Ann.

Mag. Nat. Hist. (7) xx. pp. 521-522, 1907, as Cercopithecus

hamlynt. The animal is still living in the Gardens, and owing to

change of coat and improvement in condition has altered some-

what in colour. The brownish-yellow speckling of the hairs has

changed to greenish-yellow. There is no pale brow-band except

such as is indicated by the grey bases of the anterior hairs. The

arms are for the most part black except for a speckled area below

the elbow on the outerside. The wrist and ankles, like the hands

and feet, are black.

EXPLANATION OF PLATE X.

Fig. 1. Cercopithecus nigroviridis Poc.

2. Cercopithecus ezre@, sp. n.

3. Cercopithecus hamlyni Poe.

(The attitudes are taken from photographs of living animals.)

1908. | ON A MALFORMED PLAICE. 161

March 17, 1908.

Dr. Henry Woopwarp, F.R.S., Vice-President,

in the Chair.

The Secretary read the following report on the additions made

to the Society's Menagerie during the month of February

1908 :—

‘The number of registered additions to the Society’s Menagerie

during the month of February was 171. Of these 146 were

acquired by presentation and 7 purchased, 9 were received on

deposit and 9 by exchange.

The number of departures during the same period, by death

and removals, was 189.

Among the additions special attention may be directed to :—

One male Orang Utan (Simia satyrus), from Deli, Sumatra,

presented by Dr. J. C. Graham on Feb. 5th.

A male and two female Pronghorn Antelopes (Antilocapra

americana), from North America, presented by President Roose-

velt on Feb. 4th.

A. Pallas’s Cat (Felis manul), from Tibet, deposited on Feb. 29th.

A Vaviegated Spider-Monkey (Ateles variegatus), a Red Brocket

(Cariacus rufus), and a collection of 63 Birds, including one

Scarlet Cardinal (Cardinalis phaniceus), four Desmarest’s Green

Tanagers (Calliste desmaresti), two Olive Saltators (Saltator

olivascens), and five Sonini’s Colins (Hupsychortyx sonini), new to

the Collection, from Venezuela, presented by A. Pam, Esy., F.Z.S.

Mr. G. A. Boulenger, F.R.S., V.P.Z.S., exhibited a remarkably

malformed Plaice (Pleuronectes platessa) from the London market,

which had been presented by the Secretary of the Fishmongers’

Company, Mr. J. Wrench-Towse, to the British Museum.

The fish measures 375 millim. from the end of the snout to the

end of the caudal fin, and for the first 245 millim. of its length is

perfectly normal in form, scaling, fins, and coloration. But after

that a sudden change takes place. The point where the change

takes place is indicated on the eyed or coloured side by a rather

deep chink in the dorsal and ventral (or left and right) outlines,

accompanied by a sudden rise in the length of the dorsal and anal

rays, which at that point are separated by a wider membrane ;

after this, however, the rays gradually decrease again in length

towards the caudal peduncle, which, as well as the caudal fin, is

perfectly normal. There is no solution of continuity in the

arrangement of the scales or in the course of the lateral line. The

only other difference, on this side, from a normal specimen, is a

slight raising, like a, median swelling, of the vertebral region

immediately preceding the caudal peduncle.

On the blind side, the aberrant aspect is much more striking.

At the point mentioned, ¢.¢. 50 millim. in advance of the caudal

162 MR. G. A. BOULENGER ON A | Mar. 17,

peduncle, the colourless lower surface is squarely truncate, and the

truncated portion, 45 millim, in diameter, is fringed with a series

of rays, 7 in number, connected by a membrane which forms a

continuum with the dorsal and anal fins. Just in front of this

truncature, the lateral line divides into two branches terminating

at the base of the abnormal fin. Behind the truncated part, which

Coloured side of malformed Plaice, caudal end.

is thin and contains bony supports to the fin-rays, and separated

from it by a deep groove, the normal region reappears, normal

but for the fact that the two sides are scaled and coloured exactly

alike, instead of the lower side being colourless and clad with

smaller scales.

Mr. Boulenger suggests as an explanation of this wonderful

1908. | MALFORMED PLAICE. 163

“freak” that, at an early stage of its existence, the fish lost the

posterior part of its caudal region, at the point now indicated by

the truncature on the blind side; it at once proceeded to repair

the injury by producing a secondary terminal fin, viz., the trans-

verse fin connecting the dorsal with the anal—similar structures

‘IIIV[q PoUlLoJ[VUI Jo apis pug

being well known in the regenerated tails of some fishes. But at

the same time, or soon after, the true tail asserted its rights, and

grew again, alongside the secondary fin, and this regeneration

would fall under what Prof. Giard has designated as ‘ Régéné-

rations hypotypiques” (C. R. Soc. Biol. iv. 1897, p. 315), an

164 MESSRS. 0, THOMAS AND R. C. WROUGHTON ON [Mar. 17,

atavistie phenomenon to which Mr. Boulenger was one of the first

to draw attention (P. Z.8. 1888, p. 351).

As Prof, Giard has defined it, hypotypical regeneration represents

a sort of abridgment of the processes of reintegration, by which

the regenerated part corresponds, not to the state of stable equi-

librium prevailing in the given form or type, but toa previous state

of equilibrium, usually to the maximum of stability immediately

preceding that of the present state of evolution. As flat-fishes

are undoubtedly derived from symmetrical forms in which both

sides are similarly coloured and scaled, the present case, if correctly

interpreted, is another to add to the numerous examples of this

phenomenon which have been furnished by various groups of the

animal and vegetable kingdoms.

Mr. Walter A. Kidd, M.D., M.R.C.S., F.R.S.E., F.Z.8., gave

an account, illustrated by diagrams, of observations he had made

regarding the effects of pressure on the direction of hair in

Mammals.

The following papers were read :—

1. The Rudd Exploration of South Africa —IX. List of

Mammals obtained by Mr. Grant on the Gorongoza

Mountains, Portuguese §.E. Africa. By OLpFiELp

THomas, F.R.S., F.Z.8., and R. C. Wrovueuron, F.Z.8.

[Received February 27, 1908. ]

In order to make the collection from the Pungwe River district

of Portuguese S.E. Africa more complete, Mr. Grant spent some

weeks in the Gorongoza Mountains, and there made the present

collection, which forms a useful supplement to the Beira series, of

which we gave an account in our last paper *.

The series consists of about 150 specimens, belonging to 31

species and subspecies, of which we have described three as new,

while many are rare forms of which the additional material will

be of the utmost service as the different groups are worked out.

The following are Mr. Grant’s notes on the place he worked

“'Tambarara lies on the foothills under the south-western

slope of the Gorongoza Mountains; roughly 300 m. (975 feet)

above sea-level, in a very typical stretch of country. Higher up

collecting was impossible, owing to the height of the grass.

‘“‘ Both the foothills and mountain-slopes are here clothed with

virgin forest, in parts of great beauty.

““ Many of the forest trees are of great sizeand comprise ‘native

mahogany,’ ‘ teak,’ &e., &e.

* P. Z. 8. 1907, p. 774.

1908. ] MAMMALS FROM PORTUGUESE §.E. AFRICA. 165

“The thick forest, however, does not extend far either to the

south or north-west, its place being taken by more open forest

and bush-veldt. The north-east and north of the range are dry

and stony.

“The highest point of the range is said to be 1600 m. (5200 ft.).

“The soil is very fertile, and splendid crops of maize, Kaffir corn,

rice, &c. are grown by the natives.

“The climate is healthy and the temperature never rises very

high, averaging during the trip 79° in the shade; the rainfall is

heavy, and this and the thick mists that often envelop the country

keep the trees in a continual state of verdure,

“The natives are a tribe known as the Gorongozas and many

gave great assistance, my personal ‘ boys’ taking a keen interest

in furthering the work,

“Throughout the trip collecting was difticult, owing to the density

of the grass and other undergrowth, which made it impossible to

form a really complete series.”

1. Papio CYNOCEPHALUS Geoff.

Gig AUSha, Qesotls\70)-

“Native name, ‘ Nkerua’ (‘Bongwi’ towards the Zambesi).

“ Eixcessively common everywhere and generally in large

troops.

“At the time the crops are ripe it is quite impossible to scare

them away even if several are shot.

““ Apparently not as pugnacious as the southern species, although

dogs are often killed by them. The natives say that the ‘old

men’ will sometimes turn on them.

“The flesh is eaten by the women and children, but seldom by

the men.”

2. CERCOPITHECUS ALBIGULARIS BEIRENSIS Pocock.

oe 1902. 592. 1802; 1803, 1818.

‘“ Native name, ‘ Nsimbo.’

“Very common in the forest, to which they are confined, and

often observed in very large troops.

‘“ At first they were tame and specimens were easily obtainable,

but they soon became wild and after a few weeks were seldom

seen.

“ Living on wild fruits, young shoots of trees, &c., and seldom

damaging the natives’ crops.”

3. CERCOPITHECUS PYGERYTHRUS RUFOVIRIDIS I. Geoff.

3. 1819, 1880, 1917, 1936. ©. 1842.

“Native name, ‘ Ukoro.’

“Not quite so common as the last species, but still very

plentiful and often observed in large troops.

‘“When the native crops are ripe they visit the lands and do

considerable damage.”

166 MESSRS. 0. THOMAS AND R. C. WROUGHTON ON [ Mar. 17,

4, GALAGO GRASSICAUDATUS E. Geoff.

3. 1886, 1889, 1899, 1934. 2. 1887, 1888, 1890, 1891,

1892, 1897, 1898, 1901.

“‘ Native name, ‘Gwea.’ |

“Very common in the forests, where they appear to consort

together in small parties.

“The species has a variety of calls, none of which, however, are

similar to that of Galago garnetti.

‘‘ Apparently principally vegetarian, and feeding largely on the

exudation from the bark of certain trees.

“Strictly nocturnal, passing the day im hollow trees.”

5, GALAGO GRANTE Thos. & Wrought.

©. 1905, 1906, 1907, 1911, 1912, 1913.

““ Native name, ‘ Konsiti.’

“Common ; habits similar to those of its congener at Inhambane

and elsewhere.”

6. ScoropHILus NigRirA Schreb.

6. 1919, 1920, 1921, 1922. ©. 1923, 1924.

We follow the precedent we established in dealing with the

Tnhambane and Beira collections, and provisionally accept these

specimens as S. nigrita, merely noting that they all belong to the

larger form.

‘Native name, ‘ Nyagelingwelingwe.’”

7. CROCIDURA sp.

1903.

“ Native name, ‘ Nyungeyunge.’”

8. PETRODROMUS TETRADACTYLUS Pet.

3G. 1824, 1844, 1845, 1857,1877,1895,1914. ©. 1840, 1841,

1846, 1847, 1893, 1894, 1896.

“‘ Native name, ‘Gumbwa.’

‘¢Common in the thickest part of the forest and having regular

runs in which they are easily trapped.”

9, GENETTA sp.

g. 1839, 1900, 1910, 1935, 1944. ©. 1885, 1925.

This is very possibly zambesiana, though it does not altogether

agree with the short description given by Prof. Matschie.

‘“¢ Native name, ‘ Mulimba.’

“Common, especially near kraals, where they steal quantities

of fowls.”

10. MuNnGos PALUDINOSUS RUBELLUS, subsp, n.

ey 926.

A bright rufous form of the Marsh-Mungoose (/ungos palu-

dinosus). ie

Before describing the new subspecies, however, we have to

1908. | MAMMALS FROM PORTUGUESE S8.E. AFRICA. 167

show why we use the name paludinosus for the Marsh-Mungoose

itself.

The name pi has hitherto been supposed to date from

Erxleben *, whose pr imary basis for the name was the “ Galera” of

Brown’s Ji amaica’ T, which is no doubt the Marsh-Mungoose. But

unfortunately Schreber’s plate t of ‘‘ Mustela galera Brown,” is

one year earlier, and instead of being really based on Brown’s

Mungoose, is a.mnere coloured copy of Buffon’s figure $ of the

** Vansire” of Madagascar, which, as Gray suggested in 1864 ||, is

certainly the animal usually known as Galidia elegans I. Geof?.

Consequently this latter must bear the name of (ralidia galera

Schreber, and another name be found for the Marsh-Mungoose.

The next names on the list, Viverra nems and Mustela afra of

Kerr 4], seem to be equally inapplicable, when critically examined.

The first is based on the ‘“‘Nems” of Smellie’s Buffon, which

appears to be some form allied to the common Indian Mungoose,

while the second is again primarily based on the Madagascar

“ Vansire ” (Galidia FUEED and should be considered fg =

synonym of that animal.

Next among the synonyms usually quoted comes Jchnewmon

major K. Geoftroy **, but this is based on Buffon’s “ La Grande

Mangouste” TT, which there is no reason to consider as anything

but a large individual of Mungos ichneumon.

Then follows the “ Vansire” (Atilax vansire) of Geoffroy and

F. Cuvier $f, which is undoubtedly the Marsh-Mungoose, but

though the figure and description date from 1826, the technical

name was only. given to the animal on the appearance of the

General Index in 1842.

Before this latter date there was published G,. Cuvier’s name

Herpestes paludinosus $$, which antedated Smith’s Mangusta

urinatriz ||| by one month, and appears to be the tenable title of

the animal under consideration.

The description of the new form is as follows :—

About the size of typical I. paludinosus.

Upper surface of body clothed with a close, soft fur, about

15 mm. long, under a pelage of long (50-60 mm. on the back)

hairs, somewhat harsher than in paludinosus.

General colour above near ‘‘tawny ochraceous,” obscurely

mottled with black; individual hairs, basal + slaty marked with

four rings of dirty whitish at inter vals, distal z made up of 10mm.

tawny and tip (5 mm.) black ; fur slaty. On the flanks and belly

* Syst. R. A. p. 453, 1777. + P. 4835, pl. xlix., 1756.

{ Saug. i. pl. 135, 1776 (quoted by Erxleben).

§ Hi st. Nat. xiii. p- 167, pl. xxi., 1765.

Il P. Z. S. 1864, p. 523.

€ Linn. Syst. Nat. pp. 160 & 175, 1792.

** Descr. Egypte, Hist. Nat. u. p. 139, 1812.

++ Hist. Nat. Supp. iii. p. 173, pl. <XVi, 1776.

tt H. N. Mamm. ii. pl. 198, 1826.

§§ Régne Anim. (2) i. p. 158, April 1829.

|| Zool. Journ. iv. p. 437, May 1829.

168 MESSRS. 0, THOMAS AND R. C. WROUGHTON ON [Mar. 17,

tawny, bases of individual hairs buffy ; fur asabove. Face to the

nape greyer than body, finely grizzled black and whitish ; upper

lip, chin, and throat buffy. Handsand forearm, feet to the ankles,

and chest blackish (forearm and chest grizzled, owing to the

presence of pale rings on the individual hairs). Tail coloured like

the back, except at the extreme tip (50 mm.), where black

predominates.

Skull much as in typical paludinosus; heel of last lower molar

measurably larger than in the Cape form.

Measurements of the type :—

Head and body 462 mm.; tail 375; hind foot 105; ear 36.

Skull: greatest length 102 mm. ; basilar length 85; zygomatic

breadth 50; length of m' 7.

fab. Tambarara, Gorongoza Dist., Portuguese East Africa.

Type. Young female. B.M. No. 8.1.1.53. Original number

1926. Collected 8th May, 1907.

The rich rufous colour of this Mungoose will readily distinguish

it from any of the described forms of J, paludinosus.

11. RayYNCHOGALE MELLERI Gray.

3. 1861.

12. BDEOGALE CRASSICAUDA Pet.

©, 1947.

An examination of this specimen, which we provisionally

identify as Bd. crassicauda, makes it clear that two specimens

received from Mr. J. T. Last from Zanzibar Island represent a

new species, and we take this opportunity to publish a description

of it under the name of

BDEOGALE TENUIS, Sp. 0.

A small, short-tailed form.

Covered above with a dense, soft underfur (15 mm.), mixed with

longer hairs (25-30 mm.), forming an outer coat, scarcely con-

cealing the underfur; longer hairs entirely absent on the belly.

Tail cylindrical, its individual hairs about 25 mm. in length.

General colour buffy clay, washed with black—the former being the

colour of the underfur, the latter that of the longer hairs. On

the shoulders and sides of neck, however, these longer hairs are

white-tipped and thus produce a grizzled appearance. Face, hands,

and feet tawny. ‘Tail black.

Skull small. Teeth proportionally large.

Measurements (those of the body from a dried skin) :—

Head and body (c.) 410 mm.; tail 180; hind foot 70; ear 20.

Skull: greatest length 77 mm.; basal length 69°5; greatest.

breadth 42; upper cheek-tooth series 22; palatal length 41 ;

breadth across m* 25,

Hab. Zanzibar Island.

Type. Old individual (sex unknown). B.M. No. 6.6.5.18.

Presented by the collector, Mr. J. T. Last. Two specimens.

1908. | MAMMALS FROM PORTUGUESE S.E. AFRICA. 169

13. VIVERRA CIVETTA Schreb.

2. 1828.

“« Native name, ‘ Mfungo.’”

14. CRossARCHUS FASCIATUS Desm.

g. 1941. ©. 1810 (juv.), 1942.

“‘ Native name, ‘ Ndembo.’

“Quite common, but not easy to secure.

“« Always observed in troops, often of considerable size. The

troops seem to comprise a great proportion of immature indi-

viduals; in one instance I killed five from a troop with one shot,

four of which were young.

‘“« Living almost exclusively on insects, for which they diligently

search everywhere, uttering continually the peculiar chattering

which generally betrays their whereabouts.

“ Exclusively diurnal.”

15. FuNISCIURUS MUTABILIS Pet.

6. L798; 1868) 1904. 19095 9. 1829; 1869; 1916:

‘“ Native name, ‘ Sindi.’

“Very common in the forest ; always in pairs.”

16. Funiscrurnus ceparr A. Sm.

rei LL). INS TSh.

‘““ Native name, ‘ Konkwa.’

‘‘Fairly common, more so in the open ‘ bush veldt’ than in the

real forest.

‘“‘ Feeding on various seeds and fruits—especially, when in

season, the fruit of the Mafura.

‘“‘ Diurnal only.”

17. TATERA INCLUSA, sp. n.

ic Meme, Oe ils7(9e

A large Tatera, with tail longer than head and body.

Size about as in 7’. draco.

Fur soft but short, about 10 mm. in length on the back.

Ground-colour above ‘“ ochraceous buff,” more or less modified

by black, below white. Individual hairs of the upper surface

dark slate-colour, tipped with ochraceous on the sides, sub-

terminally ringed with ochraceous and tipped with black in the

dorsal region; of the belly white from their bases. Hands and

feet white. No white tip to tail.

Skull narrower than in 7’. draco, with longer nasals and upper

molar series; the upper incisors less deeply grooved, the groove

less central.

The following are measurements (those of the body taken by

the collector) :—

Head and body 163 mm.; tail 184; hind foot 39; ear 27.

170 MESSRS. 0. THOMAS AND R. C. WROUGHTON ON [ Mar. 17,

Skull: greatest length 44 mm.; basilar length 35; zygomatic

breadth 23; length of upper molar series 7; bulle 11.

Hab. Tambarvara, Gorongoza Dist., P. EK. A. Alt. 1300 ft.

Type. Adultmale. B.M. No. 8.1.1.79. Collector’s number 1838.

Taken 20th March, 1907.

The second specimen, an adult female, is rather smaller, but the

hind foot, ear, upper molar row, and bulle are exactly as in the

type. (The collector records ‘‘ Kar 22,” but this is an evident

mistake.)

The appearance of this very distinct form in this region is

interesting. The Gerbils found at Mazoe on the high plateau to

the west and at Beira at sea-level to the south are races of

T. lobengule, while the forms north of the Zambesi are short--

tailed ones. This is apparently a species confined to the

mountainous Gorongoza country and is perhaps allied to 7. draco,

also an animal with a very restricted habitat on the Drakensberg

at Wakkerstroom.

In Wroughton’s key to the species of the genus 7atera (A. M.

N. H. xvii. p. 475, 1906) this species would come next to 7’. afra,

from which it may be distinguished by its longer hind foot and

ears (viz. 39 and 27 to 34 and 24 respectively in 7’. afra), its

longer, narrower skull, and larger teeth.

““ Native name, ‘ Mpynya.’

“ Apparently not common, the soil not being suited to its

habits.”

18. ARVICANTHIS DORSALIS Sm.

3. 1799, 1808, 1816, 1817. 9. 1800, 1804, 1806, 1807, 1811,

1812.

‘* Native name, ‘ Mhoni.’

** Very common everywhere, especially in the native ‘ lands.’

“* Diurnal only.”

19. SaccostoMUS CAMPESTRIS Pet.

6. 1822, 1832, 1835, 1863, 1883.

The series of specimens of the genus Saccostomus, like that

from Beira, seems to contain two forms separable by size, and for

these we adopt as before the names S. campestris and S. mashone.

‘* Native names, ‘ Tuda’ and ‘Suka.’

‘Common in the natives’ ‘lands’ and along the banks of the

streams.

** Nocturnal and strictly vegetarian.”

20. SaccosroMUS MASHON de Wint.

6. 1834, 1878, 1882. ©. 1836.

21. Mus rarrus Linn.

Ge We2oe «OD. 8b3ke

1908. | MAMMALS FROM PORTUGUESE S.E. AFRICA. 171

22. Mus cHRYSOPHILUS de Wint.

Ga leswle Oss:

‘“‘ Native name, ‘ Mpynya.’

“Common; habits as elsewhere in 8. Africa.”

93. Mus coucua A. Sm.

3. 1809, 1813, 1814, 1820, 1823, 1825, 1837, 1853. 2. 1805,

1815, 1831, 1856, 1884.

‘‘ Native name, ‘ Nhingo.’

“Common; habits as elsewhere in 8. Africa.”

24, CRICETOMYS GAMBIANUS CUNCTATOR, subsp. n.

Ge SOI sb 2 186d) 1867 182 Oe 849518645866)

1873.

Size and general characters as in the races viator and adventor

of C. gambianus, between which itis geographically intermediate.

On the upper surface approaching nearest, in coloration, to viator,

but the ochraceous on the flanks and shoulders brighter than in

that form, the white on the face hmited to the upper lip at the

angle of the mouth, not spreading over the cheeks as in adventor ;

white tip less than 4 the length of the tail, as in viator, it is

more than 4 in adventor. On the other hand, the colour of the

back gradually passes into that of the belly as in adventor, very

different to the comparatively sharp line of demarcation found in

viator. Belly-hairs tinged with yellowish, with pale slaty bases,

while in both the other forms they are pure white to their bases.

A patch of pure white on the chest.

Skull and teeth as in adventor.

Dimensions of the type :—

Head and body 370 mm. ; tail 454; hind foot 71; ear 43.

Skull: greatest length 76 mm.; basilar length 64; zygomatic

breadth 36; nasals 31x10; interorbital breadth 11:5; palatal

foramina 9; length of upper molar series 12.

Type. Adult male. B.M. No. 8.1.1.130. Original number

1862. Collected 6th April, 1907.

Hab. Tambarara, Gorongoza Dist., Portuguese East Africa.

Though it appears that cwnctator has certain characters in

common with viator and others with adventor, and that there is

a considerable amount of individual variation in coloration, still

the yellowish tinge and slaty bases of the belly-hairs and the

constantly present pure white breast-patch serve to distinguish

this race from either of its neighbours.

‘* Native name, ‘ Kurubini.’

“Very common, especially in the forest, where it often lives

singly.

‘“‘ Its holes are often placed at the foot of some forest tiee and :

also amongst thick vegetation.

‘“‘ Nocturnal only and considered a great delicacy by the natives,

by whom great numbers are taken,”

172 ON MAMMALS FROM PORTUGUESE S8.E. AFRICA. [ Mar. 17,

95. Acomys SELoUSI de Wint.

1848 (no skull).

‘Native name, ‘ Chedare.

‘‘ Apparently rare at ae toe as only one specimen was

secured or,seen.”

26. PELOMYS FALLAX Pet.

6. 1827, 1843. 9. 1855.

‘‘ Native name, ‘ Bungo.’

“In habits similar to Otomys irroratus, and like that species

found in the thick grass and vegetation bordering the streams.

‘“¢ Diurnal and a vegetarian.”

27. GrorycHus BEIRAZ Thos. & Wrought.

3 . 1860 (juv.).

‘‘ Native name, ‘ Mfusi.’

“‘ Apparently common, but none could be trapped and very few

runs were observed, probably owing to the thick grass and

vegetation.”

28. THRYONOMYS SWINDERIANUS Temim.

do. 1850, 1851, 1875, 1876.

‘“‘ Native name, ‘ Nsensi.’

‘* Common along the banks of the rivers and streams.

‘They do smnnelh damage to the native rice-patches, in con-

sequence of which the natives have to stockade their fields and

continually trap them.”

29. PROCAVIA BRUCEI Gray.

@- 1929, 1930, 1931. 2. 1928, 1932, 1933:

These specimens perhaps represent Hyrax mossambicus of

Peters.

‘¢ Native name, ‘ Imbile.’

“Very common in the kranzes and masses of loose boulders on

the mountain-sides.

““ Habits as the ordinary Dassie.

‘“* According to the natives, this is the only Dassie found in the

Gorongoza Range.

“ Diurnal and feeding morning and late afternoon.”

30. CEPHALOPHUS NATALENSIS A. Sim.

6. 1852, 1874.

““ Native name, ‘ Kutwa.’

“Fairly common, seems more partial to the more heavily

timbered parts of the forest.

“Ts extremely wary and trapping has usually to be resorted to

in order to secure specimens.”

1908. | ON SEROWS AND GORALS. 173

31. CeEPHALOPHUS MONTICOLA Thunb,

3. 1858, 1859. 9. 1881.

“ Native name, ‘ Nyakoro.’

‘‘Common in the forest, and often observed in pairs, but never

more together.

“Generally can only be shot in the early morning and late

afternoon, when they are feeding.

“They are extremely wary, the alarm-cry is a sharp whistle,

almost a shriek.”

32. CERVICAPRA ARUNDINUM Bodd.

1937.

‘“‘ Native name, ‘ Sengo.’”

33. TRAGELAPHUS SCRIPTUS Pall.

3d. 1949 (juv.). @. 1908, 1833. Tambarara.

‘“‘Native name, ‘ Nsome.’ ”

2. Notes upon some Species and Geographical Races of

Serows (Capricornis) and Gorals (Nemorhedus), based

upon Specimens exhibited in the Society’s Gardens. By

R. I. Pococx, F.L.S., Superintendent of the Gardens*.

[Received March 2, 1908.]

(Text-figures 30-38.)

I. Introduction.

Serows and Gorals, which, as I have recently shown t, must be

known scientifically as Capricornis and Nemorhedus, and not as

Nemorhedus and Urotragus or Kemas respectively, are very rare

animals in captivity. Within the last four years, however, the

Zoological Society has had the good fortune to exhibit no fewer

than two well-marked species or subspecies of each of these

generat. The necessity for determining these animals correctly

involved the looking up of a good deal of the literature, especially

the older literature, of the subject ; and this brought to light a

wholly unexpected amount of confusion in nomenclature, both

generic and specific, and not a few mistakes and misconceptions

as to specific characters in recent catalogues and treatises on these

ruminants.

The specimens of the two kinds of Serow (Capricornis) exhibited

* {The complete account of the new species and subspecies described in this com-

munication appears here; but, as the names and preliminary diagnoses were

published in the ‘Abstract,’ the species and subspecies are distinguished here by

the names being underlined.—Ep1rTor. | ,

+ Ann. Mag. Nat. Hist. (8) 1. pp. 183-188, 1908.

£ Not including a young example of a third species of Goral from Nepal which

was presented by the Prince of Wales, but lived only a few days atter arrival.

174 MR. R. I. POCOCK ON [Mar. 17,

in the Gardens came from widely separated localities—one from

Kalimpong, near Darjiling, and the other from Selangor, in the

Malay Peninsula. Identification of the former necessarily entailed

examination of all the available material from the Himalayas,

which resulted in the discovery of one or two undescribed forms.

The Kalimpong specimen itself proves to be a representative of

an unnamed local race, distinguishable from the typical Nepalese.

race described by Hodgson, the name of which has hitherto been

applied to all the Serows of the Himalayas. For the opportunity

to describe a second Himalayan race I am indebted to Major G.S.

Rodon, F.Z.S., who very kindly sent to me the perfect skin and

skull of a specimen shot by himself at Chamba. A third hitherto

unnamed local race from these mountains is exemplified in the

British Museum by a mounted head from Kashmir belonging to

the Hume collection. Finally, the determination of the example

from Selangor showed that it, too, was nameless, since it presented

characters not previously recorded in any race of Serows. These

characters I consider worthy of nominal recognition, when taken

into consideration with the geographical distribution of the

animal.

With regard to the two Gorals (Vemorhedus) the Society has

lately exhibited, these belonged to two very distinct species. One

of the specimens, which is still living, was presented to the

Society by Major G. 8. Rodon, F.Z.S., who brought it from

Chamba; and I am indebted to Major Rodon for kindly sending

to me for examination a series of skins of specimens shot in the

same locality by His Highness the Maharajah of Chamba. The

second specimen came from the mountains of Korea, whence

Gorals have not been previously recorded, and was presented to

the Society by Mr. C. F. 8. Bilborough, F.R.G.S.

Although neither of these Gorals represents, in my opinion, an

undescribed form, their determination revealed some hitherto

unrecorded facts touching the nomenclature, variability, and

distribution of some of the Gorals of North India and China.

II. On the Serows (Capricornis) of the Himalayas, the Malay

Peninsula, and Sumatra.

Genus CAPRICORNIS Ogilby.

Nemorhedus Hamilton Smith, Grifith’s Animal Kingdom, v.:

p- 852, 1827 (in part).

Capricornis Ogilby, P.Z.8. 1836, p. 188; Gray, List Mamm.

B.M. pp. xxvi & 166, 1843; and subsequent works.

Nemorhedus Blanford, Fauna Brit. Ind., Mammalia, p. 512,

1891; Lydekker, Great and Small Game of India, p. 128, 1900,

and id. op. cit. nov. ed. p. 139, 1907.

Capricornis Heude, Hist. Nat. Chinois, 11, pp. 222 & 234, 1894.

Nemotragus + Lithotragus + Austritragus, id. op. cit. iv. pp. 13-

14, 1898.

1908. ] SEROWS AND GORALS. 175

Capricornis Pocock, Ann. Mag. Nat. Hist. (8) i. pp. 183-188,

1908.

The principal external differences between the various kinds

of Serows that have been described are differences of colour of a

very simple kind. They consist, for the most part, in the

substitution of the three tints, black, red, and white, on definite

areas of the body and limbs—that is to say, a part which in one

form is black may be red ina second and white or grey ina third.

For example, the legs below the knees and hocks are white in

C’. sumatraensis rodonti, red in C. s. milne-edwardsi, and black in

C. s. swettenhami. The mane is hoary white in C. s. swmatraensis

and (©. s. argyrochetes, red or mostly red in CO. s. rwbidus, black

in C. s. jamrachi, a mixture of black and white in C. s. robin-

sont, and of black, white, and red in C. s. swettenhami. Similarly

the underside is almost wholly white in C. s. rodoni, and reddish

black in C. s. jamrachi. Finally, the prevailing colour of the

body in C. s. jamracht is black, while in C. s. rubidus it is red.

These three colours, black, red, and white, or a mixture of any

two of them, or of the three combined, are the commonest

variations to occur in domestic mammals. Horses, for example,

may be black, red (bay), or white, or black and white (piebald), or

bay and white (skewbald), when the colours are arranged in patches,

or roan when the coat consists of an intimate mixture of black

and white or bay and white hairs. The passage from one of these

three tints to the others 1s a common and, be the cause what it

may, an apparently simple phenomenon. It is for this reason

that I regard the differences between the various kinds of Serows

as of subspecific and not of specific importance, in spite of the

fact that there is in most cases no actual proof of the existence of

intermediates between the different forms that have been named *.

For the present, at all events, therefore I agree with M1. Lydekker

in considering all the many described forms as belonging to a

single species, which must take the name of the race that was

first made known, namely the one from Sumatra which Bechstein

desevibed as Antilope sumatraensis .

* Tt is the custom with some systematic zoologists to consider an insular form

ipso facto as a species whatever grade of difference it presents from other imsular

forms or from the form from the adjacent mainland, on the grounds that the

discontinuity in geographical distribution involves the non-existence of actual

intermediate types. Were I to follow this course with respect to the Serows, I

should be compelled to separate the Sumatran animal specifically from those from

the mainland of Malacca, while uniting the latter specifically with those from

Burma, China, and the Himalayas. Such a course, however, would, in my opinion,

be a gross contravention of common sense, because it would give a higher systematic

value to the comparatively trivial differences between the Sumatran and Southern

Malayan animals than to the comparatively important differences between the

Southern Malayan and Himalayan animals. In this and analogous cases it is surely

a mistake to make geographical isolation the criterion of the value of a character.

The character should be judged on its own merits and its importance determined by

a study of the extent of the variation to which the particular species or allied species

are liable.

+ Pocock, Ann. Mag. Nat. Hist. (8) i. p. 187, 1908. For the discovery of this

early name for the species I am indebted to Mr. C. D. Sherborn’s invaluable ‘ ladex

Animalium.’

Proc. Zoou. Soc.—1908, No. XII. 12

176 MR. BR. I. POCOCK ON [ Mar. 17,

The genus Capricornis ranges through the Himalayas from

Kashmir eastwards into Southern China and thence southwards

through Burma and the Malay Peninsula into Sumatra.

CAPRICORNIS SUMATRAENSIS Bechst.

Antilope sumatraensis Bechstein, Uebersicht vierfiiss. Thiere, 1.

p- 98, 1799 (based on the Cambing Outan, Marsden’s Sumatra,

ed. 1, p. 93).

Antilope sumatrensis Shaw, Gen. Zool. i. pt. 2, p. 854, 1801.

Nemorhedus sumatrensis H. Smith, Griffith’s An. King. iv.

p- 277, 1827; Jardine, Nat. Libr., Mamm. iv. p. Dit, ols tas Meas,

Antilope interscapularis Lichtenstein, Berlin Mag. vi. p. 165,

1814.

Nec Nemorhedus swmatrensis Blanford, Fauna Brit. India,

Mammalia, p. 514, 1891; Lydekker, Great and Small Game of

India, p. 128, 1900; id. op. cit. nov. ed. p. 139, 1907; id. in

Rowland Ward, Records of Big Game, p. 345, 1907.

The typical race of this species differs from all the known

Himalayan forms in having the mane on the neck and withers

hoary grey and constrasting forcibly with the dark coat of the

rest of the body, and also in having no sharp line of demarcation

in colour between the upper and lower portions of the legs, which

are blackish below the knees and hocks, merely fading to dark

brown upon the fetlocks. The conspicuousness of the mane

obviously suggested the name interscapularis given to this animal

by Lichtenstein. __

It is to be noted that Dr. Blanford must have omitted to look

up the original literature of this species, since he assigned the

name swmatrensis to. Serows from Moupin, Burma, the Malay

Peninsula, and other localities which are quite different from the

Sumatran form; and I have reason to think that, misled by

him, Mr. Lydekker, in the works quoted above, described as

typical C. swmatrensis the Darjiling Serow to which I have given

below the subspecific name jamrachi.

1t may be added that for many years there has been in the

British Museum a subadult specimen of a Sumatran Serow agree-

ing in all essential respects with the examples figured and

described by earlier authors as Antilope or Nemorhedus sumatra-

ensis (sumatrensis). Ib was originally presented to the Zoological

Society by Sir Stamford Rafiles. Inspection of this specimen by

later authors would have saved all the misconception as to the

characters of this race which have been so frequently repeated in

recent literature on the subject.

Subsp. THAR Hodgson.

The Bubaline Antelope, Hodgson, Gleanings in Science, i. _

p. 122, April 1831 (no scientific name).

Antilope thar Hodgson, Gleanings, iii. p. 324, Oct. 1831; id.

P.Z.8. 1833, p. 105. :

Antilope (Neworhedus) thar Hodgson, P. Z.8. 1834, p. 86.

1908.} SEROWS AND GORALS. 177

Text-fig. 30

TERZI.~

Skull of Capricornis sumatraensis thar Hodgs., from Nepal.

(B.M. Reg. 55.12.26.143.) x 4.

A, from the side; B, from below; C, from above.

12*

74)

178 MR. R. I. POCOCK ON | Mar. 17,

Yupricornis thar Ogilby, P. Z. 8. 1836, p. 1388; Pocock, Ann.

Mag. Nat. Hist. (8) 1. p. 187, 1908.

Antilope bubalina Hodgson, P. Z.8. 1832, p. 12.

Nemorhedus or Capricornis bubalinus (in part) of most recent

authors.

Hitherto only one kind of Serow has been distinguished in the

Himalayan area. Of late years this has been invariably but

erroneously cited as Vaemorhedus bubalinus, after the example

set by Blanford in his volume on the Mammalia of British India.

Blanford, however, quite candidly pointed out that the oldest

and therefore the correct specific (or subspecific) name for this

animal was thar. Unfortunately this disregard for the rules of

priority has been followed by authors who succeeded him. The

early literature dealing with this Serow has been here repeated to

emphasise the fact that thar is its proper name.

The only material of this race that I have been able to examine

are four stuffed skins in the British Museum. Two of these

belonged to Hodgson’s collection and came from Nepal. They

are the co-types or syntypes of the race; a third also came from

Nepal; and a fourth, which appears to be inseparable from the

others, was from Sikhim and belonged to Dr. Blanford, who

mentioned it in his volume on Indian Mammals.

The characters of this race are briefly but, for my present

purpose, sufliciently enumerated in the following pages.

Subsp. HuMEr Pocock.

Abstr. P. Z.S. No. 55, p. 12, March 17, 1908.

Distinguishable from the other geographical races of Serows

occurring more to the east in the Himalayas by having the whole

head a uniform pale chocolate-brown without any intermixture of

black. Some black hairs amongst the brown on the anterior part

of the neck. Anterior part of the lower jaw on each side white ;

no white throat-patch.

Loc. Kashmir.

Type. The mounted head of an adult example in the British

Museum formerly belonging to the collection of Mr. A. O. Hume.

Further evidence of the distinctness of the Kashmir form

from the typical C. s. thar is supplied by the skull of a specimen,

formerly belonging to Mr. R. Lydekker, from Pir Punjal in

Kashmir. This skull differs in a number of particulars from two

skulls of (. s. thar obtained by Hodgson in Nepal—particulars

which combine to make the general “facies” of the skulls from

the two localities very different. In the Kashmir skull the cheek-

teeth.are smaller, the palate wider, and the frontals and nasals

more convex. The last-mentioned is the first distinctive feature

that catches the eye upon .a superficial glance at the skulls.

There is, of course, no actual proof that the skull of the type of

this race resembles the skull from Pir Punjal nor that the

eclouring of the Pir Punjal specimen resembled that of the type.

1908. ] SEROWS AND GORALS. gD

TERZ! —

Skull of Capricornis sumatraensis humei Poc., from Pir Punjal in Kashmir.

(B.M. Reg. 88.3.20.16.) x4.

A, from the side; B, from below; C, from above.

180 MR. R. I. POCOCK ON [Mar. 17,

Subsp. RoDont Pocock

Abstr. P. Z.S. No. 55, p. 12, March 17, 1908.

General colour of upper parts of head and body not jet-black

but brownish black, due to the presence of a decided rufous tinge

in the black terminal portion of the hairs. This rufous tinge is

Text-fig. 32.

Flat skin of Capricornis sumatraensis rodoni Poc., from Chamba. Type.

observable even in the hairs forming the mane on the middle line

of the neck and along the spine. ‘The forehead and the summit

of the muzzle are nearly black, but the rufous colour persists as a

narrow ring on most if not all the hairs. On the sides of the

1908. | SEROWS AND GORALS. 181

head below the eye the rufous speckling is much more in evidence,

and it is particularly well marked on the corners of the upper lip

and at the base of the ear in front and up the back of that organ.

Upper lip, lower lip, and chin white; the white from the chin

extending backwards along the jaw and over the interramal area

to the upper part of the throat, where it expands into a con-

spicuous patch. In the centre of the interramal area, behind the

chin, there is a distinct elongated patch of dusky-brown hair, and

behind this patch the hairs of the interramal area and of the

throat-patch are not wholly white but apically infuscate. On

the shoulders and on the sides of the body the rufous area in the

hairs gradually increases in extent, so that the general colour

above the white of the belly is markedly browner than that of the

back. The outer side of the fore leg is rufous brown, intermixed

with white in front, down to a point two inches above the knee,

and the outer side of the hind leg down from the root of the tail

nearly to the hock is also rufous brown, becoming mixed with

white inferiorly. The chest, including the whorls of hair, the

belly, the insides of the thighs and of the upper part of the fore

legs, and the whole of the legs from above the knees and hocks are

milk-white, and sharply defined by their colour from the adjacent

rufous-brown areas. Only on the chest is the white clouded with

a faint brown tinge. The tail, which is triangular and 2 inches

long, is brownish black above, like the back.

The coat is long and shaggy, with a long and copious mane on

the neck. The hairs on the sides of the body measure about

3 inches, and on the dorsal line of the neck about 8 inches in

length. The underfur is plentiful.

The horns are 8 inches long and 4? inches in basal circumference

Total length of skin from tip of nose to root of tail 52 inches,

distance from middle line of shoulder to hoof 34. Tail (skin)

12 inches.

Loc. Chamba.

This Serow differs from examples received from Hodgson from

Nepal, and referable to typical C. s. thar, both in coloration and

the structure of the skull. In the matter of coloration the

difference lies principally in the clean whiteness of the whole

of the under side, and its sharp definition from the rufous-brown

or rufous hue of the sides of the body, and also in the backward

extension of the white of the interramal area on to the throat,

where it forms a conspicuous white patch. Jn C. s. thar the

patch on the throat is absent or represented by a few white hairs,

and these are not continuous with the white running along the

anterior portion of the lower jaw behind the chin. The under

side is not clean white, but dusky brownish grey, the hairs being

whitish at the base and sooty grey distally, and the colour of the

sides of the body gradually blends with that of the belly and

chest. C. s. thar is also apparently a shorter and thinner coated

form, and the underfur if present at all is scanty. Blanford,

indeed (op. cié. p. 513), says “ no underfur.”

182 MR. R. I. POCOCK ON [| Mar. 17,

Text-fig. 33.

TERZI.—

Skull of Capricornis swmatraensis rodoni Poc., from Chamba, belonging to the

skin shown in text-fig. 32. x i.

A, from the side: B, from below; ©, from above.

1908. ] SEROWS AND GORALS. 183

The skull also presents some differences from the skull of a spe-

cimen sent by Hodgson from Nepal (text-fig. 30). In the latter,

which measures 241 mm. in length from the occipital suture to

the premaxillz, the series of upper cheek-teeth measures 87 mm.,

the last molar being 20 mm. long and 16 wide; the width of the

palate between the last molars is 52 and between the first

premolars 38. Whereas in the skull of the specimen from

Chamba, which measures, as above, 223 mm., the upper cheek-

teeth are 93, the last molar 20 x 12, and the two palatal breadths

are 57 and 38. Thus in the specimen from Chamba the tooth-

series is longer, the last molar much narrower as compared with

its length, and the palate posteriorly broader. The skull is also

higher and has the facial pit less deep. Be it noted, however,

that it belonged to a younger animal.

The type of C. s. rodoni is the above described specimen, which

is now in the British Museum.

Subsp. sAMRACHT Pocock.

Abstr. P. Z.S. No. 55, p. 12, March 17, 1908.

General colour of head and body coal-black, the hairs being

white at the base and black distally, without any rufous-brown

tinge in the black terminal portion. On the upper lip behind

the white anterior portion there is a patch of brown, and hairs of

a similar brownish-yellow colour surround the base of the ear and

extend up the back of that organ. The white patch on the chin

extends back on each side of the jaw halfway towards its angle,

but there is scarcely any white hair on the throat. The chest also

is blackish, and the hairs forming the whorls on the front of the

chest near the base of the legs are black, with red basal portion.

The belly is a dirty grey-brown. The shoulders are black, but

on the fore leg between the elbow and knee, the hairs are a

mixture of black and rufous brown, both on the outside and the

inside of the limb. The knee itself is white like the fetlocks and

pasterns, but the area between the fetlock and the knee (the

cannon-bone) is strongly tinged with fawn or rusty yellow.

Along a line running from the tail to the stifle, the black of the

hind-quarters passes into the rusty-brown colour which pervades

the outside and inside of the hind legs. Inferiorly this rusty

brown pales to yellowish brown, the fetlocks being white.

Except on the fetlocks, knees, inside of ears, upper lip, and chin,

there is no white on the animal.

The coat is short at all seasons of the year, and not shaggy,

and there is practically no underfur.

Loc. Kalimpong, near Darjiling.

A young female specimen of this Serow was purchased by the

Society from Mr. W. Jamrach in August 1906. It was then, I.

should judge, about six months old, and it is important to record

that up to the present time (that is to say, during the sixteen

months that the animal has been under observation) she has not

184 MR. R. I. POCOCK ON [ Mar. 17,

changed at all in the matter of colour. She stands about 3 feet

high at the withers, but from the size of her horns which measure

6 inches long, with a basal circumference of 43 inches, I judge

her to be not yet fully adult.

Text-fige. 34.

Capricornis sumatraensis jamrachi Poe.

Photograph of specimen from Kalimpong, now living in the Society’s Gardens.

In the Mammal Gallery of the Natural History Museum there

is a mounted male specimen of this same race of Serow,

which was presented by the Duke of Bedford, and is labelled

Nemorhedus swmatrensis. According to Blanford’s ‘ Mammals

of British India,’ this determination is correct; but the animal

is obviously quite different from the typical Capricornis suma-

traensis of Bechstein, which has a grey mat-like mane on the

withers and much darker lower legs. From information kindly

supplied to me by the Duke of Bedford and Mr. Jamrach, I have

been able to ascertain that this animal also came from Kalimpong.

Except that there is a noticeable quantity of rusty yellow in the

hairs of the forehead, this specimen does not appear to differ in

any important particular from the female now living in the

Gardens. As in the latter, the horns are rather small, measuring

156 mm. (=6j inches) long, with a basal circumference of

1908. ] SEROWS AND GORALS. 185

100 mm. (=4 inches). Conceivably the smallness of the horns is

a racial character; but I am more inclined to attribute it to

immaturity.

The type of C. s. jamrachi is the mounted specimen above

referred to in the Mammal Gallery of the British Museum

(Reg. No. 2.10.12.1).

Of all the extra-Himalayan Serows recorded up to the present

time, this subspecies is most nearly allied to C. s. milne-

edwardsti David *, from Eastern Tibet. The colour of the body

in the latter, however, is not so black as in C. s. jamrachi, and

the tint of the legs is much more ferruginous. The darkening of

the legs is carried still further in C. s. argyrochetes Heude 7,

from Sze-chuen and Tché-kiang, for the anterior side of the

ceannon-bone is blackish. The mane, moreover, is grey. In the

coloration of the legs C. s. argyrochetes lies midway between

C. s. jamrachi and the typical C. s. swmatraensis, while in the

greyness of the mane it resembles the latter race. Of C. s.

argyrochetes the British Museum possesses two specimens, one a

flat skin from Sze-chuen, obtained from Berezowski in 1896,

the other a mounted specimen said to have come from Tibet,

which Mr. Lydekker has recently described and figured (P. Z.8.

1905, p. 329, pl. viii.).

Subsp. Roprinsont Pocock.

Abstr. P. Z.S. No. 55, p. 12, March 17, 1908.

Prevailing colour black, the hairs white at the root, usually

brownish mesially and jet-black in their distal two-thirds. Head

black, with a narrow grey rim to the upper lip and corner of the

mouth, this grey continuous with a large patch of the same

colour tinged in places with brown, which extends backwards

to a point on a level with the orifice of the facial gland; chin

black. Mane formed of a nearly equal mixture of white and

black hairs, without any red; its anterior end, forming the

occipital tuft, black with small white tips. The mane does not

form a white mat-like patch on the withers. It is continued

down the spine as a black crest. On the throat there is a small

patch formed by the rufous or white tips to the hairs. On the

tail and the outer side of the thighs there is a noticeable quantity

of reddish-brown hair, and hairs of a similar hue surround the

anus and extend along the edge of the under side of the tail,

the upper side of which is black. Lower surface a dirty dark

brown; inner side of thighs at base scantily clothed with dirty

white hair. Fore and hind legs black, with dark chocolate-brown

knees, hocks, and fetlocks.

Measurements in English inches of freshly stripped skin :—

Total length from nose-tip to tail-tip 60, tail 7 (with hair 10).

Height at withers 36, hock to heel 134, knee to heel 104, distance

* Nouy. Arch. Mus. v. Bull. p. 10,1869. Also A. Milne-Edwards, Rech. Mamm.

p. 365, pls. Ixxii. & lxxiii., 1874.

+ Hist. Nat. Chinois, i1. p. 4, 1888, and p. 228, 1894.

186 MR. R. I. POCOCK ON [ Mar. 17,

from anterior edge of eye to posterior edge of nostril 73; ear

(along back) 8, inside space 62, width 3.

Loc. Selangor, in the Malay Peninsula.

The type of this species is a male specimen kindly presented to

the Zoological Society by the Government of Selangor.

The skin and skull are now in the British Museum (Reg.

’ No. 6.11.14.1).

Text-fig. 35.

Capricornis sumatraensis robinsoni Poc.

Photograph of specimen, from Selangor, that formerly lived in the

Society's Gardens.

This Serow closely resembles the typical Sumatran form of

the species, Cs. swmatraensis, in the black colouring of the lower

half of the legs. So far, indeed, as I can judge, it differs from

the latter only in having the mane less grey and less copious

and thick on the withers. As might be expected from its distri-

bution, it is almost intermediate between the Sumatran race and

the one from the Larut Hills in Perak, C. s. swettenhami Butler *.

Of the latter I have only seen one flat skin, ticketed Biserat

* P.Z.S. 1900, p. 675. Mr. Butler, misled by Blanford, separated this form

from C. s. sumatraensis because of the blackness of the lower legs, which Blanford

erroneously described as rufous in C. s. swmatraensis. See also S. S. Flower,

P.Z.S. 1900, p. 371.

1908. | SEROWS AND GORALS. 187

in the Malay Peninsula. This is in the British Museum (Reg.

No. 3.2.6.77). Butler records the animal from the Larut Hills

in Perak. In the Biserat skin the occipital crest is composed of

hair principally white with black or red extremities; the rest of

the mane consists of a mixture of red, black, and white hair, the

red being much in evidence; also the pale patch on the anterior

portion of the lower jaw is mostly red. It is on account of the

almost complete absence of red from the mane and the small

amount on the jaw of the Selangor specimen that I regard it

as representing a geographical race of Serow, distinct from

C. s. swettenhamt.

It is important here to repeat Mr. Butler’s statement that a

dead specimen seen by Mr. L. Wray, a living specimen seen

by himself in the jungle, and a stuffed young one together

with the type in the Perak Museum are alike, to all intents

and purposes, in colour, Including, therefore, the example from

Biserat in the British Museum, no fewer than five skins have

been examined and reported upon. This is sufficient to justify

the conclusion as to the constancy of the characters upon which

C. s. swettenhami was based and to warrant the view that the

Selangor form described above must be regarded as distinct.

I learn from My. H. C. Robinson, F.Z.S., the Curator of the

Selangor State Museum, that this Serow was surprised in a patch

of scrub near Batu on the coast of Selangor, and driven by dogs

into the sea, where it was captured. There are no hills of more

than two or three hundred feet in height for many miles from

the spot where it was first discovered, the district towards the

coast being mostly swampy land. The Serow must have wandered

either from the range of hills between Negri Sembilan and

Selangor, which jut off from the main backbone of the Peninsula

of Malacea, or possibly from the latter range itself.

The animal was kept alive by natives for three months before

it was shipped for London. It arrived in a very emaciated

condition and died after a few weeks from starvation, caused by

the blocking of the pylorus with masses of thickly felted soft

woolly hair. That this was not derived from the animal itself is

rendered probable by the absence of all underfur to the coat.

It seems possible, therefore, that the animal devoured a blanket,

or a piece of woollen cloth, during the time of his captivity with

the natives.

The skulls of the typical C. s. swmatraensis and of C. s. swetten-

hami are unknown to me; but that of ©. s. robinsoni differs in

certain well-marked features from the skulls of all the Himalayan

and Burmese specimens that I have been able to examine in the

British Museum. It is long, narrow, and high, the frontals being

unusually elevated and convex both antero-posteriorly and

transversely between the orbits, so that the plane of the horns

lies in almost the same line as the plane of the face, and the

downward slope of the cranium posteriorly is somewhat abrupt.

The facial or lacrymal fossa is, shallow. The nasals also are

188 MR. R. I. POCOCK ON [Mar. 17,

Text-fig. 36.

Skull of Capricornis sumatraensis robinsoni Poc., belonging to the specimen

shown in text-fig. 35. (B.M. Reg. 6.11.41.)

A, from the side; B, from below; C, from above.

1908. | , SEROWS AND GORALS. 189

transversely convex and long. Regarding the skull from its

yentral aspect, it is evident that the basi-facial axis 18 more

bent upon the basi-cranial axis than in other specimens. The

horns are short and thick, the sheath measuring only 160 mm.

(62 inches) with a basal circumference of 125 mm. (5 inches).

Subjoined is a table of measurements of four skulls of Serows

from Kashmir, Nepal, Chamba, and Selangor.

Skull-measurements in millimetres *.

| |

| Kashmir. Nepal. Chamba. | Selangor.|

| | |

| |

+Basal length from occipital suture |

to distal end of premaxille ...| 245 | 41 | 223 240

| Width across zygomata ............ 130 e283 en ell277 els

| As Pe ate. al lie: Mecepectarcemena | 98 98 | 98 | 102

| », between orbits ............... 93 87 | 78 81

| Median length of frontal ............, 112 105 eelali 118

Fe a TESA, ake bao noo ono 94 | 93 | 73 Piss; |

Width across nasals .................. 50 | 47 41 48 |

Width across premaxille (maxi- | | |

| TUAW T) aaa meapeneaatonaaASAenaseSeasas 53 | 51 | 50 54: |

| Width across premavxille (distal | | |

CLIC) kg easing ets aie 32 | 31 | 26 27

Height from alveolus of molar 2 to | |

| Summait of f:onuals) .a).seeee ss 104 94, 98 Hale

| Height from alveolus of premolar 3 | | |

| to summit of nasals............... 94. 84 | 90 eelOs

fencu mote lee keneen ney mere ere: | amma oO i NaS nie ago Vo?

| Lenethand width of last molar...) 20,18 | 20,16 | 20,12 DilealleSaat

| Median length of palate to distal

end of premaxille ............... 162 167 | 148 161

Width of palate between last

WIOLAUS! pA eceey Siesta oe 60 | 52 57 50

Width of palate between first | |

THRSUMOI EWS occas saocdonnawosesse0o56 | 41 | 38 38 38

In spite, however, of the differences above set forth, I think it

premature to attach very much weight to them and to draw

conclusions as to their constancy until the extent of the variations

of the skull with age, possibly with sex, and in a number of

individuals from the same locality, has been ascertained. These

remarks apply more particularly to the specimens from the

Himalayas, of which the dimensions are tabulated above. The

explanation of the differences, for example, between the Kashmir

and Nepal skulls may bea matter of age if the skull becomes

flatter, narrower in the palate, and deeper in the preorbital pit

with advance of years{. It is significant that one of the skulls

% Measurements taken with a compass or dividers.

+ This measurement is taken from the basioccipital suture instead of from the

occipital foramen, because the occipital region of the skull is not infrequently

absent in prepared skulls. :

{ Implicit faith in the systematic value of cranial and dental differences, coupled

with inexperience in osteology and injudicious handling of material, seems to have

been responsible for the vast numbers of “ species ” into which the Deer, Pigs, and

Antelopes, including Serows and Gorals, of Southern China have been split by

Pére Heude.

190 MR. R. I. POCOCK ON [Mien li

brought by Hodgson from Nepal is intermediate, or nearly so,

between two others from that country and the one from Kashmir.

The skull from Selangor, however, can be at once singled out

from the others by its general aspect and dimensions.

The seven geographical races of Serow (Capricornis suma-

traensis) discussed above may be distinguished externally as

follows :—

a. Legs below the knees and hocks white, whitish, or fawn

and lighter than the area above these joints *.

6. Head pale chocolate-brown, not appreciably intermixed

with biack . gee Stes dn an Bee Lan Hataaonuecmenenete U0 UU

. Head black or ron nish Whack,

c. Breast and under side white, and sharply defined from

the dark colour of the rest of the body, which is

covered with a thick coating of long black hair tinged

with red; underfur conspicuous; a distinct and large

white patch on the throat ............. .. rodoni.

c!. Breast and under side at most dirty w mite enn blowing

with the dark colour of the rest of the body, the hair

of which is scantier and darker with little if any

underfur ; at most a small patch on the throat.

d. Under side dirty white or rufous white; the coat

less black; lower portion of legs white or faintly

THUS yale] MUNK AMT Gg agn aso ase senodu comand baasusounucdpoadeo, | UAC

d', Under side blackish red ; the coat jet-black; lower

portion of legs very decidedly tinged with fawn

on the cannon-bone .......... .. janrachi.

. Legs below the knees and hocks inlewik, raat iemen ‘dhe

area above these joints f

e. Scarcely any red in the mane and on the white patch

on the under jaw.

JF. Mane for the most part hoary grey and forming a

large thick mat on the withers .......... .. sumatraensis.

Jf’. Mane less grey and not fermi a Teel Te on 1 lhe

wilbhers) yeeeeeecens: . robinsoni.

. A considerable quantity ae toll in ihe. mane Danae on nate

DENI Oral INO WGI FEIN soatcoacseodadecconusceaeouasrcanaxocon SUBELUCIOLGIT.

III. On the Gorals (Nemorhedus) of the Himalayas, Burma,

and China.

Up to the end of 1904 only one species of Goral was known

from the Himalayas; but in the ‘Zoologist’ for March 1905,

pp. 81-84, Mr. Lydekker correctly pointed out that two types of

Goral occur in those mountains, one in the western and the other

in the more eastern portions of the range, the two meeting in Nepal.

To the eastern form, termed the “brown” Goral, the specific

name goral was restricted; while the western form, termed the

‘“‘orey” Goral, was described as a new species under the name

* Not known for certain in the case of C. s. hwinei.

+ Some Chinese Serows are intermediate in the coloration of the legs between

C. s.jamrachi and C.s. sumatraensis. For instance, C. s. milne-edwardsi from

Eastern Tibet has the lower legs rusty yellow, and an example in the British

Museum from Sze-chuen (Berezowski), referable to C. s. argyrochetes, has the

cannon-bone blackish in front and rufous elsewhere.

1908. | SEROWS AND GORALS. Ol

bedfordi. Apart from the noticeable difference in colour between

the two, the “brown” Goral is further distinguished by the

presence of a black spinal stripe which is absent, at least on the

back and tail, in the “ grey” form.

Now, Antilope goral was first described by Hardwicke (Tr.

Linn. Soc. xiv. p. 518, 1825). The description, taken from a

living animal, states that the hair was of a ‘“‘ grey mouse-colour

(but almost white about the lower part of the neck and throat),

and darker; it is longer along the upper part of the neck and

back, inclining to ferruginous about the legs.” Similarly the

Latin diagnosis says: ‘‘Corpore supra colore murino canescente,

subtus pallidiore, gula albente.”

It is, In my opinion, impossible to maintain that Hardwicke

can have described as ‘‘ grey mouse-coloured ” an animal which is

not grey, and which was regarded independently by Hodgson as

“rusty and brown”; by Dr. Blanford as “‘ brown, more or less

rufous” *; and by Mr. Lydekker as “‘ rufous brown.” Hardwicke’s

use of the terms “ grey ” and “canescent” as applied to the body and

the contrast that he draws between that colour and the inclination

to a ferruginous tint on the legs make unavoidable the conclusion

that the specimens upon which the specific name goral was based

represented a form identical with or very closely allied to the one

that Mr. Lydekker spoke of as the “grey” Goral and named

Urotragus bedford.

It must be particularly borne in mind, too, that although

Hardwicke noticed the length of the hair on the upper part of the

neck and back in the type of his species, he made no mention of

the presence of the black spinal stripe so conspicuous in adults of

the “ brown” Goral. The figure, it is true, shows such a stripe

on the neck and withers; but this is sometimes present in speci-

mens of the “grey” Goral, and is very noticeable in the living

example of the latter now in the Zoological Gardens when the

neck-hairs are parted.

The description of WV. goral was taken from a male specimen

living in the menagerie at Barrackpore, near Calcutta, which had

been previously the property of the Court at Katmandu in Nepal;

and there is a skin of a “‘ grey” specimen in the British Museum,

ticketed Nepal (Maharajah Dhuleep Singh; 55.1.20.5), which

Mr. Lydekker identified as V. bedfordi.

For the type of WV. bedfordi, a mounted specimen now in the

British Museum (Reg. no. 97.4.3.1) and at one time the property

of the Duke of Bedford, no locality was known. ‘The specimen,

however, as His Grace has kindly informed me, was imported by

Mr. William Jamrach; and [I learn from Mr. Jamrach that he

formerly procured Gorals from Dharmsala. This circumstance

and the similarity between the specimen named J. bedfordi and

a series of skins of Gorals from Chamba, shot by H.H. the Rajah

* This author adds “or greyish.” Be it remembered, however, that he had access

to the material in the British Museum containing a specimen of the “grey” Goral,

which he apparently did not distinguish from the “brown” form.

Proc. Zoou, Soc.—1908, No. XIII. 13

192 MR. R. I. POCOCK ON [ Mar. 17,

of Chamba and kindly sent to me through Major Rodon, F.Z.S.,

point to Dharmsala as the locality of the typical example of

WV. bedfordi. This specimen Mr. Lydekker described as yellowish

grey-fawn, suffused with blackish, which can be interpreted as

merely another way of describing the colour which conveyed to

Hardwicke the impression of ‘‘mouse-grey and darker,”

But, as one of the features distinctive of NV. bedfordi,

Mr, Lydekker mentioned the large extension of the white of the

interramal area up the cheek. This exists undeniably in the

stuffed specimen ; but one’s confidence in the systematic value of

the character is completely shattered by the entire absence of any

indication of it in the excellent photograph of the living animal,

taken by the Duchess of Bedford, which Mr. Lydekker has pub-

lished *. No one who looks at this photograph can for one moment

believe that the lower part of the cheek up to or even above the

level of the corner of the mouth was white or different in tint from

the rest of the cheek, The upward extension, therefore, of the

white in the stuffed specimen must be merely due to a taxidermic

distortion, uniess the colour of this region changed between the

time of taking the photograph and the death of the animal, which

is unlikely.

In view of the above-mentioned facts, the following conclusions

appear to me to be inevitable :—(1) That the type of WV. goral

was the “grey” Himalayan form; (2) that Mr. Lydekker re-

described this form as VV. bedfordi. And from this it follows that

the “brown” Himalayan Goral is up to the present time without

a specific name. From the nature of the differences separating

the “ grey ” and “brown” Gorals I think it probable that they

will be found to intergrade. Up to the present, however, there

is, so far as I am aware, no proof of the fact. Since the two forms

have been recorded from Nepal it is possible, as Mr. Lydekker has

suggested, that they occur at different altitudes in the Himalayas.

Pending additions to our knowledge in these particulars, I propose

to follow Mr, Lydekker in treating these Gorals as distinct

species.

N#®MoRHEDUS GoRAL Hardwicke.

Antilope goral Hardwicke, Tr. Linn. Soc., Zool. xiv. p. 518,

pl. xiv., 1825.

Urotragus bedfordi Lydekker, Zoologist, March 16th, 1905,

p. 83; id. Great and Small Game of India, noy. ed. p. 151, 1907;

id. in Rowland Ward’s Records of Big Game, p. 343, fig. p. 348,

1907.

Prevailing colour yellowish grey, speckled or suffused to a

varying extent with black, so that the depth of the tint varies

considerably individually, but the pale band in the hairs is always

yellowish grey and never rufous or brown, Forehead suffused

with rusty yellow, the same tint traceable on the sides of the

1907, p. 149; Rowland Ward’s ‘ Records of Big Game,’ 1907, p. 348. ~

1908. ] SEROWS AND GORALS. 193

nose; cheeks grey, interramal area and chin white; upper lip

white, stained with yellow at the sides. Throat-patch yellowish

or snow-white, sometimes set off by a darker border; ears mouse-

grey or yellowish externally. Between the horns a tuft of dark

or black hair intermixed with white; usually there is no black

spinal stripe either on the neck or body, but the hairs on the

middle line of the neck show up as distinctly darker than the

sides, owing to their partially erect position revealing the slate-

grey colour of the basal portion. In one specimen, however, the

Text-fig. 37.

Nemorhedus goral Hardw.

Photograph of specimen from Chamba, now living in the Society’s Gardens.

hairs on the middle line of the neck and withers have long black

tips, which run together to form a distinct blackish line. Tail the

same colour as the back in its basal half*, but the tip with a

longish tuft of hairs black throughout. Skin of tail white below.

Subcaudal area fringed with white hairs. No blackish stripe

extending up the buttocks from the legs. Colour of legs very

variable; fore legs usually with a well-defined black stripe

* In Mr. Lydekker’s description of N. bedfordi the “base” of the tail, instead of

the “ tip,” is described as blackish.

13%

194 MR. R. I, POCOCK ON { Mar. 17,

extending down the middle from above the knee to the fetlock ; but

in one skin from Chamba and in the type of V. bedford the stripe

is quite short and inconspicuous, except upon the knee; the sides

and posterior surface of the leg below the knee varying from a

rich fawn to greyish, sometimes even whitish on the inner side.

Hind legs similarly variable, darker or lighter fawn down the

front, sometimes showing some black hairs above the fetlock and

sometimes whitish on the inside; the posterior surface from the

hock always darker than the anterior, never fawn, and usually

blackish. Belly and inside of thighs white or greyish, sometimes

with a yellow tinge ; a dark patch on the chest.

Loe. Chamba; ? Dharmsala and Nepal.

The above-given description is taken from a single example from

Chamba now living in the Gardens, which was presented to the

Society by Major Rodon, F.Z.S., on June 3, 1904, and also from

a series of eight skins of specimens shot in Chamba by H.H. the

Maharajah, who kindly gave them to Major Rodon to forward to

me for examination and description. This series has been espe-

cially useful in showing the variation in detail, and at the same

time the constancy in general appearance presented by a number

of individuals from the same locality. From these Chamba skins

the type of WV. bedfordi, which probably came from Dharmsala,

and the above-mentioned skin in the British Museum ticketed

Nepal (Maharajah Dhuleep Singh), are not, in my opinion,

separable by a single character of systematic value.

It is interesting to record that the example of this race now

living in the Gardens was a quite young animal on its arrival

in June 1904, and that, except for increase in size of body and

length of horn, it has not appreciably altered in appearance.

Nor is there any marked seasonal variation in colour, the new

summer coat being merely a little richer in tint than the old coat

before shedding begins.

On one occasion about a year ago this animal got into a yard

where there was a well-grown specimen of a Grecian Ibex, con-

siderably larger and more heavily built than the Goral. But, in

spite of the confined space, I am quite sure that the fight that

ensued would have ended fatally for the Ibex, which, although

full of pluck and eagerness for the fray, seemed bewildered by the

agility of his antagonist. The Goral’s method of fighting was to

charge low under the guard of the Ibex’s heavy recurved horns,

then to get out of reach, never giving the Ibex a chance of one

straightforward butt, which would probably have ended the contest

in his favour. Before the combatants could be separated the

Goral had succeeded in wounding the Ibex in the nose with his

short sharp horns. The efficacy of the short pointed horn as

compared with the heavy horn in Goat-like ruminants was further

exemplified some years ago by another fight that took place,

T have been told by Mr. ‘Thomson, between a male Thar (Hemi-

tragus jemlaicus) and a Markhoor (Capra falconert). In less than

five minutes the Thar killed the Markhoor by getting past his

1908. } SEROWS AND GORALS. 195

horns and ripping open his abdomen. But in this particular

instance I have no first-hand knowledge of equality between the

contestants in the matter of age and condition, as I have in the

case of the Goral and the Ibex.

N#MORHEDUS HODGSONI Pocock.

Abstr. P, Z.8. No. 55, p. 12, March 17, 1908.

Nemorhedus (Antilope) goral Hodgson, P. Z. 8. 1834, p. 85.

Cemas goral Blanford, Fauna of Brit. Ind., Mammalia, p. 516,

1891.

CUrotragus goral Lydekker, Zoologist, March 16th, 1905, p. 83;

id. Great and Small Game of India. nov. ed. p. 151, 1907; id. in

Rowland Ward’s Records of Big Game, p. 342, 1907.

Nec Antilope goral Hardwicke, Tr. Linn. Soc., Zool. xiv. p. 518,

1825.

General colour of type darkish golden brown speckled with

black, the individual hairs dark brown at the base, black at the

apex, and with an intermediate golden-brown or rufous area.

Forehead and nose deep reddish brown, becoming blacker towards

the root of the horns, a small black patch above the muzzle:

cheeks yellowish brown, paler than body owing to the absence of

the black apical tip to the hairs; interramal area not white but

stained with yellow, a brown patch on the chin; lips yellowish

white; throat-patch yellowish white, defined laterally by an

indistinct blackish streak.

A black stripe extending from the head along the neck, where

the hairs form a short mane, down the back to the root of the tail.

On the neck it 1s about 1 inch broad, but behind the shoulder it

gradually tapers away and almost disappears upon the lumbar and

sacral regions. <A black patch on the chest; belly greyish yellow.

Tail wholly black above, except for a few pale hairs at the side.

Area below the tail white above in the adanal region, yellowish

below on the inside of the thighs. From near the root of the

tail on each side extends a blackish-brown stripe down the back

of the thigh and leg to the hock, and this is continued from the

hock to the fetlock and beneath the “ dew-claws” to the hoof;

front of hind legs below the hock golden brown, with commonly a

blackish stripe. Fore legs golden brown, with a black patch over

the knee and black hairs extending in the middle line thence both

below and above the knee, where the hairs are a bright, almost

fiery, brown. Horns but little curved, 43 inches long, corrugated

and ringed basally, basal antero-posterior width 3 of an inch

(23 mm.). Ear-cavity about 34 inches long.

Loe. (of type). Sikhim (W. 7. Blanford; no. 91.10.7.169 in

B.M.); also Nepal (4. H. Hodgson).

In addition to the typical skin above described, there are in the

British Museum three other skins referable, I think, to this

race of Goral. They are ticketed Nepal (B. H. Hodgson ; nos.

45.1.8.325-327). In the largest of these the general colour is

196 MR. R. I. POCOCK ON [Mar. 17,

browner, with the hair less noticeably speckled than in the type,

and the throat-patch and interramal area are white, the latter

being a dirtier white than the former. The horns are 33 inches

long (90 mm.), with a basal diameter of ? of an inch (20 mm.).

A second example is evidently young; the fur is softer and

thicker and also less speckled than in the first, and the face is

less richly coloured and somewhat greyer. The third specimen,

a young and hornless ‘individual, while presenting the same

general type of coloration as the others, differs from them in that

the black spinal stripe fades away on the lumbar region and the

tail is not black above. Hodgson’s remark that the young is

redder than the adult and destitute of mantle and mane is worth

repeating.

A young example of this species from Nepal was presented to

the Society by the Prince of Wales in June 1906, but died a few

days after arrival.

N#MORHEDUS RADDEANUS Heude.

Antilope (Caprina) crispa Radde, Reisen im Siiden von Ost-

Sibirien, i. pp. 262-270, pl. xii. fig. 1 (nec Antilope crispa Temm.).

Kemas raddeanus Heude, Hist. Nat. Chinois, 11. p. 240, pl. 35,

1894.

General colour of the shaggy winter coat greyish yellow-brown,

darker along the spinal area.

Upper surface of head blackish from the muzzle back to the

occiput ; sides of the head and of the lips greyish yellow mixed

with black; rest of lips white; chin black. Throat with a large

white gular patch. Ears white inside; pale mouse-grey, darker at

the base, with brown edging externally. On the nape of the neck

the hairs form a short blackish mane. On the lumbo-sacral area

the dark spinal stripe is fainter than it is anteriorly. On the

shoulder there is an ill-defined scapular stripe, which inferiorly

turns into a jet-black stripe running down to the knee and thence

over it externally nearly to the fetlock. On the inner side of the

cannon-bone this is set off by a yellowish-grey stripe, which is

continuous inferiorly with the yellowish-grey hue involving the

front and sides of the fetlock and pastern ; the posterior surface

of the leg is brown, turning to black on the back of the fetlock.

The hind leg is chocolate-brown from the hock to the fetlock,

brownish in front, and paler yeilow-brown at the sides, the pastern

being creamy yellow anteriorly and laterally, like that of the fore

leg. The breast and anterior part of the belly are blackish, but

the groin, the inside of the thighs, and the back of the thighs up

to the root of the tail are white. The tail is brown above and

white below ; the extremity of the tail is furnished with a mixture

of long black and white hairs which extend below the hocks.

The skin of the tail on its under side is about 43 inches (112 mm.),

and the total length of the organ to the tip of the hair is about

15 inches (375 mm.),.

1908. | SHROWS AND GORALS. 197

The length of the right horn, which is normal in position, is

rather less than 5 inches (125 mm.). That of the left horn,

which is bent sharply backwards in its distal half, so that the

point is unworn, is 6} inches (156 mm.). The basal circumference

is 3¢ Inches (89 mm.).

Text-fig. 38.

Nemorhedus raddeanus Heude.

Photograph of specimen, from Korea, that formerly lived in the Society’s Gardens.

The principal dimensions of the skull in mm, are as follows :—

iBasalllen ethyeeeae ee eee tres meeeeee SAP est Rein bo

Length of palate along middle line Ne Sri Ae 122

is cheek-teeth ............... Re AOC 70

Width of ea between pms' PASSION EAN 21

i 10S Sia adie Ae Ga aoe ae 39

Length from pm! to tip of premaxilla ......... DD

a OP mmasallgi' Hid a OAT eR A men, crrmat ayaa 69

Wadthrof nasalst 7) a ale Bere eera 2: 32

Tnaterorbuball yacht ke ne pase een renee 69

Greatest width across orbits ..................... 99

a is 4, anaassellll oe a PIRSA MAREE 67

a ay OE GRAUONTN eons 5 a ee . 65

198 MR. R. I. POCOCK ON [ Mar. 17,

Loc. Korea: Wonsan (2).

A living specimen of this Goral was chesenied to the Society

by Mr. C. F. Bilborough, F.R.G.8., on Feb. 5, 1907, and died on

June 30,1907, Mrs. Bilborough informed me that it ‘came

from the high chain of mountains that runs down the whole

length of Korea, rising at Wénsan to 12,000 feet high.”

T cannot distinguish the skin of this Korean specimen specifi-

cally or even subspecifically from three Goral skins presented to

the British Museum by Mr. Rowland Ward, F.Z.S., and ticketed

“Western Provinces of China” (nos. 99.3.5.1-3 in B.M. Register);

and, so far as I can judge, they are all referable to the species

from Amurland, wrongly identified by Raddeas Antilope (Caprina)

crispa of Temminck, and rightly renamed Aemas raddeanus by

Heude.

Nemorhedus raddeanus was regarded by Trouessart as a sub-

species of V. caudatus M.-Edwards, presumably because of the

length of the tail. In the British Museum there is a topotypical

example of WV. cauwdatus from Pekin, collected by Mr. F. W. Styan

(Reg. no. 90.7.8.6). This specimen has the thick, long woolly

- coat observable in the above-mentioned examples referred to

NV. raddeanus, and the tail also is practically of the same length,

as the following measurements show :

Skin of tail. Tail (including hair).

NV, caudatus (young)......... 4 inches. about 13 inches.

WN, raddeanus (1 3)......... 5 - aval (AU Cme

” (2 2) ote Ds ” ” 13 Dp

%5 (ives) ae Be oss pp ME og

Hi (Koréa) ...... 4i,, ic LD eae

Nevertheless, the example of A’. caudatus, which agrees with

M.-Edwards’s description and figure of the type, differs from those

identified as V. raddeanus in having the legs below the knees and

hocks almost wholly fawn-coloured in front, the fetlocks and

pasterns being the same tint as the cannon-bones, and also in

having the tail much blacker, both above and below. Hence

there are well-marked differences between WV. caudatus and

NV. raddeanus. Intermediates possibly, perhaps probably, exist ;

but until they come to hand I think it premature to regard

NV. raddeanus as a subspecies of A’. caudatus.

The specimens from the Western Province of China exhibit

variation in colour worth putting on record. The male specimen

(no. 99.3.5.1) closely resembles the Korean example, being dark

grey mingled with blackish brown, the upper side of the all being

of the same dark hue as the back; ; the fore leg is black in front

down to the knee, and the outer aud posterior sides of the lower leg

are also black, but the fetlock and anterior and inner side of this

region are white, the area above the outer false hoof being black

and above the inner white. Similarly the hind legs are blackish

both in front and behind, the fetlocks and pasterns being whitish

1908. | SEROWS AND GORALS. 139

with some black hairs towards the outer side. The female, on

the contrary (no. 99.3.5.2), with the same history and presumably

from the same locality, is much paler, the general colour being

yellowish brown, the upper surface of the tail bemg the same

yellow-brown colour as the hind-quarters. On the fore leg the

blackness scarcely extends below the knee, the blackness of the

outer side of this area, so noticeable in the male, being scarcely

in evidence. Similarly the hind legs are yellowish brown in

front below the hocks. I do not know whether these differences

are sexual, seasonal, or individual. The third specimen

(no. 99.3.5.3), an immature unsexed animal, resembles the

female.

So far as the coloration of the legs is concerned, the examples

I refer to WV. raddeanus ave more like specimens in the British

Museum from Southern China than they are like V. caudatus.

These Southern Chinese specimens, which appear to me to be

specifically the same as WV. griseus A. M.-Edw.*, described from

Eastern Tibet, were obtained by Berezowski at Loung-nyou-fou

in the Leaorma eat of Sze-chuen (nos. 96,11.4.7-8) and by Wie, 1.

W. Styan at Ichang on the Yangtse-kiang (nos. 1.3.2.4 and

95.7.4.1-2). A young example was also brought from the same

locality by Mr. P. Montgomery (no. 96.11.4.7—-8).

Except that these southern forms are covered with a thick

coating of comparatively short hair and have the hairs of the tail

also shorter and scantier, they do not differ very noticeably from

NV. raddeanus. The skin of the tail measures from 4 to 5 inches,

and, including the hair, the entire tail may reach 10 inches. The

hairs of the tail, however, are black both above and below, and

in this particular resemble those of V. caudatus rather than of

NV. raddeanus ; and the white throat-patch is tinged with yellow

at the margins. The outer and posterior sides of the lower

portion of the front leg are blackish, the knee, the inner side of

this area, and the feet being darker or paler fawn, The hind

legs, below the hock, are brown behind, fawn or greyish fawn in

front.

Like WV. raddeanus, NV. griseus exhibits instructive variation in

colour. Of the three above-mentioned skins from Ichang no two

are alike. In one male example shot on September 9 the coat is

poor and of a dirty yellowish brown, with a very distinct black

spinal stripe, and the feet are a warm ferruginous or rusty-yellow

tint. A second specimen, shot on the same date and also a male,

has no distinct spinal stripe, the coat being a mixture of brown

and grey. The feet are nearly white. The third specimen, also

a male but shot in December, is thicker-coated and much more

richly coloured with dark brown than the others, though more

approaching the second specimen; the spinal stripe is traceable

but less differentiated than in the first specimen, owing to the

darker tint of the rest of the body ; the feet are fawn—that is to

* Nouy. Arch. Mus. vii. Bull. p. 93; Rech. Mamm. p. 361, pl. lxxi. 1874.

200 MR. R. I. POCOCK ON [Mar. 17,

say, intermediate between the ferruginous tint of the first and the

white tint of those of the second specimen. The edges of the

throat-patch are also yellower than in the others. Since these

specimens were of the same sex and from the same locality, and

since the first and second were shot on the same day, it does not

appear that these colour-difterences are either sexual or seasonal.

They are perhaps due in part to differences of age and in part to

innate individual variability.

Berezowski’s specimens from Sze-chuen are inseparable from

the richest-coloured example from Ichang. They were shot in

January. The coat is richly coloured brown and grey. In one

the feet are fawn, in the other nearly white.

I cannot find any reliable character to distinguish these speci-

mens from JV. griseus, judging from the description and figure of

the latter.

A further point to be noted is this. A Goral from Ichang was

recorded by Dr. Henry as Kemas henryanus (P. Z.S. 1890, p. 93).

This name was quoted as having been already published by Heude.

But Heude’s description of Kemas henryanus was not issued appa-

rently until 1894, when it appeared in Mém. Soc. Hist. Nat.

Chinois, ii. p. 244; and since Dr. Henry’s citation was accom-

panied by the phrase “The Ichang animal stands as high asa

Sheep,” he must be regarded as the author*. Furthermore,

Dr. Sclater (P. Z. S. 1890, p. 94, note) refers the Ichang example

in the British Museum, collected by Mr. P. Montgomery, to

Nemorhedus henryanus. This Ichang Goral may possibly prove

to be subspecifically distinguishable from the typical WV. griseus

when topotypical examples of the latter come to hand for com-

parison; but for the present I think it must be referred to that

form.

Finally, I am convinced that Mr. Lydekker described the same

animal as Urotragus evansi (‘Zoologist,’ (4) ix. p. 83, 1905 ; id. in

Rowland Ward’s ‘ Records of Big Game,’ p. 343, 1907). Of this

there are two cotypical examples in the British Museum from

Mt. Victoria in the Pokokku district of Arakan (/ajor Hvans :

5.7.21.1-2). They are quite young animals with the horns

measuring only 3 inches in length. They differ in no important

particulars, so far as I can ascertain, from the Ichang and Sze-chuen

specimens that I refer to V. griseus. Moreover, in the summer of

1903 Mrs. Mumford sent to me for identification the skins of three

“Goats” shot by her late husband, Mr. G. E. Mumford, District

Superintendent of the Burma Police, at Kyank-pin-daung in the

Arakan Hills. When compared with the material in the British

Museum, these skins proved to be indistinguishable from those

from Sze-chuen and Ichang, mentioned above, which J could not

separate from IV. griseus. Hence, although no new name could be

introduced, the real credit of being the first to send home material

showing that the Arakan Goral is distinct from the Himalayan

* Trouessart erroneously cites K. henryanus as “nomen nudum.”

1908.] SEROWS AND GORALS. 201

animals belongs to Mr. Mumford rather than to Major Evans.

Honours, however, are divided, because Major Evans’s skins were

perfect, whereas those belonging to Mrs. Mumford had been

made into mats and were without heads and legs.

That Mr. Lydekker fell into the error of giving a new name to

the Arakan Goral must be attributed to his comparing it with

Himalayan specimens and not with examples from South China.

Identity between Burmese and Southern Chinese animals is in no

sense a surprising fact.

The following synonymy, therefore, I believe to be well

established :—

N. griseus M.-Edwards, 1874=N. henryanus Henry, 1890=

AV. evansi Lydekker, 1905.

If my supposition that the above-mentioned Gorals from Ichang,

Sze-chuen, and Arakan belong to the same species is correct, it

proves that this species, whatever its name, has a wide geographical

range, and presents very considerable individual variation in speci-

mens from the same locality with respect to the colour of the body

and of the feet. Indeed, when the variability in these particulars

exhibited by the three examples from Ichang is taken into con-

sideration, grave doubts must be thrown upon the status of some

of the many so-called species from Southern China described by

Heude.

This author, for example, described two “species” from

Western Sze-chuen, namely, Kemas [= Nemorhedus| xantho-

deiros and K. pinchonianus, and one from Eastern Sze-chuen,

namely, K. iodinus* ; but, judging from the descriptions, these

differ less from each other in colour than do the three skins from

Ichang. It is necessary to add that Heude relied in his specific

determinations largely upon characters in the skull and teeth,

many of which are, I suspect, attributable to differences of age

and to individual variability *.

Finally, I suspect that V. arnouxianus Heudet from Tche-

kiang must also be referred to the species I have determined

above as WV. griseus. So far as colour is concerned, no difference

seems to exist between them, and the chief character in the skull

Heude relied upon, namely the somewhat abrupt rise of the horns

from the frontal bone, is also, I think, untrustworthy ; for con-

siderable variation in this respect is exhibited by the skulls of

%* Mém. l’Hist. nat. Chinois, ii. p. 243, 1894,

+ Of the type of one of his “species,” WV. niger, Hende says that the discoverer

informed him “quwelle Gait rare ct qwonla voyait mélée aux troupeauc des

autres i Le (op. cit. p. 241). Hende’s apparent acceptance of this statement

in good faith, and his admission that the type of V. niger was in the same herd as

examples of WV. fargesianus, make it impossible to accept the author’s opinion as

to specific differences. Two distinct species of a genus of Antelopes and Sheep

sometimes run together; but such cases are quite exceptional, and in the present

instance it appears to me that the evidence points to the type of WN. niger being an

aged individual of a species of which the co-types of N. fargesianus were younger

forms.

t Mém. V’Hist. nat. Chinois, ii. p. 3, 1888, and tom. cit. p. 239, 1894; op. cit. iii.

pl. xxix., 1897.

202 ON SEROWS AND GORALS. (Miaxel7,

three specimens from Ichang in the British Museum. In one

specimen, a young one collected by Mr. Styan, the horns lie back

practically in the same line as the forehead ; in a second obtained

by the same collector they rise slightly more ; and in a third sent

by Mr. P. Montgomery they are still more elevated, forming an

obtuse angle with the frontal bone very much as in the type of

NV. arnouxianus.

The external characters of the Gorals (Vemorhedus), whether

they be regarded as species or subspecies, discussed in the pre-

ceding pages, may be analysed as follows :—

a. Skin of tail about 3 inches long in the adult; black stripe

on fore leg extending over the middle of the knee (carpus)

and usually continued thence down the middle line of the

cannon-bone (metacarpus) to the front of the fetlock.

(Himalayas.)

6. Prevailing colour grey or fawn-grey, more or less suffused

with black; black spinal stripe usually wholly absent,

when present not passing beyond withers (shoulders) ;

tail black at the end; no black up back of thighs ...... goral.

b'. Prevailing colour brown, more or less suffused with

black; black spinal stripe present in adult and sub-

adult examples, and extending at least on to the lumbo-

sacral area; a black stripe down the upper side of the

tail and an ill-defined black stripe running wp the back

of each thigh from the hock hodgsoni.

a'. Skin of tail about 5 inches long in adult; black stripe on

fore leg not passing over the middle line of the knee

(carpus), but turning aside at that area and commonly

continued down the outer side of the cannon-bone (meta-

carpus} to the outer false hoof and thence on to the back

of the fetlock and pastern. (North China to Arakan.)

e. Coat comparatively short and not woolly even in the

winter ; tailless bushy (tail-tuft black aboveand below ;

throat-patch more or less yellow, at least marginally)... griseus *.

c'. Coat, at least in the winter, long, shaggy, and more or

less woolly ; tail-tuft long and copious (throat-patch

without yellow).

d. Legs below knees and hocks nearly uniformly fawn in

front and externally; tail-tuft black, much darker

than the proximal portion of the organ and than the

back of the body; a narrow white fringe of hairs

bordering the tail below ...................c0cseeeeeeeeeeeee CUdatus.

d'. Front of legs below knees and hocks to a certain but

varying extent fuscous and contrasted in colour

with the white or dirty-white tint of the feet ; upper

side of the tail the same colour as the back, but

not wholly black; a broad white fringe bordering

LELNEVTREDIIL LOVE) rondsdocodesootecudsoovodedeseosan dog sae cecsoodabinan W DRARMIAMDOISs

* T suspect that NV. cinereus A. M.-Edwards (Rech. Mamm. p. 362, pl. Ixx. et seq.,

1874) from Eastern Tibet will prove to beat most subspecifically distinct from

N. griseus, in spite of the differences in the skull and teeth pointed out by the describer.

By Trouessart (Cat. Mamm. Suppl. p. 734, 1905) both of these Gorals are erroneously

classified with the Serows.

1908. ] ON THE CENTRAL AFRICAN FOREST-PIG. 203

April 7, 1908.

Dr. Henry Woopwarp, F.R.S., Vice-President,

in the Chair.

On behalf of Mr. Thomas Codrington, Dr. A. Smith Woodward,

F.RS., F.Z.S8., exhibited a collection of 168 stones, weighing

altogether 7 lbs. 13 oz., taken from the stomach of an Elephant

shot by Mr. H. Thornicroft in Northern Rhodesia. The animal

was a large male, with tusks weighing 49 lbs. each. The stones

showed no signs of attrition.

Dr. C. W. Andrews, F.R.S., F.Z.S., exhibited a restored model

of the ‘skull and mandible of Prozeuglodon atroz And. This

animal is one of the links uniting the true Zeuglodonts with the

land Creodonts. It is found in the Middle Eocene of Egypt,

where also the earlier type, Protocetws, was discovered by Fraas

at a somewhat lower horizon. The model was constructed by

Mr. F. O. Barlow, for the British Museum of Natural History.

The Secretary exhibited a photograph of two young living

Text-fig. 39.

Young Forest-Pigs (Hylocherus meinertzhageni).

examples of the Forest-Pig of Central Africa (7 ylocherus

204 DR. KNUD ANDERSEN ON BATS. [Apr. 7,

memertzhagent Thomas). The photograph (text-fig. 39) had been

taken by Mr. T. J. Morson, of Limoru, who had obtained the

pigs in the Limoru escarpment forest about 353 miles from

Mombasa, at an elevation of between seven and eight thousand

feet.

The Secretary stated that he had been informed by the High

Commissioner for New Zealand, that the Chamois presented by

the Emperor of Austria to New Zealand, and which had been

successfully taken to New Zealand by one of the Society’s staff

in the beginning of 1907, had been seen in the locality in which

they were liberated, one of the females being accompanied by a

strong, healthy-looking kid.

The following papers were read :—

1. A Monograph of the Chiropteran Genera Uroderma,

Finchisthenes, and Artibeus. By Kxup ANDERSEN.

[Received May 29, 1907. ]

(Text-figures 40-59.)

ContTENTs.

The dentition of Artibeus planirostris, p. 205.

Uroderma, p. 212. . J. lituratus, p. 272.

U. bilobatum, p. 217, Jj. palmarum, p. 278.

U. thomasi, p. 221. . J. preceps, p. 283.

Einchisthenes, p. 221. . glaucus, p. 285.

EH. harti, p. 224, - watsoni, p. 288.

Artibeus, p. 224. cinereus, p. 290.

A. concolor, p. 232. c. cinereus, p. 292.

A. planirostris, p. 234. ce. bogotensis, p. 298.

A. p. planirostris, p. 237. rosenbergi, p. 293.

A. p. trinitatis, p. 241. . toltecus, p. 296.

A. p. grenadensis, p. 241. t. toltecus, p. 297.

A. p. fallax, p. 242. t. ravus, p. 300.

A. hirsutus, p. 245. quadrivittatus, p. 302.

A, jamaicensis, p. 247, . pheotis, p. 308.

A, j. parvipes, p. 261. . aztecus, p. 306.

A. 7. yucatanicus, p. 263. . turpis, p. 307.

A. j. jamaicensis, p. 26d. . nanus, p. 308.

A. j. equatorialis, p. 270.

Wing-indices, p. 310.

Summary of characters of genera, species, and subspecies, p. 311.

DRRARRA RRR RR RRR RD

General remarks ;—

(a) Artificial and natural arrangement of the species of Artibeus,

p. d14.

(6) A. planirostris and its races, p. 316.

on a past connection of the West Indies and Central America,

p. 317.

The conclusions recorded in this paper are based on a study of

485 Bats (861 skulls) of the genera Uroderma, Enchisthenes, and

1908. } DR. KNUD ANDERSEN ON BATS. 205

Artibeus*. 272 of these specimens (218 skulls) form part of the

collections of the British Museum; the rest, 213 specimens

(143 skulls), were placed at my disposal, for inspection and

identification, by the Authorities of the United States National

Museum.

The British Museum series is particularly rich in South and

Central American, the Washington series in Mexican and West

Indian specimens. Thus the two collections admirably supple-

ment each other.

I wish to tender my grateful thanks to Mr. Oldfield Thomas

for the opportunities he has so kindly afforded me for continuing

my Chiropteran studies in the British Museum. To Dr. J.

Leisewitz, Munich, Dr. Mareus W. Lyon, Washington, and

Professor D. G. Elliot, Chicago, I am indebted for information on

typical specimens in the collections under their charge.

Tuer DENTITION OF ARTIBEUS PLANIROSTRIS.

The teeth of one species only, viz., Artibeus planirostris (subsp.

fallax) are described in detail in this paper, the description of the

dentition in the other forms being, as a rule, confined to those

points in which it differs from this paradigma.

On the denomination of the molar cusps.—The molar cusps are

named in accordance with Herluf Winge’s theory r. The three

cusps (labial in the upper, lingual in the lower jaw) forming the

tips of the W of a typical molar in insectivorous bats are termed,

in antero-posterior direction, respectively 1, 2, 3, cusp 2 being pro-

bably the oldest, homologous with the single cusp of a Reptilian

tooth ; the two cusps forming the bases of the W are named 4

and 5; the “heel” of the upper molars, when single, cusp 6,

when double, cusps 6 and 7. See text-figs. 40, 41, pp. 207, 208.

S10 pe SEAR ip Rg

Tooth formula.—, . — OE _ Se

es 2, P, Ps m, mM, m,

Remarks on the tooth formula.—No known bat has more than

two pairs of upper incisors. The generally accepted hypothesis

is that the permanently missing pair is i’; but, in my opinion,

the balance of evidence is decidedly in favour of the view that

1’, not 1’, has been lost. The former hypothesis (i' lost) is

generally supported by two arguments, viz., ‘“‘ by the corre-

spondence of the two upper teeth with the two outer of the lower

jaw when the maximum set is present,” and “ even more strongly

by the general tendency throughout the group [%. ¢., the Chiro-

ptera] for the premaxillaries to become reduced, particularly

along the inner edge” Z. As to the former argument, it proves

* Brief preliminary diagnoses of the genus Hnchisthenes and ten new forms of

Artibeus and Uroderma were published in the Ann. & Mag. Nat. Hist. for December

1906 (pp. 419-423).

+ Herlut Winge, Om Pattedyrenes Tandskifte, iser med Hensyn til Tzendernes

Former (Vidensk. Meddel. Naturhist. Foren. Kj@benhavn for 1882, pp. 15-69, pl. iii.).

£ Gerrit S. Miller, Jr., “The Families and Genera of Bats,’ p. 27 (1907).

206 DR. KNUD ANDERSEN ON BATS, [Apr. 7,

nothing or, if preferred, anything; attempts to determine the

homologies of teeth in Mammalia on the basis of the corre-

spondence of the upper with the lower teeth, or vice versa, would

in too many cases lead to obviously absurd results. As to the

latter argument, it seems to me, on closer examination, to lead to

precisely the opposite conclusion. The tendency for the pre-

maxillaries to become reduced along their inner edge is first

developed in the higher of the two suborders of Bats, the Micro-

chiroptera; in the Megachiroptera no such tendency obtains, and

nevertheless they have only two pairs of upper incisors. When,

therefore, one pair has been lost also in those primitive bats in

which the premaxillaries have not been reduced along their inner

edge, the loss of this pair must evidently be due to some other reason.

The strong lower canines in bats (or rather their ancestors) have

probably effected the degeneration, and ultimate disappearance,

of that pair of upper incisors which, if it were present (or present

in its full size), would hinder their free passage in front of the

upper canines ; in other words, in passing in front of the upper

canines the lower canines have checked the growth, and

ultimately caused the complete disappearance, of 1°. In accord-

ance with this we find in most Megachiroptera the four upper

incisors (i°1'1'1°) close together, but 1° separated by a wide

diastema from the canine; part of this diastema indicates the

former place of 1°, but it has no doubt been widened to allow of

the free action of the lower against the upper canines. In many

genera of Microchiroptera, this line of development has been

carried a step further; by a narrowing of the diastema between

i’ and ¢ the former has come closer to the latter and within reach

of the lower canines, which then cause a decrease in size (and

change in shape) of 1°. This is the casein the three genera

which form the subject of the present paper, and which, there-

fore, in showing what actually takes place, in living bats, with

regard to i°, give, so to say, an illustration of what has probably

taken place, in the ancestors of bats, with regard to the now

permanently lost 1’.

A large number of Chiropteran genera (some 60 out of the now

recognisel 173 genera) have three pairs of lower incisors; in

most of these genera the lower incisors are subequal in size;

in those few in which one pair is noticeably, or even considerably,

reduced, this pair is i, (compare f.1. Rhogeessa, Baodon, some

species of V2 yctinomus, Mormopter ws). From this it appears safe

to assume that in bats which have only two pairs of lower incisors,

the missing pair is 1, |

No bat has more than three pr emolars, above and below. As

recently pointed out by Oldfield Thomas* the permanently missing

upper and lower premolar is in all probability p* and p, (not, as

hitherto taken for granted, p* and p,).

In those Phy llostomatidee which have three upper premolars

* Oldfield Thomas, “The Missing Premolar of Chiroptera,” Ann. & Mag. N. H.

(8) i. pp. 346-848 (April, 1908).

1908. | DR, KNUD ANDERSEN ON BATS. 207

(Lonchoglossa f. i.) p' is the smallest. From this it is concluded

that the upper premolar lost in Stenodermatous bats is p* (not,

as in Rhinolophide, p’).

p,, if present in Phyllostomatide, is generally smaller than p,

and p, (compare f. i. Wicronycteris). From this it is concluded

that the lower premolar lost in Stenodermatous bats is p,, not p,.

Upper incisors (text-figs. 41 4, D).—Inner pair bifid, the two cusps

subequal in length (the inner one generally a trifle longer) ; front

face plane or faintly convex, with slightly wrinkled enamel; hinder

face strongly concave in direction from above downwards, the lower

half of the crown of the tooth therefore somewhat chisel-shaped.

Outer pair as broad as inner pair, but much shorter; cutting-

edge simple (not bifid), oblique ; front face as well as hinder face

concave from side to side, The reason why the outer is con-

siderably shorter than the inner incisor, and its cutting-edge

Text-fig. 40.

Rhinolophus trifoliatus, 2 ad. Singapore. B.M, 4.8.23.1,

A. Right upper, B. Left lower tooth-row, exclusive of incisors; as a paradigma of

structure of molars in insectivorous bats, for comparison with dentition of

Artibeus, text-fig. 41. +.

For explanation of lettering of cusps (1, 2, 3,4, 5, 6, 7) see text, p. 205.

differently shaped, will be readily understood when studying the

way in which the lower work against the upper teeth :—the long

and very strong lower canines pass in front of the outer upper

incisors, completely covering their front face, with exception of

their narrow inner margin (text-fig. 41 p); this circumstance it is

which has effected a decrease in the size of the outer incisors,

made their front face concave (by constant wear against the tips

of the lower canines), and the inner tip of the cutting-edge (next

to inner incisors), which is less exposed to the pressure of the

lower canine, longer than the outer tip of the cutting-edge (next

to the upper canines), which is most exposed to the pressure of

the lower canine.

Lower incisors (text-figs. 41 B, D).—AlIl four teeth subequal in

Proc, Zoou. Soc.—1908, No. XIV. 14

208 DR. KNUD ANDERSEN ON BATS. [Apr. 7,

breadth and height (the outer pair, if anything, faintly shorter) ;

antero-posterior diameter of crown much greater than trans-

verse diameter; front face slightly concave from side to side, as

is also the cutting-edge; this latter circumstance gives the teeth

in front view a slight indication of a bifid shape.

Text-fig. 41.

Artibeus planirostris fallax, 9 ad. Kanuku Mts., British Guiana.

B.M. 1.6.4.60.

A. Right upper, B. Left lower tooth-row ; C. Side view of skull and mandible;

D. Front view of incisors and canines. A, B, D x 4, CX 3.

For explanation of lettering of cusps (1, 2, 3, 4, 5, 6, 7) see text, p. 205.

Upper canines (text-figs. 41 a, p).—Very long; cingulum low,

terminating abruptly in front and behind, but not developing

distinct secondary cusps; on the lingual face of the canine the

cingulum is somewhat expanded, forming a noticeable shelf-lke

projection ; against this expansion of the cingulum works the

principal cusp (cusp 4) of the lower p,. Hinder margin of canine

1908. | DR. KNUD ANDERSEN ON BATS. 209

sharp, inner margin (next to incisors) rounded, lingual face almost

plane. It is the constant rubbing of the labial face of p, against

the lingual face of the upper canine which has made this latter

sharp on its hinder margin and almost plane on its lingual face.

Lower canines (text-figs. 418, D).—Very long; on the inner

margin of the tooth, next to the incisors, the cingulum extends

only as high as (or very little higher than) the level of the cutting-

edges of the incisors, and does not form a secondary cusp (compare

Uroderma); at the base of the hinder margin the cingulum forms a

conspicuous shelf-like projection, caused by the constant working of

the tip of the upper canine against this part of the cingulum of

the lower canine.

p (text-figs. 41 a, c).—Cusps | and 2 are entirely wanting. Cusp

3 is represented by a small (but quite distinct), backwardly project-

ing prominence at the postero-external base of the tooth. Cusp 4

has disappeared. The principal cusp—large, trenchant, raised to

about half the height of the canine, very obliquely triangular, its

front margin only about half the length of its hinder margin—is

cusp 5. The size and shape of this cusp are correlated to the

large, very obliquely triangular interspace between the principal

cusp (4) of p, and p,; its front margin is precisely of the same

length as the hinder margin of cusp 4 of p,, against which it

works, and its hinder margin is of the same length as the front

margin of cusp 4 of p,, against which it works.—The tip of the

powerful cusp 4 of the lower p, working against the lingual

cingulum of p* has caused this latter to develop a rather strong,

somewhat concave projection (“ heel,” representing cusp 6); also

in the unworn p° the deep, pit-like depression caused by the

tip of cusp 4 of p, is very noticeable. The anterior margin of the

heel of p* is high, prominent, sharp, acting against the trenchant

hinder margin of cusp 4 of p,.

p’ (text-figs. 41 a, c).—Longer at base, much higher and broader

(transversally) than p*, but in many important details of its struc-

ture formed after a similar pattern, though on the whole less

reduced in size. Cusps 1 and 2 entirely wanting (asin p*). Cusp 3,

rather less reduced than in p*, is represented by a low trenchant

margin at the postero-external extremity of p’. Cusp 4 (wanting

in p’) is present as a mere rudiment at the anterior extremity of

the tooth. The principal cusp—still more powerful than in p*, the

highest cusp in the upper postcanine series, trenchant, with the

outline of an equilateral triangle—is cusp 5; its shape is re-

markably like that of cusp 4 of p,, against the hinder margin of

which it works; the strong antero-external cusp (cusp 4) of m,

acting upon the lingual face of cusp 5 of p* has made this latter

somewhat concave in antero-posterior direction.—The whole an-

terior portion of m, (its cusp 4 and the rudiment of cusp 1, when

this latter has not completely disappeared) acting upon the lingual

cingulum of p‘ has caused this latter to develop a ‘ heel” (cusp 6),

broader and a little more complicated than in p’; the heel is

broader, because it is acted upon by the broad anterior portion of m,,

14*

210 DR. KNUD ANDERSEN ON BATS. (Apr. 7,

whereas in p’ the heel is acted upon only by the pointed cusp 4 of

p,; the tip of eusp 4 of m, has made a deep, pit-like impression

on the heel of p*, just at the median point of the labial base of

cusp 9; also the high, slender, coniform cusp 2 of m, rubs against

the heel of p, viz., against its postero-internal margin, w hich by

this pressure is kept low and rapidly worn somewhat concave.

The anterior margin of the heel of p*‘ is, like the corresponding

margin of p’, coisa high, prominent, and sharp, for a similar

reason : it fits into and works against the trenchant commissure

between cusps 4 and 5 of ‘he Tower p,; on the antero-internal

margin of the heel of p* is seen a small, but quite distinct,

triangular, rather blunt cusp, produced by the small cusp 5 (and

its commissure with cusp 4) of the lower p, which catches and

works against it on its labial side.

p, (text-figs. 41 B, ¢).—Small, “ diamond ”-shaped, slightly longer

than high. The size and shape of its principal cusp (cusp 4) are

correlated to the size and shape of the triangular interspace between

the tip of p’ and the upper canine; cusp 4and a rudimentary cusp

at the front end of the tooth (probably representing cusp 1) work

against the lingual face and cingulum of the upper canine and

the front margin of cusp 5 of p®. The hinder margin of cusp 4

(together with a small, pointed, straightly backwardly extending

prominence of the posterior margin of its base, perhaps repre-

senting cusp 5) work against the projecting anterior margin of

the heel of p’. The lingual cingulum of p, is slightly expanded.

p, (text-figs. 41 B, c)—Viewed from the external side rather

closely resembling the upper p* in shape and size. Cusp 4 is

large, triangular, equilateral, by far the highest cusp in the post-

canine series; it worls against the lingual face and , posterior

margin of cusp DO! 10 the front margin of cusp 5 of p*, and the

heel of p*®, on which its tip has produced a deep depression.

A triangular emargination of the commissure between cusp 4

and the rudimentary cusp 5 (this latter situated at the postero-

external corner of the tooth) has been produced by the elevated

anterior margin of the heel of p*, which fits into this emar-

gination. The small cusp 5 eatches the labial side of, and

works against, the small antero-internal cusp on the front margin

of the heel of p*. The lingual, low, and cingulum-like portion

of the tooth is rather larger than in p,, chiefly owing to its

action against the front of the heel of p*; there can be little

doubt that this lmgual portion of p,, like the corresponding

portion of p,, in fact represents the degenerated cusp 2 (com-

are the lower premolars in insectivorous bats).

m* (text-figs. 41 A, c)—Enormously expanded in transversal

direction, its breadth being about 14 its length at the labial

margin; considerably shorter at lingual than at labial margin.

Cusps 1, 2, and 3 have entirely disappeared. The external, tren-

chant margin of m‘ is formed anteriorly by the triangular cusp 4,

rising to about half the height of the principal cusp of p‘, pos-

teriorly by the much lower, obliquely triangular cusp 5; cusp 4

1908. | DR. KNUD ANDERSEN ON BATS. SAL IL

works against the posterior margin of csp 4 and the anterior

margin of cusp 5 of the lower m,, cusp 5 against the posterior

margin of cusp 5 of m, and the whole of cusp 4 of m,. At the

antero- internal corner of the heel of m* (at level with the lingual

margin of the heel of p*) is seen the low, but strong, triangularly

projecting cusp 6; it fits into a depression in m,, immediately

behind cusp 2 of this latter tooth, and has checked the growth of

cusp 3 of m,, which consequently has become quite neon.

The large postero- internal, inwardly projecting lobe of m* repre-

sents cusp 7. ‘The whole lingual portion of m', bordered externally

by cusps 4 and 5, in front. by cusp 6, inter nally by the lingual

margin of cusp 7, and behind by the very low posterior margin of

the ‘tooth, forms a large crushing surface, the enamel of which is

densely wrinkled and extends on the lingual face of cusps 4 and 5

almost to their tips, thus forming an “ inner cingulum” to these

cusps. This crushing surface consists euledy of two concavities ;

the one, bordered externally by cusps 4 and 5, internally by cusp .

wears against the whole posterior portion (cusps 3 and 5) of m,

the other, immediately behind cusp 6 and the somewhat project-

ing antero-internal margin of cusp 7, is acted upon by the high,

slender, conical cusp 2 of m,.

ta (text-figs. 41 A, c). —Broader than long, but not so broad as

; the elements are the same as in m’, but their arrangement

Bees hal different. Cusps 4 and 5 strong, but much lower than

inm'. The shape and size of cusp 4 is determined by that portion

of the lower m, against which it has to work, viz., the hinder

margin of cusp 4 and the front margin of cusp 5. Cusp 5 is not

(as in m’) situated in a line immediately behind cusp 4, but has

moved to the middle of the posterior margin of the tooth, where

it forms a strong, backwardly projecting tubercle; this shifting of

the position of cusp 5 has been necessitated, because it has to

work against the hinder margin of cusp 5 of m, and the small m,

Cusp 6 has almost exactly the same position, shape, and size as in

m'; it acts upon the anterior and external face of cusp 3 of m,.

Cusp 7, which in m* is so enormously developed, is in m* quite

small, represented by a low, but perfectly distinct shelf at the

postero-internal corner of the tooth ; its small size is easily under-

stood when seeing that it has to work only against the posterior

margin of cusp 3 of m, and the front of the very small m,. The

andes portion of m? forms a lar ge » crushing surface, the enamel

of which is densely wrinkled, as in m', and produced into a distinct

inner cingulum to cusps 4 and 5. This crushing surface is deeply

hollowed ‘out in the middle, owing to the strong pressure of the

whole posterior portion (cusps 3 and 5) of m,.

m°* (IBS -figs. 41 A, c).— Rudimentary, as small as a lower incisor

(scarcely 51. the size of m*). The tooth has been pushed postero-

internally to m*, pressed into an angular emargination between

cusps 5 and 7 of this latter tooth ; its elements cannot be discri-

minated. Quite functionless the tooth is not; its antero-internal

portion is acted upon by the posterior portion of the small lower

212 DR. KNUD ANDERSEN ON BATS. (Apr. 7,

m,; but its postero-external portion cannot, so far as I can see,

be touched. by m,.

m, (text-figs. 41 B, c).—Longer than broad, abruptly narrowed in

front. Cusp 1 practically wanting ; in some specimens of 4. plant-

rostris (and, more often, in A. jamaicensis) a faint trace of cusp |

is detectable, as an excessively small prominence, immediately in

front of cusp 2, but generally it has entirely disappeared and only

its commissure with cusp 4 been partly preserved ; it is the constant

pressure of this portion of m, against the anterior margin of the

heel of p* which has caused the disappearance of cusp !. Cusp 2

strongly developed, as a high slender cone, situated very near the

middle of the labial margin, close to, but not contiguous with,

the cingulum ; this cusp works against the postero-internal portion

of the heel of p*, which it has pressed low and concave. Cusp 3,

at the postero-internal corner, extremely small, barely projecting

above the level of the cingulum, acts against the hinder face of

cusp 6 of m’. Cusp 4 (antero-externally) long, but low, tri-

angular, trenchant, acting against the heel of p'; the action of

this cusp is the chief cause of the strong development of the heel

of p*. Cusp 5 (postero-externally) long, much lower than cusp 4,

only slightly projecting, triangular ; it works against the external

half of the crushing surface of m* (the depression bordered by

cusps 4-5 and cusp 6 of this tooth). The enamel of the crushing

surface of m' is densely wrinkled.

m, (text-figs. 41 B, ¢).—Slightly smaller than m,, subrectangular,

a little ionger than broad. Cusp | is represented by a very small

(but distinct) tubercle at the middle of the front margin of the

tooth. Cusp 2, antero-internally, quite of the same shape as the

corresponding cusp in m,, only slightly lower; the action of this

cusp is the chief cause of the strong development of cusp 7 in m’.

Cusp 8, postero-internally, quite small, but not so strongly reduced

as in m, ; it acts against the postero-external face of cusp 6 and

the front of cusp 7 of m®. Between cusps 2 and 3 the cingulum

has developed an exceedingly small supplementary cusp. Cusps

4 and 5 very similar to the corresponding cusps of m,, but much

lower; the former acts against the posterior portion of the

crushing surface of m* (lingually to cusp 5), the latter against the

large crushing surface of m* (between cusps 4 and 5 externally,

and cusp 6 internally), which it has made deeply concave.

Crushing surface of m, wrinkled as in m,.

m, (text-figs. 41 B, c).—Rudimentary, 1-75 the size of m,,.

Cusps 2 and 4 are rather easily detectable. It works against the

postero-internal margin of m’* and the antero-internal portion of

the rudimentary m’.

UropermaA Pet.

1865. Uroderma Peters, MB. Akad. Berlin, pp. 587-88, footnote-—Type: Phyllo-

stoma personatum Pet. 1865 (not Wagner) = Uroderma bilobatum Pet. 1866.

1878. Artibeus Leach (partim), Dobson, Cat. Chir. Brit. Mus. p. 514.

1901. Uroderma Pet., Rehn, Proc. Ac. Nat. Sci. Philad. 1900, p. 757 (9 Febr.

1901).—Remarks on the genus,

1908. | DR. KNUD ANDERSEN ON BATS. 213

The subjoined characterisation is confined to the points in

which Uroderma differs from Artibeus.

Skull (text-fig. 42, compare fig. 43).— Long and slender, in general

shape somewhat recalling a Putorius nivalis skull—Rostrum but

Text-fig. 42.

Uroderma bilobatum, 2 ad. Para. B.M. 1.7.19.4.

Upper, lower, and side view of skull; front view of incisors and canines.

A, B, C x 2, Dx £,

Text-fig. 43.

Artibeus cinereus cinereus, gad. Para. B.M. 1.7.19.3.

Upper, lower, and side view of skull; front view of incisors and canines

(for comparison with Uroderma). A, B,C x 3, D x 4.

214 DR. KNUD ANDERSEN ON BATS. [Apr. 7,

very slightly depressed; profile of skull, therefore, showing an

almost straight line from the highest point of the brain-case to

the tip of the nasals (text-fig. 42 c); height of rostrum at p*

greater than, or at least equal to, width of skull at ‘“ postorbital ”

constriction (immediately behind postorbital processes or their

rudiments). In Artibews the rostrum is considerably more de-

pressed and flattened ; the outline in profile, from the front of the

sagittal crest to the base of the nasals, steep (text-fig. 43 c); the

height of the rostrum at p* considerably less than the width of

the postorbital constriction.—Bony palate long: distance from

palation to front of incisors very nearly equal to zygomatic width

of skull (text-fig. 428); in Artibeus much less than zygomatic

width (text-fig. 43 B)—Median backwardly extending portion of

bony palate (behind m*) long, equal to the combined length

(externally) of m’ and m’*; in Artibews much less than this latter,

often only equal to the length of m’—Anterior nasal opening less

oblique, looking chiefly forwards; in Artibews noticeably more

oblique, looking upwards and forwards.

Teeth (text-figs. 44 and 47).—Chief characters, as compared

with the teeth of Artibeus (see text-figs. 46, on p. 215, and 48, on

p- 216) :—outer upper incisors bifid; cusp 2 in m, small and more

anterior in position; m*and m, always present; m’° situated direct

behind (not postero-internally to) m*, and almost as broad as the

hinder border of this latter.—The details are these :—

(1) Cutting-edges both of inner and outer upper incisors bifid

(text-fig. 42 D, on p. 213); in Artibeus, inner incisors bifid, outer

incisors simple (text-fig. 43 D, on p. 213).—(2) The cingulum of

Text-fig. 44.

Uroderma bilobatum, § ad. Para. B.M.1.7.19.4.

A. Rizht upper, B. Left lower tooth-row. X +.

For explanation of lettering of cusps see text, p. 205.

the lower canine extends, on the inner side, upwards to (or

almost to) the middle of the tooth, often terminating in a small

cusp-like projection (text-fig. 47); in Artibews the cingulum ter-

minates at about the level of the cutting-edges of the lower incisors,

1908. ] DR. KNUD ANDERSEN ON BATS. 215

without forming a cusp-like projection (text-fig. 48)—(3) Cusp 4

(anterior cusp) of p* more developed than in Artibeus, as a rule

forming a small, but distinct, pointed cusp in the unworn tooth ;

in Artibeus cusp 4 of p* is practically completely wanting.—

(4) m* is small, but not nearly reduced to the same degree as in

Artibeus, situated direct behind m’*, and almost (or quite) as broad

Text-fig. 45.

Enchisthenes harti, g imm. Trinidad. Type, B.M. 92.9.7.8.

A. Right upper, B. Left lower tooth-row. X #.

Text-fig. 46.

Artibeus planirostris fallax, 2 ad. Kanuku Mts., British Guiana.

B.M. 1.6.4.60.

A. Right upper, B. Left lower tooth-row. x #

For explanation of lettering of cusps see text, p. 205.

as the hinder margin of m’ (text-fig. 44 4); of the elements of m*,

cusps 4 and 6 are clearly observable, cusps 5 and 7 only present

as mere rudiments; in cross section m* is about six times the size

of a lower incisor, or 1 of m*; in Artibews m*, when present, Is

216 DR. KNUD ANDERSEN ON BATS. Aton;

quite rudimentary (equal to a lower incisor, or about ; of m°),

its elements cannot be discriminated, and the tooth has been

pushed postero-internally to m’* (text-fig. 46.4); but in most

species of Artibews m* is entirely wanting.—(5) As a consequence

of the larger size and posterior position of m* in Uroderma, cusp 5

of m* is considerably more labial in position (text-fig. 44 4); in

Artibeus the cusp has moved so far towards the lingual side as

to occupy, precisely or very nearly, the middle of the posterior

margin of the tooth (text-fig. 46 a).—(6) Cusp 2 of m, is in

Uroderma represented by a low subacutely pointed tubercle near

Text-fig. 47. Text-fig. 48.

Text-fig. 47.—Uroderma thomasi, ad. Bellavista, Bolivia. Type, B.M. 1.2.1.37.

Front view of lower incisors and canines. X 4.

Text-fig. 48.— Artibeus jamaicensis lituratus, Q ad. Villa Rica, Paraguay.

U.S. N. M. 105587.

Front view of lower incisors and canines. X #.

the front end of the tooth and close to the lingual side of cusp 4

(text-fig. 44.8); in Artibews cusp 2 is very strongly developed,

rising as a high slender cone near the middle of the lingual

margin of m, (text-fig. 46 B).—(7) In accordance with the less

reduced size of m°, also m, in Uroderma is proportionately larger,

equal to 3-z of m, (text-fig. 448); in Artibeus m, is 3-1; the

. a . . 8 .

Size of m,, or, in more than half the number of species, entirely

wanting.

External characters.—The lateral margin of the horseshoe, at

level with nostrils, is turned upwards so as to form a conspicuous

fold; when pressed downward to the muzzle this fold takes

the shape of a small, rounded lobe, slightly projecting beyond

the rest of the lateral margin. In several species of Artibeus

(A. planirostris, jamaicensis, etc.) there is a similar, though rather

less pronounced folding of the lateral margin of the horseshoe.

The difference between Uroderma and Artibeus in this respect is,

therefore, only one of degree.

The wing-structure is very similar to that of Artibews; the

fitth metacarpal averages a trifle shorter than the third, whereas

in all species of Artibeus it is generally a trifle longer than the

third. How closely in all other respects the wing-structure of

Uroderma resembles that of Artibews may be seen by reference to

the wing-indices on p. 310 (compare, for instance, the indices of

Uroderma with those of A. rosenbergi).

1908. | DR. KNUD ANDERSEN ON BATS. 217

Uroderma has a narrow line of whitish fur down the middle of

the upper side, sometimes ill-defined, but never quite obliterated ;

Artibews has no trace of a dorsal line.

Species.—Two, U. bilobatum and thomasi.

Range.—From Sao Paulo and Bolivia to Costa Rica; un-

represented in the West Indies (Trinidad excepted).

Remarks.—Uroderma was proposed by Peters (J. s. ¢.) as a

generic name for those known species of “ Artibews” which have 2

molars, viz., at Peters’s time, A. bilobatus, fallax, and concolor. The

name, in this sense, was adopted by Dobson (J. s. c.), though only

as the designation of a “subgenus” of Artibeus, and he recognised

two species only, 4. bilobatus and planirostris, the latter including

Peters’s fallax as a “synonym” and concolor as a “ variety.” The

number of species would now be five: bilobatus, thomasi, concolor,

planirostris, hirsutus.—In 1901, Rehn (J. s.c.) proposed to restrict

the name Uroderma to A. bilobatus, on account of its elongate

skull, noticeably higher rostrum, and “two rounded lobes” on

the lateral margins of the horseshoe.

Any subdivision of the genus “ Artibeus,” in its old sense,

according to the number of molars (3, 3, or 3) is artificial. The

natural subdivisions are these two: A. bilobatus and thomasi on

one side (Uroderma), all other species on the other (Artibeus).

There is a wide gap between Uroderma and <Artideus, in this

sense, whereas all species referred to Artibeus in the present

payee, irrespective of the presence or absence of the rudimentary

m* and m,, are extremely closely inter-related. Uroderma differs

both in the shape of the skull and in several important dental

characters, and it 1s in these respects not approximated by any

species of Artibeus. As to the dental characters pointed out

above (pp. 214-216), nos. 1 (ote baits outer upper incisors), 4 (posi-

tion of m®*), 5 (position of cusp 5 of m°), and 6 (no cusp 2 in m,)

constitute absolute differences between Uroderma and Artibeus,

though a slight restriction is perhaps advisable with regard to

no. 1, in so far as in some species of Artibews the outer upper

incisor can show, rarely and as a perfectly individual aberration,

a faint leaning towards a bifid shape; as to nos. 2 (cingulum of

lower canines), 3 (cusp 4 of p*), and 7 (size of m,), the difference

is one of degree only. In having a narrow line of white fur down

the middle of the upper side, Uroderma is similar to the majority

of species of Vampyrops and allied genera, but different from all

species of Artibeus.

URODERMA BILOBATUM Pet.

1842. Phyllostoma species inedita Ruppell, Verzeichniss der in dem Museum der

Senckenbergischen naturforschenden Gesellschaft aufgestellten Sammlungen,

i. p. 11, no. Il. D. 3 a.—Brazil ?

1865. Phyllostoma personatum (not Wagner) Peters, MB. Akad. Berlin, pp. 587-88,

footnote.

1866. Uroderma bilobatwn Peters, MB. Akad. Berlin, p. 394.—Sao Paulo; Cayenne.

1878. Artibeus bilobatus Pet., Dobson, Cat. Chir. Brit. Mus. pp. 518-19,

1880. Artibeus (Uroderma) bilobatus Pet., Thomas, P. Z. S. p. 396.—Sarayacu,

Keuador.

218 DR. KNUD ANDERSEN ON BATS. [ Apr. 7.

1889. Artibeus bilobatus Pet., Cope, Amer. Naturalist, xxiii. no. 26, pp. 130-31

(Febr. eae Chapada” [probably Chapadas da S. Maria, N. Minas

Geraes |.

1897. <Artibeus bilobatus Pet., J. A. Allen SA aca Bull. Am. Mus. N. H. ix.

Art. ii. p. 15 (26 Febr. 1897) .—Trinidad, W. I

1900. Uvoderma bilobatum Pet., J. A. Allen, Bull. Am. Mus. N. H. xiii. Art. viii.

p. 89 (12 May, 1900). —Santa Marta region, Colombia.

1901. Artibeus bilobatus Pet., Thomas, Ann. & Mag. N. H. (7) vii. p. 191 (Sept.

1901).—Para.

1902. Uvroderma convexum Lyon, Proc. Biol. Soc. Wash. xv. pp. 83-84 (25 April,

1902).—Colon, Panama.

1904. Uroderma bilobatum Pet., J. A. Allen, Bull. Am. Mus. N. H. xx. Art. XXXV.

p. 458 (28 Nov. 1904). —Ciudad Boliv yar, Venezuela.

1906. Ur ‘oderma bilobatum Pet., Peters, Chir. Mus. Zool. Berol. pl. xi. a (issued

Jan. 1906).

Diagnosis.—Skull small, tooth-rows short, ears small.

U. bilobatum and thomasi.—The difference between these two

species will be pointed out below, p. 221.

Hairing on limbs and interfemoral.Forearm densely haired

for the proximal two thirds of its upper surface. A tuft of

short hairs on the metacarpal of the pollex. Upper side of tibia

and foot distinctly haired. Interfemoral very short-haired, its

hinder margin almost naked.

Colowr.—General impression brown, varying in shade; four

facial stripes, a dorsal stripe. There are two colour extremes,

a dark brown and a light brown, but the contrast between them

is by no means great.

Dark-coloured examples (many skins, all of fully adult in-

dividuals, teeth unworn or slightly worn; localities: Colombia

(Cali; Santa Marta), islands off Panama, Chiriqui) :—Upper side

dark brown, darker than Ridgway’s Prout’s brown ; base of hairs

on hinder back approaching drab, on the neck lighter, varying

from light drab to wood-brown or ecru-drab. Under side a dark

shade of drab. Supraorbital and infraorbital stripes broad,

whitish. A narrow whitish longitudinal stripe from the occiput

to the interfemoral; front half of the stripe sometimes very

indistinct or quite obliterated. A more or less distinct narrow

whitish margin to the ears.

Three examples (Chanchamayo, Peru, and Brava I., W. of

Panama; adults, with unworn or very slightly worn teeth) are

noticeably darker: upper side sooty brown, under side dark smoky

grey.

Lighter-coloured examples (two skins: Hgas, Amazonas, and

Valencia, Venezuela; adults, with unworn teeth) :—Upper side

Prout’s brown (one skin) or mars-brown washed with russet (the

other), base of hairs wood-brown ; under side light drab. Facial

stripes, dorsal line, and ear-edgings as usual.

Individuals from different localities——Specimens have been

examined from localities dotted over practically the whole area

from Para, Amazonas, and Peru in the south, to Costa Rica in

the north. I am unable to see any differences, in the skull, teeth,

or external characters, between individuals from all these localities.

The subjoined table of measurements (p. 219), in which the

specimens have been arranged according to their geographical

habitat, shows that also the dimensions are the same.

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220 DR. KNUD ANDERSEN ON BATS. [Apr. 7,

Specimens examined.—-24 specimens (19 skins) and 23 skulls,

from the following localities :—

British Museum :—Brazil: Para (1); Egas (Teffé), Amazonas

(1).—Peru: Chanchamayo, 1200 and 1500 m. (2).—Kcuador :

Sarayacu (1).—Colombia: Cali, 1100 m. (1); Onaca, Santa Marta,

700 m. (2).—Venezuela: Valencia (1).—Panama: Colon (1);

Chiriqui (2); islands off Panama: Brava I, (3); Cebago I. (1);

Jicaron I. (1); Insoleta (1); Gobernador I. (3).—Costa Rica ;

Miravalles, 400-500 m. (1).—21 skulls, from all the localities

enumerated.

U.S. National Museum * :—Trinidad: Port of Spain (1).—

Panama: Colon (1).—Skulls of both specimens.

Range.—From Sio Paulo and Peru, at least as far north as

Costa Rica; unrepresented in the West Indies (Trinidad

excepted).

Peters’s Uroderma bilobatum, 1866.—The type, in the Berlin

Museum, is ‘ein jiingeres Exemplar aus St. Paulo in Brasilien” ;

Peters had also ‘‘ zwei andere ausgewachsene [| Exemplare] aus

Cayenne” in the Berlin Museum, and “ein miinnliches aus-

gewachsenes Exemplar in Weingeist” from the Frankfurt

Museum without exact locality (cf. Riippell, 7. s. c.). The whole

of the original description and all measurements (there is an

obvious error in the measurement of the second phalanx of the

fourth digit) precisely agree with the series of specimens here

referred to U. bilobatwm.—The figures in the plate belonging to

Peters’s intended Monograph of Bats (/. s. ¢.) are excellently

drawn and partly well reproduced, but the hind legs in the life-

size figure (fig. 1) are much too short, as if drawn from a damaged

specimen (with broken legs 2).

Lyon’s Uroderma convexwm, 1902.—Type: 2 young ad.; Colon,

Panama. Based on two specimens from the type locality. For

comparison Lyon had two U. bilobatwm from Chapada, Brazil

(probably Sao Joaio River, Chapada da 8. Maria, N. Minas Geraes ;

and probably the same specimens as recorded by Cope, 1889, /. s. ¢.,

and by Rehn, 1901, /. s. ¢.).—The characters of U. convexum are

summed up, by Lyon, as follows: “Similar to U. bilobatum Peters,

but with tooth-rows distinctly arcuate” (‘less nearly parallel

than those of U. bilobatwm”).

Besides a specimen from Colon in the British Museum (pre-

sented by Marquis Doria), nine specimens from islands W. of

Panama, two from Chiriqui, and one from Costa Rica, I have had

for examination Lyon’s paratype, a young adult from Colon.—

“ U. convenum” is in every respect indistinguishable from JU,

bilobatwin from Brazil, Peru, Ecuador, Colombia, Venezuela, and

Trinidad. In the whole series of skulls of U. bélobatwm examined,

23 in number, the upper tooth-rows are decidedly arcuate; by

close comparison of the skulls an excessively small variation in the

outline of the tooth-rows is, of course, observable, as is also.the

* US. N. M. nos. 22472 (37901), 111721.

1908. ] DR. KNUD ANDERSEN ON BATS. 221

case in any sufficiently large series of skulls of any species of

Artibeus, Vampyrops, &c.; but these minute variations are

entirely independent of differences in the geographical habitat of

the individuals. The straightest tooth-row I have seen is in a

skull from Colon (the type locality of U. conveauwm) and in one

from Chiriqui; the most arcuate in one from Peru, one from Para,

and one from Chiriqui; the others are, of course, intermediate.

UropEerMA THoMASI K. And.

1906. Uroderma thomasi Knud Andersen, Ann. & Mag. N. H. (7) xviii. p. 419

(1 Dec. 1906).—Bellavista, Bolivia.

Diagnosis.—Similar to U. bilobatum, but with noticeably larger

skull, longer tooth-rows, and larger ears and nose-leaves.

U. thomasi and bilobatum.—U. thomasi differs from U. bilobatwm

in the following particulars :—

The skull is precisely of the same shape as in U. bilobatum, but

in every respect larger; the largest skull (out of 23) of U. bilo-

batum measures in total length 23°3 mm., the smallest skull of

two U. thomasi 24-7 mm.; all other dimensions of the skull are

correspondingly increased.—In the whole series of U. bilobatwm

examined the length of the maxillary tooth-row varies between

7-8 and 8:5 mm. (average 8°1 mm.), in U. thomasi it measures

8-9-9 mm. ‘The ears are not only absolutely, but proportionately

larger. The lancet longer and, especially, broader.—For further

details see the table, below p. 223.

U. thomasi is, probably, the Bolivian representative of U. bi-

lobatum ; but in all the points referred to above there seems to be

a perfectly clear line of separation between the two forms; the

gap between them is not overbridged by any specimen I have

seen.

The distribution of the fur on the limbs and interfemoral is as

in U. bilobatum. ‘The colour of the pelage as in the ordinary

dark brown “ phase ” of that species ; the whitish ear-edgings are

very distinct.

Specimens examined.—Two, with skulls, viz., ¢ ad., Bellavista,

Bolivia, 1400 m., about 15°S8., 68° W. (type specimen) ; and ¢ ad.,

Reyes, Bolivia, about 13° 8., 67° W. (presented by Marquis

Doria). Both specimens in the collection of the British Museum.

Range.—As yet known from N., Bolivia only.

ENCHISTHENES K. And,

1906. Enchisthenes Knud Andersen, Ann. & Mag. N. H. (7) xviii. p. 419 (1 Dec.

1906).—Type: Artibeus harti Thos.

Diagnosis—Allied to Artibeus, but median upper incisors

simple; m* in row, as broadas the hinder margin of m*; m, com-

paratively large, equal to about + of m, ; tragus with a pointed

projection on the inner margin, near the tip.

222 DR. KNUD ANDERSEN ON BATS. [Apr. 7,

Enchisthenes, Artibews, and Uroderma.—Enchisthenes is much

more closely allied to Artibeus than to Uroderma. In all the

characters which separate Uroderma from Artibeus, Enchisthenes

agrees with the latter genus, the following points excepted: the

position and relative size of m* and the relative size of m, are as

in Uroderma; asa consequence of the former fact, also the position

of cusp 5 of m® is as in Uroderma. Hnchisthenes differs from

Artibeus chiefly in the particulars referred to above, in the

diagnosis of the genus.

Principal characters —The skull has all the characters of an

Artibeus skull: short and broad, not long and subeylindrical as

in Uroderma ; profile, from front of sagittal crest to nasals, much

concave, as in Artibeus, not very slightly concave or almost straight,

as in Uroderma; palate short, as in Artibeus, not long as in Uro-

derma; plane of anterior nasal openings very oblique, as in

Artibeus, not looking chiefly forward, as in Uroderma.

Outting-edges of median upper incisors simple, pointed in the

centre, without any indication of a median notch (text-fig. 49 c) ; in

this respect Hnchisthenes differs both from Artibeus (median upper

incisors bifid) and Uroderma (all upper incisors bifid), Outer

upper incisors somewhat narrower and much shorter than median

. Text-fig. 49.

Enchisthenes harti, g imm. Trinidad. Type, B.M. 92.9.7.8.

A. Right upper, B. Left lower tooth-row ; C. Front view of upper incisors

and canines. X 4.

3 4

incisors; cutting-edge simple. and p* as in Artibews and

Uroderma. Cusp 7 of m* well developed, but rather small, more

recalling the corresponding cusp in Uroderma than that of Artibeus.

Position of cusp 5 of m® as in Uroderma, not at (or very nearly at)

the middle of the hinder margin of the tooth, as in Artibews ; cusp 7

of m? so excessively small as to be scarcely observable without a

lens, forming only a very low postero-internal margin to the tooth.

m® situated direct behind m*, in position and shape quite as in

Uroderma, not rudimentary and situated postero-internally to m’,

as in Artibeus (compare text-fig. 45 with text-figs. 44a and 46 4

on pp. 214 and 215), Cingulum on inner side of lower canines

1908. | DR. KNUD ANDERSEN ON BATS. DD

(next to i,) low, terminating at level with cutting-edges of incisors,

as In Artibeus. Cusp 4 of p’ practically completely wanting, as

in Artibews. Cusp 2 of m, very much as in Artibeus (perhaps a

trifle more anterior in position); the cusp is situated so close to

the lingual cingulum as to appear, on cursory inspection, in direct

connection with this latter; in reality the cingulum passes the

lingual side of the cusp, as in Artibeus; between cusps 2 and 3

the cingulum has developed two very small supplementary

tubercles. Cusp 1 of m, (antero-internally, direct in front of

cusp 2) small, but quite distinct, m, of the same relative size as

in Uroderma, viz. equal to about + the bulk of m,,

A very conspicuous, pointed, upwardly directed projection on the

inner margin of the tragus, about 1 mm. below the tip. Neither

Measurements of Uroderma bilobatum and thomasi, and

Enchisthenes harti.

a a ee = -

| U. bilobatum. | U. thomasi. |E. harti.

23 adults, | 2adults, | Type,

22 skulls. | 2skulls. |¢ yg. ad.

Min. Max.

Min. Max. Mep.

mm. mm. mm. | mm. mm. mm.

Skull, total length, to front ofc ... 22 233 92°83 | 94:7 24:8 20°8

op TANEYSEONGL WGI socococeoscsancoos| Ill WS} Ts. |} 111) ill

» Width of brain-case............ 97 10 99 | 106 10°38 97

oy ANROUMCNNO WGI coosercoocee nn) ORS RY IS} | i, ie) So

» maxillary width, across m1... a8 O8 OF | 1@ iO 83

| 5, across cingula of canines...... 55 62 50 | Gail oe 58

| Mandible, to front of inc. .............. 15 162 156 | 16:8 16:8 142

| Wistorere weet, GHWT suctaosnoeecesssces)) Ye HS Pl | 8 © 72

| Lower teeth, c-m3 ..................... Sion) SO | OS o%7 8

| Kar-conch, length, immer margin... 11 = 11°5 ) U2) 18}

RS length, outer margin -.| 157 168 (el Siero)

5 WOU SIN oonesabemdenucosaunondall ya lk 12 12:8 13°7

AMPEG), MEVEtHN scocen oon son bbe ess eeecoa cas 58 5:8 le ze 7

| IDE WOVGE te, METER 3 cco nonncb dor sea anseceesal 83 9:2 1@ 10) 6

Su peel Ob igen ine (as eee iawmeee [Eb AR Oe 5 | 62 65 5

Horseshoe, width eee] 62 75 78 8 6:2

LNGIREENAIO® <Geskaopndcbencmatasaveeeeaysonocs|| 440) is) AIgy I idl 38

Bollexiare saree hs edema eeeeerl nmelelt Wee I} IB TI 102

| GL WNETEKCADEN oso snassononsbeasconsoccal| CYite) ZPRR AOS 42 Ag 345

ATT yh tee aa ah Ae Mera tanya teresa ma] 16 14°9 148 165 13

Ot Dem cece ER eee acboce dbo tatiracriie cacll nel 23°8 22°3 232 23:5 17-2

IIs posnocsobseacvgeavecscnacoreHaue noone LOM BP TLile7/ 13:2 14 12

Ath metacarpal..........................| 868 41:2 39:1 40° 41°7 33°2

ANS a peseeer rn enomaa ae aman seu ineelh ampli: pS Roe TO Ny/ 12°8 13°5 11:7

LE ea cone aeceee Gaatiaactnscnaeaudasonneliea pl dlepey) alia) 13°7 142 14:8 10°8

Hilo WDEUXCEOAM co ccococcaosvcasasconao sell OY AIS 849%) 41-7 492 33°8

ee acon ok Siete Semen ners NAc Lnneets all 2 10:8 10 Oren lell 9°2

Vi2i ieee ee ok cei ae Cerra orn meal 13 121 12:2 12:8 92

[GAETMCNOVEM |, ceoocanaccaccprcucssusensee| lees 5 142 178

Wowerslegitn 3.2. -at eae ill al Sin eel 7i Suma Grae 17. —s «18 13

MOO, WH GENS coonatancccusocosscvecee| NOD WR les 11:8 12-2 10

(Cal cane ae cet rice: Beamer aml Nee AZ eG 53 58 68 5:2

Proc. Zoou. Soc.—1908, No. REV oy eee hs

224 DR. KNUD ANDERSEN ON BATS. | Apr. 7,

in Artibeus nor in Uroderma is there any trace of a similar

projection.

The material ig unsuitable for a description of the wing-

structure, the only specimen known being a young adult. It

would seem, however, that the wing-indices do not differ very

essentially from those of Artibews; the second phalanx of the

third digit is, apparently, relatively rather short, less than 13 the

length of the first phalanx.

Species. —The type of the genus is the only species known.

Range.—Trinidad.

ENCHISTHENES HARTI Thos.

892. Artibeus harti Thomas, Ann. & Mag. N. H. (6) x. pp. 409-10 (Nov. 1892).

Diagnosis.—Upper tooth-row 8 mm.; forearm 38 mm.

Nose-leaves.—Horseshoe in front completely fastened down to

and continuous with the integument of the muzzle; lateral part

turned up into a slight fold, somewhat recalling the fold in a

Uroderma horseshoe, but not forming a small rounded lobe.

Lancet unusually short and broad, the width at base being almost

equal to 2 its length from nostrils to tip.

Interfemoral.—Extremely short, only about 3-4 mm. in the

middle line.

Fur on limbs and interfemoral.—Upper side of proximal two

thirds of forearm, the whole of the interfemoral, and upper side of

tibia and foot, densely haired. <A tuft of very short hairs on the

metacarpal of the pollex.

Colour (of an immature specimen, preserved in aleohol).—The

colour of the fur would seem to be much as in the light phase of

Artibeus toltecus ravus (below p. 300). Details as to the facial

stripes and ear-margins cannot be given from the only specimen

available, which is not in a perfect state of preservation.

Measurements.—On p. 223.

Specimens examined.—One, the type, a young adult male, in

the collection of the British Museum.

Range.—As yet only one record, from the island of Trinidad,

W.I.

ArtiBpeus Leach.

1821. Artibeus Leach, Trans. Linn. Soc. London, xiii. pt. 1. pp. 74-75.—Type:

Artibeus gamaicensis Leach.

1821. Madateus Leach, op. cit. pp. 74, 81-82.—Type: Madateus lewisii Leach

(=Artibeus jamaicensis Leach).

1827. Medateus Gray, Griffith’s Animal Kinglom, v. p. 74.—Misspelling of

Madateus.

1835. Arctibeus Gray, Mag. Zool. & Bot. ii. pp. 486-87.—Misspelling of Artibeus.

1856. Pteroderma Gervais, Expéd. Castelnau, Mamm., livr. 15, p. 34, pls. vin.

fig. 7, x. fig. 1.—Type: “Pteroderma perspicillatum L.” (=Artibeus

jamaicensis Leach).

1856. Artibeus Gervais, op. cit. pp. 34-35, pl. ix. fig. 2.—Misspelling for Artibeus.

1856. Dermanura Gervais, op. cit. p. 36, pls. viii. fig. 4, ix. figs. 4, 4a, x1. fig. 3.—

Type: Dermanura cinereum Gervais.

1892. Artobius Winge, E Museo Lundii, ii. pt. i. p. 3 (Gf p. 38).—Nomen

emendatum.

1908. ] DR. KNUD ANDERSEN ON BATS. 225

Artibeus and Uroderma.—Artibeus differs from Uroderma in

the following particulars :—

Skull short and broad. Rostrum conspicuously depressed ;

profile of skull, from front of sagittal crest to nasals, therefore

much concave (text-fig. 43, on p. 213); height of rostrum at p*

much less than width of skull at “ postorbital” constriction (imme-

diately behind postorbital processes or their rudiments). Bony

palate shorter: distance from palation to front of incisors always

less (generally very much less) than zygomatic width (text-fig, 43,

on p. 213). Median backwardly extending portion of bony palate

(behind last molar) shorter, not equal to the combined length

(externally) of m* and m*, often equal only to the length of m’.

Plane of anterior nasal opening more oblique than in Uroderma.

Inner upper incisors bifid, outer upper incisors simple. The

cingulum of the lower canine terminates, on the inner side,

about the level of the cutting-edges of the lower incisors, without

forming a cusp-like projection (text-fig. 48, on p. 216). Cusp 4

(anterior cusp) of p* practically completely wanting (for this and

the following dental characters see text-fig. 41, on p. 208). m®

either quite rudimentary or, in most species, completely wanting ;

when present the tooth is situated postero-internally to m®. As

a consequence of the small size and postero-internal position of m°,

eusp 5 of m° is decidedly more lingual in position, occupying,

precisely or very nearly, the middle of the posterior margin of the

tooth. Cusp 2 of m, very strongly developed, rising as a high

slender cone near the middle of the lingual margin of the tooth.

In accordance with the rudimentary condition, or complete dis-

appearance, of m*, also m, is relatively smaller than in Uroderma,

or, in certain species, entirely wanting.

No species of Artibeus has a narrow line of whitish fur down

the middle of the upper side.

Artibeus and Hnchisthenes.—Artibeus differs from Enchisthenes

chiefly in the following respects :—

Cutting-edges of inner upper incisors bifid, not simple, pointed

in the centre, without any trace of a median notch, as in Hnchis-

thenes. m° rudimentary, situated postero-internally to m? or,

most often, entirely wanting, not relatively large and situated

direct behind m’*,as in Hnchisthenes. m, very small, equal to a-qs

of m* or entirely wanting ; in Hnehisthenes equal to about 7 Of

m?.—In no species of Artibews is there any trace of a pointed

projection on the inner margin of the tragus.

On the principal characters subject to specific variation.—(A)

Skull.—In the fourteen species referred to the genus Artibeus

in the present paper, three types of skull can be discriminated :-—

(1) The ordinary shape of the skulls, characteristic of all species

but three, is that figured on p. 213 (text-fig. 43): rostrum

moderately depressed, profile of nasals, from base to tip, very

nearly horizontal (not slightly ascending), palate not shortened.—

(2) In one species, A. concolor, the facial portion of une skull is

226 DR. KNUD ANDERSEN ON BATS. (Apa

peculiarly shortened (see p. 233).—(3) In two species, 4. turpis

and nanus, the depression and flattening of the rostrum and

heightening of the brain-case reach a climax, the rostrum being

even very slightly bent upwards (profile of nasals rather a little

ascending than horizontal), the palate shortened (see text-fig. 57,

on p. 307).

(B) Teeth.—The species fall into two sections, probably forming

two natural branches of the genus: those in which cusp 7 of m*

is comparatively small, viz. 4. glaucus, watsoni, cimereus, and

rosenbergi (text-figs. 53, 54,55); and those in which the cusp

is comparatively large, viz. A. concolor, planirostris, hirsutus,

jamaicensis, toltecus, quadrivittatus, phieotis, aztecus, turpis, and

nanus; in their extremes (upper extreme of the former and

lower extreme of tie latter section) these two sections come very

near to each other.—The rudimentary upper posterior and lower

posterior molar (m* and m,) can completely disappear; con-

sequently the number of molars varies between # (A. concolor,

planirostris, hirsutus), 2 (A. jamaicensis, glaucus, watsoni), and 3

(A. cinereus, rosenbergi, toltecus, quadrivittatus, pheotis, aztecus,

turpis, nanus).

cross-section, therefore, triangular. A notchin the outer margin,

at level with base of inner margin; below this notch a square-shaped

lobe, the upper and lower corners of which are produced into

sharp points (this lobe is, in the following pages, called “ the basal

lobe”); above the notch a sharply projecting point (in the following

pages, “the median projection”); outer margin, above the median

projection, asa rule serrate ; inner margin of tragus perfectly simple

from base to tip. In so far the tragus of Artibeus does not differ

appreciably from that of Uroderma.—The number of serrations on

the upper half of the outer margin, above the median projection,

is practically the same in all species but one, varying between 0

and 5; the variations within these limits 1s not specific, but indi-

vidual; the usual number is 4, 3, or 2; sometimes the serrations

are sharp, very often rounded, often reduced to very small

nodules, this latter leading, in extreme cases, to complete obliter-

ation of some, or all, of the serrations. In A. concolor I have

found the number of serrations to be 7-8, but only one specimen

has been available for examination.

(D) Nose-leaves.—In all species but two the front margin

of the horseshoe is free; in A. planirostris it is sometimes, in

A. jamaicensis often fastened down to, or even perfectly con-

tinuous with, the integument of the muzzle. In no species is the

lancet so short as in Hnchisthenes hart.

(E) Wing-structure.—Broadly speaking the wing-structure is

the same in all species: fifth metacarpal averaging a trifle longer

than third, fourth slightly the longest; second phalanx of third

digit a little less than, or equal to, or a little more than 14 the

length of the first phalanx.—The specific variation chiefly affects

the proportionate length of the first phalanx of the third, fourth,

1908.] DR. KNUD ANDERSEN ON BATS. DON

and fifth digits; in A. fallax, hirsutus, and jamarcensis (and

concolor ?) the first phalanx is proportionately shorter, its indices

being: third digit 281-298, fourth digit 245-260, fifth digit 186—

201; in all other species the phalanx is proportionately longer,

its indices being: third digit 327-357, fourth digit 279-304, fifth

digit 217-240.

(F) Hairing on limbs and interfemoral—tIn most species the

posterior part of the interfemoral and the upper side of the tibia

are very thinly haired or, at least on cursory inspection, almost

naked ; in a few species, especially 4. hirsutus, toltecus,and aztecus,

they are densely furred.

(G) Colour.—The specific variation in colour is extremely small,

the individual variation considerable. As a means to separate the

species of this genus colour-characters must, therefore, be used

with great caution.—The general colour pattern is this: upper side

some shade of brown, under side lighter; very often four facial

stripes ; often narrow whitish margins to the ears; sometimes

white tips to the wings. Young (not full-grown) individuals darker

and duller than adults.—In most, if not all, species there is a darker

and a lighter colour extreme, as a rule (perhaps always) connected

by several intermediate stages ; the light-coloured extreme some-

times occurs in full-grown specimens with quite unworn teeth,

7.é. In specimens which have evidently just reached the mature

age, but I have never seen it in decidedly immature (not full-

grown) individuals. A pair of white or whitish supraorbital and

infraorbital stripes are very often present, but they vary, some-

times even in the same species (4. jamaicensis, and others), through

all stages from complete absence to very strong development; as a

rule (not always) they are strongest in lighter-coloured indi-

viduals. White tips to the wings are most conspicuous in the

larger species (4. planirostris, hirsutus, jamaicensis), more in-

distinct or, as a rule, practically wanting in the smaller species.

(H) Size.—No less than ten species (A. glaucus, watsoni,

cinereus, rosenbergi, toltecus, quadrivittatus, pheotis, aztecus, turpis,

nanus) are, externally, approximately of equal size, the forearm

varying between 36°5 and 47 mm. ‘Three species (A. planirostris,

hirsutus, gamaicensis) ave noticeably, or much, larger: forearm

535-76 mm. One (A. concolor) is intermediate: forearm about

50 mm.

(1) Conclusions.—The principal, and in most cases the only

reliable, differences between the species are cranial and dental.

No specimen of Artibews ought to be identified without a careful

examination of the skull and teeth.

Species.—In 1878, Dobson catalogued 5 forms of Artibews, viz.

A. planirostris, A. pianirostris var. concolor, A perspicillatus,

A. cinereus, A. quadrivittatus. The total number of forms

recognised in the present paper is 25 (14 species). The following

table gives, in chronological order, a view of all the forms

named, their type localities, their identification in Dobson’s

Catalogue, and their identification in this paper :—

[Apr. 7,

DR. KNUD ANDERSEN ON BATS.

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1908. ] DR. KNUD ANDERSEN ON BATS. 229

Range.—From 8. Brazil, Paraguay, and Bolivia, to Sinaloa in

North Central Mexico, including the whole of the West Indies.

Geographical review of the species and subspecies. —The subjoined

geographical review is based almost exclusively on the material

examined by myself. An asterisk before a technical name indi-

cates that the type-locality of the species or subspecies falls within

the region under consideration.

Paraguay.—A, jamaicensis lituratus.

Brazil.—4. concolor (Para, Upper Amazonas).—* 4. planirostris

planirostris (Bahia, Matto Grosso, Pernambuco, Maranhao,

Para).—A. planirostris fallax (Para).—*A. jamaicensis lituratus

(Sta. Catharina, Parana, Minas Geraes, Bahia, Para).—* A. cine-

reus cinereus (Para).—<d. quadrivittatus (Pernambuco).

Peru.—4A. planirostris (? fallax ).—* A. glaucus.

Eeuador.—* 4. jamaicensis equatorialis.—A. jamaicensis litu-

ratus.—* A. rosenbergi.—*A toltecus ravus.

Guiana.—*4. concolor.—* A. planirostris fallax.—A.. cinereus

cinereus.—* A. quadrivitiatus.

Venezuela.—. planirostris planirostris.—A. planirostris fallax

(Lower Orinoco).—<A .jamwaicensis palmarum.—A . cinereus cinereus.

—A. cinereus bogotensis (N.W. Venezuela).—4A. rosenbergi.

Trinidad.—* 4. planirostris trinitatis—*A. jamaicensis pal-

marum.—A. cinereus cinereus.

Tobago.—d. planirostris trinitatis—| A. jamaicensis palmarum. |

Grenada.—* 4. planirostris grenadensis.—[ A. jamaicensis pal-

marum. |

St. Vincent.—A. jamaicensis palmarum.

Dominica, Guadeloupe.—* 4. jamaicensis preceps.

Colombia (excluding Panama).—4dA. jamaicensis cequatorialis

(Cali).—A. jamaicensis lituratus.—* A. cinereus bogotensis.

Central America (including Panama).—4d. planirostris plani-

rostris (whole region).— A. jamaicensis jamaicensis (whole region).

—A.jamaicensis palmarum (whole region).—* A. watsoni (Panama,

Nicaragua).— A. toltecus toltecus (Costa Rica, Nicaragua, Guate-

mala).

Belize and Yucatan.—* A. jamaicensis yucatanicus —* A. pheotis.

Mexico (excluding Yucatan). — A. planirostris planirostris

(Chiapas, Guerrero).—* A. hirsutus (Michoacan, Colima, Jalisco).

—A, jamaicensis jamaicensis (Campeche, Chiapas, Tehuantepec,

Oaxaca, Morelos, Vera Cruz).—A. jamaicensis palmarum (Oaxaca,

Vera Cruz, Jalisco).—* A. toltecus toltecus (Oaxaca, Vera Cruz,

Jalisco, Durango).—A. pheotis( Vera Cruz).—* A. aztecus (Morelos).

—*A. turpis (Tabasco).—* A. nanus (Guerrero, Vera Cruz, Colima,

Sinaloa).

Cuba (perhaps including Key West).—*A. jamaicensis par-

vipes.

St. Andrews, Old Providence, Jamaica, San Domingo, Porto

Rico, St. Martins, St. Kitts.—* A. jamaicensis.

Proposed subdivisions of the genus.—Gervais (1856, 1. s. ¢.) was

230 DR. KNUD ANDERSEN ON BATS. LAN oe C6

the first to make an attempt to subdivide dArtebews mto three

genera characterised by their number of molars, as follows :—

(1) Artibeus, molars 2; species three, viz. “A. gamaicensis,”

according to the figure of the teeth clearly not A.jamaicensis Leach,

but A. planirostris Spix ; the localities given by Gervais, “de la

Jamaique, de la Guadeloupe et de Cuba” are undoubtedly wrong ;

further, ‘A. lineatus ” (= Vampyrops lineatus), and “ A. undatus”

(= Stenoderma rufumy) ;

(2) Pteroderma, molars 3; species one, “ Pt. perspicillatum,”

“répandue au Pérou, au Brésil, et a la Guayane”; this is, as

shown by the figures of the teeth, A. jamaicensis Leach (sensu

lato) ;

(3) Dermanura, molars 3; species one, “ D. cinerea,” 1. e.

A, cinereus of the present paper.

In July 1865*, Peters divided <Artibeus into two ‘“ groups”

(subgenera), viz. :—

(1) Artibeus Leach, molars 3 or 2; species four: “A. per-

spicillatus Geoff.” (. e. A. gamarcensis lituratus of this paper),

“4, jamaicensis Leach” (i.e. A. jamaicensisjamaicensis), A. fallax

(i. e. A. planirostris fallax), andA. concolor ;

(2) Dermanura Gervais, molars 3; species three: D. cinerea,

? D. tolteca, and D. quadrivittata.

Peters placed the species with 2 and 2 molars together in one

group, ‘weil dieser kleime Zahnstumpf allem kei Grund sein

kann, Arten, die sonst im Schadel- und Zahnbau, so wie in jeder

anderen Beziehung ganz mit einander iibereinstimmen, generisch

von einander zu trennen.” But he was not quite consistent; if

“dieser kleine Zahnstumpt” (m"*) is not sufficient reason to

separate, as different groups, species with } and 2 molars, it is

difficult to see why the other, equally rudimentary tooth (m,)

furnishes a valid reason to separate, as a distinct section, the

species with % molars from those with 2 molars. From Peters’s

standpoint there would seem to be two alternatives only, either

not to subdivide the genus, acknowledging that the presence or

absence of a perfectly rudimentary tooth is a character of specific,

but not of subgeneric or generic importance, or to subdivide it

into three groups.— Peters himself has probably felt the incon-

sistency of his classification. At all events, in spite of his own

argument, that the species with = and 3 molars “sonst im

Schidel- und Zahnbau, so wie in jeder anderen Beziehung ganz mit

einander tibereinstimmen,” he, only a few months later 7, proposed

a new subgeneric name, Uroderma, for the species with 3 molars.

And, finally, in June 1866 {, he evidently regarded the sections no

more as subgenera, but as genera.—Thus Peters had now, in 1866,

practically adopted Gervais’s view, that Artibeus is to be divided

into three genera, according to the number of molar S, VIZ.

(1) Uroderma Peters 1865 (synonym: Aricbeus en 1856,

?es|bo

* Peters, MB. Akad. Berlin, 13 July 1865, p. 356, footnote.

+ Peters, MB. Akad. Berlin, 13 Nov. 1865, p. 588, footnote.

{ Peters, MB. Akad. Berlin, 25 June 1866, p. 394.

1908. | _ DR. KNUD ANDERSEN ON BATS. 231

not Artibeus Leach 1821), molars 3; species U. bilobatwm Pet.,

“U. fallax Pet.,” and U. concolor ;

(2) Artibeus Leach 1821 (synonym: Pteroderma Gervais 1856),

molars $; species “A. verspicillatus Geoftt.” and “A. jamaicensis

Leach ”

(3) Der manure Gervais 1856, molars 3; species as above.

By this arrangement Peters, as already said, had practically

gone back to Gervais’s standpoimt; these wor ds are true also in

the sense that his arrangement is in no respect an improvement

upon the older one ; both of them are typical exampies of artificial

classification. Gervais selected as the only leading character for

his subdivisions the presence or absence of a vanishing tooth ;

Peters did precisely the same. Gervais proved the fallacy of the

taxonomic character selected by him, in so far as he placed together

in one “ genus” (Artibeus) a true Artibeus, a Vampyrops, and a

Stenoderma, because they, though difterent in many important

respects, happen to have 3 molars; and, on the other hand, sepa-

rated into two genera (Artibeus and Pteroder ma) two species so

closely related as to be sometimes extremely difficult to distin-

guish (4. planirostris and jamaicensis). Peters proved the same,

by putting together in one “ genus” (Uroderma) two generically

widely different forms (U. bilobatum and ‘+A. fallax”), because

they both happen to have 3 molars, at the same time separating

into two “sections” or genera “A. fallax” and A. jamaicensis,

which differ in next to nothing but the presence or absence of a

rudimentary tooth.

In the description of the genus Uroderma (above, p. 217) I

have given my reasons for keeping U. bilobatum and thomas

generically separate from drtzbeus. The next question, therefore,

is, if, having removed these two species from Artibews, it might me

convenient to divide it into three subgenera or genera, according

to the number of molars. Also in this modified shape I am unable

to accept Peters’s proposal, for the following reasons :—

(1) The series of species here referred to the genus Artibeus

form one natural group the members of which are perfectly

sunilar in all essential cranial, dental, and external characters.

(2) A. planirostris has # molars; but of 73 skulls examined of

this species, two lack m* on one side, two on both sides, and one

of these latter aiso lacks m, on one side. A. hirsutus has 3 molars ;

but of 8 skulls, two lack m* on one side. A. jamaicensis has 2

molars; but of 182 skulls, two lack m, on one side, four on both

ailest 4. poses gi has } molars, but of the only two individuals

known, the one has an m, on one side. A. toltecus has 2 molars,

but of 26 skulls, one has an m, on one side.—None of the in-

dividuals here referred to are aberrant on account of very young

or very high age. Some of them, it will be noticed, have lost the

rudimentary molar * (m* or m,) which is normally present in

* It is hardly necessary to say that in all the aberrant individuals referred to

above m? or m3 (or both) are entirely lost, 7. e. no trace of their alveoli has

been left.

232 DR. KNUD ANDERSEN ON BATS. (Pasor si,

individuals of their species; others, in which the absence of m,

is normal, show, individually, a tendency to reversion to the

more primitive stage in which this small tooth was present.—A

character which is not only in itself very insignificant, but not

even individually perfectly constant is evidently unsuitable for

the separation of groups of generic or subgeneric rank.

(3) By subdividing Ar lane according to the number of molars,

the “ genera” or « subgenera > would be these three :—molars 3 =

A. Genwalirn, planirostris, hirsutus; molars 2, A. jamaicensis,

glaucus, watsoni; molars 3, A. cinereus, rosenbergi, toltecus, qua-

drivittatus, pheotis, aziecus, turpis, nanus.—But A. planirostris is

much more closely related to dA. jamaicensis, which is placed in a

different genus or subgenus, than to A. concolor, with which it is

associated in one group. A. glawcus and watsoni are put together

with A. jamaicensis, solely because they like this latter have a

rudimentary m,, but in all other respects they are much more

closely related to dA. cinereus, which has permanently lost m,.

The eight species with 3 molars constitute a strangely hetero-

geneous section; dA. cmerews is nearer to A. glaucus and watsoni

than to any of the forms with $ molars with which it is put

together; A. rosenber gi iS unique in the genus in the strong

reduction of m*; A. toltecus, quadrivittatus, and aztecus are,

probably, rather more closely allied to A. Jamaicensis than to any

species with 3 molars; and, finally, 4. turpis and nanus form

a small natural group characterised by the unusually strongly

depressed and slightly upwardly directed cranial rostrum.— From

this it will be evident that a subdivision of the genus based on the

presence or absence of m* or m, would give only a very distorted

view of the mutual affinities of the species.

(4) A study of the species of Artibews has led the writer of this

paper to the conclusion that they, probably, fall into two natural

groups, which have nothing to do with the hitherto proposed sub-

divisions of the genus, viz., those species in which cusp 7 of

m' is relatively small (A. g glaucus, watsoni, cinereus, and rosen-

bergi), and those in which it is relatively large (all the other

species). This point, which has more theoretical than practical

interest, will be discussed in the last section of the present paper,

pp. 314-316.

ARTIBEUS CONCOLOR Pet.

1865. Artibeus concolor Peters, MB. Akad. Berlin, p.357.—Paramaribo (Surinam).

1878. Artibeus planirostris (not Spix), var.«, Dobson, Cat. Chir. Brit. Mus. p. 518.—

Upper Amazons.

1892. Artibeus concolor Pet., Thomas, Ann. & Mag. N. H. (6) x. pp. 409-410,

footnote (Nov. 1892).—Some cranial measurements of the type.

1901. Artibeus concolor Pet., Thomas, Ann. & Mag. N. H. (7) viii. p. 191 (Sept.

1901).—Para.

Diagnosis—An Artibeus with 3 molars, the maxillary tooth-

row measuring about 7°2, the forearm about 50 mm.

Teeth.—The teeth of A. concolor are proportionately very much

smaller than in the two other species with molars, A. plani-

1908. | DR. KNUD ANDERSEN ON BATS. 233

rostris and hirsutus. A. concolor is externally not very inferior

in size to a small A. hirsutus or A. planirostris trinitatis; the

forearm in the only specimen examined of 4. coniilees measures

50 mm., in the smallest A. hirsutus 53-7 mm.; but the length

of the maxillary tooth-row is in concolor only 7-2 mm., the

smallest-toothed A. hirsutus 95 mm. A comparison with

A. planirostris shows a similar contrast.—In structure the teeth

of A. concolor do not differ from those of A. planirostris and

hirsutus.

Skull.—Rostrum (probably owing to small size of teeth) rela-

tively much shorter than in 4. hirswtus or planirostris. The ratio

between the length of the rostrum (from front of sagittal crest to

front of alveolus of a median incisor) and the length of the brain-

case (from front of sagittal crest to median posterior point of

lambdoid crest) is in concolor 68: 100, in hirsutus and planirostris

83: 100; or, expressed in another way, the length of the nasal

region, from front of sagittal crest to front of nasal bones, is in

A. concolor equal to the least interorbital width of the skull, m

hirsutus and planirostris equal to 13 this width. The rostrum

in A. concolor is a trifle less depressed than in hirsutus and

planirostris, but the difference in this respect is inconspicuous.—

The short rostrum makes, of course, the total length of the

skull much smaller: in concolor 22°4, in the shortest-skulled

hirsutus available 26°8 mm., although, as mentioned above, the

animal in external dimensions is only a trifle larger than concolor.

The difference in the length of the forearm, between concolor and

a small hirsutus, is only 3°7 mm., but in the length of the skull

4-4 mm.—In every other respect the skull of concolor is similar

to that of planirostris and hirsutus.

Nose-leaves.—Front margin of horseshoe free ; both front and

lateral margins quite plain (not crenulate). It remains to be

ascertained if these characters are perfectly constant (compare

the individual variation in A. planirostris, hirsutus, and jamat-

censis).

Tragus.—? or 8 sharp, but short, serrations on the upper half

of the outer margin, above the median projection ; the highest

number of serrations found in any other species of Artibews is 5.

A large series of A. concolor will, no doubt, show some variation

both in the number and shape of the serrations.

Hairing on limbs and interfemoral.—Kssentially as in A. plani-

rostris : wpper side of proximal half of forearm, upper side of inter-

femoral (the extreme posterior margin excepted) and of femur (the

distal part excepted) densely haired.

Colouwr.— Upper side from shoulders backward yellowish brown

(rather browner than Ridgway’s ‘“ wood-brown”); base of hairs

almost ecru-drab. In front of the shoulder region the darker

hair-tips are short or altogether wanting, exposing the white

or yellowish-white ground-colour of the fur. Under side light

reyish drab. Supraorbital stripes distinct, infraorbital stripes

almost obsolete. There seems to be no light margins to the ears.

234 DR. KNUD ANDERSEN ON BATS. [ Apr. 7,

Tips of wings (region of third phalanx of third digit) lighter-

coloured.

The above description is taken from an adult female with

unworn teeth, preserved in alcohol. There can scarcely be any

doubt that the specimen represents a ‘‘ light phase”; the type in

the Berlin Museum is, judging from Peters’s short description,

considerably darker. Similar contrasts in the coloration of the

fur occur in many other species of Artibews.

Measurements.—On p. 246.

Specimens examined.— 2 ad. (alc.), Para; with skull; British

Museum.—I have been unable to find the specimen (@ ad.,

Upper Amazons) catalogued by Dobson (J. s. ¢.) as A. planirostris

var. a.

Range.—Surinam ; Para; Upper Amazons (probably).

Peters’s A. concolor,1865.—Type locality: Paramaribo, Surinam.

A. concolor was described by Peters as being in every respect

similar to A. fallax [i.e. A. planirostris fallax of the present

paper |, but much smaller; molars e m’* in position and relative

size as in A. fallax; “ Unterarm” 47 mm. (Peters probably

measured the radius, not the ‘‘ forearm”), tibia 18 mm. There

are no measurements of the skull in the original description, but

according to Prof. Matschie (in a letter quoted by Oldfield Thomas

in 1892, 7. s.¢.) the maxillary tooth-row measures 7°5 mm., the

maxillary width across m'—m' 9-9 or 10 mm.—These details seem

to exclude all doubt as to the identification of A. concolor.

From the above description of 4. concolor it will be evident that

Dobson was mistaken in regarding this species as a mere var lety

of A. planirostris ; it is far more different from 4. plamirostris

than is this latter. from dA. jamaicensis (A. perspicillatus” in

Dobson’s Catalogue).

A. concolor seems to be very rare in collections, the type in the

Berlin Museum, two specimens in the British Museum, and one

in the Para Museum being, to my knowledge, the only examples

on record.

ARTIBEUS PLANIROSTRIS Spix.

Diagnosis.—Molars 2. Maxillary tooth-row 9°8-12 mm. Fore-

arm 55-73 mm. ‘Tibia and distal part of interfemoral so short-

haired as to appear almost naked. Colour of fur of upper side

not drab.

Teeth.—The teeth of this species have been described in detail

and figured above, pp. 207-212, text-fig. 41.

The rudimentary m’, situated postero- internally to m°, partly

pressed into a sharp angular emargination in the posterior margin

of this latter, between its cusps 5 and 7, is very rarely wanting

in adult individuals. 67 skulls of fully adult individuals have

been examined, representing all the races of dA. planirostris re-

cognised in this paper; in two skulls (4. p. fallaw, 3 ad. and

@ ad., British Guiana, teeth unworn and slightly worn, B.M

nos. 6.4.8.7 and 8) is m’ present on one side, while the tooth

1908. ] DR. KNUD ANDERSEN ON BATS. 235

and its alveolus are wanting on the other side; in two skulls

(A. p. fallax, 2 ad., British Guiana, teeth almost unworn, B.M.

no. 6.4.8.11; and 4. p. planirostris, 2 ad., Bahia, teeth unworn,

U.S. N. M. no. 102457) is m’* entirely wanting on both sides, and

in one of these latter skulls (102457) also m, is entirely lost on one

side, Thus, only 3 per. cent. of the large series of skulls of adults

examined have entirely lost m’ on both sides.

It is of some importance to emphasise that the disappearance of

m*in 4. planirostris is a very rare individual aberration, inasmuch

as the presence of this small tooth is in many cases the only

character by which A. planirostris can be safely discriminated

from A. jamaicensis.

Tragus.—In most individuals there are 4 or 5 small serrations

on the outer margin of the tragus, above the median projection ;

the serrations may be sharp, but as a rule they are more or less

rounded off, often reduced to inconspicuous nodules, sometimes

almost obliterated. The variation is quite individual.

Nose-leaves.— According to Dobson, the anterior margin of the

horseshoe in 4. planirostris is “free, separated from the muzzle,

straight, unnotched,” and the author of the British Museum

Catalogue of Chiroptera lays much stress on this character as a

difference between A. planirostris and A. jamaicensis (“ A. per-

spicllatus”). Ina majority of individuals of A. planirostris the

front margin of the horseshoe is distinctly “free,” but there is

every intermediate stage from this condition, through a margin

clearly “bound down,” though still more or less projecting, to a

margin so completely fastened down as to be almost continuous

with the integument of the muzzle. As a similar (or, if anything,

still greater) variation in this respect occurs in A. jamaicensis, the

character is quite useless for a discrimination of these two species.

—The margin of the horseshoe is sometimes simple, sometimes

crenulate in front, sometimes crenulate all round; the crenulation

occasionally extends to the margins of the lancet.—The lateral

margins of the horseshoe are not rarely bent up so as to form a

fold, suggesting the condition characteristic of Uroderma.

Hairing on limbs and interfemoral.—Vhe proximal half or two-

thirds of the forearm, the metacarpal of the pollex, the upper side

of the femur (the tip, as a rule, excepted), and the base of the

interfemoral next to the body and the femur, are densely haired.

The tip of the femur and the whole of the tibia covered with so

short and sparse hairs as to appear almost naked. The toes, from

the tarsus to the base of the claws, clothed with rather long,

coarse hairs.

Colour.—Y oung individuals :— Upper side from shoulders back-

ward dark and dull smoky brown, this colour confined to the

distal third or fourth of the hairs; base of hairs slate. On the

anterior part of the upper side, from the shoulders forward, the

hair-bases are distinctly lighter, almost smoke-grey. Under side

dark smoke-grey, with a peculiar mottled appearance, due to the

very short, almost greyish-white tips to the hairs. Tips of wings

236 DR. KNUD ANDERSEN ON BATS. | Apr. 7,

(region of third phalanx of third digit) more or less whitish. A

air of whitish or greyish-white supraorbital stripes, as a rule

indistinct, sometimes altogether wanting.—This is the usual colour

in young, not full-grown individuals; it never occurs in the

mature A. planirostris.

At a somewhat later stage, the colour of the hinder back is less

smoky, more approaching dark brown in tinge, with the hair-

bases almost drab. The rest of the upper side, from the shoulders

forward, much of the same general colour, but the dark hair-tips

shorter, the hair-bases considerably lighter, varying from wood-

brown to greyish white; very often the hair-tips in this region

of the upper side are so short as to more or less (or almost

completely) expose the light ground-colour, in which case there,

consequently, is a contrast between the anterior and posterior part

of the upper side. The under side essentially asim immature indi-

viduals, though as a rule a shade lighter. Supraorbital stripes

often completely wanting, often rather indistinct, rarely strongly

developed; there is sometimes, though rarely, an indication of

infraorbital stripes.—This is the commonest colour in the adult

A, planirostris, very often occurring also in specimens with much

worn teeth.

A considerably lighter colour is acquired by some adult indi-

viduals :— Upper side approaching Prout’s brown, base of hairs

almost ecru-drab. Anterior portion of upper side, from shoulders

forward, as a rale noticeably lighter, owing to the dark hair-tips

being shorter and the wood-brown hair-bases showing through.

Under side almost drab, with short greyish-white tips to the

hairs. Supraorbital stripes as a rule well marked, often strongly

developed, wood-brown or whitish; there is often a more or less

definite indication of infraorbital stripes.—This stage evidently

represents the “light phase” so common in many bats: I never

saw a young A. planirostris im this colour-stage ; it is apparently

confined to the fully adult age, and it is only acquired by a limited

number of individuals.

Thus there are three stages of colour: a dark and dull smoky

brown, a dark brown, and a Prout’s brown; the two former come

very near to each other, the third, when fully developed, is

different ata glance. ‘The first is confined to the immature age ;

the second is characteristic of a majority of adults; the third

seems to occur only in some fully adult and aged individuals ; it

is especially common in the largest race, A. p. fallax, but not rare

in A. p. planirostris and trinitatis.

Range.—From Central Brazil (Bahia, Matto Grosso) and

S. Bolivia (Caiza) to 8. Mexico (Chiapas, Guerrero), including the

Venezuelan coast islands (Trinidad, Tobago) and the southern

Windward Islands (Grenada), but excluding the rest of the West

Indies.

Remarks.—By the combination of the five characters given in

the brief diagnosis above, p. 234, A. planirostris (all races) is

readily distinguishable from all other species of the genus. The

1908. | DR. KNUD ANDERSEN ON BATS. 237

first character (molars 3) excludes all species, except A. concolor

and hirsutus; the addition of the second and third characters

(large skull and teeth, large external dimensions) excludes A. con-

color; the addition of the fourth and fifth (tibia and distal inter-

femoral almost naked, general colour of fur of upper side not drab)

exclude also A. hirsutus.

Forms.—Four races of A. planirostris are described below:

A. p. planirostris, trinitatis, grenadensis, and fallax. The three

former come very near to each other, the fourth is rather more

completely differentiated, but cannot be specifically separated.

ARTIBEUS PLANIROSTRIS PLANIROSTRIS Spix.

1823. Phyllostoma planirostre Spix, Simiarum et Vespertilionum Brasiliensium

species nove, p. 66, pl. xxxvi. fig. 1—Bahia.

1826. ? Phyllostoma obscurum Wied, Beitr. Naturg. Bras. ii. pp. 203-205.—Rio

de Janeiro.

1840. Phyllostoma perspicillatum (partim, nec L.) Wagner, Schreber’s Siugthiere,

Suppl. i. pp. 403-405 —Re-deseription of Spix’s type of Ph. planirostre.

1865. Phyllostoma planirostre Spix, {Peters, MB. Akad. Berlin, p. 587.—Spix’s

type of Ph. planirostre re-examined.

1878. Artibeus planirostris Spix (partim), Dobson, Cat. Chir. Brit. Mus. pp. 515-517.

1901. Artibeus planirostris Spix, Robinson & Lyon, Proc. U.S. Nat. Mus. xxiv.

p. 148.—La Guaira, Venezuela (specimens examined).

1904, Artibeus intermedius All. (errore), J. A. Allen, Bull. Am. Mus. N. H. xx.

Art. iv. p. 79 (29 Webr. 1904); cf Allen, t.c. Art. xx. p. 233 (29 June

1904).—Chiriqui.

1904. Artibeus planirostris Spix, Thomas, P. Z. 8. 1908, ii. p. 234 (1 April 1904).

—Chapada, Matto Grosso (specimen examined).

Diagnosis.—Votal length of skull 27°5-30 mm. (average 28-5

mm.); zygomatic width 16°8-18°5 mm. (average 17-6 mm.)

forearm 57:°8-65°2 mm. (average 61°8 mm.).

A. p. planirostris and trinitatis.—A. p. planirostris can only be

discriminated from its nearest relative, 4. p. trinitatis, by average

characters. In A. p. planirostris the forearm and metacarpals

average about 4 mm., the tibia 1:5 mm. longer; the ears are,

generally, a little larger; the average difference in the size of the

skull and teeth is very small.—For further details see the table,

p. 246.

Specimens from different localities—In the subjoined com-

parative table of measurements (p. 240) I have divided the material

examined into three groups, viz. specimens from Brazil, Venezuela,

and 8. Mexico. ‘The table shows that the size of the skull and

teeth and the external dimensions are identical in individuals

from these three regions.

Specimens examined.—26 specimens (12 skins) and 20 skulls,

from the following localities :—

British Museum :—Brazil: Chapada, Matto Grosso, 700-900 m.

(1); Pernambuco (2); 8. Lourengo, Pernambuco, 28-60 m. (8) ;

Igarapé, Assii, Para, 50 m. (2); “ Brazil” (1).—10 skulls, from

all the localities enumerated.

U.S. National Museum * :— Brazil: Bahia (1); Anilo, Maranhao

* US. N. M. nos, 100201, 100203, 102457, 102894-96, 104565, 104567-69, 104574,

126554,

238 DR. KNUD ANDERSEN ON BATS. [ Apr. 7,

(5).—Venezuela: Macuto, La Guaira (3).—S. Mexico: Palenque,

Chiapas (2); Papayo, Guerrero (1).—10 skulls, representing all

these localities.

Range.-—From Central Brazil (Bahia, Matto Grosso) to 8. Mexico

Chiapas, Guerrero). As yet no record from Guiana (see 4. p.

fallax).

Spix’s Phyllostoma planirostre, 1823.—Type locality: “ im

suburbiis Bahie.” From Spix’s description and figure so much

only can be decided with certainty that his Ph. planirostre is a

large species of Artibews. There being only two large species

known, the point to be settled is this: is Spix’s type the form

called A. planirostris in the present paper (3 molars), or is ib

A. jamaicensis (2 molars). Both of these species occur in the

Bahia region.—Three authors, Wagner in 1840, Peters in 1865,

and Dobson in 1878, have discussed this question and arrived at

different conclusions :—

Wagner’s description of ‘“ Phyllostoma perspicillatum” (1840,

J. s.c.) was based on what he considered Spix’s types of Ph. plani-

rostre :-—‘* Von seinem Phyllostoma planirostre hat Spix 3 Kxem-

plare in Weingeist hinterlassen,” he writes; and “die nachstehende

Beschreibung ist nach den Spix’schen Exemplaren entworfen.”

The only additional information of importance contained in

Wagner's description is this: “ Backenzihne finden sich + vor,”

i. e. Wagner found only 3 molars in the presumed types.-—From

this it might be inferred that Spix’s Ph. planirostre is Leach’s

A. jamaicensis.

Peters writes (1865, l.s. ¢.): “Nach Untersuchung des einzigen

Originalexemplars in Weingeist [von Ph. planirosire| kann ich

nur die Uebereinstimmung desselben mit Ph. perspicillatwm

Geoffroy bestiitigen,” and found the specimen which he considered

the type to have § molars.— From this, again, it would seem that

Ph. planirostre Spix (1823) is a synonym of A. jamaicensis Leach

(1821), this latter being the only large species of the genus with

2 molars. There is, it will be noticed, a discrepancy between

Wagner and Peters with regard to the number of typical speci-

mens; according to Wagner there are three, according to Peters

one only.

Dobson (1878, 7. s. ¢.), though he had no opportunity of ex-

amining the type (or types) of Ph. planirostre, rejected Peters’s

identification on account of the following words in Spix’s original

description : ‘vexillum nasale ... . inferius lateraliter et antice

libere pendens”; he regarded this statement of Spix, that the

horseshoe is “free” in front, as decisive evidence that Ph. plan-

rostre is the large species with # molars, not the large species

with 3 molars ; and the difficulty that, according to Peters, the

type of planirostre has 3 molars only, he overcame by arguing

that the type might be immature, or very old, or in this respect

abnormal.-—But the fact is, it must be said at once, that the con-

dition of the front margin of the horseshoe is thoroughly unreliable

1908. | DR. KNUD ANDERSEN ON BATS. 239

as a differential character between the two large species of Artibeus

(see pp. 235 and 253 of this paper), so that Dobson’s way of settling

the question was exceedlngly unsafe. That, nevertheless, the con-

clusion was right is proved by the following information kindly

forwarded to me by Dr. W. Leisewitz, Munich (in litt., 15 Sept.

1906) :—

TH register of the Munich Museum (“ Zoologische Sammlung

des Bayerischen Staates”) for 1830 has this entry: “No. 65,

Phyllostoma planirostrum (Sp.), 1 Exemplar”; the specimen is

labelled “ Bahia, Spix coll.” ; this settles the question as to the

number of typical specimens; there is one only. When Wagner

mentioned three typical examples, the reason was, I am informed

by Dr. Leisewitz, probably that Spix brought back from Bahia

not only one Ph. planirostre but also two “ Ph. perspicillatum ”

(i.e. A. jamaicensis litwratus of the present paper), both of

which latter are also in the Munich Museum; Wagner evidently

considered all three examples to be one species (A. planirostris

and jamaicensis are difficult to discriminate externally), and his

statement that “ planirostre” has 2 molars is undoubtedly taken

from one of the two 4A. jamaicensis, not from the true type of

Ph. planirostre.—This latter has, Dr. Leisewitz writes, a distinct

m’ on both sides of the upper jaw; the anterior margin of the

horseshoe is (as said by Spix) free; the forearm measures 58:5,

third metacarpal 57, first phalanx of third digit 17-7, second

phalanx of third digit 28°5 mm. This settles, beyond all doubt,

the identification of Spix’s type: it is Ph. planirostris planirostris

of this paper, not A. jamaicensis lituratus (molars 5, forearm

64-76 mm.), the only other large form of Artibews known from

Bahia.—There remains Peters’s wrong statement about the

number of molars of the “type” (2 according to Peters, not 3 as.

in fact is the case): On Oct. 17, 1865, Siebold sent Spix’s.

Chiroptera to Peters for inspection, among these the type of

Ph, planirostre and one example of “A. perspicillatus L.” (A. gamai-

censis lituratus Licht.); in the list accompanying the specimens,

Siebold unfortunately entered these two bats as “2 Phyll. plani-

rostre, Bahia.” When, therefore, Peters wrote that Spix’s type

of Ph. planirostre has # molars only, he no doubt examined the

wrong specimen (A. 9. litwratus), not the true type; this ex-

planation is further strengthened by the fact that the true type

(Dr. L. writes) shows no trace of having had the mouth opened

for examination of the molars; finally, when Peters wrote that

there is only “ein einziges Originalexemplar,” it was, Jrom his

standpoint, a mistake, for on sending Spix’s bats back to Munich

he wrote (letter dated 10 Dec. 1865): “2 St. Artibeus perspicil-

latus Geoftr.= Phyll. planirostre Spix! Original.”

As a final result: there is one type only of Spix’s Ph. planirostre

still in the collection of the Munich Museum; this specimen has

3 molars, and the forearm 58°5 mm.: both facts are decisive

evidence that it is the bat called 4. p. planirostris in this paper ;

Wagner's and Peters’s statement that it has 2 molars is a’

Proc. Zoou. Soc.—1908, No. XVI. 16

240 DR. KNUD ANDERSEN ON BATS. [Apr. 7,

mistake, due to their having examined the teeth not of the true type,

but of specimens of A. jamaicensis lituratus also collected by Spix

at Bahia. It has been of importance, from a purely nomenclatural

point of view, to have this question definitely settled; if Wagner

and Peters were right, Spix’s Ph. planirostre would have been a

synonym of A. jamaicensis litwratus, whereas the species hitherto

called A. planirostris would have had to stand as A. fallax Pet.

Maximilian of Wied’s Phyllostoma obscurum, 1826.—Type from

“Villa Vicosa am Flusse Peruhype,” 7. e. Parahyba, province of

Rio de Janeiro. Judging from the description there can only be the

question whether this is A. planirostris planirostris or A. gamai-

censis lituratus. The number of “ Backenziihne im Oberkiefer ” is

Brazil | Venezuela S. Mexico

(Matto Grosso, (La Guaira). (Chiapas,

Bahia, Per- | Guerrero).

nambuco, Ma- |

ranhao, Para).

| 16 adults, 3 adults, 3 adults,

| 11 skulls. 3 skulls. 3 skulls.

Min. Max. Min. Max.|Min. Max.

mm. mm.) mm. mm. | mm. mm.

Skull, total length, to front of ec...) 27°5 30 27°5 29 27 28°3

Pieinastoidewacdtly: qe. eee en lcs 16 | 15:2 15°3 | 14°8 14:9

» width of brain-case ......... 12°2 13:2 | 12:8 13 12 12°6

» Zygomatic width \.......... | 168 185 | 17 17-7 | 16:8 175

» Maxillary width, across m1) 12°2 132 | 128 - 13 12:1 12:7

»» across cingula of canines.... 7 82) 81 84 | 7:8 8

Mandible, to front of ine. .......... 193 20°8 | 19°5 20 19 19:2

Upper teeth, cm? ..................) 99 10°8 | 101 10°7 | 1071 10:2

ower teeth; coms .........-..-.--.- | 10°8 11:8 | 11:2 lls | 11 11

Har-couch, length, imner margin .| 15°2 16°8 | 15°7 16°5

a length, outer margin .) 21 23°6 | 215 295

s Wide) .cee es meee| Lato 16 | 14 14°5

TUNIS, HOME joocoscsntecoscancwussss| Oe 78) 68

ancetalenethy weseree- eee eee cree 8 105; 9 105

ss Tal Cit Wapesercchottek pastes eee a 68 78) 6 68 |

Horseshoe, width ................---+: 72 Q | ay Pe

orearmiay ee ee OS 652 | 588 612 | 60 62°2

Pollex meee eis: Mee ee Reels 156 | 13 145 | ... 158

yal WATAVGAMEDENM ooo coo ca vodacor ee ogssue 50°8 59 =| «54 548 | 54°7 552

PTT aD eles, caval crmstevetern allele, 20:2 | 17 188 | 168 185

PUAN Ls the A So dete de RB ees tele One 32°8 | 29°5 30 28°5 312

MOTT 5 acer Se A cae saath taiciae dod 15 18°8 | 147 162 | 15 16

At mMetacarpalleesane es eeee-eeeee eee OO 578 | 52-2 54 53 53

TINY RSS RR eptetee beg = Sa aatien $455 0st seb Gad fel a) 17°8 | 147 162 | 15 16°2

TOV 2UIR Ve. MAN ROTO Me eee call ail 7eo Py} | AS 20 18 19

5th metacarpal........................, 618 597 | 54:2 552 | 543 56

A ee eee REE EE Ee Geetnoll Lat 14 | 113 I) we 13°5

2a Buen set Gh oie WeRie en ANN) 17°7 | 14:2 152 | 13% 16

imbertemoralle ee eee eee eel Ol | 12 12°8 |

TOO SP Meee soo cnonca ace sca soo osavcaceoasel| ZL) Ps |) 2nle7/ 22°5 | 21°8 23°2

Hoot, with claws ......¢-0-..s.c2-4:..) Lo 17:2 | 14:7 158

[2@alear iets favs come cee | 65 82) 75 AS |) BS 65

1908. ] DR. KNUD ANDERSEN ON BATS. 241

stated to be “ vier auf jeder Seite”; if this is correct, the bat is

A. j. lituratus. But three points make me hesitate to draw this

conclusion :—first, Prince Maximilian describes the incisors and

canines of Ph. obscurum tolerably well, but passes very lightly

over the molars; if his examination of these latter has been

similarly cursory, he may very easily, indeed, have overlooked the

rudimentary m*: second, the length of the head and body is

stated to be “3'"”; it would seem to be too small for an 4A. 7.

lituratus, but would agree very well with dA. p. jlanirostris:

third, he describes in the same book a ‘“ Ph. superciliatum,” also

from Rio de Janeiro, which probably is A. 7. litwratus, and it

might seem rather unlikely that he has described, a few pages

later, a specimen of the same form as Ph. obscurum; this latter

argument is, however, rather weak ; the possibility is not quite

excluded that Ph. superciliatum might be the light ‘“ phase,”

Ph. obscurum the dark “phase” of one species; but the other

evidence speaks against this assumption. Only a re-examination

of the type of Ph. obscurwm, if it still exists, can place the identi-

fication beyond doubt.

ARTIBEUS PLANIROSTRIS TRINITATIS K. And.

1893. Artibeus planirostris Spix, Thomas, Journ. Trinidad Field Nat. Club, i. no. 7,

p. 6 (April 1893).—Trimidad.

1897. Artibeus planirostris Spix, J. A. Allen & Chapman, Bull. Am. Mus. N. H.

ix. Art. i. p. 15 (26 Feb. 1897).—Trinidad.

1906. Artibeus planirostris trinitatis Knud Andersen, Ann. & Mag. N. H. (7) xviii.

p. 420 (1 Dec. 1906).—T ype locality : Trinidad.

Diagnosis.—Similar to A. p. planirostris, but averaging smaller,

with shghtly smaller skull and teeth.

A. p. trinitatis and planirostris—A. p: trinitatis can only be

discriminated from its nearest relative, A. p. plainirostris, by

average characters. In 4. p. trinitatis the forearm and meta-

carpals average about 4 mm., the tibia 1°5 mm. shorter; the ears

are, generally, a little smaller ; the average difference in the size

of the skull and teeth is very small. For further details see table

of measurements, below p. 246.

Specimens examined.—135 specimens (6 skins) and 9 skulls, from

the following localities :—

British Museum :—Trinidad (5).—Tobago (2).—5 skulls, repre-

senting both localities.

U.S. National Museum * :—Trinidad (6).—4 skulls.

Range.—The islands of Trinidad and Tobago, W.i.

ARTIBEUS PLANIROSTRIS GRENADENSIS K. And.

1906. Artibeus planirostris grenadensis Knud Andersen, Ann. & Mag. N. H. (7)

xviii. p. 420 (1 Dee. 1906).—T ype locality: Grenada, W.I.

Diaginesis.—In the size of the skull and teeth very similar to

* U.S. N. M. nos. 101898-99 104016, 104018, 104022-93.

16#

42 DR. KNUD ANDERSEN ON BATS. [evan 0

A. p. planirostris, in external dimensions rather intermediate

between A. p. trinitatis and planirostris.

A.p. grenadensis and closely allied forms.—The skull, teeth,

and external dimensions of A. p. grenadensis average somewhat

larger than in its nearest relative, A. p. trinitatis. The size of

the skull and teeth is almost quite as, or if anything still a trifle

larger than, in A. p. planirostris, but externally A. p. grenadensis

averages somewhat smaller than this latter race. For details see

the table of measurements, p. 246.

Though undoubtedly an offshoot of the Trinidad-Tobago race,

this form has almost reversed to the size of the continental

A. p. planirostris; practically it is difficult to discriminate

A. p. grenadensis from this latter. But even if it were proved

that A. p. grenadensis is, also in average characters, completely

similar to the Venezuelan and Brazilian A. p. planirostris, it

would be reasonable to keep it separate; there is only one other

alternative, viz. to unite A. p. planirostris, trinitatis, and grena-

densis into one ‘‘race,” as opposed to the unquestionably much

more different A. p. fallax; but this would obliterate the two

facts that passing from Venezuela to Trinidad-Tobago there is a

decrease in the average size of the individuals, and passing from

Tobago to Grenada there is, again, an increase in the average

size,—facts which seem to me worth recording and, if so, are to

be expressed in the technical names of these bats.

Specimens examined.—11 specimens (6 skins) and 8 skulls, from

the following localities :—

British Museum :—Grenada, W.I. (5).—2 skulls.

U.S. National Museum * :—Grenada, W.I. (6).—6 skulls.

Range.—As yet recorded only from the island of Grenada,

Windward Isles, W.i. :

ARTIBEUS PLANIROSTRIS FALLAX Pet.

1865. Artibeus fallax Peters, MB. Akad. Berlin, pp. 355-57.—Type locality : Guiana.

1878. Artibeus planirostris Spix (partim), Dobson, Cat. Chir. Brit. Mus. p. 517.—

British Guiana.

1898. ? Artibeus planirostris Spix, Thomas, Boll. Mus. Torino, xiii. no. 316, p. 3

(18 April, 1898).—Caiza, ‘larija, S. Bolivia.

1901. Artibeus planirostris Spix, Thomas, Ann. & Mag. N. H. (7) vii. p. 143

(Aug. 1901).—Kanuku Mts., B. Guiana (specimens examined).

1901. Artibeus planirostris Spix, Thomas, Ann. & Mag. N. H. (7) viii. p. 191

(Sept. 1901).—Para (specimen examined).

1902. ? Artibeus hercules Rehn, Proc. Acad. N. Sci. Philad. pp. 688-39 (12 Oct.

1902).—K. Peru.

1904. ? Artibeus planirostris Spix, J. A. Allen, Bull. Am, Mus. N. H. xx. Art. xxix.

(8 Oct. 1904).—Ciudad Bolivar, Venezuela.

1907. Uroderma validum D. G. Elliot, Field Col. Mus., Publ. fno. 115, Zool. Ser.

vol. vill. pp. 587-88, fig. 74 (skull).—Cayenne.

Diagnosis.—Similar to A. p. planirestris, but averaging con-

siderably larger.

A. p. fallax as compared with the other races.—A. p. fallas is

the most completely differentiated of the four races of Artibeus

* U.S, N. M. nos. 111510, 111518-14, 111517, 111519, 111523.

1908. | DR. KNUD ANDERSEN ON BATS, 243

planirostris described in this paper. The largest skull available

is 3 mm. longer, 2°3 mm. broader (zygomatic width) than the

largest skull of A. p. planirostris; the upper tooth-row, in the

largest specimen, is 1:2 mm. longer than in the largest A. p. plani-

rostris ; there is approximately the same difference between the

smallest skull of 4. p. fallax and the smallest of 4. p. planirostris,

and the average difference in the size of the skull and teeth 1s,

consequently, very well marked, The difference in the external

dimensions is equally pronounced, 4. p. fallax being, as a rule,

noticeably larger than 4. p. planirostris. See the table of measure-

ments, p. 246.

L have had no real difficulty in discriminating any individual

of dA. p. fallax, in the whole large series examined, from A. p.

planirostris ; only one unusually small specimen of the former

race from Demerara (B.M. no. 75.11.3.17) caused me some hesi-

tation. Nevertheless, A. p. fullax cannot, in my opinion, be

separated as a distinct species, but only as a local race of Artideus

planirostris, for the following reasons :— First, there is no struc-

tural difference, in any respect, between A. p. fallax and the other

races; second, small individuals of A. p. fallaa come so exceedingly

near to large individuals of dA. p. planirostris that there can be

no doubt that, occasionally, the two forms will prove to be practi-

cally quite indistinguishable ; third, some examples of A. p. fallax

from the Lower Orinoco Valley (Ciudad Bolivar) and Para show

decidedly leanings tewards 4. p. planirostris, and there at least,

the border districts between the areas occupied by the two races,

they will, no doubt, be found to intergrade.

Specimens eaamined.—55 specimens (42 skins) and 36 skulls,

from the following localities :—

British Museum :—Para (2).—French Guiana: Cayenne (8).

British Guiana: Demerara (6); Comachka, Demerara River

(5); EHssequibo River (20); Kanuku Mts., about 59° W., 3° N.

(11); B. Guiana (1).—Lower Orinoco: La Vuelta, Ciudad Bolivar

(2).—36 skulls, from all the localities enumerated.

Range.—Guiana, extending southward to Para, where it meets

A. p. planirostris, northward to the Lower Orinoco Valley, where

it also meets A. p. planirostris.

If Rehn’s Artibeus hercules, from KE. Peru, and a specimen of

A. planirostris recorded by Oldfield Thomas (J. s. c.) from Caza,

S. Bolivia, neither of which has been examined by me, are referable

to A. p. fallaw, the range of this form is considerably more

extensive.

Peters’s A. fallaa, 1865.—The species was based on “ einem

weiblichen Exemplar in Weingeist” from Guiana in the Berlin

Museum, and “anderen trockenen Exemplaren ” in the Leyden

Museum (probably specimens } and ¢ in Jentink’s ‘ Cat. Syst.

Mamm.’ p. 208, 1888). According to Peters (J. s. c.) 4. fallaa is

in size, in the form of the ears and nose-leaves, and in colour

“dem A. perspicillatus (A. jamaicensis of the present paper |

244 DR. KNUD ANDERSEN ON BATS. [Apr: 7,

tiiuschend iihnlich und gewiss auch schon oft mit ihm verwechselt

worden,” but differs in the following respects: ‘‘ Der untere Rand

des Hufeisens ist langer, deutlicher abgesetzt und fein gekerbt.

Das Gebiss ist namentlich dadurch verschieden, dass der zweite

obere wahre Backzahn am hinteren Rande hinter dem Zacken

des Cingulums [cusp 5 of this paper] viel tiefer eingebuchtet ist,

und dass hier ein sehr kleiner fiinfter Backzahn sich hineilegt.”

-—This statement, taken together with the locality, leaves no

doubt as to the identification of A. fallax.

Rehn’s 4. hercules, 1902.—Type locality: Eastern Peru; two

specimens (one skull). Its ‘general characters” were summed

up by Rehn (/. s.¢.) as follows :—“ Allied to A. planirostris (Spix),

but differing in the larger size, the much larger foot, and in

numerous dental characters.” These latter are thus described :—

“Second upper premolar subquadrate in basal outline, quite

different from the subpyriform tooth of A. planirostris, the in-

ternal node well developed and forming a conspicuous cusp. First

upper molar broad, deep, the anterior inner angle more developed

than in A. planirostris, which species has this portion rounded.

Second lower premolar very heavy and broad, the posterior in-

ternal border with the dentate ridge low. First lower molar

subquadrate in outline.” Upper tooth-row 12, lower tooth-row

12:5, maxillary width across m’* 14:5 mm.; forearm 65:2, 3rd

metacarpal 61°5, tibia 24 mm.

Rehn had for comparison two A. planirostris from ‘“ Chapada

| Matto Grosso], Brazil” ; so it was quite natural that he found the

large, large-skulled and large-toothed Peruvian bat considerably

different in size. But if he had been able to compare it with

A. p. fallax, he would have seen that the size of the two Peruvian

examples is in every respect precisely as in the Guianan form

described long ago by Peters. If, further, he had had a tolerably

good series of skulls of A. p. fallax and A. p. planirostris, he would

have realised that the dental characters as derived from the

single skull of A. hercules examined by him have neither specific

nor subspecific importance, but are individual peculiarities, found

in any form of A. planirostris (as, indeed, they are found also

in the various forms of A. jamaicensis). As to this latter point

T subjoin the following details :—p*, in a series of six skulls of

A. p. fallax (B.M. nos. 3.4.5.22—-27), all of adult individuals,

none with the teeth much worn, all from Cayenne and taken

almost on the same date: basal outline subpyriform in two skulls ;

subquadrate with strongly rounded inner margin, in one; sub-

quadrate with slightly rounded inner margin, in one; subquadrate

with straight inner margin, in one; strongly subquadrate (nearly

quadrate) with straight inner margin, in one. The antero-internal

cusp on the heel (‘internal node,” Rehn) of p* varies, in this

series, from small, through strong, to very strong; the cusp is not

always most conspicuous i in quite unworn teeth. m_’, in the same

series from Cayenne :—one extreme (two skulls): short antero-

posteriorly, broad from side to side, antero-internal angle quite

1908. | DR. KNUD ANDERSEN ON BATS. 245

rounded (this would be, so far as this character is concerned,

Rehn’s A. planirostris); the other extreme in the same series of

skulls (one specimen): m’ long antero-posteriorly, especially in its

lingual half, antero-internal angle very sharp, practically rect-

angular (this would be Rehn’s 4. hercules); in the three remaining

skulls the tooth is intermediate in shape. As to p, and m,, I

must confess I do not quite understand Rehn’s description, unless

“second lower premolar” is a lapsus for first lower molar, and

“first lower molar” for second lower molar; there is, of course,

no “dentate ridge” on the postero-internal border of p, in any

bat of the genus Artibews, but there is on m,, and the absolute

size of the cusps of this dentate ridge (cusps 2 and 3) is subject

to notable individual variation.—When entering upon such

minute details as here under consideration, we shall scarcely find

two skulls of A. p. fallaa (or any other form of 4. planirostris)

precisely alike ; we have passed from the characters useful for a

specific or subspecific discrimination to the field of individual

variation, and, judging from what I have seen in a large series of

skulls of A. p. fallax, I should think it highly probable that if

Rehn extracted the skull of his second specimen of 4A. hercules,

he would find the form of the lingual portion of p* or m! or both

of them slightly differing from that of the corresponding teeth in

the single skull described.

It is, of course, quite possible that Peruvian individuals differ

in some minor details (or some average characters) from the

Guianan A. p. fallax; for the present it is at least certain that

there is not in Rehn’s description one single character by which

A. hercules can be discriminated from 4. p. fallax.

Elhiot’s. Uroderma validwm, 1907.—'Type locality: Cayenne.

From the figures of the skull, the description and measurements,

clearly an A. p. fallaw. Elliot has apparently been unaware of

the fact that the type of U. validwm (Cayenne) is practically a

topotype of A. p. fallax Peters (Guiana).

ARTIBEUS HIRSUTUS K. And.

1906. Artibeus hirsutus Knud Andersen, Ann. & Mag. N. H. (7) xviii. p.420 (1 Dee.

1906).—Type locality: Michoacan, Mexico.

Diagnosis.—Like a small form of A. planirostris (though

averaging still smaller), but tibia and interfemoral densely haired,

and colour of fur of upper side of body in adults drab with a

silvery tinge. Maxillary tooth-row 9°5-10°-4 mm.; forearm 53-7-

59:7 mm.

Skull and teeth.—In the skull and teeth there is no essential

difference between A. hirsutwsand a small form of A. planirostris,

f. i. A. p. trinitatis; the rostrum of the skull may, perhaps,

average somewhat narrower.—Molars 3, as in A. concolor and

planirostris, and as in these species the presence of the small m*

is almost constant. Hight skulls of A. hirsutus have been

examined ; in two (@ ad., teeth unworn, Michoacan, U.S. N. M.

(Apr. 7,

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1908. | DR. KNUD ANDERSEN ON BATS. 247

no. 126451; and ¢ ad., teeth slightly worn, Colima, U.S. N. M.

no. 52091) m3 and its alveolus are wanting on one side, in none

on both sides.

Nose-leaves.—Of eight specimens examined, four are preserved

in alcohol ; in one of these latter the front margin of the horse-

shoe is free, but narrow; in three it is almost continuous with

the integument of the muzzle. Thus there is in this species the

same variation in this respect as in A. planirostris (p. 235) and

A. jamaicensis (p. 253).

Tragus.—About three small, rounded, sometimes almost obsolete

serrations above the median projection.

Wings.—The metacarpals and phalanges (especially the proximal

phalanges) are proportionately a little shorter than in A. plam-

rostris. The wing-indices on p. 310 and. the table of measure-

ments, p. 246, show the details.

Hairing on limbs and interfemoral.—Difterent from A. plani-

rostris. 'The whole of the interfemoral (above and below) right

to the posterior margin, and the upper side of the femur, tibia,

and foot are densely haired.

Colour.—General impression: much more drab above and

lighter beneath than in any phase of A. planirostris.

Upper side dark drab or brownish drab, with very short, almost

greyish-drab tips to the hairs, giving the whole of the upper side

a peculiarly silvery tinge; base of hairs of hinder back grey with

a tinge of ecru-drab, in the shoulder region and on the neck

noticeably lighter, whitish ecru-drab. Under side smoke-grey

with conspicuous white tips to the hairs. Supraorbital stripes

indistinct or none.

Specimens examined.—8 specimens (4 skins), with skulls, from

the following localities:—Michoacan: La Salada (3); Colima :

Colima (3); Jalisco: Etzatlan (1); W. Mexico (1).—All from the

collection of the U.S. National Museum *.

Range.—As yet only known from the States of Michoacan,

Colima, and Jalisco, Mexico.

ARTIBEUS JAMAICENSIS Leach.

Diagnosis—Molars %. Maxillary tooth-row 9°3-12°2 mm.

Forearm 54-76 mm.

SkullSimilar in shape to that of A. planirostris. In the

largest race of A. jamaicensis (viz. A. j. litwratus) the size of the

skull is practically as in the largest form of A. planirostris (A. p.

fallax), though averaging still a trifle more heavily built; so

complete is the resemblance that certain skulls of A. 7. litwratus

would be indistinguishable from those of A. planirostris, were it

not for the absence of the small m*. In the smailest races of

A. jamaicensis (A. j. parvipes and yucatanicus) the skull is as

small and delicately built as (or, if anything, still smaller than)

in A. planirostris trinitatis or A. hirsutus.

The skull, especially its facial portion, is in certain races of

* U.S. N. M. nos. 9052 (36860), 52063, 52091, 52092, 52101, 126448-49

126451.

248 DR. KNUD ANDERSEN ON BATS. (Apr. 7,

A. jamaicensis subject to considerable modification dependent on

the age of the individuals; one of these modifications has served as

type for the description of a distinct species (J. A. Allen’s A. anter-

medius). The text-figures below show, approximately, the two

Text-fig. 50.

Artibeus jamaicensis palmarum, 9 ad. Macuto, Venezuela, July 14, 1900.

U.S. N. M. 102845.

A, Lateral, B. Upper view of skull. X 3.

Text-fig. 51.

Artibeus jamaicensis palmarum, g ad. Macuto, Venezuela, July 14, 1900.

US. N. M. 102843.

A. Lateral, B. Upper view of skull. X 3.

extremes in the shape of the skull in adult individuals (in some

skulls examined the old age modification is carried still a little

1908. | DR. KNUD ANDERSEN ON BATS. 249

farther than represented in fig. 51):—the two skulls of 4. We

palmarum figured are from the same place (Macuto, La Guaira,

Venezuela); the one (fig. 50) is of an adult female with the distal

epiphyses of the metacarpals ossified, but the teeth unworn, 7. ¢. a

mature but young individual; the other (fig. 51) an older male

with somewhat worn teeth: these two skulls have been selected as

paradigmata, because they belong to the same geographical race

of A. jamaicensis and were obtained precisely at the same locality ;

but perfectly similar extremes are found in other skulls of the series

examined, and not only in A. j. palmarwm but in A. 7. litwratus

as well (f. i. in a British Museum series of this latter race, from

Sapucay, Paraguay). The skulls figured are, the one of a female,

the other of a male, but the modifications have nothing to do with

sexual differences. In the rather younger individual (fig. 50) the

brain-case makes the impression of being lower, the naso-frontal

depression (on the dorsal face of the rostrum, in front of the

sagittal crest) is shallow; the supraorbital ridges, where starting

from the anterior point of the sagittal crest, form an angle with

this latter of about 125°-135°, the postorbital processes are very

inconspicuous (in still younger individuals scarcely indicated). In

the somewhat older individual (fig. 51) the brain-case makes the

impression of being higher, more vaulted, but the difference

is more apparent than real, chiefly due to the higher and more

forwardly extending sagittal crest. This latter fact, that the

sagittal crest has been produced farther forward, has two

other effects—first, that the naso-frontal depression (viewed in

profile) is more abrupt; second, that the supraorbital ridges

are not directed forward and outward, but almost straightly

outward, forming an angle with the sagittal crest of very little

more than 90°; the postorbital processes are very conspicuous,

and a pair of comparatively large anteorbital processes has been

developed.

The adult and old age modification of the skull as described

above is characteristic of the forms called in this paper the

“southern races” of A. jamaicensis (viz. A. j. lituratus, palmarum,

preceps). The transition from the young to the old stage takes

place earlier in some individuals than in others; I have seen

individuals with somewhat worn teeth which have still almost

the young type of skull, or are only in a transitional stage,

while others with almost quite unworn teeth have already

reached far on the way towards the old age type.—The “ northern”

races (A. j. parvipes, yucatanicus, jamaicensis, cequatorialis)

never, or exceedingly rarely, reach that degree of old age modifi-

cation of the skull attained by the southern races.

A. j. parvipes, yucatanicus, jamaicensis, and wquatorialis retain,

throughout the whole life, a shape of the facial portion of the

skull not very different from that of immature individuals of all

races; in A. j. lituratus, palmarum, and dominicanus the skull of

aged individuals is very conspicuously modified. From this it

is concluded that the former group of races, in this particular

250 DR. KNUD ANDERSEN OW BATS. [Apr. 7

respect, is slightly more primitive than the latter group. Other

facts, to be mentioned hereafter, point to the same effect.

Teeth.—As in A. planirostris, with the following differences :—

m*, which is rudimentary in A. planirostris, has definitely

disappeared in A. jamaicensis. 185 skulls, of the seven sub-

species recognised in this paper, and of practically all ages, from

half-grown ‘to very old individuals, have been ex: amined ; all of

them lack m*, and in none is there any trace of the tooth having

been present.

m, ison the whole still somewhat smaller than in 4. plant-

rostris. When, as is the case in A. jamaicensis, the development

has reached the point that m’® is invariably wanting and m,

reduced to a mere rudiment, it might be anticipated “that this

rudimentary m, would, probably, show some tendency towards

complete disappearance. But it must be said at once that indi-

viduals lacking m, on both sides of the jaw, without any trace of

its alveoli, are extremely rare; the remarkable fact is not that

such exceptions do occur, but, in view of the minute size of the

tooth, that they do not occur more often. In two individuals

(a young adult A. 7. jamaicensis from San Domingo, B. M.

no. 50.7.8.43 ; and a fully adult male, with unworn teeth, of the

same race from Peten, Guatemala, U.S. N. M. no. 37912) m, is

entirely wanting on one side; in four individuals (an adult

male, with somewhat worn teeth, of A. 7. palmarum from Costa

Rica, B.M. no. 98.10.9.4; a young female of A. 7. jamaicensis

from Oaxaca, Mexico, U.S. N. M. no. 73255; an aged male, with

much worn teeth, of A. j. gamaicensis from Morelos, Mexico,

U.S. N. M. no. 64482; and an adult male, with unworn teeth,

ot the same race from Old Providence Island, U.S. N. M.

no. 37811) it is wanting on both sides. Thus in 97 p. ct. of the

185 skulls examined m, (or its alveolus) is present on both sides,

but averaging a little smaller than in A. planirostris, in 3 p. ct.

it 1s completely wanting either on one side or on both sides.

In those few species of Artibeus which have preserved a rudi-

mentary m*, this tooth is situated postero-internally to m*, partly

fitting into a sharp, subrectangular emargination in the posterior

border of this latter tooth, between its cusps 5 and 7 (text-fig. 41 A,

on p. 208). What becomes of this angular notch, ven as 1s

invariably the case in 4. Jamuicensis, m0” completely disappears g

It is an interesting fact that in the races of northern origin

(A. j. parvipes, ypucubanicus, jamaicensis, equatorialis) the emar-

gination is, in 77 p. ct. of the individuals, preserved quite or

almost as conspicuous as in any 4. planirostris, sometimes (in

about 19 p. ct.) it is decidedly reduced in size, rarely (4 p. ct.)

almost or quite disappeared; whereas in the races of southern

origin (A. 7. kitwratus, palmarum, dominicanus) the emargination

has only been preserved, as conspicuous as in A. planirostris, in

about 10 p. ct.; in 38 p. et. it is decidedly on the way towards

disappearance, in 52 p. ct. it has practically disappeared. Thus,

the northern races, though having like their southern relatives

1908. } DR. KNUD ANDERSEN ON BATS. 25]

lost m’, have in the large majority of individuals preserved the

notch in m* into which m’ fitted; they have, consequently, also

in this respect remained in a slightly more primitive stage than

the southern races, in which the notch, in no less than 90 p. ct.

of the individuals, is either conspicuously reduced or quite

obliterated.

The disappearance of the sharp angular notch between cusps 5

and 7 of m* is not effected by a reduction of the large, posteriorly

projecting cusp 5 (which would imply a decrease in the area

of m*), but, on the contrary, by a filling out of the notch, conse-

quently by a slight increase in the volume of m*; it is as if the

loss of m* has been compensated by a corresponding, or partly

corresponding, addition to that part of m* against which the

missing m’*® was pressed: in other words, the function of the

missing m* has, in the more highly developed races of A. jamat-

censis, been transferred, to a certain extent, to the postero-

internal border of m° (text-fig. 52).

Text-fig. 52.

A. Artibeus jamaicensis jamaicensis, f ad. St. Andrew’s I. B.M. 92.12.20.6.

Right m?, to show strong emargination of hinder margin of tooth. +.

B. Artibeus gamaicensis lituratus, 6 ad. Morretes, Parana. B.M. 3.7.1.127

Right m2, to show slight emargination of hinder margin of tooth. x +.

Reference has been made above to the fact that of 185 skulls

only 6 lack m, either on one side or on both sides of the mandible,

and it may be worth the while drawing attention also to the fact

that of these 6 aberrant individuals no less than 5 belong to the

race A. j. jamaicensis. The number of skulls examined of this

race is 75, the number of aberrant individuals 5 (about 7 p. ct.) ;

the number of skulls of all other races together is 110, of which

only one single individual is aberrant (1 p.ct.). It is probably not

quite accidental that in the large series examined the loss of m,

is less rare in the northern group of races. m,, in A. planirostris,

works against the whole surface of the small m? and a very

narrow postero- -internal margin of m*. When now, as is the

case in the large majority (about 80 p. ct.) of individuals of the

northern races “of A. jamaicensis, there is no compensation at all

for the loss of m* (7. e., no filling up of the notch in m? into

which m’? fitted), then m, has exceedingly little or nothing at all

to work against in the upper jaw, and it appears quite conceivable

that in such circumstances it shows a rather more pronounced

tendency to disappearance. In the sowthern races, on the other

252 DR. KNUD ANDERSEN ON BATS. [Apr. 7,

hand, the loss of m’ is, inthe large majority of individuals (about

90 p. ct.), more or less compensated by a slight increase in the

postero-internal portion of m* (by a filling up, partly or completely,

of the notch between cusps 5 and 7), and, consequently, m, has

almost as much to work against in the upper jaw as in those

species which possess an m?; the disappearance of m, is, probably

for this reason, of extreme rarity in the southern races.

So far as m’ and the notch in the posterior margin of m° are

concerned, the various stages represented by A. planirostris and

jamaicensis may be tabulated as follows :—

(A) m? present on both sides: almost all A. planirostris (94 p. ct.).

(B) m$ present on one side, entirely wanting on the other: 3 p. ct. of A. plani-

rostris.

CENSUS.

(1) Notch in hinder margin of m2, between cusps 5 and 7, perfectly preserved

(i. e. as distinct as in any Artibeus which possesses an m3): the large

majority (77 p. ct.) of the individuals of the northern races of A. jamai-

censis (A. 7. parvipes, yucatanicus, jamaicensis, equatorialis) ; a small

minority (11 p. ct.) in the southern races (A. 7. lituratus, palmarum

preceps).

(2) Notch in hinder margin of m? decidedly reduced: a minority (19 p. ct.) in

the northern races; a large number (38 p. ct.) in the southern races of

A. jamaicensis.

(3) Notch in hinder margin of m? almost or completely filled up: a vanishing

minority (4 p. ct.) in the northern; half the number of individuals

(51 p. ct.) in the southern races of A. jamaicensis.

The subjoined table and diagram are intended to give a view

of the gradual reduction of the notch between cusps 5 and 7 of

m* in the various races of A. jamaicensis (one race, A. 7. preeceps,

as being too poorly represented in the collections examined,

is excluded from the table; it apparently agrees with its nearest

relatives, A. 7. palmarwm and lituratus) :—

Angular emargination between cusps 5 and 7 of m2

|Total number |

of skulls quite or almost | aliaoeheor

examined, as conspicuous | decidedly ite

as in any | reduced. | Pete oncal

A. planirostris. | Sapp. E

parvipes ......... 12 92 p.ct. | Sma | 0 p. ct.

yucatanicus ...... 10 0) ep | FAD) op Ove

Jjamaicensis ...... 74 75 36 2s 3) 5

e@quatorialis...... 8 US ep 12°5 ,, 12°5 ,,

lituratus ......... 42 UP oy 36 Ca, BAP

palmarwin......... 30 ORs 40 ,, 50,

Northern races... 104 ep. Cte 19 p. ct. 4 p. ct.

Southern races .. 72, LL Opa? BB sg Gil 55

1908. | DR. KNUD ANDERSEN ON BATS. 253

p.ct. Parvipes | yucatanicus| jarmaicensis |aequatoria/(s | /ituratus «

palmarum

100 |

20 |

80 | |

Diagram showing the percentage of individuals, in the races of Artibeus jamaicensis,

in which the angular emargination between cusps 5 and 7 of m2 (former place

of m*) (1) is quite, or almost, as pronounced as in any A. planirostris (upper

curve), (2) has almost, or quite, disappeared (lower curve).

Tragus.—In most individuals there are 4 or 5 quite small

serrations on the outer margin of the tragus, above the median

projection ; the serrations may be sharp, but as a rule they are

more or less rounded off, rather often reduced to inconspicuous

nodules; very often the number is reduced to 3 (generally by

obliteration of the lower ones), rarely to 2, and still more rarely

all serrations are almost obliterated, the margin from the median

projection to the tip of the tragus being practically simple. There

is every transition between these stages, and they occur in all

races.

Nose-leaves.—According to Dobson A. jamaicensis (A. perspi-

cillatus in his ‘Catalogue’) is ‘‘at once distinguished” from

A. planirostris by having the front margin of the horseshoe

“completely bound down to the muzzle, in some individuals even

confluent with its integument,’ whereas in A. planirostris the

margin is “ free, separated from the muzzle, straight, unnotched.”

This character is thoroughly untrustworthy. First, in the same

geographical race of A. jamaicensis, often in individuals obtained

on the same spot, all transitions can be found, from a horseshoe

with the front marginas free as in any planirostris, to a horse-

shoe with the front margin completely continuous with the

integument of the muzzle, as shown in the table, p. 254 (based

exclusively on specimens preserved in alcohol). Second, in

A. planirostris the front margin is, as a rule, more or less free,

but individuals occur in which it is so completely bound down as

to be almost confluent with the integument of the muzzle.—The

front margin of the horseshoe is often simple, sometimes more or

less crenulate ; the crenulations extend not rarely over the whole

of the lateral margins, sometimes even over a smaller or greater

254

part of the lancet.

DR. KNUD ANDERSEN ON BATS.

[ Apr. 7,

The lateral margins of the horseshoe are

sometimes turned up so as to form a fold; in some individuals

they even show some indications of an emargination.

Front margin of horseshoe

Em aleneet ie rather more definitely | almost completely, or

aes Sgt some ct fastened down, but quite completely, fastened

a as nee still distinctly down to, or confluent

See eS: projecting. with, muzzle.

PAPVIPES ...-..00 Cuba (2). Cuba (10). Cuba (4).

GOCCTRDICLIS coool con voonds Yucatan (3). Yucatan (1).

jamaicensis ...... | Honduras (1). Nicaragua (1). Nicaragua (1).

Old Providence I. (2). | Honduras (1). Chiapas (2).

Jamaica (1). Guatemala (1). Oaxaca (2).

Porto Rico (6). Cainpeche (8). Vera Cruz (1).

| Chiapas (1). Morelos (8).

Vera Cruz (1). San Domingo (1).

Old Providence I. (2). | Porto Rico (1).

Jamaica (5).

Porto Rico (5).

St. Kitts (1).

@quatorialis......) ssveseeee | Heuador (1).

lituratus ......... Paraguay (1). Paraguay (1). Paraguay (5).

| Parana (1). Sta. Catharina (1).

Para (1).

palmarwm......... Venezuela (2). Venezuela (2). | Trinidad (2).

St. Vincent I. (1). Costa Rica (1). Costa Rica (3).

Nicaragua (1). | Guatemala (3).

| Guatemala (1). | Jalisco (1).

FURGOGOS weacooteacce|| ~ ——— goneno nce Guadeloupe (2). | Dominica (4).

All races (94) ... 18 p. ct 48 p. ct. 34 p. ct.

Wing-indices.—The proportionate length of the metacarpals

and phalanges is quite as in A. planirostris; so complete is the

similarity that the wing-indices of the latter species could be

substituted for those of A. jamaicensis without any appreciable

error (see the table on p. 310).

Hairing on limbs and membranes.— Upper side of proximal two

thirds (or half) of forearm, upper side of lateral membranes next

to body, of metacarpal of pollex, of femur, tibia, foot, and inter-

femoral (a narrower or broader distal portion excepted), under

side of proximal half of forearm and of proximal portion of inter-

femoral, very distinctly haired.

Tn the northern races the upper side of the tibia and inter-

femoral are generally more sparsely haired than in the southern

races, sometimes almost naked.

Colour.—There is often a certain colour difference between

1908. | DR. KNUD ANDERSEN ON BAS. 255

younger and aged individuals; further, some variation in the

development of the supraorbital and infraorbital stripes, partly

quite individual, partly dependent on age, and partly on the race

to which the individuals belong; and last, there is a certain

colour difference between A. 7. parvipes, yucatanicus, jamaicensis,

and wquatorialis (7. e. the northern races) on one side, A. 7. litu-

ratus and palmarum (the southern races) on the other side. It

has therefore proved convenient to give the description of the

coloration under the following five headings :—dark-coloured

individuals ; lighter-coloured individuals; indication of dorsal

stripe and white ear-edgings ; facial stripes ; concluding remarks

on the colour.

Dark-coloured individuals—Upper side, from the shoulder

region backward, dark smoky brown, almost blackish brown,

this colour confined to the distal third or fourth of the hairs;

base of hairs slate. On the anterior part of the upper side, from

the shoulder region forward, the hair-bases are as a rule dis-

tinctly lighter, almost smoke-grey. Under side dark smoke-grey

or brownish smoke-grey, with a peculiarly grizzled appearance,

owing to the short whitish or greyish-white tips to the hairs.

Tips of wings (region of third, or second and third phalanges of

third digit) generally light-coloured (whitish or yellowish white).

This is the extreme of the dark colour-type in fully adult

individuals (young, not full-grown individuals are still a shade

darker or duller). It occurs in ali races, but is especially common,

and much more frequently retained (or retained in a slightly

lighter shade) throughout the whole life, in the northern

than in the southern races; about 75 p.ct. of the fully adult

specimens examined of the northern races are “dark,” as against

only 25 p. ct. in the southern races. By advancing age (the

precise period varying considerably) the colour becomes gradually

of a somewhat lighter shade, even in those individuals which

never assume the proper ‘“ light phase ” described below.

Lighter-coloured indiwiduals.— Upper side, from shoulder region

backward, Prout’s brown, base of hairs almost wood-brown. On

the shoulder region and neck the Prout’s brown hair-tips are

generally very short or altogether wanting, the wood-brown

ground-colour of the fur therefore more or less, or completely,

exposed, producing an often very strong contrast between the

anterior and posterior parts of the dorsal surface. Under side dull

brown, base of hairs very little, or not at all, lighter; extreme

tips of hairs generally whitish or greyish white. Tips of wings

as in dark-coloured individuals.

This is very nearly the extreme of the light colour. The hair-

bases on the shoulder region and neck are in some specimens still

lighter than ‘“ wood-brown,” almost yellowish white.

The dark extreme described above and the light extreme here

under consideration are very different indeed, but there is

absolutely no sharp line of separation between them; they are

connected by many intermediate stages. Putting aside all

Proc. Zoou, Soc,—1908, No. X VII, 17

256 DR. KNUD ANDERSEN ON BATS. [ Apr. 7,

variations of shade, the principal stages are these three: dark

smoky brown (dark type), Vandyck-brown (intermediate stage),

Prout’s brown (light type).

The light colour type occurs in all races, but much more fre-

quently in the southern than in the northern races; about

75 p. ct. of the fully adult specimens examined of the southern

races (representing many different stages of wear of the teeth) are

light-coloured, as against only 25 p. ct. of the northern races.

The subjoined table (p. 257) gives a statistical view of the

number of dark and light coloured individuals in the large series

examined. It is based exclusively on fully adult individuals (distal

epiphyses of metacarpals ossified), and exclusively on dried skins,

no spirit-specimens, however well preserved, having been taken

into account. It will be noticed that 62 p. ct. of the available

individuals of the southern races have the teeth from slightly

worn to much worn, whereas the same is the case with a some-

what smaller percentage (50 p. ct.) of the northern races ; it may,

perhaps, have slightly exaggerated the final results, but a glance

at the table will be sufficient to prove that it cannot have had

any essential influence on the conclusions, which may be epito-

mised as follows :—(1) Light-coloured individuals are much more

common in the southern than in the northern races: (2) of 30

individuals of the northern races which, though fully adult, have

the teeth unworn or practically unworn, 28 are dark-coloured ;

of 20 individuals of the southern races, of corresponding age, only

half the number are dark-coloured: (3) of 20 specimens of the

northern races with the teeth slightly or somewhat worn, 12 are

dark-coloured ; of 18 specimens of the southern races of corre-

sponding age, none are dark-coloured: (4) of 9 specimens of the

northern races with the teeth well worn or much worn, 4 are

dark-coloured ; of 15 specimens of the southern races of corre-

sponding age, 2 only are dark-coloured. These two conclusions

therefore, would seem to be well founded: that there is a well-

marked average difference of colour between northern and southern

races; and that the light colour type, though sometimes (par-

ticularly in the southern races) occurring in adult individuals

with unworn teeth, is especially characteristic of the somewhat

more advanced age.

Immature individuals are always dark-coloured ; of the northern

races the large majority of adult and aged individuvls are dark-

coloured, of the southern races a minority only. From this it is

concluded that the northern races have also in this respect

remained in a slightly more primitive stage than the southern

forms. One of the following paragraphs will show that a closer

study of the development of the head-stripes in these two groups

of races leads to the same conclusion.

Indication of dorsal stripe and white ear-edgings—One speci-

men of A. 7. jamaicensis (2 ad., San Vicente, Chiapas, Mexico,

teeth much worn, fur rather light-coloured, U.S. N. M. no. 133044)

has a short longitudinal stripe on the hinder part of the back.

1902. | DR. KNUD ANDERSEN ON BATS. Q57

Dark and light coloured individuals.

Teeth

ee a Revs wil) aleve!

of skins | unworn or slightly or! well worn | fully. adult

; < : 3 | Individuals.

of adults. practically somewhat or 1]

unworn. worn. much worn |

| PATUUDES eyes. 9 darker 3, darker 3, | darker 2, | darker 8,

| lighter 0. lighter 1. | lighter 0. | lighter 1.

yucatanicus ..... 9 darker 5, | darker 1, a4 darker 6,

lighter 0. | lighter 3. ae | hghter 3.

Jamaicensis ...... 36 | darker 19, | darker 7, darker 1, | darker 27,

lighter 2. | lighter 3. lighter 4. || lighter 9.

equatorialis...... 5 | darker 1, | darker1, darker1, || darker 3, |

| lighter 0. | lighter 1. lighter 1. || lighter 2. |

Liturgies) 30 darker 9, | darker 0, darker 1, | darker 10,

, lighter 6. | lighter 10. lighter 4. |) lighter 20.

/palmarum......... 23 || darker 1, | darker 0, | darker 1, || darker 2,

| | lighter 4. lighter 10. lighter 9. | lighter 21.

Siig We Tee eS | 2 SHER ie) fete area Ce | af

| Northern races... 59 darker 28, | darker 12, | darker 4, || darker 75 p. ct. |

| | lighter 2. lighter 8. lighter 5. |) lighter 25 p. ct. |

| | lighter 10. | lighter 18. lighter 13.) lighter 77 p. ct. |

The anomaly is worth noticing, because it recalls the white

dorsal stripe characteristic of many species of the more primitive

genus Vampyrops (and of Uroderma).

A few individuals show some slight trace of a narrow white

margin to the ears, a feature of common occurrence in those

species of Artibeus which possess 3 molars.

Facial stripes.— Many individuals have two pairs of whitish

or yellowish-white, longitudinal stripes on the head, viz. a pair

of supraorbital stripes from the base of the lancet, passing above

the eye, to a point above the inner side of the ears; and a pair

of infraorbital stripes from the angle of the mouth to the outer

base of the ears.

There is every stage of transition, from complete absence to

very strong development, of these facial stripes. A certain corre-

lation exists between the development of the supraorbital and

infraorbital stripes; very strong supraorbital stripes are almost

always associated with well-marked infraorbital stripes, faint

supraorbital stripes with completely wanting or indistinct infra-

orbital stripes. The infraorbital stripes are, almost invariably,

markedly fainter (narrower, or more indistinct) than the supra-

orbital stripes.

Immature individuals have the facial stripes less well marked

than adult or aged examples. As to fully adult and aged indi-

viduals, there is a difference in the development of the facial

stripes between the northern and southern races, and a difference

258 DR. KNUD ANDERSEN ON BATS. [EaypiE

between dark-coloured and light-coloured specimens :—Of 59

ful y adult examples (skins) of the northern races, 53 have the

supraorbital stripes perfectly wanting or but very faintly mdi-

eated; in 4 they are distinct, in 2 strong; compare with this

the southern races: of 53 fully adult examples (skins) 3 have

these stripes faint (or quite undeveloped), | distinct, and no less

than 49 strong or very strong. In all individuals of the northern

races the infraorbital stripes are quite wanting or, at most, very

faintly indicated; in more than half the number of skins of the

southern races (32 of 53) they are also faint or undeveloped, but

in 15 they are distinct, and in 6 strong.—That strong facial

stripes are much more frequently associated with light than with

dark colour of the fur is proved by the following statistics :—

Of the dark-coloured individuals of the northern races scarcely

5 p. ct., of the light-coloured individuals of the same races about

25 p. et., have the supraorbital stripes well developed; of the

dark-coloured individuals of the southern races 75 p. ct., of the

light-coloured individuals 100 p. ct., have the supraorbital stripes

well developed.—More extensive material than I have been able

to bring together may, of course, alter these figures somewhat,

in one or other direction, but it is not likely that it will alter

the general conclusion to any essential degree-—The subjoined

table gives the details (compare diagram p. 259).

From the two facts, viz. (1) that the facial stripes are less

developed in immatures than in adults, less developed in the dark

Facial stripes.

| Supraorbital stripes Infraorbital stripes |

| Total | sh Died, al

| | number | |

| of skins | none strong | none | strong

faint. strong. | faint. strong.

izes | |

parvipes, dark ...... 8 8 ® | @ 8 | @ 0

ew iliaht wee Ieee 0 ek a a nO 0

yucatanicus, dark ...| 6 6 Onna aetO GB | © 0

55 light ...| 3 2 1 0) 3 0) 0

jamaicensis, dark soll 26 | iL CO 27 (0) (0)

$5 light ...} 9 8 al Oo | 9 0) 0)

equatorialis, dark...| 3 2 0 Lees 0 0

fs light...) 2 1 0 1 2 0 ® |

lituratus, dark ......| 10 3 0) 7 9 1 0)

| eo hibeate babi 20 0 1 19) ||) We 4 A

palmarum, dark......| 2 OT oO 2 1 1 Onn

| an light...... 21 ® | oO 21 10 9 2

La | es =| | x

N. races, dark ...... | 44: 42 | 1 iL 44 | O 0

ee elit. 15 11 3 eh 1B 0

S. races, dark......... | 12 3. 0) 9 10 | 2 0)

ieeiioht ie 41 ONM ee! AO || 22 | 13 6

| | 1 |

1908. ] DR. KNUD ANDERSEN ON BATS. 259

than in the light colour type, and (2) much less developed in

the northern than in the southern races, I conclude that the

former, also in this respect, occupy a rather lower stage in the scale

of evolution *.

Concluding remarks on the colour—The facts recorded above

may be briefly summed up as follows :—Adult and aged indi-

viduals are dark smoky brown on the upper side (dark extreme),

or Vandyck-brown (intermediate stage), or Prout’s brown (light

extreme), or some shade of brown intermediate between these

three. Dark colour is, in all races, more common in younger

adults than in aged adults. Dark-coloured individuals are pre-

dominant in 4.7. parvipes, yucatanicus, jamaicensis, and wqua-

torialis (‘‘northern” races), light-coloured in A. j. ditwratus and

palmarum (“southern ” races). Facial stripes are commoner and

more strongly developed in aged individuals than in immature

and young adults; commoner and more strongly developed in

light-coloured than in dark-coloured individuals ; commoner and

stronger in the southern races than in the northern. There is a

certain correlation in the development of supra- and infraorbital

stripes: when the former are strong, the latter are as a rule

rather well marked or, at least, not quite obsolete; when the

former are rather indistinct, the latter are as a rule wanting.

Diagram showing the percentage of individuals, in some races of Artibeus jamai-

censis, in which the supraorbital stripes are distinct, strong, or very strong.

* Generally speaking, facial stripes are no doubt a very primitive feature among

Vertebrates, going back, as it does, to Reptiles (and being present in a vast number of

Birds). But each particular case cannot, of course, be considered only from this

general point of view. Asa matter of fact, the young individuals of A. jamaicensis

s. lat. have, as pointed out above, the facial stripes much less developed than the

adults, and we are therefore compelled to assume that this primitive feature has, for

some reason or other, been lost in that type of Bat (or its predecessors) from which

the living A. jamaicensis originated, and that now there is again a tendency to

reversion to the old feature in adult individuals.

260 DR. KNUD ANDERSEN ON BATS. [Apr. 7,

range.—From 8. Brazil, Paraguay, and Peru, to Central

Mexico (Jalisco), including the West Indies; no indisputable

record from Guiana.

Races.—Seven races of Artibeus jamarcensis are described below,

viz., A. j. parvipes, yucatanicus, jamaicensis, and equatorialis ;

these four, as being probably of Central American and West

Indian or agin, may be conveniently called the “ northern ” races ;

further, A. 7. litwratus, palmarum, and preeceps ; these three, as

es probably of South American origin, may be called the

« southern ” races.

Remarks.—A. jamaicensis is at once distinguishable from all

other species of the genus by the combination of these two

characters: molars 2, maxillary tooth-row 9:3-12°2 mm.; the

first character excludes all species with lhe exception of A. glaucus

and watsont, which possess a small m,; the second character

excludes the two latter species, A. glanens and watsont being very

much smaller (maxillary tooth-row 6°5-7 mm.)

There is no reliable eaternal character by which A. jamaicensis,

all races taken together, can be discriminated from A. planirostris

s. lat.; the nose- tenes, ears, wing-structure, coloration, even the

dimensions, are practically identical; to the larger races of

1. jamancensis (lituratus, palmarwm) correspond the large A. plani-

rostris fallax; to the small races of A. 7. jamaicensis (parvipes,

yucatanicus, jamaicensis) the small races of A. planirostris (plani-

rostris, trinitatis, &e.); A. jamaicensis might properly be described

as an “ A. planirostris” which has permanently lost m*. Dobson’s

way of discriminating the two species externally—the front

margin of the horseshoe free in A. planirostris, bound down to the

muzzle in “4. perspicillatus” (A. jamaicensis)—would in 66 p. ct.

of cases lead to wrong or doubtful identification.

But the matter becomes rather different when considering each

race of A. yamaicensis separately. Brazil is inhabited by the

large A. 7. ituratus, whereas the race of A. planirostris occurring

in the same region (A. p. planirostris) 1s considerably smaller, so

that only in very rare instances is there any difficulty at all in dis-

criminating these two forms. Guiana is inhabited by the largest

race known of A. planirostris (A. p. fallax); A. jamaicensis is as yet

not recorded with certainty from Guiana, and perhaps it has not

spread to that country. In Venezuela much the same contrast

obtains as in Brazil, A. 7. palmarum being as a rule considerably

larger than A. p. planirostris. A. jemaicensis has spread over the

whole of the West Indies (parvipes, jamaicensis, preeceps) ; A. plani-

rostris 18 totally absent from the West Indies proper, having

spread only over the Venezuelan coast islands as far as Grenada.

Only Central America and Mexico are inhabited by certain small

forms of A. jamaicensis (jamaicensis, yucatanicus) which ex-

ternally, as a rule, are so completely like the small race of

A, planirostris (planirostris) living in the same region, as not to

be distinguishable without an examination of the teeth.

1908. ] DR. KNUD ANDERSEN ON BATS. 261

ARTIBEUS JAMAICENSIS PARVIPES Rehn.

1828. Phyllostoma Jamaicense Leach, Horsfield, Zool. Journ. ii. (April to July,

1827) pp. 238-40.—Macleay’s specimens from Cuba.

1872. ? Artibeus perspiccilalune (sic), C. J. Maynard, Bull. Essex Inst. iv. no. 10,

p. 144 (Oct. 1872)—Key West, Florida.

1878. Artibeus perspicillatus (partim, not L.), Dobson, Cat. Chir. Brit. Mus.

pp- 519-20,

1878. <Artibeus macleayii Dobson (from Gray’s MS.), op cit. p. 520, speci-

mens h-k.—Cuba.—Nomen nudum.

1902. <Artibeus parvipes Rehn, Proc. Ac. N.Sci. Philad. pp. 639-40 (12 Oct. 1902),

—Type locality: Santiago de Cuba.

Diagnosis.—The smallest form of A. jamaicensis: zygomatic

width of skull 15°5-17 mm. (average 16:1); maxillary tooth-row

93-10 mm. (average 9°7); forearm 54-60 mm. (average 56:8).

General characters.—The angular emargination between cusps

5 and 7 of m?is generally as well marked as in any 4. plant-

rostris. The coloration of the fur of the upper side is generally of

the dark type, but light-coloured individuals occasionally occur.

The facial stripes are usually obsolete or faint.

A. j. parvipes and yucatanicus.—There is only a rather small

average difference between A. 7. parvipes and A. 9. yucatanicus.

The skull of parvipes is generally a trifle smaller (total length

26°9 mm., as against 27:4 mm. in yucatanicus) and more deli-

cately built (zygomatic width 16:1 mm., against 16°8 mm.); the

difference in the size of the teeth is infinitesimal (maxillary tooth-

row 9°7 mm., against 9°9 mm.). The forearm and metacarpals

of parvipes average 2°5 to 3 mm. shorter than in yucatanicus ;

also the phalanges, the tibia and foot average a little shorter.

(See the detailed measurements, p. 264.) Being in every respect,

save their average dimensions, perfectly similar, 4.7. parvipes,

and yucatanicus are, in many cases, practically indistinguishable,

if the precise locality in which the specimens were obtained is

unknown.

A. j. parvipes and jamaicensis.—Although there is no absolute

difference between 4. j. parvipes and j. jamaicensis, the former is

as a rule easily distinguished by its smaller and slenderer skull

and smaller teeth. Externally there is the same average difference

in dimensions between parvipes and jamaicensis as between

parvipes and yucatanicus.

Specimens from different localities —There is no difference

between examples from Eastern and Western Cuba.

Supposed occurrence at Key West, Florida.—While at Key West

Island, south of Florida, in the early winter of 1870, Mr. C. J.

Maynard watched several large bats flying about the city; the

single specimen secured has since been lost, but a drawing made

by Maynard enabled the late Dr, Harrison Allen to identify the

species as “ Artibeus perspicillatus Linné.”* The Key West

bats, Maynard writes, “closely resembled in flight a species

which I had seen in Northern Florida two years before.” If

* Gerrit S. Miller, Jr., Proc. Bost. Soc. N. H. vol. xxviii. no. 7, p. 214, March

1898.

262 DR. KNUD ANDERSEN ON BATS. (Apr. 7,

the identification is correct, it appears natural to suppose that

the Key West bat was A. 9. parvipes, and that, consequently,

the range of this form extends to the islets between Cuba and

Florida, perhaps to peninsular Florida, the flora and fauna of

the southern part of which has, as well known, a subtropical

character:

Specimens examimed.—25 specimens (9 skins) and 12 skulls,

from the following localities :—

British Museum :—Cuba (4). Skulls of 3 specimens.

U.S. National Museum *:—W. Cuba: El Guama (3); San

Diego de los Banos (5); Guanajay (6); Mariel (1).—E. Cuba:

Baracoa (2); El Cobre (4).—9 skulls from practically all the

localities enumerated.

Range.—Cuba, perhaps extending to Florida.

Rehn’s 4. parvipes, 1902.—Stated to differ from the Jamaican

A. jamaicensis “in the smaller forearm and tibia, and the narrower

and lighter built foot”; forearm on average “53°7 mm.” (in

jamaicensis “583 mm.”), tibia ‘ 20°8 mm. " ( jamaicensis

“21-9 mm.”), foot in the type “14 mm.” (in a jamaicensis

“15 mm.”). ‘The second upper premolar is broader and with

a heavier internal shoulder, the teeth being actually broader

than [in] yamaicensis, though the latter possesses a slightly larger

skull.”

As to the external characters given by Rehn, it must be said,—

first, that there is no definite line of separation between the

Cuban A. parvipes and the Jamaican dA. 7. jamaicensis: I have

seen examples of jamaicensis with the forearm only 57 mm.

long, and examples of parvipes with the forearm 60 mm.; in

jamaicensis the lower leg is occasionally only 22 mm., in par wipes

sometimes 23 mm.; in some jamaicensis the foot measures only

15-2 mm. in length, in some parvipes as much as 16°8 mm.

Second, the average measurements calculated by Rehn from

SIx parvipes and six jamaicensis give a somewhat exaggerated

idea of the difference in size between the two races; compare

the table of measurements, p. 284. Third, since especial stress

was laid by Rehn on the smaller foot in parvipes (see his

paper), 1b may be well to emphasise that this is perhaps the

least conspicuous of the external differences ; the average differ-

ence in this respect between the two forms 1 is so small (0: 7 mm.),

and the actual measurements so frequently overlapping each

other, that it would only in a small minority of cases be pos-

sible to distinguish the two forms by the size of the foot; the

smaller foot in parvipes is simply a consequence of the smaller

size of the animal; a closer study of the table of measure-

ments (p. 284) will show that proportionately the foot of parvipes

is precisely of the same size as in jamaicensis.—The dental

difference mentioned by Rehn must either have been derived

from an individual aberration in the specimen examined by him,

* U.S. N. M. nos. 103621-22, 102627, 103631, 103640, 103643, 103670, 103692-95,

103725-26, 103735-34, 113758, 113761, 113823, 113834-36,

1908.] DR. KNUD ANDERSEN ON BATS. 263

or it must be a mistake ; as a matter of fact, the structure of the

teeth in parvipes is exactly as in yamaicensis, but the size of the

teeth on an average slightly smaller; and the heel of p* 1s, as

might be expected from this, not larger, but on average a trifle

smaller than in janwicensis.

ARTIBEUS JAMAICENSIS YUCATANICUS J. A. Allen.

1888. Artibeus perspicillatus (partim, not L.) Thomas, P. Z.S. p. 129 (21 Feb.

1888).—Cozumel Island (Yucatan).

1897. Artibeus perspicillatus (partim, not L.) J. A. Allen & Chapman, Bull. Am.

Mus. N. H. ix. Art. 1. pp. 3-5 (23 Feb. 1897).—Yucatan.

1904. Artibeus yucatanicus J. A. Allen, Bull. Am. Mus. N. H. xx. Art. 20,

pp. 232-33 (29 June, 1904).—T ype locality: Chichen Itza (Yucatan).

Diagnosis.—Similar to A. 9. parvipes, but skull and external

dimensions averaging a little larger. Zyeomatic width of skull

16°3-17-2 mm. (average 16°8) ; maxillary tooth-row 9°7—10°2 mm.

(average 9°9); forearm 56-61 mm. (average 59-6).

General characters.—The angular emargination between cusps

5 and 7 of m? is usually as well marked as in any 4. planr ostris,

but individuals occur in which it is distinctly reduced in size.

The coloration of the fur of the upper side is generally of the dark

type, but light-coloured examples are not rare, especially among

individuals with somewhat worn teeth. The facial stripes are

usually obsolete or faint.

A. 7. yucalanicus and parvipes.—The skull of A. J. yucatanicus

is generally a trifle larger (total length 27:4, as against 26°9 mm.

in parvipes) and, especially, broader (ay gomatic width 16-8 mm.,

against. 16:1 mm.) ; the difference in the size of the teeth is

infinitesimal. The forearm and metacarpals of yucatanicus

average 2°) to 3 mm. longer than in parvipes ; also the phalanges,

tibia, and foot average a little longer. See the detailed measure-

ments, p. 264.—The two races come so extremely close to each

other, and are practically so difficult to discriminate that, were it

not for their different habitat, they ought not to be kept separate.

A. 9. yucatanicus and jamaicensis.—A. j. yucatanicus forms a

transition between the Cuban 4. j. parvipes and the Central

American (and Mexican, and West Indian) 4. 49. jamaicensis.

Externally yucatanicus is indistinguishable from jamaicensis,

there being not even an average difference in size; but in the

size of the skull and teeth it is decidedly nearer to parvipes than

to jamarcensis.

Specemens examined.—l4 specimens (7 skins) and 11 skulls,

from the following localities :—

British Museum :—N. Yucatan (1); Cozumel I. (1).—Skulls of

both specimens.

U.S. National Museum *:—Yucatan (4); Merida, Yucatan (1);

Progreso, Yucatan (3); Chichen Itza, Yucatan (3); Belize (1).—

9 skulls, from all the localities enumerated.

* US. N. M. nos. 11445 (87547), 108153-55, 108489-91, 143119-22

264 DR. KNUD ANDERSEN ON BATS. | Apr. 7,

Range.— Yucatan, including the coast islands. The only speci-

men examined from Belize is referable to dA. 7. yucatanicus.

Certain specimens from Campeche come very near to this race.

J. A. Allen’s A. yueatanicus, 1904.—In 1897 (J. s. ¢.), Allen and

Chapman recorded four specimens of “ Artibeus perspicillatus”

(A. jamaicensis Leach; A. carpolegus Gosse) from Chichen Itza,

Yucatan. In 1904 (7. s. ¢.), Allen selected one of these examples

as the type of a new species, A. yuwcatanicus. In size and colour,

he writes, it resembles ‘“‘the West Indian forms of the genus,

especially 4. parvipes of Cuba, from which it is not readily distin-

guishable”; “four skulls of 4. parvipes measure the same [ viz.

27 mm.] in total length, but a little less in zygomatic width.”

Allen does not enter into further details as to the differential

characters of the new species. (The forearm measures, according

to Allen, 55 mm., the third metacarpal 57 mm.; the latter must,

of course, be a misprint, if the former is correct.)

Measuremenis of A. j. parvipes and yucatanicus.

A. j. parvipes. || A.J. yucatanicus.

25 adults, 14 adults,

12 skulls. 12 skulls.

Min. Max. Mep.|Miy. Max. Mep.

| mm. mm. mm.|/mm. mm. mm. |

| Skull, total length, to front of c... 26 273 26:9 || 262 28:3 27-4) |

> Mastoidiwadth =... 1... 14 15 14:4 | 14-2 15 146

» width of brain-case... ..... 118 12;'8 12:1 || 11-7 12°38 = 12°3 |

» Z“ygomatic width ......... 15°5 17 161 , 16°3 172 #8168

| 4, maxillary w ea 112 12:3 = 11°8 || 11°5 12°8 12°71

| across cingula of canines ...._7°2 8 75 73 8 ea

Mandible, to front OL WNC coococoee| 1} 19 184 18 197 189

Upper teeth, c-m2 ..................| 93 10 97 || 9:7 102 99

Lower teeth, cam, ....... 102 10°7 10'4 || 10°2 11 10°7

Kar-conch, leneth, inner margin. 13°2 155 14-4 || 14 145 |

eS leneth, outer margin .| 19 21:5 20:3 || 19°8 21

an width See eee ae meme lulls, 155 14°2 | 13°7 13°7

MrRAS, NEMEXN oocsscosoosecsvsevevcan| (OA 7-2 68 | 68

| Wenm@etir, WEEN on conccooscooHosecnsana| oD 10 973 || 85 9

ae A ATGU ELIA AMER mma alec tert 9) a 64 || 7 75

Horseshoe, width ..................) 652 8 DM WS 78

JIOVKEEWEN IA. ko. Roe ncomaumenecrmacdsnaessnal ie! 60 568 | 56 61 59°6

Pollexcyiewervae | Meni ECR Eyes eal eal azo 145 13°9 | 13:2 14:3 13°7

3rd metacarpal. ae ete En AS 55 51:3 50 552 53°8

1 US CLL aa vale aaa aoe We ee Ps 18 163 162 18°8 17-2

VEE) I Sa a Ae? A Ue Ra are i ee el es) 29°5 27°3 | 26°7 80°2 28°8

TRS ess eee Ace ABER copped ee cs asolN Lene! 168 145 |, 13°2 15°2 146

4th metacarpal... ees hana ere ll ATR 538 502 | 488 547 53

NYA age eae Be te, oles Genet Seat a 158 145 | 14 16:3 15°3

11\c2 pene bob haeaan dtandocuae Nloko 20°5 179 | 17:8 20°7 19

5th metacarpal .. Feag lonaddadcereacuel| Rens 56 51°9 | 50 56 54°5

vl Weg ee. oan en 10°7 12°2 11°3 | 10°8 12°8 117 |

W222 TS RRO ery ort lth Aled ot on rin Mal 15 13°6 | 13 16 14°6

JUN FesPETTAO TA co nos. scessa00e00505cea00n, LO) 17 141 | 148

TW Ose Weer .gagsaee osc 620080 590 opnc00 ens) PAUPY/ 23 21°6 | 21:3 23 22°2 |

IDOLE WAU CIENYS arecans-scencanonnesell 1 168 154) 142 17 15°7

(OE INGEIES ndreamBanemeRen atoadungh <tueeaeck lie anes 73 64 58 62

1908. | DR. KNUD ANDERSEN ON BAS. 265

As already pointed out above, d. 7. yucatanicus 18 similar

to A. 7. parvipes and jamaicensis in every respect but a small

average difference in size; in the size of the skull and teeth it

comes extremely near to the former race, in external dimen-

sions it is indistinguishable from the latter. 4.7. yweatanicus

therefore, cannot be considered a distinct species; it forms, in

its characters as in its geographical habitat, a transition between

the two races, and is in some cases practically inseparable from

either.

ARTIBEUS JAMAICENSIS JAMAICENSIS Leach.

1821. Artibeus Jamaicensis Leach, Trans. Linn. Soc. xni. pt. i. p. 75 (read 7 March,

1820).—T ype locality : Jamaica.

1821. Madateus Lewisii Leach, t. c. pp. 81-82.—Type locality; Jamaica.

1851. Artibeus carpolegus Gosse, A Naturalist’s Sojourn in Jamaica, pp. 271, 272

(footnote), pl. vi. fig. 5—Type locality : Jamaica.

1861. Arctibeus perspicillatus (not L.) Tomes. P. Z. 8. (26 Feb. 1861) p. 64.—

Jamaica (W. Osburn’s specimens).

1865. Arctibeus carpolegus Gosse, W. Osburn, P. Z. 8. (24 Jan. 1865) pp. 64-67.—

Jamaica (habits).

1865. Arctibeus jamaicensis Leach, Peters, MB. Akad. Berlin (13 July 1865),

p. 356. f

1878. Artibeus perspicillatus (partim, not L.) Dobson, Cat. Chir. Brit. Mus.

pp. 519-20.— Jamaica, 8. Domingo, Mexico, Central America (partim).

1889. Dermanura eva Cope, American Naturalist, vol. xxi. no. 266, p. 130 (Feb.

1889).—Type locality: St. Martins, W.I.

1890. Artibeus coryi J. A. Allen, Bull. Am. Mus. N. H. ii. no. 1. p. 173, ef.

pp. 171-72 (14 Noy. 1890).—Type locality: St. Andrew’s I.

1896. Artibeus perspicillatus (not L.) Elliot, Field Col. Mus., Zool. Ser. vol. i.

no. 3, p. 82 (May 1896).—San Domingo, W.I.

1902. Artibeus intermedius J. A. Allen (partim), Outram Bangs, Bull. Mus. Comp.

Zool. xxxix. no. 2, p. 50 (April 1902).—Bogava, Chiriqui (the smaller

specimens).

1904. Artibeus insularis J. A. Allen, Bull. Am. Mus. N.H. xx. Art. 20, pp. 231—

32 (29 June, 1904) —Type locality: St. Kitts, W.I.

Dagnosis—Similar to A. 7. yucatanicus, but the skull, on

average, a little larger and more heavily built, the teeth a little

larger. Zygomatic width 16°2—-18-2 mm. (average 17-4) ; maxillary

tooth-row 9°8-1]1 mm. (average 10°3); forearm 56°5-65 mm.

(average 60:1).

General characters.—The angular emargination between cusps

5 and 7 of m* (reminiscent of the missing m’) is in 75 p. ct. of

the large number of skulis examined as strongly pronounced as

in any of those species (A. concolor, planirostris, hirsutus) which

possess an m*; in only about 20 p. ct. it is distinctly reduced in

size, and in 4 or 5 p. ct. it has almost disappeared. The colora-

tion of the fur of the upper side of the body is generally of the

dark type, even in individuals with somewhat worn teeth ;

specimens of the light colour type are rare among full-grown

individuals with unworn teeth, become rather more common

among those with somewhat worn teeth, and appear to be pre-

dominant among those with much worn teeth. The supraorbital

stripes are almost always undeveloped or faint, rarely distinct,

never (so far as the available material goes) very strong; the

infraorbital stripes are always undeveloped or faint.

A. j. jamaicensis and closely allied forms.—Neither in structure

266 DR. KNUD ANDERSEN ON BATS. [Apr. 7,

nor in colour is there any difference between A. j. jamaicensis and

its closest allies, 4. 7. yucatanicus and parvipes; but there is a

distinct average difference in size, small though it is. The details

have been briefly commented upon in the descriptions of the two

latter races (pp. 261 and 263), and are expressed in the table of

measurements, p. 284.

Specimens from different localities ——A. j. jamaicensis covers a

much wider area than the two foregoing races (yucatanicus,

parvipes). It is distributed over the whole of Central America,

including the outlying small islands (St. Andrew’s, Old Providence),

and S. Mexico, at least as far as Morelos; further, over Jamaica,

San Domingo, Porto Rico, and the smaller islands east of this

latter, at least as far as St. Kitts. I have carefully compared

specimens (skulls, spirit-specimens, skins) from all these places,

and am unable to find the slightest indication of a difference. It

is very easy, indeed, to contrast a smaller-skulled (smaller-toothed)

individual from one place with a larger-skulled (larger-toothed)

from another continental place or another island, a smaller-winged

with a larger-winged, a darker-coloured with a lighter-coloured,

or a specimen with short with a specimen with long interfemoral.

When material is scarce, differences of this kind may very easily

lead (and, in fact, have led) to separation of different forms or

even species; but whenever the various localities have been

represented by a sufficiently extensive series in the material

examined by me (as has been the case with Central America,

S. Mexico, Jamaica, Porto Rico, St. Andrew’s Island, Old Provi-

dence Island), I have found the range of individual variation to

be precisely the same within each particular locality ; there is no

difference whatever in structure nor in coloration, and there is

not even an average difference in size.

In the comparative table of measurements below (p. 269) the

specimens have been arranged under seven headings, viz., Central

America (Panama, Nicaragua, Honduras, Ruatan I., Guate-

mala), S. Mexico (Campeche, Chiapas, Tehuantepec, Oaxaca, Vera

Cruz, Morelos, ‘‘ Mexico”), Jamaica, Porto Rico, San Domingo,

St. Kitts, St. Andrew’s and Old Providence Islands. From each of

these seven areas are given the minimum, maximum, and average

measurements (the latter not for the few available specimens

from San Domingo and St. Kitts). The table shows the complete

accordance in the size of individuals from these seven areas.

To prevent wrong identification it is important to emphasise

that Central America is inhabited by two races, which ought not

to be (but hitherto have always been) confused, viz. the smaller

(truly indigenous) A. 7. jamaicensis and the larger A. j. palmarum

(an immigrant from south).

Specimens examined.—105 specimens (58 skins) and 76 skulls,

from the following localities :—

British Museum :—Nicaragua : Corinto (1).—Honduras: Half-

Moon Key (1); Ruatan Island (5).—Guatemala: Duenas (1).—

““ Mexico” (1).—St. Andrew’s Island (3).—Jamaica (17).—San

1908. | DR. KNUD ANDERSEN ON BATS. 267

Domingo (2).—Porto Rico (2).

enumerated.

U.S. National Museum * :—Panama: Colon (1).—Nicaragua:

Greytown (1); Escondido River, 50 miles from Bluefields (1).—

Guatemala: Peten (1).—Campeche : Apazote, near Johallun (4).

—Chiapas : Palenque (7) ; San Bartolomé (4); San Vicente (4).—

Tehuantepec (1).—Oaxaca: Santo Domingo (4).—Vera Cruz:

Tuxtla (1); Mirador (1).—Morelos (3).—Old Providence Island

(5).—Jamaica, various places (9).—San Domingo (1).— Porto Rico,

various places (23).—St. Kitts Island (1).—47 skulls, from all the

localities enumerated.

Range.—Central America and 8. Mexico, as far north as Morelos,

and exclusive of Yucatan; St.. Andrew’s and Old Providence

Islands; Jamaica, San Domingo, Porto Rico, as far east as

St. Kitts.

Leach’s A. jamaicensis, 1821.—-The probable type, the skin

(unregistered) of an adult individual from Jamaica (‘“a@” in

Dobson’s Catalogue, p. 520), with skull (“q’ ”) is in the British

Museum.

Leach’s Madateus lewisti, 1821—The type, in the British

Museum (skin, with skull; unregistered ; “>” in Dobson’s Cata-

logue, p. 520), 1s a very young A. j. jamaicensis, sent from

Jamaica by W. Lewis. The peculiarities which led Leach to

regard it as a distinct species (and genus) are due to the

immaturity of the individual.

Gosse’s 4. carpolegus, 1851.—Based on a ¢ ad., obtained by

Gosse at Content, Jamaica, preserved in alcohol ; Brit. Mus.

no, 47.12.27.13. Indistinguishable from Leach’s 4. jamaicensis.

The Bat now called Aritews achradophilus was described by Gosse

under three names :—4d. jamaicensis Leach (of which he had only

Horsfield’s description in Zool. Journ. iii. (1828) p. 238 for com-

parison), 4. achradophilus Gosse, and A. sulphureus Gosse (light

phase); the true 4. jamaicensis Leach he described under a new

name, 4. carpolegus Gosse.

Cope’s Dermanura eva, 1889.—Founded on two adult males,

from the island of St. Martins, West Indies, now in the collection

of the Academy of Natural Sciences, Philadelphia. They were

stated by Cope to have 3 molars only, and therefore referred by

him to the “genus” Dermanura ; the rest of the description

‘(lip tubercles, nose-leaves, ears, interfemoral, fur, colour, dimen-

sions) is that of an ordinary A. j. jamaicensis; forearm 59,

tibia 21, foot 17 mm.; no description nor measurements of

the skull.—The types were re-examined by Rehn, in 19007,

29 skulls, from all the localities

* US. N. M. nos.:—8671 (9387), 11187, 13220 (37912), 14305, 14410, 14412-13,

14415, 14753 (37811), 16332 (23360), 53063, 6148283, 70431, 70153, 73225, 73265,

86280, 86282-84, 86310-15, 86349, 86352-53, 86392-93, 96182, 100192, 100199,

100202, 100204, 102458, 108025, 108232, 108234, 108236, 108238-39, 108242, 110939,

112122, 112124, 112131-34, 112136-37, 113442, 113927-29, 114038, 122430-32,

133042-45, 133050, 133052-54, 147135-36.

+ James A.G. Rehn, “ Notes on Chiroptera,” Proc. Ac. Nat. Sci. Philad., Dec. 1900,

pp. 758-59 (9 Feb. 1901).

268 DR. KNUD ANDERSEN ON BATS. (Apr. 7,

who, on extracting the skull of one of the specimens, found it to

have 3 molars; thus the only difference between D. eva and

A. 7. jamaicensis to be derived from Cope’s description proves to

be imaginary. According to Rehn, the skull measures 27-6 mm.

(total length), the zygomatic width 17, the forearm 62, tibia 24,

foot 14, measurements which fall completely within the limits of

A.J. jamaicensis. Cope found the interfemoral ‘“ notched to a

line opposite the middle of the tibia”; Rehn writes that it is

‘‘of much greater expanse than either [in] perspicillatus or

planirostris,’ and this is evidently his only reason to keep

“D), eva” separate; but there is in this respect a very great

individual variation: in 7 spirit-specimens of A. 7. jamaicensis

from Jamaica the length of the interfemoral, measured in the

middle line, varies between 11:2 and 19 mm. ; a similar variation

is found in specimens of A. j. jamaicensis from other places, as well

as in other races of the species. A. j. jamaicensis occurs west

of St. Martins (Porto Rico) and east of St. Martins (St. Kitts),

so that also from this point of view there is not much proba-

bility that St. Martins individuals differ from the true 4. j.

jamaicensis.

J. A. Allen’s 4. coryi, 1890.—Based on a single specimen, taken

on St. Andrew’s Island, Caribbean Sea, Feb. 12, 1887, by one of

My. C. B. Cory’s collectors. It was described by Allen (J. s. c.)

as a distinct species on account of its colour :—‘‘ General color

above dark seal brown (brownish black), but very little lighter

at the base than at the surface; no trace of white streaks on the

face.” Forearm ‘50:1 mm.” (sic); third metacarpal “56 mm.”

(or ‘55 mm.,” see p. 171 of Allen’s paper).

There are three skins of ‘A. coryi” in the British Museum,

from the same island as the type specimen, taken by the same

collector, on the same date. Their coloration is as described by

Allen, but on examination of their teeth they prove to be

youngish, though full-grown, individuals (teeth perfectly un-

worn); this accounts sufficiently for their dark coloration, which

is indistinguishable from that of youngish specimens of A. 7

jamaicensis from Jamaica, Porto Rico, 8. Mexico, &e. (and of

any other race of A. jamaicensis, the South American forms not

excluded). The absence of facial stripes is due, partly to the

fact that St. Andrew’s individuals belong to the northern group

of races of A. jamaicensis, in which these stripes are generally

wanting or but faintly developed; partly to the fact that the

individuals obtained by Cory are young adults, in which the

facial stripes are generally wanting or still more faintly indicated

than in specimens of more advanced age.—Allen’s measurement

of the forearm, viz. 50-1 mm., is incorrect, and the explanation of

the error is no doubt this: im the three British Museum speci-

mens the proximal part of the radius has been cut away by the

taxidermist ; presumably the same is the case in Allen’s specimen,

since it is from the same collector and place; the true length of

the forearm, judging from the length of the metacarpals, would

be about 60 mm., as in an average dA. j, jamaicensis.—On p. 171

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270 DR. KNUD ANDERSEN ON BATS. | Apr. 7,

of the paper quoted above, Allen writes that “A. coryi” “ differs

greatly from all others fall other forms of the ‘ perspicillatus’

section] in colour and in the distribution of the fur on the

forearms, which on the dorsal surface extends densely in a broad

band along the humeri and over about the proximal third of the

forearm bones.” There must be some mistake here ; the distribu-

tion of the fur on the forearm in St. Andrew's specimens is as

described by Allen, but so it 1s also in all other individuals of

A. 9. gamaicensis, from any place, I have seen, and, in fact, in all

other races of A. 7. jamaicensis.

J. A. Allen’s A. insularis, 1904.—Type locality: St. Kitts, West

Indies. ‘‘ Based on a single alcoholic specimen, a very old male,

in excellent preservation, but with the skull badly broken. In

external measurements it agrees very well with A. jamaicensis,

..... The skull, however, in A. insularis is much larger than

in A. jamaicensis, the width across m*’-m° being 1 mm. (about

one tenth) greater, and the rostruin at the base of the canines is

also a millimetre wider, or about one eighth wider. This indi-

cates a much larger and more massive skull than in 4. jamai-

censis, while the external measurements are about the same.”

“ Dorsal surface pale reddish brown, paler below; membranes

pale brown; no head stripes.” Width of skull across m® 13;

width at base of canines 8°6; upper tooth-row (c-m’) 11; lower

tooth-row 11; forearm 61; third metacarpal 58 mm.

The specimen described by Allen is, in certain respects, a little

above the average size, but all the measurements recorded fall

quite within the limits of individual variation reached by 4. j.

jamaicensis; the specimen is matched, or surpassed, by a good

number of A. 7. jamaicensis from many different places. The

measurement of the lower tooth-row must have been taken from

the front of the canine to the back of m,, not to the back

of m, (or it would be greater than, not the same as, that of

the upper row, from ¢ to m*). The measurement of the second

phalanx of the third digit, stated to be 40 mm., is obviously

wrong. ‘The light colour of the fur is due to the specimen being,

as said by Allen, “‘ very old.” The absence of facial stripes is a

common feature in A. j. jamaicensis.— An example from St. Kitts

examined by me (U.S. N. M. no. 110939) is in every respect

(external characters, skull, teeth) indistinguishable from 4. 7.

jamaicensis ; it Shows no approximation to A. 7. preceps.

ARTIBEUS JAMAICENSIS ZQUATORIALTIS, K. And.

1906. Artibeus jamaicensis equatorialis Knud Andersen, Ann. & Mag. N. H. (7)

xviil. p. 421 (1 Dec. 1906).—T ype locality: Zaruma, $. Ecuador.

Diagnosis.—Similar to A. j. jamnaicensis, but skull, teeth, and

external dimensions averaging somewhat larger. Total length

of skull, to front of canines, 28-8-30 mm. (average 29-7); zygo-

matic width 18-2-18°5 mm. (ave rage 18:3); maxillary tooth-row

10:8-11:2 mm. (11:0); forearm 60: 8-66 mm. (62°9).

1908. | DR. KNUD ANDERSEN ON BATS. 271

General characters.—The angular emargination in the posterior

margin of m°, between cusps 5 and 7, is generally as well marked

as in any A. planirostris, sometimes reduced in size or almost

obliterated. The coloration of the fur is, in the only adult

individual available with quite unworn teeth, of the dark type, in

more aged specimens often light. The supraorbital stripes are

sometimes well marked ; the infraorbital stripes in all specimens

examined undeveloped or faint.

A. Jj. equaterialis and jamaicensis.—The skull of A. j. equa-

torialis averages larger, and especially broader, than in A. /.

jamaicensis, the zygomatic width being 18°3 mm., as against 17-4

in jamaicensis, the maxillary width (externally, across m'‘) 13°6,

against 12°6, the width across the cingulum of the upper canines

8-9, against 8:1; the teeth are a little larger, the maxillary

tooth-row being 11:0 mm., against 10:3. In all these respects

A. 7. equatorialis is as much larger than A. 7. jamaicensis as this

Measurements of A. }. jamaicensis and equatorialis,

A. j. gamaicensis. A. j. equatorialis.

95 adults, | 7 adults,

65 skulls. 6 skulls.

Min. Max. Mep.| Min. Max. Men.

| mm. mm mm.|/mm. mm. mm.

Skull, total length, to front ofe ...; 27 29:2 28°3 | 28:8 30 29°7

SeeTN AS holla Ohh ese eee | 14-2 1671 15:1 | 15:2 16°2 15°8

>» width of brain-case ......... | 12 ie LO NALD SP weiss 131

» zygomatic width ..........| 162). 18:2 174|182 185 18°3

» maxillary width across m!) 12 13°7 12°76 | 133 14 13°6

across cingula of canines. Pi 9 Sai esis 9-2 89

Mandible, to front of ine. 188 20:7 IGS |) BOB “BIAS | API

Upper teeth, can? ese OG lil 10°33 | 10°8 = =11°2 11

Lower teeth, c-m, ........ LOS UES AS | UES 12:6 1271

_ Ear-conch, length, Inner margin .| 13°8 16 14°8 | 15 15°8

3 length, outer margin | 19°7 22:2 21:3 | 22°7 9 23°5

ee ridt herent. ee 1135 16 143|148 15:7

TURNS, STEIN coocpsasoooseeiber cae cou), © ail 7 T2 75

iiancelmcnatha ee ee ree ee rs ORT O.S 9:61". O;erat at

ma 8S eric g lee Mckee esto | RHC 78 68) 74 75

| Horseshoe, width esl OS 85 76) 85 9

THORS .35 nonsschascooosenorcoplooness|| Gm (5 60:1 | 608 66 62°9

Wola xcs \Siyecy: eee re eee nea ealted 157 14:2 | 14 15°2 14-4

| 3rd metacarpal .. So dsaaecack cae ieoOs 61 53°7 | 5455 5 9°5 569

1 Ud Os arte Rh Saeed cpenoaccecd ial uaiteiy mmeAe) Iy/3) j| lee EB} 18°4

Ne) Gl bl erence soca sino Sid mena ces actalt 2740) 32°8 28°8 | 303 33 30°9

11D RAE pes oe es Remo Soccooedancalmuleriin) nillyfte; 155 | 162 178 16°9

4th metacarpal. geet tere AO 59°8 52:9'| 53:7 57-2 55°4

| IV2... ee ee ene else) ly 2! LONI S MMe eomee) 166

| IIW2re she Se eee tee RELL 215 19 198 22 19°9

| 5th metacarpal .. Nepean ie aera Any Oilh 62°5 547 | 545 59°7 57-2

Vile i as oe BER RS ee ee ee OS TS 11°9 | 12 13°2 12°8

Vv? . Ber Eee nccnensocnonacl lian me 145 | 142 17:2 1574

Interfemoral .. coset ere led 19 LSromielmomd So

Lower leg ........ Me tases St ball elk 24:2 22°38 | 22°38 26 24c1

Foot, with claws‘! coleeee ee NAS e el 27) 1671 | 16 17 16°8

Galcare en fet 78 Gi5y aed 8 72

@ |

Proc, Zoot. “Soo.1908, No. XVIII. l

272 DR KNUD ANDERSEN ON BATS. | Apr. 7,

latter is larger than A. 7. parvipes. The forearm and metacarpals

average 2°5 to 3 mm. longer than in A. 7. jamarcensis ; thus, also

in external dimensions it bears much the same relation to A. ).

jamaicensis and yucatanicus, as these latter to A. j. parvipes.

For further details see the tables of measurements pp. 271 and

284.

Geographically there is, in the series of jamaicensis and equa-

torialis available, a break between the former and the latter. I

have had no specimen of jamaicensis from any place south of

Panama (Colon), and none of equatorialis north of 8. Colombia

(Cali); but there cannot be much doubt that further investiga-

tions will fill up this gap, so that there, also geographically, will

prove to be a perfect transition between 4.7. yamaicensis and its

southern representative, 4.7. equatorialis.

Specimens examined.—9 specimens (8 skins) and 8 skulls, from

the following localities :—

British Museum :—Zaruma, 8. Ecuador (6). Cali, 8. Colombia

(2).—7 skulls, representing both localities.

U.S. National Museum * :—Corondelet, N. Ecuador (1, with

skull).

Range.—As yet known only from the region between Zaruma,

S. Ecuador, and Cali, S. Colombia.

ARTIBEUS JAMAICENSIS LItuRATUS Licht.

1801. Chauve-souris premicre ou Chauve-sowris obscure et rayée, Azara, Hssais

sur histoire naturelle des Quadrupédes de la province du Paraguay, ii.

pp. 269-70.— Paraguay.

1815. Phyllostomus lituratus Mliger, Abh. Akad. Berlin, 28 Feb. 1811 (issued

1815), p. 109—Nomen nudum.

1823. Phyllostomus lituratus Ill., Lichtenstein, Verz. Doubl. Berlin. Mus. p. 3—

Brazil.

1826. ? Phyllostoma superciliatum Wied, Beitr. Naturg. Bras. ii. pp. 200-202.—

Type locality; Rio de Janeiro.

1830. Phyllostoma superciliatum Wied, Rengger, Naturg. Saugeth. Paraguay,

pp. 74-75.—Paraguay.

1840. Phyllostoma perspicillatum (partim, not L.) Wagner, Schreber’s Saugthiere,

Suppl. i. pp. 403-5.—Spix’s type of “Ph. planirostre” compared with

Spix’s specimens of A. 7. lituratus.

1878. Artibeus perspicillatus (partim, not L.) Dobson, Cat. Chir. Brit. Mus.

pp. 519-20.—Brazil, Bolivia, Colombia.

1878. Artibeus grandis Dobson (from Gray’s MS.), op. cit. p. 250, specimen &1.—

Nomen nudum.

1892. Artobius perspicillatus (not L.) Winge, Jordfundne og nulevende Flagermus

fra Lagoa Santa, Minas Geraes, Brasilien; E Museo Lundii, ii. pt. 1, p. 10,

pl. 1. fig. 13.—Minas Geraes.

1900. Artibeus lituratus Licht., Thomas, Ann. Mus. Civ. Genova (2) xx. p. 547

(4 July, 1900).—Parana.

1901. Artibeus jamaicensis Leach, Thomas, Ann. & Mag. N. H. (7) viii. p. 192

(Sept. 1901).—Para.

1901. Artibeus lituratus Licht., Thomas, Ann. & Mag. N. H. (7) viii. p. 441 (Nov.

1901).—Paraguay.

1904. Artibeus rusbyi J. A. Allen, Bull. Am. Mus. N. H. xx. Art.20, pp. 230-31 (29

June, 1904).—Type locality: Yungas, “ Peru ”’ (probably aslip for Bolivia).

Diagnosis.—In all essential respects similar to A. 7. jamaicensis,

but considerably larger. Total length of skull 30°5-34 mm.

* U.S. N. M. no. 113363.

1908. | DR. KNUD ANDERSEN ON BATS. 273

(average 32:2); zygomatic width 18°8-20°2 mm. (average 19-4) ;

maxillary tooth-row 10°7-12°2 mm. (11°7); forearm 64-73:5 mm.

(70-2).

General characters.—This is the first of the three very closely

related southern races of A. jamaicensis described in this paper.

The southern group (4. j. lituratus, palmarum, preeceps) differs

from the northern (4. 7. parvipes, yucatanicus, jamaicensis, equa-

torialis) chiefly in the following points :—

The angular notch in the posterior margin of m*, between its

cusps 5 anil 7,18 rather rarely as well saanlzad as in A. plant-

rostris; generally (in about 90 p. ct. of 68 skulls examined) it is

either noticeably reduced or completely filled up; in the latter

case the bulk of m* has been increased by an area more or less

corresponding to that of the missing m*. By advancing age the

sagittal crest of the skull is produced considerably forward, the

supraorbital ridges directed almost straightly outward, the post-

orbital and anteorbital processes conspicuously developed,—as

described in detail and figured on p. 248. The dimensions are,

generally, in every respect larger, sometimes (especially in

A, j. lituratus and palmarum) considerably larger. The colour in

full-grown individuals with unworn or practically unworn teeth

is, most often, of the dark type, rather often, however, light; at

a higher age the majority of individuals are light-coloured. The

supraorbital stripes are, as a rule, well marked, often very strong,

rather rarely faint or obsolete ; the infraorbital stripes not rarely

well developed.

A.j. lituratus, jamaicensis, and wequatorialis.—Vhere is very

ravely any difficulty in discriminating A. 7. lituratus from A. 7.

jamaicensis. The skull averages almost 4 mm. (14 p. ct.) longer,

and 1:5-2 mm. (14 p. et.) broader, the tooth-rows 1:4-1:7 mm.

(about 14 p. ct.) longer than in A. 7. jamaicensis ; the forearm and

metacarpals average nearly 10 mm. (16-18 p. ct.) longer. Even

the very smallest individuals of 4. j. lituratus have, almost always,

the skull and teeth in some direction or other more heavily built

than in the largest individuals of A. 7. jamaicensis. When,

further, it is taken into consideration that there is absolutely no

overlapping of the geographical areas occupied by these two races, a

confusion becomes practically impossible.

A. j. equatorialis is generally a little larger than A. 7. jamai-

censis, and, consequently, comes a little nearer to A. j. anietibis

in size; but the average difference between cquatorialis and

lituratus is still very great, the absolute difference as a rule well

marked, A. 97. litur «tus extends into the area inhabited by

A. 7. equatorialis, but individuals which cannot be referred with

certainty to either the one or the other form seem, even in the

region common to both, to be very rare.

A, j. lituratus cannot be separated as a distinct species. First,

individuals do occur, though rarely, that cannot, by their characters

alone (7. e. apart from their habitat), be discriminated with cer-

tainty from some individuals of 4. 7. equatorialis and jamaicensis ;

13*

274 DR. KNUD ANDERSEN ON BATS. | Apr. 7,

second, A. 7. palmarum and preeceps completely overbridge the

gap (if gap it be called) between litwratus and jamaicensis.

On the difference between A. j. litwratus and palmarwm see

this latter form.

Specimens examined.—54 specimens (42 skins) and 42 skulls,

from the following localities :—

British Museum :—Paraguay: Sapucay (7); Asuncion (1).—

Sta. Catharina (1).—Parana: Morretes, Serra do Mar, 10 m. (5) ;

Palmeira (1).—S. Paulo: Cruzeiro, 530 m. (5).—Minas Geraes :

Sete Lagoas (1).—Bahia; Samariao, 300 m. (2).—Para (2).—

“ Brazil” or uncertain localities (6).—Colombia : Cali (4); Bogota

region, various places (4); ‘ Colombia” (2).—36 skulls, from all

the localities enumerated.

U.S. National Museum*: Paraguay: Sapucay and Villa Rica (9).

—Sta. Catharina (1).—N. Ecuador : Pambilar (1).—S. Colombia :

Cali (2).—6 skulls, representing all these localities.

Range.— Paraguay, Brazil, Ecuador, 8. and Central Colombia.

Linné’s Vespertilio perspicillatus, 1758 .—As pointed out by

Oldfield Thomas in 1892 and 1901 &, Linné’s V. perspicillatus was

based primarily on Seba’s “ Vespertilio Americanus vulgaris” §,

which is the bat commonly called Carollia brevicauda.

Schreber’s V. perspicillatus || is probably also Carollia; the

presence of a tail (“ganz in die Schwanzhaut eingeschlossen, und

so kurz, dass er noch nicht an die Hilfte ihrer Mitte reicht”’)

excludes at all events the genus Artzbeus.

Geoffroy’s Phyllostoma perspicillatum 4] from Guiana is either

A. planirostris or one of the large races of A. jamaicensis, which

of these two species cannot be decided from the description. The

species commonly brought by collectors from Guiana is 4. plani-

rostris; if Geoftroy’s bat is an A. jamaicensis, it is the only record

known to me of this species from Guiana. The figure has the

front of the horseshoe completely confluent with the integument

of the muzzle.

Wagner’s description of “ Phyllostoma perspicillatum ” (1840,

J. s. c.) was based on Spix’s type of Ph. planirostre and two speci-

mens of A. jamaicensis lituratus, all of which had been obtained

by Spix at Bahia and were by Wagner taken to be “ types” of

Ph. planirostre ; see pp. 238-239 of this paper.

Dobson’s Artibeus perspicillatus (1878, 1. s.c.) is the species

here called A. yamaicensis.

Azara’s Chauve-souris premiére, 1801.—In 1901**, Oldfield

* U.S. N. M. nos. 105587, 105625-26, 113362, 113364-65, 115065, 121445, 121447,

121466-67, 121469, 122139.

+ Syst. Nat. ed. 10, i. p. 31 (1758).

{ Thomas, P. Z. 8.1892, p. 315; Aun. & Mag. N. H. (7) viii. p. 192 (Sept. 1901).

§ Seba, Thesaurus, i. p. 90, pl. lv. fig. 2 (1734).

|| Schreber, Saéugthiere, i. pp. 160-61, pl. xlvi. a (1775).

{| Geoffroy, Ann. Mus. d’Hist. Nat. xv. pp. 176-77, 186, pl. xi, upper right figure

(1810).

** Thomas, Ann. & Mag. N. H. (7) viii. pp. 441, 443 (Noy. 1901).

1908. | DR. KNUD ANDERSEN ON BATS. 275

Thomas called attention to the identity of Azara’s “‘ Chauve-souris

premiere ou Chauve-souris obscure et rayée” with Lichtenstein’s

Phyllostomus lituratus. The essence of Azara’s description

(Azara did not see this bat, but copied the description from manu-

script notes by Noséda) is this: the “ Chauve-souris premiére ”

isa bat, (1) of large size (length 110 mm., expanse 525 mm.),

(2) with no tail, (3) with a white stripe from the nose to the ear,

and (4) inhabiting Paraguay. A. 7. litwratus is the only tailless

bat of this size known from Paraguay.

Lichtenstein’s Phyllostomus lituratus, 1823.—Based, without

description, on Azara’s ‘“‘ Chauve souris obscure et rayée.” As the

identity of Azara’s bat is unquestionable, the name litwratus is

technically valid.

Maximilian of Wied’s Phyllostoma superciliatwm, 1826.—Based

on a single specimen found dead and much decomposed “ bei der

Fazenda von Tapebugt . . . . nordlich von Cabo Frio zwischen den

Fliissen 8. Joao und Macahé,” 7. e. in the province of Rio de

Janeiro. The author placed this bat (‘“‘ Der Vampyr mit weissem

Augenstreif”) in a section of ‘“ Phyllostoma” headed “ Unbe-

stimmte Arten, deren Gebiss nicht untersucht werden konnte,

welches aber wahrscheinlich mit dem der vorhergehenden Ab-

theilung tibereinstimmt ” ; the preceding section has 2 ‘‘ Backen-

zihne” (i.e. 3 molars). The statement is, of course, without any

value, as the author admits it to be mere conjecture. In the rest

of the extremely vague description there is nothing which prevents

the identification of Ph. superciliatum with A. 9. lituratus.

Rengger’s “ Ph. superciliatuwm Wied,” from “Jhu,”’ Paraguay,

is no doubt 4.7. lituratus. His specimen had no ‘‘weisse Spitzen

der Fliigelhaut”; but the amount of white at the tip of the wing

(region of third, or second and third, phalanx of third digit) is

individually very variable; examples occur in which it is but

slightly indicated (very restricted, and clouded with dark colour).

Dobson’s (Gray’s) A. grandis, 1878.—The name “ Artibeus

grandis,” quoted in Dobson’s Catalogue from a manuscript label

by Gray, but apparently never published by this latter author,

refers to an unregistered spirit-specimen in the British Museum,

an adult male without history; maxillary tooth-row 12 mm.,

forearm 70°38 mm. The specimen is indistinguishable from an

ordinary A. j. hituratus.

J. A. Allen’s A. rusbyi, 1904.— Based on the skin and skull of an

individual from Yungas, Bolivia *, 6000 feet. A. rusbyi is stated,

by Allen, to be “nearly related to 4. palmarum of Trinidad and

adjoining parts of north-eastern South America, but differs in the

* Allen writes, “Yungas, Peru” [%.e. the coast region round Trujillo, N.W.

Peru]; but the specimen made by Allen, in 1904, the type of A. rusbyi had been

mentioned, by the same author, on two previous occasions (Bull. Am. Mus. N. H.

ii. (1890) pp. 170 & 172, and ix. (1897) p. 16) as beg from “ Yungas, Bolivia”

[i. e. the eastern slopes of the Bolivian Cordilleras to the Amazonian plains]. I take

the latter to he correct, but whether my conjecture is right or wrong is, for the

identification of A. rusbyi, of no consequence.

276 DR. KNUD ANDERSEN ON BATS. [ Apr. 7,

lower, less convex, and more spreading brain-case, broader palate

and heavier dentition, the upper tooth-row (canine and molar—

premolar series) having a length of 11-5 mm. against 10-2 in

A. palmarum; also in more prominent face stripes and darker

coloration.” Forearm 71 mm. ; third metacarpal 69 mm.

IT have had no adult specimen of the Artibews jamaicensis

section from Bolivia*, but nine from Ecuador, S. and Central

Colombia; they accord in every respect with Allen’s description

of A. rusbyi, being at the same time perfectly indistinguishable

from A. 7. ditwratus. Allen gives as length of the maxillary tooth-

row in his single specimen of 4. rusbyi 11-5 mm., as against 10-2

in “ dA. palmarum”; in so far he had good reason to separate the

former. But in 6 skulls of adult individuals of 4. j. lituratus

from the type locality, viz. Paraguay, the average length of the

maxillary tooth-row is 11°6 mm.; in 19 adult skulls of the same

race from various localities in Brazil 11-7 mm.; thus precisely as

in the type of A. rwsbyi.—The explanation, why Allen regarded

A. rusbyi as an undescribed species, is evidently this: Allen

compared d. rushby: not with a series of A. j. lituratus from

Paraguay or Brazil, but with an wnewsually small-toothed A. J.

palmarum. I have examined 28 skulls of A. 7. palmarwm, and

10°2 mm., given by Allen as the length of the upper tooth-row

in this form, is precisely the actual minimum in the whole series.

Allen found the skull of dA. rusbyi “lower” (less convex, &c.)

than in palmarwm; it only means that the skull of the type

of A. rusbyi is in the adolescent stage, whereas the skull (or

skulls) of palmarwm with which he compared it must have been

in the stage characteristic of adult and aged individuals; that

these differences in the shape of the skull are dependent on

age has been pointed out elsewhere in this paper (p. 248, text-

figs. 50, 51). The colour characters mentioned by Allen are

of no taxonomic value; individuals of lituratus and palmarum

may be dark or light, their facial stripes strong or obsolete ;

these variations are partly dependent on age, partly purely

individual.

In the table of measurements below, p. 277, I have arranged

the adult specimens of 4. j. litwratus in three sections according

to their geographical habitat, viz. Paraguay, Brazil, and Ecuador—

Central Colombia. In each of these sections are given the abso-

lute minimum and maximum and the average, chiefly in order to

show that individuals from these three regions are in every

respect identical in size, 7. e. vary within the same limits. It

will be noticed that the average measurements of skulls (including

teeth) from the three regions are perfectly alike; that averages

of external measurements of individuals from Paraguay and

Brazil are alike; and that the actual measurements of skulls,

teeth, and external dimensions, within all these categories are

* Specimen “7!” in Dobson’s Catalogue (p. 520), stated to be from Bolivia and

“adult,” is quite young; the correctness of the locality is questionable.

1908. ] DR. KNUD ANDERSEN ON BATS. 277

practically alike, when due consideration is taken of the fact that

the number of individuals examined from each region is not the

same. The external average measurements of the nine individuals

from Kceuador, 8. and Central Colombia, are somewhat larger

than in the two other sections, but there is no reasonable doubt

that this is quite accidental; in Paraguay individuals the dif-

ference between the absolute maximum and minimum of the

length of the forearm is 9°5 mm., in the still larger Brazilian

series 11 mm.; so that it is not very likely that in individuals

from Ecuador and Colombia the amount of variation in this respect

should be 4-5 mm. only; all of these latter individuals evidently

happen to be rather large, none of them representing the mini-

mum size, and consequently the average, as calculated from this

series, is also above the true normal.

Measurements of Artibeus jamaicensis lituratus.

| Paraguay. Brazil Ecuador,

(detailed localities | S. & C. Colombia.

see p. 274). |

12 adults, 20 adults, 9 adults,

6 skulls. 19 skulls. | 8 skulls.

SAE EUE a Sep ee ne

|Min. Max. Mep. Min. Max. MeEp.| Min. Max. Mep.

mm. mm. mm. mm mm. mm.)mm mm mm

Skull, totallength, totrontofe...) 315 33 3271 | 30:5 34 3271 | 3p BS 32

» mastoid width ...............)163 172 169/168 18 173/168 175 173

PMauthrotbrain-cascess saa loon milan slag ssi sls 14: hy 138 148 14

» zZygomatic_width ............1188 20 192/188 202 194/195 196 19:4

5 maxillary width across m!/1382 147 141/136 152 145/133 °142 13:9

across cingula of canines. 86 9-2 9h S:Seiaao 93 | 87 97 9

Mandible, to front ot ine. 2G) 23s | DPB GA, DBI. 227 | 22 29°3 22-9

Upper teeth, c-m? ee esx ele LSC LOS ae WS 7 2) et 115

Lower teeth, Cams es Tiles} als} IG} ) These IB ey aR) ae 13 12°5

Zar-conch, length, inner margin Ge ls Ilyas) |) ess ily}

33 length, outer margin | 23 26 24 | 225 24

° width Siete tye mGaM ise ay |

AB RTS, HEWEUN cost oscscpeseccosnsoswool|. 2 88 78 | ie, 78

Manceta) lene: bhi. sees me nearest 12 ES O83. elles |

pee) WAC DIN 9 oe cere ne | ed 9 81] 7 85 |

Horseshoe, width | 82 9 88) 8 9°5 |

Forearm .| 64 73:5 70-4) 65 76 70A| 70:5) 75 73'3

Pollex pcan eee hee el Gio EO NIE ES Si alls 16°6| 142 17 16°4

srdimetacanpal.......)-44--2"..-..| 001) 167m» 63:9 i582 69 63:7) 655 682 665

dG) (0 EL heed es oan a tena RSyO Ay tele SNe Mii es B75) 21°8) 22 245 23:2

88 30°38 40 361 | 325 41 355 365 = AL 38°5

NEL (Gs Pee Seo Bee earn eee eee aeneel| lis) Dili lO eG Sauces) ils Osels 21 19°3

4th metacarpal... Ga dopiondianass sasse) DD 667 6295 | 568 67 62°5 | 63 67 65°2

NP ce ascaoeee Hel ke ee 21 18°6 | 17 20:3 «19 |S Bal il }er/

YE) POE TeS ore ty ts ree eee, omen ee 111) 24 29°5 | 198 252 22°3| 22:2 2a 23°9

sth metacarpal... 57 70 65:1 | 58 70 644.) 65 70 67:9

127 158 144) 13 16 144; 142 16 166

v2. 15 LOS 7-H eS Orie AN EViEA, 20 18°8

Interfemoral .. Nn aster erell besten + rik sven #740) 15 20 |

NGG Werpl Comer cee conse oye ee OD 78 262 | 24-3 28 269 | 26 27 26°7

Foot, with claws ... Renee oe ALAS inne) Osetia Steyn nel li 20 1825 117-2) 20 18°3

Calcar Ueda Sot ty CRS NSD MIR ah 87) 10 10 85|/ 8 10 89

88 | 78

278 DR. KNUD ANDERSEN ON BATS. [Apr. 7,

ARTIBEUS JAMAICENSIS PALMARUM All. & Chapm.

1878. Artibeus perspicillatus (partim, not L.) Dobson, Cat. Chir. Brit. Mus.

pp. 519-20, specimens », p, 7, s, t, v, y, 2—Venezuela, Costa Rica,

Guatemala.

1893. Artibeus fe spicillatus (not L.) Thomas, Journ. Trmidad Field Nat. Club,

i. no. 7, p. 6 (April 1893).—Trinidad.

1893. Ar ace sp. n., J. A. Allen & Chapman, Bull. Am. Mus. N. H. v. Art. 13,

208 (21 Sept. 1893).—Tiimidad.

1897. Uapribeis palmarum J. A. Allen & Chapman, Bull. Am. Mus. N. H. ix.

Art. 2, p. 16 (26 Keb. 1897).—Type locality : Trinidad.

1897. Artibeus intermedius J. A. Allen, Bull. Am. Mus. N.H. ix. Art. 3, pp. 33-

34 (11 March, 1897).—Type locality : San José, Costa Rica.

1899. Artibeus femur villosum Outram Bangs, Proce. New Engl. Zool. Club, 1.

pp. 73-74 (24 Nov. 1899).—Type locality: Santa Marta, Colombia.

1900. Artibeus palmarum All. & Chapm., J. A. Allen, Bull. Am. Mus. N. H. xiii.

Art. 8, p. 89 (12 May, 1900) Santa Marta, Colombia.

1901. Artibeus palmarum All. & Chapm., Robinson & Lyon, Proc. U.S. Nat. Mus.

xxiv. pp. 148-49.—La Guaira, Venezuela (specimens examined).

1902. Artibeus intermedius All. (partim), Outram Bangs, Bull. Mus. Comp. Zool.

xxxix. no. 2, p. 50 (April 1902).—Bogava, Chiriqui (the larger specimen

recorded by the author).

Diagnosis.—Precisely similar te A. 7. litwratus, but skull and

teeth averaging slightly smaller.

General char vacters.— As in A. J. lituratus the notch in the

hinder margin of m® is comparatively rarely (in 10 p. ct. of the

skulls examined) as distinct as in A. planirostris, in the large

majority either noticeably reduced in size or quite filled up. The

skull is subject to the same modifications depending on age (text-

figs. 50, 51,0n p. 248). The light colour type is predominant,

especially i in aged individuals, but common also in adults with

unworn teeth. The supraorbital stripes are as a rule, the infra-

orbital stripes often, well developed.

A. }. palmarum and litwratws— Externally A. 7. palmarum is

indistinguishable from dA. j. litwratus; there is no structural

difference in any respect; the dimensions vary within the same

limits, and even the average dimensions are practically quite the

same. But there is a small average difference in the size of the

skull and teeth :—the length of the skull averages 1:3 mm. (only

about 4 p.ct.!) shorter, dhe width of the skull from 0°4 to 0-9 mm.

narrower, the maxillary tooth-row 0-7 mm. shorter.

I should not have tried to keep this form separate from 4d. 7.

lituratus, if the name palmarwm had not been available. The

separation is artificial rather than natural; the trifling average

difference pointed out above will, I believe, hold good also for

much more extensive series of both forms; but it is a matter of

fact that in the large majority of cases A. 7. palmarwm cannot

practically be discriminated from litwratus.

A. j. lituratus and palmarum taken together (and as mentioned

above it would be both more natural and more convenient to

unite the two ‘“ forms”) are distributed from S. Brazil to S. Mexico,

including the coast islands of Trinidad and St. Vincent, but

excluding the whole of the West Indies proper.

In Central America and 8. Mexico A. j. palmarum meets the

considerably smaller 4.4. jamaicensis. There is no doubt what-

1908. | DR. KNUD ANDERSEN ON BATS. 279

ever that the latter race is the truly indigenous form in the

region north of Panama, and that A. 7. palmarwm is a late

intruder from south into the same region. This point will be

further discussed in the ‘General Remarks,” below p. 317.

Size.—In the table of measurements p. 282, the specimens

examined have been arranged in four sections, viz. individuals

from (1) Venezuela, (2) Trinidad and St. Vincent Islands,

(3) Central America, and (4) 8. Mexico. The table shows that

measurements of individuals from these four areas are identical.

The four specimens from 8. Mexico (Oaxaca, Vera Cruz, Jalisco)

do not show the extremes of individual variation in size; hence

the average measurements are not given.

Specimens examined.—47 specimens (26 skins) and 32. skulls,

from the foliowing localities :—

British Museum :—Trinidad (4)—St. Vincent Island (1).—

Venezuela : Caripé (1) ; Tachira (1); ‘“‘ Venezuela” (1).— Panama :

Bogava, Chiriqui, 250 m. (4).—Costa Rica: San José (3); Los

Cuadros, 8. Pedro (1); Costa Rica (2).—Nicaragua : Matagalpa

1).—Guatemala : Duefas (4); Cahaban (1).—“ Central America”

(1).—Jalisco: Huajimic, Tepic (1).——18 skulls, from all the localities

enumerated.

U.S. National Museum* :—Trinidad (1).— Venezuela : Macuto,

La Guaira (12).—Costa Rica: San José (3),—Nicaragua : Escon-

dido River, 50 miles from Bluefields (1).—Guatemala: Peten (1).

—Oaxaca: Santo Domingo (2).-—Vera Cruz: Mirador (1).—14

skulls, representing all these localities.

Range.— Venezuela, including Trinidad and St. Vincent Islands,

through Central America, to Vera Cruz and Jalisco, Mexico.

Allen and Chapman’s A. palmarwm, 1897.—In 1898 (J. s.c.),

Allen and Chapman recorded an “Artibeus sp. nov.?” (skin with-

out skull) from Trinidad ; it differed ‘‘in coloration and in the

distribution of the fur on the wing-membranes from any of

the currently recognised species of Artibeus ;’ forearm 63, third

metacarpal 61, tibia 25-4 mm.; ‘‘color above and below light

brown, much lighter on the head and anterior half of the body,

the hairs nowhere tipped with gray; a broad white stripe above

and a faint whitish line below each eye.”

The same specimen, together with six others, also from Trinidad,

formed, in 1897 (/.s.c.), the basis for Allen and Chapman’s

A, palmarum :—‘“‘ From true Artibeus perspicillatus,” the authors

write, “the present species differs notably in colour, particularly

in the presence of two prominent broad white head stripes, and

two narrower and shorter whitish cheek stripes. It is also very

much larger, the forearm measuring 68 mm. against 56 in true

perspicillatus, with all the other dimensions proportionately

larger. The skull is much more massive, at least one-third

* U.S. N. M. nos. :—6973 (37809), 7222, 13778, 13809 (87551), 51571, 73257-58,

101331, 102843, 102845-46, 102855-56, 102860-62, 102873-74, 102877, 102879,

103964.

2806 DR. KNUD ANDERSEN ON BATS. [ Apr. 7,

larger in general bulk, and about one-sixth larger in linear

measurements.” Forearm of type 68, third metacarpal 64;

length of skull 31, zygomatic width 19 mm.

Later on, in 1900 (J. s.c.), d. palmarwm was recorded by Allen

from Bonda, Santa Marta region, Colombia, and Cali, 8. Colombia;

in 1901 (@.s.c¢.), by Robinson and Lyon from La Guaira,

Venezuela.

To understand what led Allen and Chapman to separate

A, palmarum as a distinct species the following must be borne in

mind :—Allen and Chapman identified Linné’s Vespertilio per-

spicillatus (1758) with the bat later on (1822) desevibed by Leach,

on the basis of a Jamaica specimen, as Artibews jamaicensis

(A. 7. jamaicensis of the present paper)*; having found that the

Trinidad representative of this type of bat differed [to a certain

degree] in size and colour from the Jamaica bat and, consequently,

required a name of its own, they called the Trinidad form

A. palmaruwm. In so far all is lea: the Trinidad bat is, in fact,

as a rule distinguishable from the West Indian form. But Allen

and Chapman were mistaken in their identification of Linne’s

V. perspicillatus (Seba’s V. Americanus vulgaris), which, as

pointed out by Oldfield Thomas, is not Leach’s A. jamaicensis,

but the bat commonly called Carollia brevicauda; further,

although there is a very well-marked average difference in

size and colour between Trinidad and Jamaica individuals of

A. jamaicensis, there is absolutely no “ hard-and-fast” line

between them, so that they cannot be separated specifically ;

again, the Trinidad (Venezuelan, Colombian) bat comes so ex-

ceedingly near to the common Brazilian form of the species that

it, for all practical purposes, is completely indistinguishable from

this latter; and, last, this Brazilian form had already a name,

viz. A. 7. lituratus Licht.—The infinitesimal average difference

in the size of the skull between 4. 9. lituratus and A.j. palmarum

is the only reason (if reason it can be properly called) on the

strength of which the latter can be kept separate as a “race.”

J. A. Allen’s A. intermedius, 1897.—Type locality: San José,

Costa Rica. According to Allen, A. intermedius is “ rather smaller

than dA. palmarum,” % apparently intermediate between dA. pal-

marum and A. perspicillatus fi. e. A. 7. jamaicensis |,” but “ much

darker, with the head stripes narrower and much less distinct,

and the cheek stripes obsolete” ; “‘ brain-case narrow and high, the

dorsal outline remarkably convex ;” forearm 65, third metacarpal

57; length of skull 29, zygomatic width 19 mm.

A. intermedius was based on two adult and five “ nearly full-

grown” young individuals. If Allen had had a larger series of

adults he would have found that the colour characters on which

he laid stress are of no diagnostic importance ; in Costa Rica, as

elsewhere throughout the whole area occupied by A. 7. lituratus

* Allen & Chapman, Bull. Am. Mus. N. H. ix. Art. i. pp. 3-5 (23 Feb. 1897).

1908. ] _DR. KNUD ANDERSEN ON BATS. 281

and palmarwm, some individuals are of the dark, others of the

light colour type, some have the superciliary stripes less distinct

or even obsolete, while others (and these the majority, in Costa

Rica and elsewhere) have them well marked or even very strong ;

the infraorbital are always less pronounced than the supraorbital

stripes; the colour characters given by Allen must therefore be

left out of consideration when judging the validity of “4. inter-

medius.”—When Allen found the type of 4. intermedius “ appa-

rently intermediate [in size] between 4. palmarum and A. per-

spicillatus,” it is only because the specimen happens to be a small-

sized individual ; in the series from the type locality and adjoining

regions of Central America examined by myself, there are several

examples as small as (and slightly smaller than) Allen’s specimen,

but these represent unquestionably the minima of size; the other

extreme 1s shown by the following measurements of the forearm :

in two specimens from Chiriqui 70°5 and 72 mm., two from Costa

Rica 71 and 73 mm., two from Nicaragua 71 and 73 mm., three

from Guatemala 70,72, and 72°5mm.; the rest of the individuals

are, of course, intermediate in size between these two extremes.

Thus, also the size-character given by Allen must be dropped.—

There remains the shape of the skull; but as pointed out above

(p. 248, text-figs. 50, 51), this “remarkably convex” brain-case ~

(sagittal crest produced forward, supraorbital ridges directed

almost straightly outward, well developed post- and anteorbital

processes) is an age character in A. j. litwratus and palmarum,

not a specific character ; it is by no means peculiar to Central

American individuals, but occurs, to the same degree, in individuals

from Paraguay, Brazil, Colombia, Venezuela, Trinidad, &e. I have

carefully compared Central American (and Mexican) specimens

with Trinidad and Venezuelan specimens of 4. 7. palmarum, and

am unable to find any difference whatever.

In 1902 (7. s. c.), Outram Bangs recorded three ‘4. intermedius”

from Bogava, Chiriqui, one old 2, one youngish ¢, and one

youngish 2, and wrote :—“ The younger specimens are more

sooty, with the facial stripes less well indicated, and have smaller

skulls .... The difference in size is great, and the skulls do not

show the degree of immaturity that one would expect with the

difference in size.” The explanation is this: the two youngish

specimens (provided they are really full-grown) were, no doubt,

A. J. jamaicensis, the old female an A. j. palmarum. It cannot

be too strongly emphasised that Central America and 8S. Mexico

are inhabited by two forms of A. jamaicensis—the one, and small,

is the truly indigenous race, 4. j. jamaicensis, occurring also in

Jamaica, San Domingo, Porto Rico, eastwards at least to St.

Kitts; the other, and larger, is A. j. palmarwm (or, if preferred,

A, j. lituratus), which has come from south,—two forms which

till now have been mixed together. As the two races are not

perfectly differentiated ‘“ species,” one cannot expect them to be

separable in all particuiar instances ; but many individuals can

be identified at a glance, and whenever the external characters

282 DR. KNUD ANDERSEN ON BATS. [ Apr. 7,

leave the identification unsettled, a careful examination of the

skull and teeth will, with rare exceptions, decide the matter. It

is acase parallel to that of Hipposiderus caffer caffer and H. c. cen-

tralis in British and German Hast Africa * ; two very well marked

modifications of one common type, differmg in size only, and

originally, no doubt, occupying quite separate areas, but the

latter has in the course of time spread over a part of the region

of the former, as has A. 9. palmarum over that of A. 7. jamaicensis;

in the one instance as in the other the races are, however, on

careful examination almost always separable, even when found

together on the same spot.

Outram Bangs’s A. femurvilloswm, 1899.—Type locality : La

* Knud Andersen, Ann. & Mag. N. H. (7) xvii. p. 281; March 1906.

Measurements of Artibeus jamaicensis palmarum.

| Venezuela Trinidad, | Central America | Mexico

(Macuto, Caripé, | St. Vincent (Panama,Costa Rica, (Vera Cruz,

Tachira). Island. Nicaragua, Jalisco,

Guatemala). Oaxaca).

| 15 adults, 4, adults, 20 adults, 4 adults,

10 skulls. 3 skulls. 15 skulls. 3 skulls.

| Min. Max. Mep.| Mry. Max. | Min. Max. Men.) Min. Max. |

| |

Skull, total length, to front of ¢ ...) 30 32 31 ee el 29°7 318 30°6 | 30 30°7 |

| 4, mastoid width ..................) 162 17:5 162 158 176 168 | 162 17

5, Width of brain-case............| 13 14 13°38 | 1386 145 14 13°8 14

| 5, “Zygomatic width . ne) LPI PAORS} 20 18 20 189} 188 19:2 |

/ 4, maxillary width across m}.. | 13 14 14 146 | 13 14 135 | 13:2 13°8

| across cingula of canines ...| 85 9 | fri @ ay Pil sy) 8s ®

| Mandible, to front of ine. Beep AOS) es) 22°2 | 20 22°6 21:4 | 20°77 21:2 |

| Upper teeth,c-m? ............|108 118 '47 118 -| 102 112 109] 103 117 |

| Lower teeth, c-m3 ....... cool) LLL LP 7 | 12 12°8 Wale ey le) ale) alles

| Ear-conch, length, imner margin ...| 16 17-7 laure aig 1155 173 167 165

35 lensth, outer margin ...| 21 24-5 23. 25 21 26 23°6 24-5

| a WAGGNs ek a eee 163 155 | 16 16°5 | 152 177 16 17

| IIEREAUS, METER ccaosocgaovenenmesaacasnces| dS SIONS 7 Bi5en| 077 8 79 a

(Abaneets length) ores sheer OO mm owen | Opie anil) pe le omlOs7 11:2

e width SRC eee eee | S38 7) 7H BS | ¥ 88 79 82

“Horseshoe, width | 82) OR OK 88 OOD | BB 8B 89

| Forearm 5 682 758 71 | 675 725 | 64 75 70:6 | 68 71

| Pollex Seat Ur cucg ine, Pack call W7 171) 152 175 | 15 168 161/157 15:7

| 3rd nnetacerpal 617 685 646 | 60 68 | 588 665 638 | 685 63 |

(Palelie he as 21 24°8 22°38 | 19:2 23:2 | 20 24 22°4 | 20 22°2

| II? 31 39°8 3864 | 33 38°2 33 39 36°7 | 33°7 358

_ I BO Re deEEEMen eT onnt cavene ase || Kone! 202 182/177 21 | 172 208 18°77 | 17 18°7

4th mnetecatpal | 597 665 633 585 66 | 58 662 62°6 | 58 615

TOV ee 17°8 21 191 175 19:2 | 172 «215 «19 175 =618'8

fu Nea ees ROM a Aa A et tcl tetena|| P40) 24:2 22°5 20°38 24 | 21 25°38 23:3 | 208 23

5th mctacarpaley es eke eed 617 69 65°8 618 685 | 595 673 645 | 59° 65

WRU Rr erree renee c 8 ac nctaree meee ML aKS 16 148 128 16 142 17 14°7 | 14 148

v2. Pee cee Oe By TEA IA NSS 20. TS 1G. 1s

Interfemoral.. | as eS} 17 22 19'6 | 15

Mowreriecge etree cats: ¥. seat [24 287 25: 1/25 27:5 | 24/268) 25:6)| 2918 24s

Foot, with claws ........................| 17 198 184 178 19 165 20 18 162 178

Calcar i We NO Sri 9H GP | 72 9 86 | 7:2

1908. | DR. KNUD ANDERSEN ON BATS. 283

Concepcion, Sierra Nevada de Santa Marta, Colombia, 3000 feet.

—The essential points in the original description are these :—

“ About the size of A. palmarum, differing from that species in

having but one pair of face stripes (no cheek stripes) .... these

stripes narrow and not very conspicuous. The new species differs

from all others of the genus I have seen in having the upper

surfaces of legs, feet, and interfemoral membrane clothed with

short fur. In the allied species these parts are naked.” Length

of forearm, metacarpals and phalanges not given by the author ;

it is unnecessary to quote the measurements of the skulls, as they

are precisely as in many 4. 7. palmarwm.—A. femurvillosum

was half a year later (1900, J. s. c.) put down by Allen as a

synonym of ‘A. palmarwm.”

The colour characters given by Bangs need no comment; they

are valueless for diagnostic purposes. The statement that the

upper surfaces of the legs, feet, and interfemoral are naked in

“the allied species,’ is a mistake; in all the specimens I have

seen of A. jamaicensis, of any race, they are hairy. Thus

nothing is left by which A. femurvillosum can be discriminated

from 4. 7. palmarum.

ARTIBEUS JAMAICENSIS PR@CEPS K. And.

1906. Artibeus Jamaicensis preceps Knud Andersen, Ann. & Mag. N. H. (7) xviii.

p. 421 (1 Dec. 1906).—T ype locality ; Guadeloupe, W. I.

Diagnosis.—Similar to A. 7. palmarum, but forearm and hand

averaging shorter.

j. preceps and palmarwm.—tindividuals of A. jamaicensis

from Trinidad and St. Vincent Island are indistinguishable from

the continental A. 7. palmarum (or, if this form is not recognised,

from A. 7. litwratus). From the Windward Islands between St.

Vincent and Dominica I have had no specimens.—In Dominica

and Guadeloupe A. j. palmarum is replaced by the slightly dif-

fering A. j. preeceps. Three skulls of this race are practically

almost indistinguishable from the ordinary palmarwm skull ; they

show a tendency to go slighty below the minimum size in the

large number of palmarwm skulls examined, so that there can

scarcely be any doubt that, in a more extensive series, skulls of

preceps will prove to average a little more slenderly built. The

teeth will probably also average slightly smaller. There is a

similar indication of a decrease in the external dimensions,

especially noticeable in the length of the forearm and hand: in

the three adult examples of A. j. praceps the forearm measures

60, 65:5, and 66:2 mm., in 43 adult examples of A. 7. palmarwm

the average length of the forearm is 70-9 mm., and none has the

forearm less than 64 mm.; of the whole series of palmarum two

only (5 p. ct.) have the forearm less than 66°5 mm., whereas in all

examples of preceps available the forearm falls short of that

length; for further details (metacarpals, proximal phalanges,

tibie) see table of measurements below, p. 284.—From the

[Apr. 7,

DR. KNUD ANDERSEN ON BATS.

284

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1908. } DR. KNUD ANDERSEN ON BATS. 285

islands between Guadeloupe and St. Kitts no specimens have

been available. St. Kitts is inhabited by 4.7. jamaicensis, the

range of which, so far as the West Indies are concerned, ex-

tends from here westwards over Porto Rico and San Domingo to

Jamaica.

A.j. preeceps comes in every respect considerably nearer to its

southern neighbour, 4. 7. palmarwm, than to its western neighbour,

A. 9. jamaicensis. From this it seems reasonable to conclude that

it is anorthern offshoot of A. 7. palmarwm, not an eastern offshoot

of A. 7. jammicensis.

It would be practically impossible to discriminate 4. 7. preceps

from 4d. j. equatorialis; but the latter is a large southern

representative of A. j. jamaicensis inhabiting EKeuador and §.

Colombia, the former a slightly diminished insular representative

of A. j. palmarum, inhabiting, as just pointed out, certain islands

between St. Vincent and St. Kitts. Their extremely close re-

semblance is a coincidence; the descent of the two races is

different, and they occupy widely separated areas; they have

acquired similar features, but by different lines of development.

Specimens examined.—Dominica (1), Guadeloupe (2). With

skulls. From the collection of the U.S. National Museum *.

Range.—As yet known only from Dominica and Guadeloupe,

W-.

ARTIBEUS GLAUCUS Thos.

1844. ? Phyllostoma pusillum (not Natterer) Tschudi, Fauna Peruana, pp. 63-64.

1893. Artibeus glaucus Thomas, P. Z. 8. 1893 (18 April) pp. 336-37, pl. xxix.

figs. 7-9.—Type locality: Chanchamayo, Peru.

Diaguosis.—Molars ?. Cusp 7 of m' small. Forearm 43:8 mm.

Skull.—In shape the skull is almost precisely similar to that

of A. hirsutus, planirostris, ov yamaicensis, the only appreciable

difference being the slightly lower brain-case; but it is much

smaller: in linear dimensions #, in bulk less than 4, the size of

an A. hirsutus skull. [

Teeth.—Difter from those of A. concolor, planirostris, and

hirsutus, and accord with those of A. jamaicensis, in the complete

disappearance of m*; differ from those of all the species mentioned

chiefly in the less developed lingual parts of the premolars and

molars (particularly of m*), and in the much smaller size of all

the teeth.

The inner heels of p* and p* are proportionately slightly smaller

than in the foregoing species. In all of these latter there is a

small, but perfectly distinct, cusp rising from the antero-internal

margin of the heel of p‘; in glawcus this cusp is practically

wanting (an exceedingly faint indication of the cusp is detect-

able by the aid of a strong lens). Cusp 7 of m'‘ (the postero-

internally projecting portion of the tooth) 7s considerably less

developed than in any of the foregoing species; in the single

skull available it is not much more than a distinctly projecting

* U.S. N. M, nos, 1135034, 113628,

286 DR. KNUD ANDERSEN ON BATS. [Apr. 7,

ledge, whereas in jamaicensis (and in concolor, planirostris, and

hirsutus) it is equal to from one-third to one-fifth the area of the

tooth. The angular emargination in the posterior border of m*

(into which m’* is pressed in those species which possess this

rudimentary tooth, and which, as pointed out above, pp. 250—252,

is very often preserved in A. jamaicensis, although this species

has lost m’*) is in glaweus but faintly indicated, the whole postero-

internal portion of m* being formed by the slightly projecting

cusp 7.—The lower teeth do not differ in structure from those of

the foregoing species; the small m, is still more reduced in size.

Text-fig. 53.

A. Artibeus glaucus, 9 ad. Chanchamayo, Peru. Type, B.M. 94.8.6.13.

Right upper tooth-row. xX 4.

B. Artibeus gamaicensis jamaicensis, § yg. ad. Jamaica. B.M. 7.1.1.677.

Right upper tooth-row. X 4.

Nose-leaves.—Front margin of horseshoe free, simple.

Tragus.—Two indistinct serrations on the outer margin above

the median projection ; ina larger series some individual variation

will be found in this respect.

Vings.—The third, fourth, and fifth digits are proportionately

longer than in the planirostris and jamaicensis sections; in

planirostris the indices of these digits (including the meta-

carpals, but excluding the terminal cartilaginous rods of the

distal phalanges) are, respectively, 1945, 1477, and 1346; in

glaucus and watsoni (which are similar in the wing-structure

as in almost all other respects), 2025, 1497, and 1405; as

proved by these figures, particularly the third and fifth

digits are lengthened*, to a less degree the fourth. <A

* “Tenethened” means here simply longer as compared with the digits of bats of

the planirostris and jamaieensis type; it does not imply that the writer is of

opinion that the wing-structure of glaucus and watsoni can, phylogenetically, be

derived from that of the planirostris-jamaicensis type. We have no means to

determine, with any degree of probability, which of these wing-structures is the

more primitive, 7. e, comes nearest to that of the prototype of the genus,

1908. | DR, KNUD ANDERSEN ON BATS. 287

closer examination shows that not all parts of the digits have

been lengthened in glaucus and watsoni; the metacarpals are

practically quite as in planirostris and jamaicensis (the fifth

metacarpal very slightly lengthened), but the first and second

phalanx of the third digit, and the first phalanx of the fourth

and fifth digits are noticeably longer. The wing-indices on p, 310,

and the diagram below show the details. One fact resulting from

this modification is worth noticing: both the first and the second

phalanx of the third digit are lengthened, but the former con-

siderably more than the latter (lengthening of first phalanx 57,

of second only 27 mm., for an assumed length of forearm of

1000 mm.). The result is that, whereas in the planirostris and

jamaicensis type the second phalanx of the third digit is very

distinctly more than 14 the length of the first, it is in glaweus

and watsoni always less than 14 of the first, a peculiarity easily

ascertained on careful examination of the wings of these bats.

f |

planirostris

glaucus, watsoni 6%

planirostris dt Pia sted Lice ale @ (tikes eR iP S|

glaucus, watsoni 44 |

planirostris a? |

glaucus, watsoni 6? | d u s aeallie. shire |

f, forearm. Given this length of the forearm, the third, fourth, and fifth digits

have in A. planirostris the lengths indicated by the lines d%, d‘, and d, in

A. glaucus and watsoni the lengths indicated by the lines 4%, 4, and 6°, The

subdivisions of d# and 4%, in direction from left to right, indicate the meta-

carpal, first, second, and third phalanx; those of d, 64, and d°, 6 the meta-

carpal, first and second phalanx.

Hairing on limbs and membranes.—Above, the proximal two

thirds of the forearm densely haired; a tuft of hairs on the

metacarpal of the pollex ; the interfemoral, femur, tibia, and foot

to the claws, covered with very short sparse hairs. Below, the

interfemoral hairy along the middle, almost naked laterally.

Colouwr.—The fur of the only specimen on record, a young

adult female (full-grown, but epiphyses of metacarpals separate)

preserved in alcohol, has the following colour :—Upper side greyish

drab witha slight tinge of fawn, base of hairs lighter, washed with

ecru-drab. Under side light greyish drab. White supraorbital

and infraorbital stripes distinct. Apparently no white margins

to the ears.

Proc. Zoou, Soc.—1908, No. XIX. 19

288 DR. KNUD ANDERSEN ON BATS. (EAT! 5

Measurements.—On p. 289.

Specimens exanvined.—One, the type, in the collection of the

British Museum.

Range.—Chanchamayo, Junin, Central Peru.

Tschudi’s Phyllostoma pusillum, 1844.—Indeterminable from

the description ; the author refers only to the colour, the nose-

leaves, ears, membranes, and general size; neither to the skull

nor to the teeth. The only measurement of any practical value is

that of the forearm, “1 5'"'” (i.e.37 mm.) ; if this measurement

was taken according to the same method as used in the pre-

sent paper, and if the specimen measured was full-grown, then

Tschudi’s bat was not an A. glawcus; if an Artibeus (not a

Vampyrops), it may have been A. rosenbergi.—Kven if Tschudi’s

Ph. pusillum were proved to be A. glaucus, the latter name would

have to stand, the former being preoccupied by Natterer’s

Ph. pusilluwm, which is Vampyressa pusilla*.

ARTIBEUS WATSONI Thos.

1901. Artibeus watsoni Thomas. Ann. & Mag. N. H. (7) vii. pp, 542-43 (1 June,

1901).—Type locality; Bogava, Chiriqui.

1804. Artibeus watsoni Thos., J. A. Allen, Bull. Am. Mus. N. H. xx. Art. iv.

p. 79 (29 Feb. 1904).—Boqueron, Chiniqui.

Diagnosis.—Similar to dA. glaucus, but smaller. Forearm

37°2-40°5.

A. watsoni and glaucus.—A. watsoni is a Central American

representative of the 4. glaweus type, differing from the Peruvian

species only in the rather smaller size, proportionately slightly

longer ears, and longer interfemoral.

The skull is quite of the same shape as in glaucus, but a trifle

smaller, especially narrower; of nine skulls of watsoni none

reaches the single skull of glawcus in size.

The teeth are in every respect (including the size) as in glaweus.

There is, asa rule, a faint remnant of the notch in the hinder

margin of m’, between its cusps 5 and 7, but in some individuals

also this trace has disappeared, the margin being perfectly simple.

The horseshoe is free all round, the margin sometimes simple,

sometimes finely crenulate.

The ears are, apparently, proportionately a little longer, and

rather narrower in their upper half (less broadly rounded off)

than in glawcus; but of nine specimens examined of watsoni two

only are preserved in alcohol, and only one specimen of glawcus

is available for comparison. Three small serrations on the outer

margin of the tragus above the median projection.

Both the alcoholic specimens have the interfemoral markedly

longer than in A. glaweus: 11:8 and 13°5 mm., as against 8 mm.

On the wing-structure, see A. glaucus (above, p. 287). Hairing

on limbs and membranes as in 4. glaweus.

% Thomas, Ann. & Mag. N. H. (6) iv. pp. 169-70 (Aug. 1889) ; and (7) v. p. 270

(March 1900).

1908. } DR. KNUD ANDERSEN ON BATS. 289

Measurements of Artibeus glaucus and watsoni.

| A. glaucus. A. watsoni.

2 yg. ad. 9 adults,

Type. 9 skulls.

Skull, total length to front ofc ... 21°2

Soh IMARL OLA eV L Resse eo oe ne 11 98 408 101

1 i 95

» width of brain-case ............ 87 8&8 87

» zygomatic width ............... BED ES ANG

;, maxillary width across m! ... 82 86 84 |

across cingula of canines ... 58 5d 6 57

Mandible, to front OGUINC ec cc ae 14 AZ, ASD i

Upper teeth, Cae ee ee een 68 65 7 67 |

Lower teeth, Cag ee ek 72 68 75 71

Ear-conch, length, inner margin ... 105 11 12

length, outer margin ... 14 15 162

» width. st Sake 12 12 «192

fracne:lencthy 5.05 4. eee 4°83 55 8662

Lancet, iL aA ate A 92 Sia) -9

ee Wid Lh tee ee ho. d a2 62

IOTRESHOE, WIERD | 20 onc soe cece ond 65 C3 a. 7.

Horearnt sc ac ee pee ee 438 372 405 39

PollexiNirssa0 SE. e Vt eed 11 Vase tf | 102

oa metacarpal... Se ACN etd 2 ees 39°4 33°77 362) «35°38

15°8 13 15 13°9

} te Se Nicer oe Meet hoes a a RN la ia ee 23 187 «21 20

eel Es Per ee Berd ata aE 12°8 | 88 112 102

4th metacarpal... = es eee ee 39 | 832 362 347

[vi boas sanaters «ohare aeee 13 11 12°22 116

Iv2.. Be cease 3B bee ee yee 15 11 135 123

5th metacarpal... acca eee 41 33°83 375 361

yi Eee eee euicete rencucee 11 8 9°5 87

NN tan mare acer OT 12 9 112 102

intertemoralec ete, 0.5 eos 8 118 135

SHOWED eo eae orc. tenets tam 145 Di Asa Ay,

Foot, with claws So eee eee BLS 103 92 10 oil

CHES OS Aisaeg a are aes 45 4Q 52 47

Colour.—Of six skins, four are dark, two lighter coloured.

Darker stage; 4 skins, Bogava (Chiriqui) and Cebaco I.;

full-grown, teeth unworn or slightly worn, epiphyses of meta-

carpals separate in one, ossified in the others :—Upper side from

the shoulders backward sepia~brown (browner than Ridgway’s

“sepia”), this colour confined to the narrow tips of the hairs;

base of hairs drab. Front part of upper side, from the shoulders

forward, of a rather lighter shade, more drabbish brown, owing

to the darker hair-tips being extremely short or wanting ; base

of hairs approaching ecru-drab ; there is no contrast between the

hinder and anterior part of the back, the colour of the former

shading gradually intc that of the latter. Under side dark grey.

White supra- and infraorbital stripes strongly pronounced. LEar-

conch very narrowly margined with white. Tip of the third

digit and adjoining membrane not lighter-coloured.—The four

specimens are almost alike in the shade of the colour.

1D*

290 DR. KNUD ANDERSEN ON BATS. [Apr. 7,

Lighter stage; 2 skins, Bogava (Chiriqui) and Sevilla I.;

full-grown, teeth unworn, epiphyses of metacarpals ossified.—

Upper side wood-brown, washed with a slightly darker shade on

the hinder back; base of hairs ecru-drab on the hinder back,

whitish washed with ecru-drab in front of the shoulders. Under

side lighter wood-brown. Facial stripes strong. Narrow white

edgings to the ears. No light-coloured tips to the wings.—The

two specimens are almost alike in colour, rather strongly con-

trasting with the four described above.

Measurements.—On p. 289.

Specimens examined.—9 specimens (two in alcohol, one skin in

alcohol, six dried skins) with skulls, from the following localities :-—

British Museum :—Bogava, Chiriqui, Panama (5); Cebaco L.,

Panama (2); Sevilla I., Panama (1).

U.S. National Museum* :—Escondido River, Nicaragua (1).

Range.—Central America (Panama, Nicaragua).

ARTIBEUS CINEREUS Gervais.

Diagnosis.—Molars 2. Cusp 7 of m' relatively small. m* equal

to about $ or ? the area of m'. Forearm 39-44 mm.

Skull.—Not differing in shape from that of A. glawcus and

watsoni; also the linear dimensions are, in every respect, very

nearly the same as in those species.

Teeth (text-fig. 54).—The teeth bear still more decisive evidence

of the very close relationship between 4. cinerews and A. glawcus

and watsoni. As in these latter, cusp 7 of m’ is relatively small,

Text-fig. 54.

Artibeus cinereus cinereus, 6 ad. Para. B.M. 1.7.19.3.

Right upper tooth-row (note small cusp 7 of m!). X 4.

equal to from one ninth to one seventh the bulk of the whole

tooth ; at least on average it is, no doubt, a trifle more developed

than in glaucus and watsoni, but not nearly as in Zoltecus and

aztecus.—The teeth differ only in the following points, of minor

importance :—

m, (rudimentary in glawcus and watson) has completely dis-

appeared ; 16 skulls, representing both of the races of A. cinereus

* US. N. M. no. 51544.

1908. ] DR. KNUD ANDERSEN ON BATS. 291

recognised in this paper, have been examined, in none of them is

m,, or any trace of its alveolus, present.—The heel of p* is a trifle

larger than in glaucus and watsoni, but the difference is so small

as only to be ascertained on very close comparison of the teeth.

The small increase in the size of the heel of p* corresponds to the

small increase (on average) mentioned above in the size of the

“heel” (cusp 7) of m’.

As in glaucus and watsoni there is only a very faint trace (or,

often, no indication at all) of a notch in the hinder margin of m’°,

between its cusps 5 and 7.

Tragus.—2, 3, or 4 serrations on the outer margin above the

median projection, always small, sometimes rather sharply defined,

sometimes so obsolete that their number cannot be counted with

certainty.

Nose-leaves—The margin of the horseshoe is free all round,

simple or finely crenulate.

Wings.—The peculiarities in the wing-structure described above

in A. glawcus and watsoni (p. 287) are also found in their closest

relative A. cinereus; aS in the former species the second phalanx

of the third digit is not fully 14 the length of the first. The

only appreciable difference is a slight lengthening of the meta-

carpals and of the phalanges of the fourth and fifth digits; by

this modification the wings have become on the whole slightly

longer and, because the increase falls chiefly on the fourth and

fifth digits, somewhat broader than in glaucus and watson.

See the wing-indices, below p. 310.

Hairing on limbs and membranes.—As in A. glaucus and watson

(above p. 287).

Colour.—There is no difference in colour between the two

geographical races of A. cinereus. The only skin available of a

young (not full-grown) individual is very dark-coloured, almost

precisely as young individuals of A. jamaicensis. Some of the

adult specimens with unworn teeth come extremely near in colour

to this young one, while the majority are a shade lighter, being

quite indistinguishable in colour from 4. watsoni ; a few specimens

are washed with drab-brown on the upper side ; none are as light

as the light-coloured stage of A. watsoni described above (p. 290).

Young, nearly full-grown (San Julian, Venezuela; U.S. N. M.

105432; A. c. bogotensis):—Upper side from the shoulders back-

ward dark smoky brown, almost blackish brown, this colour

confined to the tips of the hairs; base of hairs slate. On the

anterior part of the upper side, from the shoulder region forward,

the hair-bases are considerably lighter, nearly smoke-grey. Under

side dark grey, approaching hair-brown. White supraorbital and

infraorbital stripes strong. Har-conch narrowly margined with

white. No light tips to the wings.

A majority of adult individuals, teeth unworn or worn (both

races) :—Precisely as the darker stage of A. watsoni (p. 289).

Two adults (Kanuku Mts., Guiana; and Merida, Venezuela ;

teeth unworn; A. c. cinerews):—Upper side washed with drab-

292 DR. KNUD ANDERSEN ON BATS. [Apr. 7,

brown ; hair-bases on the hinder back light greyish drab, on the

neck ecru-drab. Under sidedrab. Facial stripesstrong. Narrow

white edgings to the ears. No light tips to the wings.—These

are the lightest coloured individuals I have seen; very likely a

still lighter phase occurs, similar to that of A. watson (p. 290).

Range.—Northern part of S. America: Para, Guiana, Vene-

zuela (including Trinidad), Colombia.

Remarks.—-This species cannot be discriminated with certainty

from 4. glauwcus and watsoni without an examination of the teeth

(a rudimentary m, present in glawcus and watsoni, wanting in

cinereus).

Two geographical races are separable, differing only in size.

The one, 4. ¢. cinereus, has probably its centre of distribution in

Guiana, having spread southward at least to Para, northward

to N.W. Venezuela; the other, 4. c. bogotensis, seems to have its

centre in Colombia, having spread northward to N.W. Venezuela,

where consequently both forms meet.

ARTIBEUS CINEREUS CINEREUS Gervais.

Stenoderma cinerewn Blainville, MS. label in Paris Museum (fide Gervais,

Us Os @5)¥

1856. Dermanura cinerewn Gervais, Exp. Castelnau, Mamm. 2° Mém. livr. 15,

sheet 5*, p. 36; pl. vii. figs. 4, 4a; pl. ix. figs. 4, 4a; pl. xi. fig. 3—Type

locality : “ Brésil.”

1901. Artibeus cinereus Gerv., Thomas, Ann. & Mag. N. H. (7) viii. p. 143 (Aug.

1901) —Kanuku Mts.

1901. Artibeus cinereus Gery., Thomas, Ann. & Mus. N. H. (7) viii. p. 192 (Sept.

1901).—Para.

Diagnosis.—Teeth, skull, and external dimensions averaging

smaller. Forearm 39-42 mm.

A.c. cinereus and bogotensis—There is only an average dif-

ference between the eastern form of dA. cinerews here under con-

sideration and the western form of the same species described

below (4. c. bogotensis). The skull, in 4. c. cinereus, is on the

whole slightly narrower, the maxillary width ranging between

8 and 86 mm. (average 8°3 mm.), in 4. ¢. bogotensis between

8°5 and 8-9 mm. (average 86 mm.). The teeth are slightly

smaller; the length of the upper tooth-row varies between 6:4

and 6°8 mm. (average 6°7 mm.), in A. c. bogotensis between 6:7

and 7-2 mm. (average 6°9 mm.). The forearm and metacarpals

average 2°2—3°5 mm. shorter.

Measurements.—On p. 295.

Specimens examined.—10 specimens (4 skins) and 8 skulls, from

the following localities :—

British Museum :—Para (1). Kanuku Mts., British Guiana,

about 59° W., 3° N. (4). Trinidad (1). Merida, Venezuela (2).

N.W. Venezuela (1).—7 skulls, representing all these localities.

* The title-page of the volume is dated 1855; on the probable dates of publica-

tion of the livraisons see C. Davies Sherborn and Bb. B. Woodward, Ann. & Mag.

N. H. (7) vin. p. 164 (Aug. 1901).

1908. ] DR. KNUD ANDERSEN ON BATS. 293

U.S. National Museum* :—Merida, Venezuela (1), with skull.

Range.—Guiana, southward at least to Para, northward to

N.W. Venezuela, including the island of Trinidad—In N.W.

Venezuela it meets A. c. bogotensis ; in this region the two races

apparently merge into one another.

Gervais’s Artibeus cinereus, 1856.—Type locality: Brazil.—

Notwithstanding the rather defective description there cannot be

much doubt as to the identification of Gervais’s specimen ; it is a

small Artibeus with 2 molars, a small cusp 7 of m’ (see Pl. ix.

fig. 4), and the locality is, as mentioned, “ Brésil.”- The combi-

nation of the habitat and the two characters quoted exclude all

other known species of the genus. The coloured figure on PI. ix.,

stated to be of natural size, is about 16 p. ct. too large in linear

dimensions (compare the measurements of the forearm and tibia

as given by Gervais); the metacarpals and phalanges are very

carelessly drawn.

ARTIBEUS CINEREUS BOGOTENSIS K. And.

1880. ? “Artibeus quadrivittatus Pet.,’ Dobson, P. Z.S. p. 465.—Popayan,

N. Colombia (Paris Museum ; specimen not examined). ;

1901. Dermanura quadrivittatum (not Peters) Robinson & Lyon, Proc. U.S. Nat.

Mus. xxiv. (no. 1246) p.510.—San Julian, Venezuela (only specimens W.I.

1586 and 1617 examined).

1906. Artibeus cinereus bogotensis Knud Andersen, Ann. & Mag N. H. (7) xvii.

p. 421 (Dec. 1906).—T ype locality : Curiche, nr. Bogota, Colombia.

Diagnosis—Similar to A. ¢. cinereus. but teeth, skull, and

external dimensions averaging larger. Forearm 41:2-44 mm.

A. c. bogotensis and cinereus.—The differences between this

form and its eastern representative, A. c. cinereus, have been

pointed out above, p. 292.

Measurements.—On p. 295.

Specimens examined.—9 specimens (8 skins) with skulls, from

the following localities :—

British Museum :—Colombia: Bogota region, various localities

(5). N.W. Venezuela (1).

U.S. National Museum t:—San Julian, 8 miles east of La

Guaira, Venezuela (2). Merida, Venezuela (1).

Range.—¥rom Central Colombia to N.W. Venezuela, where

it meets A. c. cinereus. In the latter region the two races

apparently merge into one another.

ARTIBEUS ROSENBERGI Thos.

1897. Artibeus (Dermanura?) rosenbergi Thomas, Ann. & Mag. N. H. (6) xx.

pp. 545-46 (Dec. 1897) —Type locality : Cachavi, N. Keuador.

Diagnosis.—Molars 2. Cusp 7 of m* small, m* equal to about

3 the area of m’. Forearm 37°8-39°8 mm.

Skull—Very similar in shape to that of A. watson and

A, cinereus cinereus.

Teeth (text-fig. 55).—Different from those of any other species of

* US. N. M. no. 123348.

+ U.S. N. M. nos. 105432-33 (nos. W.R. 1586 and 1617), 123344.

294 DR. KNUD ANDERSEN ON BATS. [EAvpresrf,

Artibeus, chiefly in the strong reduction in the size of m*. m/' not

essentially different from that of A. watsoni; cusp 7, if anything,

still less developed, represented only by a narrow, slightly pro-

jecting shelf, therefore as narrow as in any Vampyrops, its basal

outline almost perfectly quadrate. m’ little more than 4 (in all

other species of the genus % or 4) the area of m’, ian all the

elements of the tooth (cusps 4, 5, 6, 7) are distinct, and all of

them have been very nearly equally reduced in size, though

perhaps cusp 7 a little more than the others.

Text-fig. 55.

A B

Artibeus rosenbergi, g ad. Cachavi, N. Ecuador. Type, B.M. 97.11.7.76.

Right upper (A), left lower tooth-row (B). x 4.

On mz in this species see text below.

The lower m, is proportionally a little smaller than usual in

the genus, but by no means reduced to the same degree as m*;

the area of m, is about 4 (in other species about + 8) that of m,

In the type specimen at A. rosenbergi an. excessively small m, is

present on the left side, entirely wanting on the right side;

the normal condition is no doubt that m, is wanting. In the

material sent for identification from the U.S. National Museum

I find a second specimen of A. rosenbergi (no. 62635 dy. in which

there is no trace of m,; and in A. toltecus, which has 2 molars,

one skull, out of 27, has an m, on one side ‘of the mandible, thus

showing an individual anomaly apparently perfectly like that

of the type skull of A. rosenbergi.

Tragus and horseshoe-—The tragus has one or two serrations

on the outer margin above the median projection. The horse-

shoe is free all round, the margin simple or finely crenulate.

Wings.—The most noteworthy peculiarity in the wing-structure

is a conspicuous lengthening of the metacarpals; the phalanges

are very nearly of the same relative length as in A. glaucus and

watsont, the first phalanx of the third digit perhaps not quite as

long as in those species. See the wing-indices, p. 310.

airing on limbs and membranes.—Kssentially as in the nearest

relatives, A. glaucus, watsoni,and cinereus. Above, the proximal

half of the forearm densely haired; a tuft of short hairs on

first metacarpal; femur, tibia, foot to the claws, and interfemoral

1908.]

DR. KNUD ANDERSEN ON BAYS.

295

to its posterior margin covered with rather short and sparse

hairs.

to its median portion and posterior margin.

Colour (of the type specimen, an adult male with slightly worn

teeth, preserved in alcohol) :—Brownish drab above, base of hairs

much lighter ; under side greyish drab. White facial stripes

strong.

wings.

Measurements.—See table below.

Specimens examined. —Two specimens (in alcohol) with skulls,

viz., one from Cachavi, N. Ecuador (British Museum, the type),

and one from La Guaira, Venezuela (U.S. National Museum *).

Measurements of Artibeus cinereus and rosenbergi.

Skull, total length to front of ¢

>» across cingula of canines

Mandible, to front of incisors

Upper teeth, c-m? .....................

Lower teeth, c-mg ........

Kar-conch, length, inner margin ...

length, outer margin ...

width

Brac ussslene: Ghee eee eee ree eerer

hancetelenc ihre ee eee |

>» width

Horseshoe, width

Forearm ye a

Pollexeeeee sere:

3rd metacarpal 0... ...ccccseccseeeos

Ili?

4th

OWA

ih a ent ree

KN TACHA. soos coces nae vsoanb oe 902

ve.

Itniertemorals heey a

MG Oy ele Le Oeyec yesh secrete teers

WOO, WALD. GEMS coo docondone season csenne

Calcar

TAME LFOVOL VAKGHAN GoSoon bon con ase ceo]

width of brain-case ............

zygomatic width ...............|

maxillary width across m!...|

metacarpal... 0.cccccseeees|

Below, the hairing on the interfemoral chiefly confined

No white margins to the ears, no white tips to the

A. cinereus.

ie aa

cinereus.

8 skulls.

‘Mr. Max. Mup.

| 10 adults,

mm.

19°8

—

Oo OX GO OV FA Ot Fs O SD WO OX GO Ht CO

WINOEDMS cw

bogotensis.

8 adults,

8 skulls.

mm,

Bre

WD D> DCO DN TW TAH oN

ron)

mm.

41-2

385

14°8

| 10°7

375

13°2

| 39°5

9°83

112

14:8

| 103

4°8

Min. Max. Mep.

mm. mm.)

NT G> G2 ©1200 BO ¢

EO aw

|A. rosenbergi.

2 adults,

2 skulls.

Min. Max.

mm.

20°7

mm.

10°8

ee CO qo

ODI NOTA MHD N

loo olfe ole ow Ue elie e) Ou @

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M. no. 62635.

296 DR. KNUD ANDERSEN ON BATS. [Apr. 7,

Range.—The two specimens examined are the only known.

Remark.—A., rosenbergi cannot be discriminated, with certainty,

by any external character from other species of Artibews of

similar size, f. 1. A. watsoni, cinereus (cinereus), toltecus. But it

is unique in the strong reduction of m’.

ARTIBEUS TOLTECUS Saussure.

Diagnosis—Molars 3. Cusp 7 of m' large. Bony palate not

shortened. Forearm 37°5-43°5 mm.

A. toltecus and cinereus.—A. toltecus cannot be discriminated

with certainty by any external character from A. cinereus. The

interfemoral in A. tolfecus is probably, at least on average,

markedly shorter, but of one of the races, A. ¢. ravus, only skins

have been available for examination. The skull of A. toltecus is

almost precisely, in shape as in dimensions, like that of A. cinereus,

the only difference being a proportionately greater maxillary

width in A. ¢oléecus. The length of the tooth-rows is practically

the same in both species.

Nothwithstanding this close similarity, 4. toltecus and cinereus

are not only distinct species, but evidently representatives of two

distinct sections of the genus. In A. glaucus, watsoni, and cinereus

cusp 7 of m* is proportionally small (text-fig. 56 B); in A. toltecus

and aztecus (as well asin 4. quadrivittatus, turpis, and nanus) this

cusp is largely developed, being equal to about 1 to 7 (in glaucus,

Text-fig. 56.

A. Artibeus toltecus ravus, 6 ad. Corondelet, N.W. Ecuador. B.M. 1.6.5.3.

Right upper tooth-row (cusp 7 of m! large). X +.

B. Artibeus cinereus cinereus, g ad. Para. B.M. 1.7.19.3.

Right upper tooth-row (cusp 7 of m! small). X +.

watsoni, and cinereus to about 4 y to +) of the whole tooth (text-

fig. 56 4). Also cusp 7 of m” is in toléecus and allied species lar ger

and more distinctly projecting.~-The larger cusp 7 of m’ (and m°)

increases, of course, the breadth of this, tooth ; there cannot be

much doubt that the proportionally slightly larger maxillary

width of the skull of toltecus (and aztecus) mentioned above is

a direct consequence of the slightly increased breadth of the

molars.

1908. } DR. KNUD ANDERSEN ON BATS. 297

The proportionate length of the bony palate is quite as in

all the foregoing species ‘of Artibeus, the distance from palation

to the hinder margin of the incisive foramina being larger than

the distance from palation to basion (compare A. ee and

nanus, characterised by a shortening of the bony palate, p. 307).

A. toltecus and astecus.—The difference between these two

species is pointed out below, p. 306.

m, in A. toltecws.—In one skull, out of 27 examined, m, is

present on the right side, entirely wanting on the left ; the

individual is a young adult male of 4. ¢. ravus (Brit. ‘Mus.

no. 1.6.5.6). In all the other skulls representing both races and

different ages, from immature to very old, m, and its alveoli are

wanting.

Wings——The wing-structure very closely resembles that of

A. cinereus, the only appreciable difference being the slightly

shorter proximal phalanges of the third, fourth, and fifth digits ;

in A. cinereus the indices of these phalanges are, respectively,

357, 304, and 239, in A. foltecus 342, 289, and 227. For further

details see the wing-indices on p. 310.

Hairing on limbs and membranes.—As in A. cinereus, but the

upper side of the interfemoral more strongly haired, the fur

forming a distinct fringe along the posterior margin of the

membrane.

Colour.See A. t. toltecus (p. 298) and A. t. ravus (p. 300).

Range.—From N. Kecuador to Central Mexico (Durango), but

Keuador individuals (4. ¢. ravus) differ in some respects from

Central American and Mexican individuals (4. t. toltecus).

ARTIBEUS TOLTECUS TOLTECUS Saussure.

1860. Stenoderma tolteca H. de Saussure, Rev. & Mag. de Zool. (2) xii. pp. 427-28,

pl. xv. fig. 4 (Oct. 1860).—Type locality : Mexico.

1878. Artibeus cinereus (not Gervais) Dobson, Cat. Chir. Brit. Mus. pp. 520-

21.— Costa Rica, Guatemala, Mexico.

1882. Artibeus cinereus (not Gervais) Thomas, P. Z.8. p.371.—Durango (Mexico).

Magnosis— Averaging larger : forearm 39-43°5 mm. Facial

stripes and white edgings to the ears as a rule wanting or

indistinct, rarely well developed.

A, t. toltecus and ravus.—A. t. toltecus can only be discriminated

from A.t. ravus by average characters. The skuli of A. ¢. toltecus

averages In every respect a trifle larger (especially broader), but

small skulls of ¢oltecws are indistinguishable from large skulls

of ravus. The external dimensions average larger, the forearm

and metacarpals being from 2-7 to 3:2 mm. longer. As in

A. t. ravus there is a dark and a light phase, but the dark phase

in A. ¢. toltecus 1s noticeably darker “than in any specimen I have

seen of A. t. ravus. In most specimens of A. ¢. ravus, both in

its dark and light phase, the facial stripes and light edgings to

the ears are sharply pronounced; in <A. ¢. toltecus they are but

rarely well marked, as a rule indistinct or quite undeveloped.

From this it will be seen that it is impossible to draw a

ee)

DR. KNUD ANDERSEN ON BATS.

[ Apr. 7,

definite line between -A. ¢. toltecus and ravus ; they are evidently

a northern and southern representative of one species.

facial stripes and ear-edgings).—Dark

phase :—General colour of upper side very dark, approaching

blackish brown (being, rather, a blackish shade of drab), this

colour confined to the tips of the hairs ; ; base of hairs drab in the

posterior, almost ecru-drab in the anterior part of the upper side.

Under side drab.—This is the extreme of the dark phase in adult

specimens ; it is rather more blackish than the dark phase of

Colour (excluding

A, watsoni (p. 289).

Light phase:—In its extreme the light phase is indistinguishable

from that of A. ¢. ravus (p. 300).

The dark and light phases are connected by numerous

transitional stages, but a majority of the individuals examined

are more or less dark-

coloured.

Facial stripes and ear-edgings—In a majority of adult

individuals there is no trace of facial stripes nor of white

but individuals occur in which these ight

markings are more or less distinct, and sometimes, though

rarely, they are fully developed. The subjoined table will show

the amount of variation in this respect in 17 adult individuals

of A. ¢. tolfecus examined, as well as the stronger development

edgings to the ears ;

Facial stripes and ear-edgings of adult indiwiduals of

A. t. toltecus and ravus.

| Al, te foltecus...| Costa Rica (1).

| Nicaragua (1).

Guatemala (1).

| Oaxaca (2).

| Jalisco (9).

| Vera Cruz (3).

Supraorbital stripes

none.

indistinct.

none.

none (5).

indistinct (2).

distimet (1).

strong (1).

none.

Infraorbital stripes |

none.

indistinct.

none.

none (4).

indistinct (2).

distinct (2).

strong (0).

none.

LA t. toltecus.. | jeu localities (17).

A. t. ravus .../ All localities (11).

none (70 p. ct.).

| indistinct (18 p. ct.).

distinct (6 p. ct.).

| strong (6 p. ct.).

none (0).

indistinct (18 p. ct.).

distinct (86 p. ct.).

strong (46 p. ct.).

none.

| hone.

| none.

| distinct (2).

| none.

White ear-edgings |

none (3).

indistinct (4).

strong (0). |

none.

none (70 p. ct.).

indistinct (18 p. ct.).

distinct (12 p. ct.).

strong (0 p. ct.).

_ strong (0 p. ct.).

none (65 p. ct.).

indistinct (23 p. ct.).

distinct (12 p. ct.).

none (0).

indistinct (27 p. ct.).

distinct (86 p. ct.).

strong (87 p. ct.).

none (0).

indistinct (0).

distinct (27 p. ct.).

strong (78 p. ct.).

1908. ] DR, KNUD ANDERSEN ON BATS. 299

and apparently much greater constancy of these markings in

the southern race, 4. ¢t. ravus. It is a rather strange fact

that, so far as the facial markings are concerned, there is much

the same difference between the northern and southern races of

A, toltecus as between the northern and southern races of A. jamai-

censis; also in this latter species the facial stripes are stronger

and more constant in the southern forms.

Specimens from different localities —Central American are in

every respect indistinguishable from Mexican individuals. The

comparative table of measurements below, in which I have

arranged the adult individuals examined in four sections according

to their habitat, shows this as far as the cranial and external

dimensions are concerned.

Measurements of A. toltecus toltecus.

| Costa Rica, |

Nicaragua, Oaxaca. | om US Vera Cruz.

| Guatemala. UTaNE O°

| Zadults, | 2Qadults, | 9 adults, 3 adults,

3skulls. | 1 skull. | 5 skulls. 3 skulls.

Min. Max. | Mrn. Max. | Min. Max. | Min. Max.

mm. mm. | mm mm. | mm. mm. | mm. mm. |

Skull, total length to front of ¢ 20/5) 21 ie 20;6 20°2 21°2 20: 21:2 |

| 5, mastoid width hate eel aes} ata Ose elcAr 11 Ti

5, width of braim-case ~......... 98 10 ), Os 95 102 96 10

>» zygomatic width .. ae ee ari 12:3 W273} 113} 12:7 13

5 maxillary width across sml, 92 9:8 Oi 2 9 92 95 98 |

across cingula of canines ... 6 61 | 5 uf 62 538) 6 |

“Mandible, to fr ont of incisors ...... lee? ee Ale 13: 4 13°2 14:2 138 14 |

NUpperteethwc=m2e see | Oe aegee 68 68 7 a cla

Lower teeth, Camp ee oe 7 74, i 7 72 TCO)

Ear-conch, length, inner 1 margin ... 113 118 TL ly We} eR 7/ 1972 12:5

3 length, outer margin 15°8 155 158 16:2 18 168

sy width... eae eke rat mei 12:2 11-8 12°8 12 13°5 12 135 |

Tragus, length oe: 6 | *@ 6 Gi5y Cet Wey sees |

ID BWC NOES 5 onc nacnonecksoacesesebss its} S55] tes 88 G5 SF = ORS |

| gee We fe ede, we NA er Wit oe este: 55 62 Gee Cn

Eorseshoeawidthem Mee) et mn 6.5 Colne aoisme 5:8 62 72 HOE |

Forearm Yahi 39°38 42 39 39°8 39 42°5 39°2 43°5

Pollex 1 OD ORS A ONES 8) Toy aI

| 3rd metacarpal 378 39 36°38 38 358 39°8 37.0 4.05

Hee ae 13% 142 125 1352 13 14-7 13 15

Iii? 20 21 LOSSieea 19°2 22 20:'2 23°38

ITI3 Wha ial 10 118 10 12 102 12:3 |

4th metacarpal Hc nae Wea eenell acolo Male) 35 368 aD at) 36:2 39 |

IWR ae SA ee ies Rene aioe St Me Ne Semel abit 118 11 12'°8 IL 25)

HET rs 13 14 13 14 12-7 14:3 12°7 148

5th metacarpal — RAE eRe ena saeed | Melton Veh 368 37°2 368 40°2 37 = 40°8

IV ELRPr ne sa ste a nie sic ataocecttctenin Rv aaee Sacre 9 9°8 Simao 82 98 88 10

NaS pan hea Rea eens oe IL 112 10:2 11:3 102 11:8 10 12°2 |

Interfemoral Thee 65 75 5 105 a OS |

Wo Wierd ecaenire ce chi tases. nese Lio W5ien Mine aeece: We iG} 13°8 15:7 |

Foot, TriPIMC asi Wee OO OSs, TO Gn ie ane 10 «118 Tak) alley |

Calear 42 43 4:2 Ae2) 557 45 AS

300 DR. KNUD ANDERSEN ON BATS. [ Apr. 7,

Specimens examined.—24 specimens (3 skins), 15 skulls, from

the following localities :—

British Museum :—Costa Rica (2). Nicaragua: Jinotega,

1100 m. (1). Guatemala: San Geronimo (1). Jalisco: Ambas

Aquas, Tepic (2). Durango: Ventanas (2 juv.). ‘‘ Mexico” (1).

Uncertain locality (1).—7 skulls, from all the localities represented

by adult specimens.

U.S. National Museum *:—Oaxaca: Juquila (2). Jalisco:

Plantinar (7), Teuchitlan (1). Vera Cruz: Mirador (3). Uncertain

locality (1).—8 skulls, from all these localities.

Range.—Central America, 8. and Central Mexico, as far north

as Durango.

Saussure’s Stenoderma toltecwm, 1860.— The essence of the

original description is this:—WS. toltecwm is an Artibeus, with

3 molars, very short interfemoral (45 mm.), the forearm

measuring 41 mm., and inhabiting Mexico. The molars

exclude all Mexican species of the genus except A. toltecus,

pheotis, aztecus, turpis, and nanus. Of these, A. pheotis and

nanus are excluded by their small size, A. aztecus by its larger

size, A. turpis by its longer interfemoral ; leaving only the species

here called A. t¢oltecus, and, from the locality (Mexico), the race

A. t. toltecus.

Artibeus cinereus 77 Dobson's Catalogue, 1878.—I have examined

the four specimens catalogued by Dobson (J. s.¢.) as Artibeus

cinereus; all of them are A. toltecus toltecus.

we)

ARTIBEUS TOLTECUS RAVUS Miller.

1902. Dermanura rava Gerrit S. Miller, Jr., Proc. Acad. N. Sci. Philad. p. 404

(12 Sept. 1902).—Type locality : San Javier, N. Ecuador.

Diagnosis.—Similar to A. ¢. toltecus, but facial stripes as a

rule distinct or strong, and white edgings to the ears always

distinct ; general size averaging smaller: forearm 37°5-39°7 mm.

A. t. ravus and toltecus——The differences between these two

forms have been pointed out in detail above, under the description

of A. ¢. toltecus (pp. 297-2399).

Colour.—Adult individuals show a darker and lighter phase,

closely resembling those of A. watsoni :—

Darker phase: one skin, fully adult, teeth slightly worn (Brit.

Mus. no. 1.6.5.5.) :—Upper side as in darker-coloured individuals

of A. watsoni (see p. 289). Under side distinctly darker than

in A. watsoni, almost broccoli-brown. Facial stripes strong.

Whitish ear-edgings very distinct. No light tips to the wings.

Lighter phase: eight skins, fully adult, teeth slightly worn,

well worn, or much worn:—Precisely as in light-coloured

individuals of A. watsoni (p. 290). Facial stripes as a rule strong,

or at least distinct, sometimes indistinct, in none completely

* U.S. N. M. nos. 5203, 6979, 6281, 11216, 52082, 52085, 52087-89, 52095-96,

52098, 76512, 76515.

1908. ] DR. KNUD ANDERSEN ON BATS. 301

wanting (see table, p. 298). Whitish ear-edgings strong, or at

least distinct. No light tips to the wings.—The shade of colour

is not quite the same in all of the eight skins; some of them

evidently show traces of a darker stage, but none are truly

intermediate.

Measurements.—On p. 309.

Specimens exanvined.—12 skins, with skulls, from the following

localities :—

British Museum :—N, Eeuador: Pambilar (3); Corondelet (5).

U.S. National Museum *:—N. Ecuador: Parbilar (2) ;

Corondelet (2); all specimens paratypes of ‘‘ Dermanwra rava.”

Range.—As yet only known from N. Ecuador.

Miller’s Dermanura rava, 1902.—Type locality: San Javier,

N. Ecuador.—Miller compared D. rava with D. cinerea (i. e.

probably Dobson’s description of 4. cinerews, which, however, is

taken from examples of 4.7. toltecus) and D. tolteca (1.e. A. aztecus

of the present paper; I have seen the actual specimen referred

to by Miller, U.S. N. M. no. 52051), and found it differing in the

following particulars: smaller, the ‘‘ colour much paler,” ‘the

palatine foramina much more numerous,” and persisting “as

two conspicuous rows of small perforations even in very old

individuals.’—The type was collected by G. Fleming; all the

British Museum examples are from the same collector and

practically the same locality ; and by the kindness of Mr. Miller

and the Authorities of the U.S. National Museum I have had

for examination four of his paratypes, so that all doubt as to the

identification of D. rava is excluded.

Miller apparently laid much stress on the pale colour of

A. t. ravus, and the four specimens sent from the Washington

Museum are, in fact, all light-coloured; but the British Museum

series shows that also a dark phase occurs in perfectly adult

individuals. Of twelve skins examined, three must be put aside

as being either immature or young adults; of the remaining

nine, one represents the dark phase, eight the light. Taking in

consideration that the specimens were obtained in three different

places in N. Keuador (between August 10th and October 26th),

viz., San Javier, Corondelet, and Pambilar, there seems to be

some reason for supposing that this strong preponderance of

light-coloured individuals is not quite accidental; it may be that a

majority of adult individuals of this form are light-coloured. It

has been mentioned above (p. 299) that so far as the development

of the facial stripes is concerned, there is much the same difference

between the northern 4. ¢. tolfecws and the southern 4A. ¢. ravus

as between the northern and southern races of A. jamaicensis ;

if it proves true that a majority of individuals of A. ¢. ravus

are light-coloured, there is another parallelism to 4. jamaicensis ;

as pointed out above (p. 256) 77 p.ct. of individuals of the

southern races of jamaicensis are light coloured, as against only

* U.S. N. M. nos. 113333-34, 113337, 113339.

302 DR. KNUD ANDERSEN ON BATS. [Apr. 7,

25 p. ct. in the northern races. But even if this be so, there is

at all events no absolute difference in this respect between

A, t.ravus and the true A. ¢. toltecws, in which latter dichromatism

also occurs (see above p. 298). As to the number and persistency

of the small palatine perforations, it is a character of no diagnostic

value; it varies from individual to individual quite as much as it

does from species to species.

ARTIBEUS QUADRIVITTATUS Pet.

1865. Artibeus (Dermanura) quadrivittatus Peters, MB. Akad. Berlin (13 July,

1865) p. 358.—Type locality: Surimam.

1878. Artibeus quadrivittatus Pet., Dobson, Cat. Chir. Brit. Mus. p, 521.—

Surinam; Pernambuco. i

1888. Artibeus quadrivittatus Pet., Jentink, Cat. Syst. Mamm. p. 209.—Surinam

(type specimen).

Diagnosis.—Similar to A. toltecus, but upper side of interfemoral

and tibia more sparsely haired, and interfemoral averaging

somewhat longer. Forearm 41°7—44 mm.

A. quadrivittatus and toltecus—In the shape and size of the

skull, in che structure, number, and size of the teeth, and in all

external characters, A. quadrivittatus is similar to A. toltecus,

with these two exceptions :—First, in 4. toltecws the upper side of

the interfemoral and tibia is densely haired; in A. quadrivittatus

the hairs are so short and sparse as to make the interfemoral

membrane appear almost naked. Second, in the single alcoholic

specimen examined of A. guadrivittatus the interfemoral measures,

in the middle line, 12°3 mm., whereas the maximum found in a

series of A. t. toltecws is 10°5 mm.; it probably indicates that the

interfemoral averages longer in quadrivitiatus.

The three specimens (two skins) examined are not sufficiently

well preserved for a detailed description of the colours; in one

the facial stripes are strong, in the others rather indistinct,

Measwrements.—On p. 309.

Specimens examined.—Surinam (one, with skull) ; Pernambuco

(two, with skulls); from the collection of the British Museum.

Peters’s Artibeus (Dermanura) quadrivittatus, 1865.—Type

locality : Surinam; type in the Leyden Museum.—The essence

of the short original description is this:—‘‘ Von der Grdsse und

dem Ansehen des Sé. toltecwm Saussure, aber mit etwas breiterer

und weniger behaarter Schenkelflughaut,” and “ mit vier weissen

Liingsbinden auf dem Kopfe.” Forearm 40, tibia 14, interfemoral

in middle line 9 mm.

From the above there can be no reasonable doubt as to the

identification of A. guadrivittatus. Peters’s measurement of the

interfemoral, viz. 9 mm.,if compared with the length of this mem-

brane in a British Museum specimen, viz., 12°3 mm., seems to

show (which indeed was to be expected) that in the length of

the interfemoral there is only an average difference between

A. quadrivittatus and A. toltecus toltecus; ina series of this latter

I find it varying from between 5 and 10:5 mm. (average 7-9 mm.),

1908. | DR. KNUD ANDERSEN ON BATS. 303

so that the only, apparently reliable, difference between the two

species is the denser hairing of the interfemoral and tibia in

A. toltecus.

Dobson’s Artibeus quadrivittatus——Dobson’s description of

A. quadrivittatus (1.s.¢.) may be epitomised in the following four

sentences :—First, it is ‘‘ very similar to 4. cinereus (i. e. A. toltecus

toltecus|, but the nose-leaf is narrower and not so abruptly

narrowed at the summit”; Dobson gives as breadth of the nose-

leaf in A. gquadrivittatus 0''-25, in “A. cinereus” 0°28, making a

difference of only 0':02 or 0°5 mm.; considering the not in-

conspicuous variation in the breadth of the nose-leaf in all other

species, it would, @ priori, appear highly improbable that such a

small difference would prove to be reliable; and as a matter

of fact there is no difference at all in this respect between

A. t. toltecus and A. quadrivittatus; in the former the lancet is

5:5-6°6 mm. broad, in the only alcoholic specimen I have seen

of the latter (also examined by Dobson) 6 mm.; also the shape

of the lancet is the same in the two species. Second, “ inter-

femoral membrane much deeper”; this is only correct, if for

“much” we substitute “on an average somewhat.” Third, the

upper surface of the interfemoral “thinly clothed with fine

hairs,’; this is correct (but there is no corresponding character

in Dobson’s description of his A. cinereus). Fourth, “ the head

with four longitudinal, not very distinct, white streaks, arranged

as In A. perspicillatus [i.e. A. jamaicensis lituratus|”, whereas in

his A. cinereus | A. t. toltecus| there are ‘“‘ no white streaks on the

face”; but the character is (as might be expected) individually

variable in both species (as it also is, more or less, in other forms

of the genus) ; in 4. ¢. tolfecus the facial stripes are, as a rule,

wanting or rather indistinct, but specimens occur in which they

are well developed, and, on the other hand, of three examples of

A. quadrivittatus two have the facial stripes rather indistinct,

one strong.—This perusal of Dobson’s description, based on the

same material as examined by him, leads to the same result as

emphasised above: there is, probably, an average difference in the

length of the interfemoral, and an apparently well-marked

difference in the hairing of this membrane and the tibia, but I

am unable to find any other character by which these two ex-

tremely closely related species can be discriminated from each

other.

ARTIBEUS PHXOTIS Miller.

1902. Dermanura pheotis Gerrit 8. Miller, Jr., Proc. Acad. N. Sci. Philad. p. 405

(12 Sept. 1902)—Type locality: Yucatan.

1906. ? Dermanura jucundum D.G. Elliot, Proc. Biol. Soc. Wash. xix. p. 50 (1 May,

1906).—Type locality: Vera Cruz.

Diagnosis.—Similar to A. toltecus ravus, but with somewhat

shorter tooth-rows, and no whitish edgings to the ears. Forearm

about 38 mm.; third metacarpal about 37 mm.

I have not seen this species, which was described by Miller

Proc. Zoou. Soc.—1908, No. XX. 20

304 DR. KNUD ANDERSEN ON BATS. > [PA‘pELia

from a single example (skin and skull) obtained at Chichen Itza,

Yucatan. The following notes are based partly on Miller's

published account, but chiefly on more detailed information,

photographs of the skull (upper, lateral, and lower views), camera

lucida outlines of the molars and the profile of the skull, and

measurements of skull and external dimensions kindly given me

by Dr. Mareus W. Lyon, Jr., Washington.

Skull.—General shape quite as in A. t. ravus. Judging from

photographs of the type skull of pheotis, I am unable to see

any appreciable difference from a series of skulls of ravus, but

Miller, who in the original description of pheotis compared its

skull with that of ravus, found ‘the rostrum broader and flatter

and the median backwardly extending portion of the bony palate

wider.” —The measurements, as taken by Dr. Lyon (see table

below, p. 309), are practically quite as in ravus.

Teeth.—The teeth of the type are very much worn down, but

from photographs (twice natural size) and camera lucida outlines

of the molars it is quite clear that they accord with those of

A. toltecus ; cusp 7 of m’*is large, as in this latter species. The

tooth-rows are somewhat shorter than in A. ¢. ravus: upper teeth,

cm’, 6 mm., according to Dr. Lyon, as against 6°D-7 mm. ina

series of ravus, measured by myself.

Colour.—Miller describes the colour of the fur as closely

similar to that of A. é ravus, but the ears are ‘‘ much darker and

without the whitish border.” All the specimens of ravus I have

seen have distinct or strong whitish edgings to the ears (see

table above, p. 298).

Haternal dimensions.—I am indebted to Dr. Lyon for measure-

ments of the forearm, third digit, first phalanx of fourth digit,

and first phalanx of fifth digit; they are much as in 4. ¢. ravus,

only the metacarpals would seem to be a trifle longer (see below,

p. 309).

Measurements.—On p. 309.

Material examined.—Photographs of the type skull.

Range.—Yucatan; % Vera Cruz (see below, p. 305, under

“« Dermanura jucunda”’).

Remarks.—From the available information it appears that

A. pheotis is very closely related to A. t. ravus, differing chiefly

in the somewhat shorter tooth-rows and lack of white edgings

to the ears.

A species of Artibeus described below (p. 308) under the name

of A. nanus, has like pheotis 4 molars and cusp 7 of m’ large;

the forearm measures 36°5-38 mm. (in pheotis 38), the upper

tooth-row 5:°8-6:1 mm. (in pheotis 6); in other words, in the

form and number of the teeth and, so far as the length of the

forearm is concerned, also in external dimensions, the two species

are alike; further, nanws is known from the Mexican States of

Guerrero, Colima, Sinaloa, and Vera Cruz, pheotis from Yucatan

and, probably, Vera Cruz, so that the distribution of the species

is, partly at least, the same. Some words are therefore necessary

1908. ] DR. KNUD ANDERSEN ON BATS. 305

to prevent a confusion of these two bats which, though similar in

the points mentioned, are widely distinet species :—

The skull of A. phewotis is probably of the ftoléecus pattern ;

in A. nanus (and its larger relative A. turpis) the rostrum

is peculiarly flattened and slightly bent upwards; the profile

of the nanus skull is therefore very different from that of

the pheotis skull. In A. pheotis the proportionate length of the

bony palate is quite as in ¢oltecus and allied species : measured

from palation to hinder border of incisive foramina, longer than,

or at least equal to, distance from palation to basion; in nanus

(and turpis) the palate is shortened: its length (palation to

incisive foramina) shorter than the post-palatal portion, from

palation to basion. The skull of nanws is, on the whole,

distinctly smaller than that of pheotis. All these differences

have been confirmed by Dr. Lyon (in litt.), who kindly compared

a skull of my 4. nanus (U.S. N. M. no. 51765; Colima) with

the type skull of Miller’s A. phwotis.—Also externally the two

species, in spite of all similarity, are distinguishable: although

the forearm in pheotis appears to have the same length as in a

large nanus, the metacarpals are conspicuously longer; third

metacarpal 37°3 mm., against 32°2-35 mm. in a series of “anus ;

also the proximal phalanges are a little longer in pheotis.

Elliot’s Dermanura jucunda, 1906.—Type locality: Achotal,

“State of Vera Cruz, Mexico. Described from one example, with

skull.—According to Elliot, it is ‘allied to D. quadrivittatum

from South America, but is smaller, with a considerably smaller

skull,” and “the nose behind the nose-leaf whitish”; supraorbital

stripes ‘‘ very conspicuous,” infraorbital stripes ‘‘ very indistinct.”

Professor Elliot has kindly informed me that the distance from

palation to the hinder border of the incisive foramina is slightly

greater than the distance from palation to basion (thus quite as in

the ordinary Artibeus skull, not as in 4. nanus and turpis), and

given me some measurements of the skull, teeth, and wing (see

table p. 309). From these measurements it is evident that the

size of the skull and teeth is precisely as in A. pheotis. The

forearm measures ‘‘41°8 mm.” (Elliot in litt.; not 43 mm. as

stated in the published description), as against “37-9 mm.” (Lyon)

in pheotis, a discrepancy of 3°9 mm.; but Lyon has undoubtedly

measured the forearm of pheotis to the distal end of the radius

(if not, the difference between the length of the forearm and the

third metacarpal, respectively 37-9 and 37°3 mm., would certainly

be greater), and if Elliot, on the other hand, has measured the

forearm of jucunda to the front curve of the carpus, then

the discrepancy is reduced to about 3 mm., a very reasonable

amount of individual variation in an Arétibeus of this size;

further, it should be noticed that Ellot’s measurement of the

third metacarpal is exactly (to a fraction of a millimetre!) like

that of A. pheotis, viz. 37°3 mm., and also the length of the

phalanges of the third, fourth, and fifth digits practically as in

-pheotis—The white patch behind the nose-leaf mentioned by

20*

306 DR. KNUD ANDERSEN ON BATS. [ Apr: 7,

Elliot is of no importance as a specific character ; it only means.

that the supraorbital stripes are fused together in front; the

same is often the case in other species, whenever these stripes are

strongly developed. The fur is described as “dusky brown,

whereas the type specimen of pheotis, according to Miller, does

not differ in colour from his series of A. ¢. ravus, and therefore

probably is pale-coloured ; but also this proves nothing about the

distinctness of D. jucunda; dichromatism is the rule in the

species of Artibeus.

In brief, neither in the original description of D. jucunda

nor in the additional information given me by Professor Elliot,

am I able to find a single character by which D. gucunda can

be discriminated from 4A. pheotis. So long as it has not been

proved that such characters exist, | must regard the former name

as a synonym of the latter.

ARTIBEUS AzTECUS K. And.

1902. Dermanura tolteca (not Saussure) Gerrit S. Miller, Jr., Proc. Acad. N. Sci.

Philad. p. 404, footnote (12 Sept. 1902).—Morelos, Mexico (specimen

examined).

1906. Artibeus aztecus Knud Andersen, Ann. & Mag. N. H. (7) xviii. p. 422

(1 Dec. 1906).—Type locality: Tetela del Volcan, Morelos, Mexico.

Diagnosis.— Allied to A. toltecus, but in every respect somewhat

larger ; metacarpals unusually long; interfemoral strongly haired.

Forearm 45—46°8 mm.

A. aztecus and toltecws.—A. aztecus has no closer known relative

than A. toltecus. As in this latter species cusp 7 of m* (and m*)

is largely developed, the maxillary width of the skull propor-

tionately large, the number of molars 3. But the skull is in

every respect slightly larger and more heavily built, the teeth a

little larger. The exter rnal dimensions are creater : ; in the

smallest available specimen of dA. aztecus the forearm is 6 mm.

longer than in the smallest 4. ¢. ¢oltecus, in the largest specimen

3°3 mm. longer than in the largest A. ¢. toltecus. The metacarpals

are unusually lengthened ; in 4. toltecuws the indices of the third,

fourth, and fifth metacarpals are, respectively, 912, 898, and

923; in A. aztecus 946, 928, and 954; the first phalanx of

the third digit is of the same length as in 4. foltecus, but the

second phalanx so much lengthened as to be more than 14 the

length of the first ; also the second phalanx of the fourth digit is

proportionately longer than in 4. toléecuws (compare wing-indices,

on p. 310). All these modifications of the wing-structure make,

of course, a proportionately longer wing; in 4A. foltecus the

indices of the three principal digits are 2037, 1516, and 1419, in

A, aztecus 2088, 1565, and 1442.

The interfemoral is very short (as in 4. foléecus) and unusually

strongly haired, as is also the upper side of the tibia.

Colour.—The general colour of the fur, in all the four specimens

examined, is quite as in the dark-coloured phase of 4. ¢. toltecus

(above p. 298). Facial stripes very indistinct or completely

1908. | DR. KNUD ANDERSEN ON BATS. 307

wanting. No light edgings to the ears. No light tips to the

wings.

Measurements.—On p. 309.

Specimens examined.—Tetela del Volean, Morelos, Mexico

(4, with skulls); from the collection of the U.S. National

Museum *,

Range.—As yet only known from Morelos, Mexico.

ARTIBEUS TURPIS K. And.

1906, Artibeus turpis Knud Andersen, Ann. & Mag. N. H. (7) xviii. p. 422 (1 Dec.

1906).—Type locality: Teapa, Tabasco, S. Mexico.

Diagnosis.—Molars 3. Cusp 7 of m* large. Rostrum of skull

amusually depressed and distinctly bent upward ; palate shortened.

Larger than A. nanws: maxillary tooth-row 6°7 mm., forearm

40°5 mm.

Affinities —A. turpis and nanus (below, p. 308) are rather

closely related to 4. toltecus and quadrivittatus. As in these

Text-fig. 57.

A, B. Artibeus nanus, 2 ad. Colima. U.S. N.M. 51765. X 3.

Text-fig. 58.

A, B. Artibeus toltecus toltecus, 2 ad. Jalisco. U.S. N. M. 52038. x

tales

‘species, cusp 7 of m* is large, m* and m, wanting, the rostrum

-conspicucusly broadened. In some specimens of 4. ¢. toltecus,

* U.S. N. M. nos. 52050-51, 53769, 53772.

308 DR. KNUD ANDERSEN ON BATS. [Apr. 7,

particularly in aged individuals, there is a tendency to flattening

of the rostrum and vaulting of the brain-case ; in 4. turpis and

nanus (text-fig. 57) this tendency has been carried to an ex-

treme, and at the same time the rostrum is bent slightly upward ;

probably as a consequence of this latter, the bony palate has

become shortened. These are the chief peculiarities of A. tarps

and nanus as compared with 4. toltecus and quadrwittatus.

Skull (compare text-fig. 57, of A. nanus).—Short and broad,

as in A. foltecus and quadrivittatus. Rostrum more depressed

and brain-case more vaulted than usual in the genus. Rostrum

with a slight, but distinct, upward trend, the alveolar border

of the maxillary bone, therefore, in profile more abruptly

ascending; in A. foltecus (text-fig. 58) the profile of the nasal

bones is slightly descending rather than horizontal, in A. turpis

it is slightly ascending. Bony palate shortened ; in A. ftoltecus

(as in all the foregoing species of the genus, with exception

of the short-faced A. concolor) the length of the palate, from

palation to posterior border of incisive foramina, is larger than

(extremely rarely almost equal to) the length of the post-

palatal portion of the skull, from palation to basion ; in 4. turpis

the bony palate is shorter than the post-palatal portion.

Teeth As in A. toltecus and quadrivittatus. Cusp 7 of m*

large ; cusp 7 of m* conspicuous, shelf-like, projecting.

In the nose-leaves, ears, wing-structure (see wing-indices,

p- 310), and hairing on tibia and interfemoral, A. twrpis does not

differ appreciably from 4. foltecus.

Colour (Q ad., preserved in alcohol; teeth almost unworn ;

type of species)—General colour of upper side approaching

Prout’s brown, but with a distinct tinge of drab; base of hairs on

hinder back almost wood-brown, on the neck and shoulder region

ecru-drab ; under side light wood-brown. Facial stripes strong.

Narrow light margins to the ears. No white tips to the wings.—

The general colour of this specimen has probably not remained

quite uninfluenced by the preserving-fluid.

Measurements.—On p. 309.

Specimens examined.—One adult female, Teapa, Tabasco, 8.

Mexico; with skull; the type; British Museum.

Range.—As yet only known from the type specimen described

above.

ARTIBEUS NANUS K. And.

1906. Artibews nanus Knud Andersen, Ann. & Mag. N. H. (7) xviii. p. 423 (1 Dee..

1906).—Type locality: Tierra Colorada, Sierra Madre del Sur, Guerrero,.

Mexico.

Diagnosis.—Similar to A. turpis, but in every respect smaller.

Maxillary tooth-row 5-8-6-1 mm.; forearm 365-38 mm.

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reoccur in A. nanus, if anything in a still more pronounced

degree :—the depression and slightly ascending trend of the.

rostrum, the shortening of the bony palate, the high brain-case,.

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310 DR. KNUD ANDERSEN ON BATS. [Apr. 7,

the very broad skull (text-fig. 57 on p. 307); cusp 7 of m'and m?

are relatively large, as in A. toltecus, quadrivittatus and turpis ;

m, absent (five skulls examined); facial stripes strong. But

A. nanus is conspicuously smaller than 4. turpis; so far as my

material goes, there seems to be a perfectly clear line of separation

between the two species ; in the largest example (among 12 spe-

cimens and 5 skulls) of A. manus the skull is 1-3, the maxillary

tooth-row 0:6, the forearm 2°5, and the third metacarpal 2 mm.

shorter than in 4. turpis. For further details see the table of

measurements, p. 309.

Colour (adult skin; Buena Vista, Vera Cruz; U.S. N. M.

no. 112791).—Upper side dark brown, with a tinge of drab;

under side broccoli-brown. Facial stripes strong. Conspicuous

hight margins to the ears. No light tips to the wings.—A series

of alcoholic specimens are noticeably lighter-coloured ; in all the

facial stripes are very distinct; the light ear-edgings sometimes

obsolete.

Specimens examined.—12 specimens (one skin) and 5 skulls,

from the following localities :—

British Museum :—Guerrero: Tierra Colorada, Sierra Madre

del Sur (3). Sinaloa: Presidio, near Mazatlan (1).—3 skulls,

representing both localities.

U.S. National Museum * :—Colima: Hacienda Magdalena (7).

Vera Cruz: Buena Vista (1).—2 skulls of adult specimens of the

series from Colima (the skull of the single example from Vera

Cruz has been lost).

Range.—Mexico, as far north as Sinaloa. Sinaloa is the most

northern locality from which any species of Artibeus has been

recorded.

Remarks.—The differences between A. nanus and A. pheotis

have been pointed out above, pp. 304-305.

Wing-indices.

BR BR BRA

3rd digit. | 4th digit. 5th digit.

Forearm. | an

|| Mte.'1 ph.|2 ph.!3 ph.|| Mte. |1 ph.|2 ph.|/ Mte. /1 ph. 2 ph.

| U. bilobatum and | |

thomasi (25 specim.)?.........| 1000 938 | 347 | 518 | 275 || 912 | 294 | 317 || 926 | 231 | 280

. planirostris (85 specim.) ...| 1000 903 | 297 | 483 | 262 || 883 | 260 | 324 || 905 | 197 | 244

. hivsutus (8 specim.) ......... | L000 887 | 281 | 481 | 261 || 868 | 245 | 313 || 891 | 186 | 231

. jamaicensis (225 specim.)...; 1000 || 900 | 298 | 492 | 259 || 884 | 259 | 318 || 913 | 201 | 245

. glaucus and

watsoni (10 specim.) ......... 1000 897 | 354 | 510 | 264 || 884 | 296 | 317 || 920 | 224 | 261

. cinereus (18 specim.) ...... 1000 912 | 357 | 510 | 266 || 896 | 304 | 329 || 932 | 239 | 278

. toltecus and | |

quadrivittatus (32 specim.).| 1000 || 912 | 342 | 501 | 272 || 898 | 289 | 329 || 923 | 227 | 269

. aztecus (4 specim.) ......... | 1000 || 946 | 342 | 582 | 268 || 928 | 288 | 349 || 954 | 222 | 266

. turpis and |

nanus (9 specim.) ............, 1000 903 | 341 | 505 | 263 || 890 | 293 | 323 || 914 | 240 | 277

* U.S. N. M. nos. 51765-67, 51771, 51773-74, 51776, 112791.

1908. | DR. KNUD ANDERSEN ON BATS. 311

Summary of characters of Genera, Species, and Subspecies.

a. Skull long and slender; rostrum but very slightly

depressed : height at p* greater than, or equal to,

width of skull at postorbital constriction ; bony

palate long: distance from palation to front of

incisors about equal to zygomatic width; median

backwardly extending portion of bony palate long,

equal to combined length of m! and m?; anterior

nasal opening less oblique. Outer upper incisors

bifid; cusp 2 of m, small; molars 2; m® larger

(than in Artibeus), situated in row behind (not

postero-internally to) m?, almost as broad as

hinder border of m?; cusp 5 of m? situated near

the labial margin of the tooth. A narrow line of

whitish fur down the middle of the back ............ UroprEr™4, p. 212.

a. Length of skull (to front of c) 22-23°3 mm. ;

upper teeth (c—m?) 7°5-8°5. Length of ear-conch

(outer margin) 15°7-16°8; width of horseshoe

6°2-7°5. (S. Brazil and Peru to Costa Rica.) .... U. bilohatum, p. 217.

61, Length of skull (to front of c) 24°7-24°8 mm. ;

upper teeth (e-m*) 8-9-9. Length of ear-conch

(outer margin) 18-18°5 ; width of horseshoe

Tooele in. (EXCLUDES) Mere ye dia tart lle eta eat a eke

-6. Skull short and broad; rostrum considerably de-

pressed : height at pt much less than width of

skull at postorbital constriction ; bony palate

short: distance from palation to front of incisors

much less than zygomatic width; median back-

wardly extending portion of bony palate short,

much less than combined length of m! and m?;

anterior nasal opening more oblique. Outer upper

incisors simple (not bifid) ; cusp 2 of mj, strongly

developed, raised as a high slender cone near the

middle of the lingual margin of m!; molars 3, 3,

or 2. No white longitudinal dorsal stripe.

c!, Median upper incisors simple (not bifid) ; molars

3; m3 and ms larger (m3 equal to about 4 of my) ;

m? situated in row behind (not postero-in-

ternally to) m?; cusp 5 of m? situated near the

lateral margin of the tooth. Tragus with a

pointed projection on inner margin near tip...... ENCHISTHENES, p. 221.

a1, Median upper incisors bifid; molars 2, $, or 3;

m? and m3 (when present) smaller (m%, if not

wanting, equal to 4; of mg); m*, if not want-

ing, situated postero-internally to m?; cusp 5 of

m? moved lingually so as to occupy (precisely or

nearly) the middle of the posterior margin of the

tooth. No pointed projection on inner margin

Oliiracusimean tips ene eee eee ER TLB BUS) Da 224s

a*. Molars 3.

a’. Teeth small: upper row (c-m?) about

772 mm. Rostrum shorter. Forearm

Oia, ((Civpenae, VIVAL) oc aoc ccanseoes neces A. concolor, p. 232.

63, Teeth larger : upper row (c—m2?) 9°5-12 mm.

Rostrum longer. Forearm 53°7-73 mm.

a‘, Tibia and distal part of interfemoral so

short-haired as to appear almost naked ;

colour of fur of upper side not drab.

Upper teeth (cm?) 9°8-12 mm.; forearm

SID A7/S) TCDS, coo aanpsbononaeaonescnes cduecodsecssoee | 24le (DUNG AOSAPES 14 ZED

a. Smaller. Length of skull (to front of c)

27-30 mm.; zygomatic width 16:2—

19°2; upper teeth (c-m?) 9°8-11. Fore-

arm 55-65°2.

a, Average length of skull (to front of c)

28'°5 mm.; average zygomatic width

17°6. Average length of forearm 61°8.

(Continental: Brazil toS. Mexico)... 4. p. planirostris, p. 237.

U. thomasi, p. 221.

312 DR. KNUD ANDERSEN ON BATS. (Pano i

68, Average length of skull 27°8 mm. ;

average zygomatic width 16°8. A ver-

age leneth of forearm 57°7. a

dad, Tobago.)... A. p. trinitatis, p. 241.

°&, Average length of skull 28°9 mm.

average zygomatic width 18. Av er-

age length of forearm 59°4. (Gren-

ada.)....... : A. p. grenadensis, p. 241.

6°. Very large. Length of skull (to. front

of c) 29°5-33 mm. ; zygomatic width

18°3-20°8 ; upper teeth (cm?) 10°4—-12.

Forearm 62°8-73. (Guiana, Para,

Lower Orinoco.)........ A, p. fallax, p. 242.

64, Tibia and interfemoral densely haired

above; colour of fur of upper side in

adults drab with a silvery tinge. Upper

teeth (c-m?) 9°5-10°4 mm.; forearm

53° 7-59" ZY was (ONICSEO,) anasonaacngosn coc A, hirsutus, p. 245.

62. Molars 3.

ce’, Large: forearm 54-76 mm. ..... severe. A, Jamaicensis, p. 247.

e#, Smaller races : length of skull’ (to front

of c) 26-80 mm.; forearm 54-66 mm.

Angular notch in hinder border of m2,

between cusps 5 and 7 (place of lost m3),

as a rule distinct, rarely quite obliterated ;

darker colour phase predominant ; facial

stripes as a rule wanting or faint, rarely

strongly developed. (Northern + races :

Greater Antilles, as far east as St. Kitts;

S. Mexico, C. America; extending to

Colombia and Ecuador.)

o>. Average: length of skull (to front of c)

26°9 mm.; upper teeth (c— He Ora 5

forearm 56° 8. (Cuba.)... A. J. parvipes, p. 261.

d>, Average: length of skull 27-4 mm.

upper teeth 9: 9; forearm 59°6. (Yuea-

tan, Belize.) A. 7. yucatanicus, p. 263..

e. Average : length ‘of skull 28°3 mm.

upper teeth 10:3; forearm 60: iL

(Greater Antilles, except Cuba, as far

east as St. Kitts; Caribbean Islands;

S. Mexico, except Yucatan; C.America.) 4. 7. gamaicensis, p. 265.

f°. Average: length of skull 29:7 mm. 3

upper teeth 11; forearm 62°9. (Ecua-

dor, Colombia. Wes see A.J. equatorialis, p. 270..

d4, Larger races : length ‘of skull (to front

of c) 29°5-34 mm.; forearm 60-76 mm.

Angular notch in hinder border of m?,

between cusps 5 and 7 (place of lost m‘ 3),

as a rule reduced or obliterated, rarely

perfectly preserved ; lighter colour phase

predominant; facial stripes distinctly or

strongly dev eloped, rarely wanting.

(Southern races: S. America, extending

to S. Mexico; from Trinidad to Guade-

loupe; unrepresented in the Greater

Antilles.)

g°®. Averaging larger: forearm 64-76 mm.

(S. America, “extending to S. Mexico; et J lituratus, p. 272.

Trinidad and St. Vincent. ese . A. 7. palmarum,* p. 278.

A®, Averaging smaller: forearm 60-66" 2mm.

(Dominica, Guadeloupe.) .................. A. 7. preceps, p. 283.

d3, Small: forearm 37:2-43°8 mm.

e4, Forearm 43°8 mm.; third metacarpal

39°4 mm. (Peru. Nee . A, glaucus, p. 285.

f+. Forearm 372-405 mm.; third metacar ‘pal

33°7-362 mm. (C. America.) NEES Eee ee COMESONT. De 28S.

* Practically indistinguishable from A. 7. litwratus.

1908. } DR. KNUD ANDERSEN ON BATS. 313

c?, Molars 3. All species small: forearm 36'5-

46°8 mm.

e°, Cusp, 7 of m! less developed.

g'. m? not reduced in size; equal to 3—$

Oi TY Conc snasdsesonpoovnesdessoous | Ale COMERGIS Wo ZO:

#, Averaging smaller : “maxillary width of

skull (acr oss m!) 8-8°6 mm.; forearm

39-42 mm. (Para, through Guiana to

N.W. Venezuela, incl. Trinidad) ...... A. c. cinereus, p. 292.

gj. Averaging larger: maxillary width of

skull 85-89 mm.; forearm 41°2-

44 mm. (Colombia, to N.W. Vene-

AWE) coos iu... A. @. bogoténsis, p. 293-

h*. m2? considerably. reduced in size: > equal to

only about 4 of m!, Forearm 37°8-

39°8 mm. (Ecuador, Venezuela.) ......... A. rosenbergi, p. 293.

F?. Cusp 7 of m! more developed.

a4. Rostrum not unusually depressed and not

bent upward; bony palate not short-

ened; distance from palation to hinder

border of incisive foramina greater than

(or equal to) distance from palation to

basion.

k°®, Rather smaller : length of skull (to front

of c) 19°2-21'°2 mm.; upper teeth

(cm?) 6-7°2; forearm 37°5-44; third

metacarpal 33-40°5.

d6, Upper teeth (cm?) 6°5—7°2 mm.

a’, Upper side of interfemoral densely

haired . Spsncpbetsasoseaces | Lalo WOUGAEHIDS To ANG

as, Averaging Jarger : “forearm 39-

43°5 mm. Darker colour phas2

predominant ; facial stripes and

white ear-edgings as a rule want-

ing or indistinct, rarely strong.

(S. America, S. and C. Mexico.). A. ¢. toltecus, p. 297-

68. Averaging smaller: forearm 37°5—

39°77 mm. Light colour phase

predominant ; facial stripes and

white ear-edgings as a rule

distinct or strong, rarely want-

Tine, | (EREGCIOE,)) scp sovceacousenecs A. t. ravus, p. 300.

. Upper side of interfemoral sparsely

haired. Forearm 41°7-44 mm.

(Guiana, Pernambuco.) ............ A. quadrivittatus, p. 302.

e®. Upper teeth (c-m2)6 mm. No white

ear-edgings. Forearm about 38 min.

(Yucatan, Vera Cruz.) .. A. pheotis, p. 303.

1, Rather larger: length of skull (to front

of ¢) 22-99'8 mm. ; upper teeth (c—m?)

7-5-7°6; forearm 45-A6'8 ; third meta-

carpal 42-45. Interfemoral very short,

densely haired. Facial stripes in-

distinct or wanting; no white ear-

edgings. (S: Miexico:))--0-0--sesneee ae A. aztecus, p. 306.

j*. Rostrum “unusually depressed and slightly

bent upward; bony palate shortened :

distance from palation to hinder border

of incisive foramina less than distance

from palation to basion.

m®, Larger: length of skull (to front of ¢)

20 mm.; upper teeth (c—m2) 6°7; fore-

arm 40°5; third metacarpal 37. le

Mexico.) ..... . A. turpis, p. 307

n°, Smaller: length of skull 182-187 mm.

upper teeth 5°8-6°1 ; forearm 36°5— “38.

third metacarpal 32'2-35. (S. and C.

Mexico.) yds: ekwinseeascnesacenuaa neers A. nanus, p. 308.

314 DR. KNUD ANDERSEN ON BATS. [ Apr. 7,

General Remarks.

(a) Artificial and natural arrangement of the specics.—The

arrangement of the species of Artibews given in the foregoing

pages is based primarily on their number of molars: 3,3, or 2; it

has the practical advantage of facilitating the identification of the

species ; it is easier to cownt the teeth than to study their detailed

structure. But I have no doubt that it is thoroughly artificial.

Briefly epitomised the arrangement is this :—

A. 3 molars. ‘

a. Cusp 7 of m! small: no living species.

b. Cusp 7 of m! large: concolor, planirostris, hirsutus.

B. 3 molars.

a. Cusp 7 of m! small: glaucus, watsoni.

6. Cusp 7 of m! large: jamaicensis.

C. 3 molars.

a. Cusp 7 of m! small: cinereus, rosenbergi.

b. Cusp 7 of m! large: quadrivittatus, toltecus, pheotis, aztecus :—turpis.

NaANUS.

This and any other arrangement of the species based primarily

on the number of molars is, I believe, open to the following

objections :—Is it likely that the presence or absence of a rudi-

mentary tooth (m* and m,) is more important, in taxonomic respect,

than the general level of development at which the structure of

the upper molars (small or large cusp 7 in m') has arrived? It

has been pointed out in the foregoing pages that a certain small

percentage of individuals of the species with normally # molars

have 2 only (see 4. planirostris and hirsutus); that in the species

with normally 2 molars individuals occur which have 3 only

(see A. jamaicensis); and that a few individuals of species with

normally § molars have 3 (see A. rosenbergi and toltecus): bearing

this in remembrance, is it then likely that a character which is

vacillating among individuals of the same species and geographical

race is of primary, and a character which is constant within the

species (the structure of the upper molars) of secondary import-

ance? If the nwmber of molars were of fundamental importance

in this genus, is it then likely that we should find in the lowest

section (3 molars) species which have the most advanced structure

of the upper molars (planirostris, hirsutus), and among the

species of the highest section (2 molars) such as have retained a

low character (small cusp 7) in the upper molars? Is it likely

that A. jamaicensis, because it has lost the rudimentary m*

(although some of the races have as a rule retained the notch in

m° indicative of the former place of m*), is more closely related to

A. glaucus and watsoni, from which it differs in the structure of

the upper molars, than to A. planirostris, with which it accords in

this as in every other respect except the loss of a rudimentary

tooth? These and similar considerations seem, with necessity, to

lead to the conclusion that the old and till now universally

accepted arrangement of the species according to their number of

1908. | DR. KNUD ANDERSEN ON BATS. o1d

molars does not give an adequate idea of their true mutual

relationships. If, however, we take as the leading character, not.

the presence or absence of the vanishing m® and m,, but the

smaller or greater development of cusp 7 of m', all the objections

indicated above, as far as I can see, are removed, and we then

arrive at the following scheme :—

A. Cusp 7 of m! relatively small.

a. Molars 2: no living species.

6. Molars :

c. Molars

[to

: glaucus, watsoni.

os]

ro|to

a. m? of normal size: cinereus.

B. m2 reduced: rosenbergi.

B. Cusp 7 of m! large.

a. Molars 3.

a. Rostrum short: concolor.

6B. Rostrum normai: planirostris, hirsutus.

6. Molars 3 (species closely allied to planirostris) : jamaicensis.

ce. Molars 5 (none of the species closely allied to those of sections Ba and BA).

a. Rostrum and palate normal ; guadrivittatus, toltecus, pheotis, aztecus.

3. Rostrum unusually flattened, palate shortened ; turpis, nanus.

Or expressed in the form of a diagram :—

2 .

; molars: rosenbergi

;

| nanus

turpis § 2

molars: cinereus

€

ws)

\ aztecus 7}

x pheotis

toltecus

quadrivittatus

vu |

. 2 1

Jamarcensis 3

Ww bo

watsoni

a /

ahaa glaucus |

planirostris ) 3

\ hirsutus } 3.

totes danani

3 : pi

3 molars: conde aoe Va

316 DR. KNUD ANDERSEN ON BATS. [Apr. 7,

According to this arrangement there are two principal branches

of the genus: in the one, and more primitive, cusp 7 of mis pro-

portionally small, in the other it is largely developed. The

former, as being the more primitive, is quite naturally but poorly

represented among recent species; the latter is flourishing. Of

the lowest section ‘(molars s 3) of the primitive branch (small cusp 7)

no living species are known, but there are species with 3 molars

(glaucus, watsont), and 2 molars (cinereus, rosenberg?r) ; the latter

species, rosenbergi, has attained an unusually high development

(not only loss of m* and m,, but also reduction of m?and m,). Of

the higher branch (large cusp 7) all sections are represented by

recent species : % molars in concolor (peculiarly short-faced),

a is and ‘hirsutus, the two latter species very closely

allied ; = in MEISE SS otherwise differing in next to nothing

from planirostré is; 3 1m quite a number of species (quadrivitiatus,

toltecus, pheeotis, aztecus, turpis, nanus), among which turpis

and ranws mark the highest stage of development, in so far

as the rostrum has become unusually depressed and the palate

shortened.

(b) Artibeus planirostris and its races.—A. planirostris plani-

rostris has spread over the whole continental area from Central

Brazil to S. Mexico; the complete resemblance, even in average

size, between specimens from Mexico and S$. America, and the

absence of the race from any of the West Indian Islands, are

evidence that it has reached Central America (N. of the Nicaragua

depression) and Mexico in a very recent epoch, at all events at a

time when the Greater Antilles were separated from the mainland

by water of sufticient breadth to constitute an absolute barrier for

the spreading of the race from the latter into the former. A. p.

trinitatis is apparently confined to the Venezuelan coast islands,

Trinidad and Tobago, A. p. grenadensis to Grenada. These three

races are so extremely closely inter-related, being distinguishable

only by small average differences in size, that from a broader

point of view they may be regarded as one form. In relatively

strong contrast to this form stands the large-skulled, large-toothed,

and as a rule also externally larger-sized A. p. fallax, the true

home of which seems to be Guiana, a part of S. America which in

a late geological epoch constituted an isolated imsular area ; all the

Guianan specimens of A. planirostris I have seen (51 in number,

collected at different places, by different collectors, and at different

times) are A. p. fallax ; if, therefore, Guiana is the place of origin

of this race, it has spread from there to the Lower Orinoco

(Ciudad Bolivar) and Lower Amazons (Para) ; along the numerous

southern affluents of the Amazons it has probably made its way

to Peru (“ A. hercules,” apparently indistinguishable from 4A. p.

fallax).

1908. ] DR. KNUD ANDERSEN ON BATS. 317

tribution, and its bearing on a past connection of the West Indies

and the Central American mainland (see map, p. 319).—(1) The

seven races of Artibeus jamaicensis recognised in this paper fall

into two natural groups :— A. j. parvipes, yucatanicus, jamaicensis,

and equatorialis on the one side; A. j. lituratus, palmarum, and

preceps on the other side. The former group, which may be called

the “northern,” is distributed over Central America, South and

Central Mexico, and the Greater Antilles, and has sent a branch

southward, through the Cordilleras, as far as Kcuador. The

latter group, the “southern,” ranges from Paraguay and 8. Brazil

northward through South and Central America to Central Mexico,

and has sent a branch to the Windward Islands, as far north as

Guadeloupe.

(2) In the northern group of races the individuals are generally

considerably smaller; the angular notch in the posterior margin

of m’*, between its cusps 5 and 7, is generally as well marked as in

A. planirostris, sometimes reduced, rarely completely filled up ;

the coloration of the fur is chiefly of the dark type, the facial

stripes as a rule obsolete or less distinct. In the southern group the

individuals are generally considerably larger; the angular notch

in the posterior margin of m* is rarely well preserved, generally

more or less reduced, often completely filled up; the coloration

chiefly of the lighter type, with the facial stripes, particularly the

supraorbital stripes, as a rule well marked or even very strong.

(3) The close resemblance between A. j. parvipes, from Cuba,

and A. j. yucatanicus, from Yucatan,—closer than between A. j.

porvipes and its nearest eastern and southern neighbour (in San

Domingo and Jamaica), A. 7. jamaicensis—is evidence of a past

closer land connection (or approximation) between Cuba and

Yucatan, than between Cuba and San Domingo, or Cuba and

Jamaica,

(4) A. j. jamaicensis ranges from Central Mexico to Panama,

from Honduras to Jamaica, San Domingo, Porto Rico, as far

east as St. Kitts. The perfect resemblance between indivi-

duals from all these places is evidence of a past complete, or

nearly complete, connection between Jamaica and the coast of

Honduras and Nicaragua. I do not see any other reasonable

explanation of the fact that precisely the same race occurs in

Central America, Jamaica, San Domingo, and Porto Rico, whereas

Cuba is inhabited by a clearly different race, the relationships

of which are with the Yucatan, not with the Jamaican, race.

The past connection between Jamaica and the mainland may have

been complete ; but this assumption is not necessary to explain

the present distribution of A. j. jamaicensis; the Mosquito Coast

and Jamaica are, as well known, connected by extensive submarine

banks, the Mosquito and 8. Pedro banks (part of the latter above

the surface of the water); an elevation of about 100 fathoms

318 DR. KNUD ANDERSEN ON BATS. Lesion (i

would leave between the Mosquito Coast and Jamaica only two

channels, the wider 75 miles ; these channels would be so narrow:

as to form no serious barrier for the spreading of a bat.

The geological line of Seipadiauuon between the Larger and’

Lesser Antilles runs, probably *, between the Virgin ‘elses and

Anguilla; the Virgin Islands therefore belong geologically to the

Larger, Anguilla to the Lesser Antilles. Though Antillean and

Central American in origin, 4. 7. jamaicensis has in the course of

time crossed this line ; coming from west (Porto Rico) it has

spread a little east of the line, at least as far as St. Kitts. But

there, or in some neighbouring island, the range of the race seems

to terminate ; farther southward in the chain of the Lesser Antilles

(Guadeloupe, Dominica) is found a race (4. 7. preceps) which has

no direct phylogenetic connection with 4. 7. jamaicensis, but isan

offshoot of the 8. American A. 7. palmarum.

(6) The direct connection between the northern and southern

American continent has given 4.7, jamaicensis the opportunity of

spreading southward, through the Cordilleras, as far as Eenador,

where it has developed into the comparatively large-sized, large-

skulled, and large-toothed 4. 7. equatorialis.

(7) While it is safe to assume that 4. 7. parvipes, yucatanicus,

and jamaicensis are of “ northern,” i.e. Central American-

Antillean, origin, and A. 9. equatorialis an Andean offshoot of

this northern branch, it is equally beyond doubt that 4.7. lituratus

and palmarum are of 8. American origin, for the following

reasons :—first, 4.7. palmarum, from Venezuela, is so extremely

closely related to A. 7. lituratus, from Brazil and Paraguay, as to

be, for all practical purpose, indistinguishable ; given that 4. j.

lituratus is of 8. American origin, A. j. palmar um is therefore the

same; second, the range of A. 7. palmarwm extends from Vene-

zuela, through C. America to $8. Mexico, but the race is completely

absent from the Larger Antilles, an evidence that it has: reached

C. America and Mexico at a time when the Larger Antilles were

definitely separated from the mainland; third, whereas 4. je

jamaicensis 1s common (and equally common) everywhere in

rarer farther north in C. America, until in 8. Mexico it is

very vare, an additional evidence that 4.7. yamaicensis is the

indigenous race, 4. j. palmarum an immigrant from the south.

(8) The southern group has spread from the mainland over the

Windward Islands. Individuals from the Venezuelan coast

islands, Trinidad and St. Vincent, are indistinguishable from the

continental A.j. palmarum. But farther north, in Dominica and

Guadeloupe, they have developed into a slightly different race,

A, j. preeceps.

* A, Agassiz, Three Cruises of the ‘ Blake,’ i. p. 112 (1888).

DR. KNUD ANDERSEN ON BATS. 319

1908.]

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320 SIR RAY LANKESTER ON THE [Apr. 7,

2. On certain Points in the Structure of the Cervical

Vertebre of the Okapi and the Giraffe. By Sir Ray

Lankester, K.C.B., F.R.S., F.Z.8.

| Received March 14, 1908. ]

(Text-figures 60-71.)

Among the material relating to the Okapi which has heen

acquired by the British Museum (Natural History), is a fine

skeleton of a nearly but not quite adult male, obtained from

Major Powell Cotton. It is the skeleton of the individual the

skin of which was presented by that gentleman and is exhibited

in the public gallery.

I have made some study of this skeleton, comparing the bones

with those of the Giraffe. Since I commenced this study,

Professor Fraipont of Liége has published his finely illustrated

account of the specimens of Okapi preserved in the Museum of

the Congo State at Tervueren near Brussels.

The most important difference between the general bony

skeleton of Okapi and that of Giraffe—as distinct from the

cranium—is one which is presented by the last three cervical and

first dorsal vertebree of the two animals. A certain difference in

the form and proportions of the cervical vertebre—as between

Okapi and Giraffe—is what one expects as the necessary

correlative of the much greater length of the neck in Giraffe.

But the difference goes a good deal beyond this—as a glance at

the drawings given in text-figs. 60 and 61, of the vertebre,

cervical 5, 6, 7, and dorsal 1, at once shows.

The neural spines (ze. in the figures) of the cervical vertebree

of the Giraffe are much shorter proportionately than are those of

the Okapi—and this is especially the case in cervical 7. Further,

the inferior transverse processes (éi. in the figures)—lateral out-

growths which in the mammalian vertebral series are peculiar to

the cervical region—are very different in the Giraffe from those

of the Okapi. In the Giraffe they are of small proportional size,

entirely anterior in position on each vertebra (see text-fig. 60).

In the Giraffe a right and a left inferior transverse process exist

on the seventh cervical vertebra—as well as on the vertebre in

front of it.

Not so in the Okapi (see text-fig. 61). Whilst cervical 5 (as well

as 4 and 3) hasa large inferior transverse process (I speak of the side

view as given in the drawing and therefore of one only of the pair

of lateral processes) which grows downwards (abaxially) from the

anterior part of the vertebra—and is larger than the correspond-

ing process in Giraffe—cervical 6 has its inferior transverse

process in the form of an enormous flange or plate extending the

whole length of the vertebra. This does not exist in Giraffe:

in that animal the inferior transverse process (é. text-fig. 60) of

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322 SIR RAY LANKESTER ON THE [Apr. 7,

cery. 6 is but little bigger in depth and extension than is that of

cerv. 5 or cerv. 7.

Text-fig. 61.

OKAPL

The last three cervical vertebre and the first dorsal vertebra of the Okapi

(Okapia johnstoni), seen trom the left side.

Lettering as in text-fig. 60.

Note the great size and backward extension of ¢i. of Cerv. 6, the complete absence

of ¢. from Cerv. 7, and the approximation in general proportion of Cerv. 7 to

Dors. 1.

The next difference is that in the seventh cervical of Okapi

there is no inferior transverse process at all; whilst the superior

transverse process és. is greatly enlarged, expanded and flattened

at its free end. No such appearance is presented by the seventh

cervical of the Giraffe, which contrasts very strongly with that

ofthe Okapi. In fact, the seventh cervical of the Giraffe has the

general appearance and character of the cervical series, whilst the

seventh cervical of the Okapi is, in all its characters except that

of actually giving articulation to a bony rib, a dorsal. We may

describe this by saying that the last cervical is “‘ dorsalized.”

1908. } OKAPI AND GIRAFFE. 323

The contrast in this respect between Giraffe and Okapi is very

great, as text-figs. 60 and 61 clearly show.

Text-fig. 62.

BOS TAURUS.

‘The last three cervical vertebre and the first dorsal of a domestic Ox (Bos taurus),

seen from the left side.

Letters as in text-fig. 60, except /.z.p.(r.), which points to the rudimentary

posterior lateral zygapophysial process of the first dorsal vertebra.

Note the close agreement of these five vertebrae with those of Okapi and

their difference from the same group in Giraffe.

It may perhaps be possible to show that these differences of

proportion in the neural spines and transverse processes are

connected with the special mechanism of the very long neck of

Giraffe. The greater size and breadth of these bony processes is

merely an expression of the greater size of the muscle-tendons and

of the ligaments, of which they may be regarded as but denser

324 SIR RAY LANKESTER ON THE [Apr. 7,.

ossified parts. The fact, however, as we have seen, is that the long-

necked Giraffe presents a smaller development of these processes of

the vertebra of the cervico-dorsal region than does the shorter-

necked Okapi; the Giraffe’s cervicals (from cerv. 3 backwards) are

practically all alike, and even the first dorsal is not so marked in

the contrast it presents to the cervicals as is the first dorsal of Okapi

to the cervicals in front of cerv. 7, which is, itself, strongly

dorsal in character in the contrast it presents.

The sloping forwards of the neural spine of the first dorsal

of Giraffe and the inclination of the plate formed by the superior

transverse process and anterior zygapophysis in that vertebra is

similar to that of the vertebree in front of it (see text-fig. 60, ts.

& 1.z.a.).

The condition of the vertebre cerv. 5, 6,7, and dors. 1, in

Okapi, thus seen in lateral view, is however not in any way

peculiar to Okapi. It is the condition common to the Artiodactyle

Ruminants, as is shown by the drawing of the same four vertebre

of the common Bovine (Bos sp.) given in text-fig. 62. In all the

features above noted, in which the vertebre of Okapi differ from

those of Giraffe, Okapi agrees with the Bovine. The large flange-like

development of the inferior transverse process of the sixth cervical,

giving it a wide posterior extension, is present in all Cavicorn and

Cervine genera. It is, moreover, as well marked in the long-

necked Camel as in the short-necked Bovines, and is present

in the non-ruminant forms, the Pigs and the Hippopotamus.

In the Camel (text-fig. 63) the inferior transverse process of

the fifth, fourth, and third cervicals is large, plate-like and tri-

angular, wanting the large posterior growth characteristic of the

sixth cervical.

But in the Pigs—presumably a more primitive stage of Artio-

dactyle development than that presented by any of the Ruminants—

the inferior transverse process has a broad square outline (with

posterior region well developed), in the fifth and fourth cervicals

as well as in the sixth, though the process is biggest in the sixth

and totally absent in the dorsal-like seventh. In the third

cervical the inferior transverse process is much less in lateral out-

growth, but still has a strongly developed posterior region project-

ing backwards below and behind the superior transverse process.

The inferior transverse process of the cervical vertebre is well

developed in the Carnivora as in the Pig, the special enlargement.

and plate-like character of that of the sixth cervical being as in the

Ungulata.

In the Insectivora—as shown more especially by Centetes—the

superior transverse process of cervicals 7, 6, 5, 4, 3, 2, and even I,

is well developed, and it is only on cerv. 6 that the inferior trans-

verse process is developed to any size; cervicals 5 and 4 have

a small development of it. In the Hedgehog the inferior

transverse process of cervical 6 is enormous and grows downward

and backward on each side of the neck as a very obvious and

striking pair of plates.

325

OKAPI AND GIRAFFE.

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te) ON

326 SIR RAY LANKESTER ON THE | Apr. 7,

The inferior transverse process is to be regarded as a bifurcation

of (or an accessory inferior plate of) the one transverse process

which is usually recognised (in the cervical region) as the superior

transverse process. On the present occasion it is not convenient

to discuss further its morphology.

Text-fig. 64.

OKAPT. OKAPT.

POSTERIOR FACE. ANTERIOR FACE

The posterior face of the seventh cervical vertebra and the anterior face of the

first dorsal vertebra of the Okapi (Okapia johnstoni), to show the duplicated

character of the articular facets.

m.2.po., posterior median articular facet of Cerv. 7; m.z.a., anterior median articular

facet of Dors. 1, which articulates with the foregoing; J.z.p., posterior lateral

articular facet of Cerv. 7, raised on a distinct zygapophysis; U.z.a., anterior

lateral articular facet also raised on a zygapophysis, seen fully in Dors. 1 (and

partially in Cerv. 7); U.z.a. of Dors. 1 articulates with l.z.p. of Cerv. 7;

ts., Superior transverse process.

There are a number of interesting details to be observed and

discussed in regard to these minor processes of the vertebra in

different groups of mammals. My purpose is not now to enter on

that subject, but merely to show briefly what is the value of the

1908. } OKAPI AND GIRAFFE 327

difference between Okapi and Giraffe in regard to the inferior

transverse process of the cervical region—when the chief facts as

to this structure in other mammals are taken into view. Clearly

enough it is Giraffe which is altogether exceptional,-novel and

specialised, not archaic or atavistic. Giraffe has not even the great

plate-like inferior transverse process on its 6th cervicals, which

is obvious and prominent in such widely separate forms as the

Text-fig. 65.

GIRAFFE. GIRAFFE.

POSTERIOR FACE. ANTERIOR FACE,

The same view of the same vertebra in Giraffe as that given in the case of Okapi

in text-fig. 64. The figure shows the single pair of articular facets raised on

zygapophyses. No median facets on the sides of the neural arch are developed.

Letters as in text-fig. 64, except ¢7., inferior transverse process of Cerv. 7.

Hedgehog, the Carnivora, and the commoner Ungulata. Okapi

merely agrees with other Ruminant Ungulates in the matter

of its adjacent cervical and dorsal vertebre, and they seem to

be a little more specialised, than the Pigs and Perissodactyles, in

having a large inferior transverse process only on the 6th cervical

and quite small ones on the vertebre in front; whereas Pigs and

Perissodactyles have that process more equally developed on all

the cervical series 3, 4,5, 6. The emphasis of the inferior trans-

verse process on cerv. 6 appears to be the rule in Mammalia and

328 SIR RAY LANKESTER ON THE [Apr. 7,

is carried very far in Hrinaceus. Whether the existence of an

inferior portion or inferior transverse process should be regarded

as a primitive feature of all these vertebree in Mammalia is not

clear. It certainly seems to belong as an original element of

structure to cervicals 3, 4, 5 and 6—to be increased greatly

in size on cerv. 6 in most groups, and sometimes to practically

disappear from the other cervicals whilst remaining there.

Text-fig. 66.

OKAPI. oO KAPI

POSTERIOR FACE. ANTERIOR FACE. POSTERIOR FACE.

View of the articular surfaces which connect dorsal 1 and dorsal 2 of the Okapi, and

of the anterior half of the articulation between dorsal 2 and dorsal 3. The

articular facets are seen to be single pairs and to consist of an oval surface

(m.z.p., m.z.a.) placed entirely on the neural arch. The left anterior articular

facet of dorsal 2 (middle figure) shows a tendency to divide into two.

Letters as in text-fig. 64, with the addition of cos., process for the articulation of the

tubercle of the first rib; ¢.p., concave facet for the head of the second rib ;

x., hypapophysial tubercle.

In Giraffe, as a marked exception, it is small in all the cervicals

and smaller in cerv. 6 than in the cervicals in front of that one,

indeed so much reduced that it has no posterior extension at all

but is represented by a small triangular anterior growth only.

1908. | OKAPI AND GIRAFFE. 329

DousLE ZYGAPOPHYSIAL ARTICULAR SURFACES IN OKAPT.

The posterior face of cerv. 7 and the anterior face of dors. 1 of

the Powell Cotton skeleton of Okapi are drawn in text-fig. 64 in

order to show a remarkable condition of the articular surfaces which

bring these two vertebrz into relation. Whilst ceryv. 6 of Okapi

has only the usual single pair of anterior and posterior articular

facets, common in cervical vertebre, and situated on the

zygapophyses themselves, the seventh cervical of this specimen of

Okapi shows on the posterior face an additional mediad pair of

Text-fig. 67.

GIRAFFE. GIRAFFE GIRAFFE.

POSTERIOR FACE. ANTERIOR FACE. POSTERIOR FACE.

A similar view to that given in text-fig. 66 of the corresponding vertebre of the

Giraffe. The posterior face of dorsal 1 and the anterior face of dorsal 2 show

duplicated facets (l.2.p. and m.z.p., l.z.a. and m.z.a.) wnlike dorsal 1 and

dorsal 2 of Okapi, but dike cervical 7 and dorsal 1 (see text-fig. 64) in that

animal. The lateral element, however, entirely disappears from the articular

surfaces connecting dorsal 2 and dorsal 3, as shown by the view of the posterior

face of dorsal 2.

Letters as in the preceding text-figures.

articular facets quite distinct from the lateral pair (text-fig. 64,

m.z.p., cerv. 7), whilst the anterior face of dors. 1 drawn in the same

figure shows two articular surfaces on the anterior face (/.2.a.,

M.2.0.).

If we pursue the enquiry as to the articular surfaces, we find

on examining the posterior face of dors. 1 and both faces of dors. 2

330 SIR RAY LANKESTER ON THE [Apr. 7,

as shown in text-fig. 66, that itis obvious that the mediad articular

surfaces which co-exist in cerv. 7 (posterior face) and dors. 1

(anterior face) with the lateral articular surfaces, are new and

independent morphological entities and are identical with the

articular surfaces of the dorsal vertebra, whilst the laterally

placed articular surfaces of the cervicals raised upon distinct

‘‘ 7ygapophyses” have no existence in the dorsal series exeept on-

the anterior face of the first dorsal.

It seems to me that we are entitled to conclude from the

specimen here figured (text-figs. 64 and 66) that the zygapophyses of

the cervical vertebrae of the Mammalian series are not merely in

a different position from that occupied by the articular facets of

the dorsal vertebre, but that the cervical and the dorsal articular

surfaces are distinct morphological entities. The articular facet is

not bodily “ shifted” in position, when we pass from cervical to

dorsal, but a distinct and independent mediad facet is substituted

for the lateral facet. At the same time it must be recognised

that the two articular facets can become confluent, and that one

is to be regarded as an extension and “ pullulation” of the other.

So far as I am aware, this is a new observation.

I hasten to say that in the cerv. 7 and dors. 1 of another

skeleton of Okapi (the property of Mr. Walter Rothschild) the

two articular facets of each side drawn in text-fig. 64 are not

marked off from each other, but confluent and ill-defined. The

vertebre of that skeleton differ in many remarkable points of

size and proportions from those of the Powell Cotton skeleton.

Though the Rothschild skeleton is that of a very young animal

far from complete in growth, whilst the Powell Cotton skeleton

is that of a full-grown animal with nearly adult dentition, yet

many parts of the vertebre of the younger animal are much

larger than the same parts in the older animal. At the same

time in other details the latter shows the greater size. It

is possible that the Powell Cotton specimen is exceptional and

abnormal, or that it belongs to a local race differing from that to

which the Rothschild skeleton belongs. Or again, and this I think

to be the most likely case, it seems from the variability of Okapi

in regard to the striping of the skin and various proportions of

the skull, also as to the symmetry of the horns of either side

and as to the molar teeth, that there is great range of variation in

the species—and that this variability extends even to such points

as the exact form of the vertebree and the development of articular

facets connecting successive vertebra. It is also possible that the

absence of a distinct second pair of facets at the articulation

between cerv. 7 and dors. 1 in this particular specimen of Okapi

is due to its immature stage of growth.

A comparison of these vertebre of Okapi with those of Giraffe—

in regard to the zygapophyses and articular surfaces or facets—

becomes now especially interesting.

In text-fig. 65 the same views of the two vertebre (cerv. 7 and

1908. ; OKAPI AND GIRAFFE. 331

dors. 1) of Giraffe are shown as are shown in the case of Okapi in

text-fig. 64. It is at once seen that there is no question of the

presence of a second pair of articular surfaces, of mediad position,

in Giraffe, so far as this articulation is concerned.

Let us now go on to examine the posterior face of dors, 1, and

both faces of dors. 2 in Giraffe. These are drawn in text-fig. 67

for comparison with similar views of the same vertebrae of Okapi

drawn in text-fig. 66.

The remarkable fact is at once obvious that the articulation

between dorsal 1 and dorsal 2 in Giraffe shows some of the

characters of the articulation between cervical 7 and dorsal 1 of

the Powell Cotton Okapi. There is on the posterior face of the

Giraffe’s dors. 1, a lateral and a median articular surface which

are not separate from one another but confluent. And the same

is true as to the corresponding articular surface on the anterior

face of the Giraffe’s dorsal 2.

The lateral articular facet belonging to the true zygapophysis of

the cervical vertebre is in Giraffe continued into the dorsal

series—and does not disappear until we come to the articulation

between dorsal 2 and dorsal 3 (see below as to Rhinoceros). In

fact the first dorsal of the Giraffe is in this respect drawn (as it

were) into the cervical series. The break in the vertebral series

which occurs in Okapi (and normal Ungulata) between the anterior

and posterior faces of dorsal 1 is in Giraffe pushed down the

series and shows itself in the contrast between the anterior and

posterior faces of dorsal 2. There is, what has been called in

regard to such serial metameric elements of structure, ‘ homceosis ”

of the first dorsal of the Giraffe—assimilating the articular facets

of that vertebra to those of the seventh cervical.

The complete investigation of this question of the characters of

the last cervicals and first dorsals—and the transition from the

one group to the other and the greater or less abruptness of the

break between them in the whole Mammalian series, would form

an interesting enquiry.

At present I must content myself with formulating the facts,

Jirstly, that in both Okapi and in Giraffe there is (as an exception

in Ungulata) a co-existence of lateral and mediad articular facets

of independent morphological value—at one of the vertebral

articulations at the base of the neck: and secondly, that the

articulation at which this occurs is in Okapi that between

cervical 7 and dorsal 1—whereas in Giraffe it is shifted one place

backwards in the series and occurs between dorsal 1 and

dorsal 2.

The relation of these peculiarities to the elongation of the

cervical region or to any other peculiarities of the animals in

question, is a matter for further enquiry.

I am able to add to this the following additional observations

which I have made in the Museum of the Royal College of

332 SIR RAY LANKESTER ON THE [Agora i,

Surgeons*, In the Ruminants generally the substitution of

Text-fig. 68.

Dors.|. Left side

ANTERIOR FACE,

Cerv. 7. Left side

POSTERIOR FACE.

The articular facets of the left side on the posterior face of Cervical 7 and

anterior face of Dorsal 1 of Rhinoceros sumatranus. 2 nat. size.

N.C., neural canal. To show single lateral facet.

/|\ Dors.2. Left side

Dors.!. Left side |

) ANTERIOR FACE,

POSTERIOR FACE.

The articular facets of the left side on the posterior face of Dorsal 1 and

anterior face of Dorsal 2 of Rh. sumatranus. 2 nat. size.

To show elongated facet, including lateral and mediad factors.

the mediad pair of facets for the more laterally placed pair

* J desire to take this opportunity of pointing out that though the osteological

series of the Hunterian Museum is a most valuable and useful one, it is yet not very

large and does not contain second and third specimens for comparison. Such a

collection as the zoologist really requires ought to be formed at the Natural History

Museum. The osteological collection of that museum is at present very small

and not such as is necessary for reference and comparison. A large room in the

basement was arranged by me to receive such a collection, which I had intended to

keep mounted on flat boards and movable: the boards to be fixed as sliding-shelves

in dust-tight cabinets. An assistant was appointed in 1907 for the purpose of

making and looking after this proposed collection.

1908. | OKAPI AND GIRAFFE. 333

takes place abruptly and sharply at the articulation between

dors. 1 and dors. 2. Occasionally a trace of the lateral process is

retained on the posterior surface of dors. 1, as shown in text-fig.

62, l.x.p. on dors. 1. Inthe Pigs and the Hippopotamus the same

is the case.

Text-fig. 70.

Dors 2 Left side | Dors.3. Left side

POSTERIOR FACE. | i ANTERIOR FACE.

Similar drawing, to show the duplicate facets connecting Dorsal 2 and Dorsal 3

in the same animal. # nat. size.

Text-fig. 71.

Dors.4. Left side

ANTERIOR FACE,

Dors.3. Left side

POSTERIOR FACE.

Similar view, to show the single facets (now the median not the lateral factor)

connecting Dorsal 3 and Dorsal 4 in the same animal. nat. size.

Tn Tapir and Horse the articulation between dorsal 1 and dorsal 2

does not exhibit an abrupt change, but the facets are intermediate

in position to those shown in the joints in front and behind.

In the Carnivora (Cats, Dog, and Fox) the break occurs at the

joint between dorsal 2 and dorsal 3: the anterior joint (that

between dorsal 1 and dorsal 2) resembling that of the cervicals.

334 ON THE OKAPI AND GIRAFFE. [PAwors aie,

The Rhinoceros is the only case in which I found actually two.

pairs of facets marked out, almost but not quite as separate

from one another as in Okapi. In several specimens of this

genus I find that the facets between cervical 7 and dorsal 1 are

purely lateral: between dorsal 1 and dorsal 2 the facets are very

large and each is incompletely separated into a lateral facet

and a mediad facet. Between dorsal 2 and 3 this is even more

marked: there are two facets on each side of each of the

articulating vertebree. On the posterior face of dorsal 3—and

the anterior face of dorsal 4—we find only the mediad pair of

facets. These articulations are shown in the four drawings, text-

figs. 68, 69, 70, & 71. In Rhinoceros then the transition is quite

gradual—from the “cervical” condition of the joint to the “dorsal”

condition—the articulations dors. 1 : dors. 2, and dors. 2 : dors. 3

showing clearly the lateral or cervical facet as well as the mediad

or dorsal facets on each side. This observation requires of course

further confirmation and may prove not to indicate a rule without

exception when a larger series of Rhinoceros is examined.

Thus we may tabulate the conditions in regard to this matter

in Ungulata as follows :—

Normat 1n RUMINANTS :—

med.

sue Q

Cerv. 7<—>dor s. 1. dors. Vicar s. 2. dors. 2<—-—>dors. 8. dors. 3.

OKAPI :— v

rs) es a a

3 2 ro) ®

=e a =

Cerv. 7<-->dors. 1. dors. 1<—>dors. 2. dors. 2<-->dors. 3. dors. 3.

GIRAFFE :—

: 3s mS ro

Cerv. 7<—>dors. 1. dors. 1<-—sdors. 2. dors. 2<-—>dors. 3. dors. 3<_>dor Ss. 4.

RHINOCEROS :— B_ 3 = =

Ee Se zz Z

Cerv. 7<—>dors. 1. dors. 1<-—>dors. 2. dors. 2<-—> dors. 3. dors. 3<-—>dors. 4.

The multiplication of articular facets between successive

vertebre is of course well-known in another region of the

Mammalian vertebral column. At the lumbar end of the dorsal

series in the Great Anteater (/yrmecophaga jubata) we find that

dorsal 12 exhibits no less than three pairs of zygapophysial facets—

the most dorsal and median of which, though horizontal in dorsal

12, becomes tilted to a vertical position in dorsal 13 and the

following vertebree

1908. | MR. H. R. HOGG ON AUSTRALASIAN SPIDERS. 33D

3. Some Australasian Spiders.

By H. R. Hoae, M.A., F.Z.8.

[Received March 16, 1908.]

(Text-figures 72-75 *.)

Genus Missunena Walck.

Missunena (Erropon) occatoria Walck.

Mr. W. J, Rainbow, F.L.S., of the Australian Museum, Sydney,

was good enough to send me a female of the above genus, one of

three found by him at Enfield, a suburb of Sydney.

Text-fig. 72.

Missulena occatoria Walck.

a. Eyes from above. ec. Tarsal claws.

6. Under side of body. ~X 2. d. Profile. Nat. size.

This agrees so closely with the description given by M. Lucas

in 1865 (Ann. Soc. Ent. Fr. ser. 4, vol. v. p. 309, pl. 8) of the

specimen identified by him as the above, the type species of the

genus, by comparison with Walckenaer’s original type, that I have

no doubt of its being the same.

I give the following particulars as so little is known of the

species; and as the two specimens (the type and M. Lucas’s)

* Nore.—The figures given in the eye-diagrams are ;4, millimetre.

Proc. Zoot. Soc.—1908, No, X XII, 99,

336 MR. H. R. HOGG ON AUSTRALASIAN SPIDERS. [Apr. 7,

formerly in the Museum of the Jardin des Plantes, Paris, have

disappeared, this is probably the only specimen in Hurope now

available for reference.

The cephalothorax, mandibles, legs, and palpi are uniformly dark

red-brown, smooth and shiny, the abdomen dark yellow-brown.

The thoracic fovea is long, deep and procurved, and three deep

radial indentations on either side of the thoracic part.

The four lateral eyes and front median pair are of the same

size * (3), the laterals being raised, the medians sessile, their

diameterapart. The intermediate row sessile and slightly smaller

than the others (24). They lie just above the level of the front

row, which is straight and two-thirds of an eye-breadth from the

margin of the clypeus. The distance between the posterior

laterals is 43; between anterior and posterior laterals at each

side 8; between the posterior laterals and intermediates at each

side 9 ; between the intermediates 28; between the intermediates

and the anterior median at each side 104; and between the

anterior laterals and the medians at each side 20.

The rastellum consists of three rows of short stout spines. On

the inner margin of the falx-sheath are about 9 large teeth. A

few small intermediate and two large teeth, with thick fringe, on

the outer margin.

There are numerous club-shaped spines on the fore part of the

lip and on the maxille ; the latter very square with a protuberance

on the outer upper corner. The lip is rounded in front and twice

as long as broad.

The sternal! sigille are large and away from the margin.

The legs areshort and stout. The three tarsal claws and female

palp-claw are short, stout and well curved, with one long tooth

near the base in each.

The abdomen is oval, high, slightly overhanging the cephalo-

thorax, and thickly covered with smooth downlying hair. The

spinnerets are short, the first joint of the superior pair is twice

the length of the second; the inferior pair are as long as the

second joint of the superior.

The measurements (in millimetres) are as follows :—

Long. Broad.

Cephalothorax ... 83 ; Hy Eno

Abdomen ......... 1S 11

Werchinec 33 horizontally. 4 broad each.

Diedks, 63 vertically.

Trochanter Patella Metatarsus

Coxa. &femur. & tibia. & tarsus.

Hepes) $0 208tue. l. 34 6 5 t= 19

De | hes 6 5 4i = 19

oe MES 63 5 5 = 20

4, 5 8 6 Co) oe

Paonia seat ae 3 5 LET ee 1S)

Superior spinnerets ...... 14, #=23. Inferior 3.

* The figures are tenths of a millimetre.

1908. | MR. H. R. HOGG ON AUSTRALASIAN SPIDERS. 337

These leg measurements are correct; they do not agree with

L. Koch’s, but he did not have a specimen and does not give his

authority.

Genus HexATHELE Ausserer.

Our knowledge of the small and little-known genus Hewathele,

whose sole habitat lies in New Zealand, has recently been added

to by two specimens from Auckland sent to the British Museum

by the Rev. W. H. Webster; these and some others herein referred

to, by the courtesy of Mr. A. S. Hirst, in charge of the collection

of Arachnida, I have been able to examine.

The want of precision in the descriptions of some of the known

specimens from different localities seemed at one time to render it

doubtful whether they were really different from the type species

H. hochstetteri Auss., of which the locality is only known as New

Zealand ; but, on revising those I have been able to see, with what

has been written about others, I think it may now be taken that

there are four separable species. Mr. Goyen’s H. petreii from

Otago is certainly very like the type species, but he specifies three

points which if correctly described will make his species good.

The male and female described by myself (Proc. Zool. Soc. 1901,

276) from Pahiatua, Wellington, N.Z., as H. hochstetterr, will

form another, and J now record it as Hexathele huttona, sp. n.,

after the sender the late Capt. Hutton. Those of Mr. Webster’s

are a fourth. They may be distinguished as follows :—

a. Front median eyes one half their diameter apart.

Cephalothorax longer than patella cwm tibia iv. ............ HL. huttoni, sp. n.

6. Front median eyes their diameter apart.

Cephalothorax not longer than patella cam tibia iv.

a, Front median eyes the same distance from the front

side eyes as from one another.

a*, Front and rear rows of eyes of equal Jength ; no spines

on femoral or patellar joints of legs (sec. L. Koch).

Cephalothorax shorter than patella cwm tibiaiv. ... H.hochstettert Auss.

62. Rear row of eyes shorter than front row; spines on

femoral and patellar joints of all legs (sec. Goyen).

Cephalothorax as long as patella cum tibia iv. ...... H.. petreii Goyen.

61, Front median eyes not more than half their diameter

from side eyes of same row. pea ee shorter

than patella cwm tibia iv. .......... ponoovgctepesscooss Saks GAPARIAP I, Bo 5b

The description of H. huttoni I have already furnished (Joc. cit.)

and now append that of H. websteri.

HEXATHELE WEBSTERI, Sp. n.

Cephalothorax pale yellow-brown, almost bare, with a few short

brown hairs at sides, slightly darker in the depressions and dark

brown between the eyes. The mandibles are red-brown with

long brown bristles on the inner side. Fangs black-brown. Pale

orange fringes.

Maxille, lip, sternum, legs, and palpiyellow-brown. Abdomen

yellow-brown, thinly covered with yellow-brown downlying hairs,

longer and dar ker brown anteriorly; no discernible pattern.

Hairs on under side darker yellow-grey, spinnerets the same.

QDO*

338 MR. H. R. HOGG ON AUSTRALASIAN SPIDERS. Pore

The cephalothorax is flat, nearly bare, only slightly shorter than

patella cwm tibia iv., 2 mm, longer than broad ; three side streaks ;

the transverse depression at end of cephalic part deep, short, and

recurved; sides slightly rounded. The head part is all clearly

raised above the thoracic, highest in front, and the eyes ona

raised hillock about twice as long as broad. ‘The first row of eyes

is straight when seen from above but clearly procurved viewed

from in front; the side eyes being almost close to the margin of

the clypeus. ‘The middle eyes of the front row are their diameter

apart and half that distance from the side eyes and rear middle.

The rear row is recurved; the oval laterals being as large as the

same of the front row. The middle eyes of the rear row are-

nearly round.

Hewxathele websteri, sp. u.

a. Eyes. | b. Profile. | c. Under side of abdomen.

The mandibles are prominent, longer than front patelle without.

rastellum ; a row of eleven large and two smaller teeth on inner

1908. ] MR. H. R. HOGG ON AUSTRALASIAN SPIDERS. 339

side of falx-sheath, a thick fringe on outer, and a median row of

6 or 7 quite small teeth at lower end.

The lip is broader than long, slightly hollewed in front, with a

moderate number of dark brown short club-shaped spines in front.

The latter also occur on inner lower corner of maxille; on the

upper inner corner is a rounded apophysis. Light yellow-brown

upstanding hair on both maxille and lip.

Sternum longer than broad, hollowed in front and pointed at

rear end. Two moderately large sigille in hollow by lip, and

three smaller each side nearly marginal.

The abdomen is oval. The superior spinnerets two-thirds the

length of metatarsus of fourth pair of legs, tapering and rather

divergent; the third pair of spinnerets being situated in a line

with them just at their base, and the median pair lying between

them.

The first, second, and third pairs of legs are of equal length.

Hight or nine small teeth le slopingly across the superior claws ;

inferior claws without teeth.

There are no scopula on any of the legs in the female.

This would appear to differ from H. hochstettert Ausserer, besides

being a good deal larger, in having the front middle eyes their

diameter apart but less than half that distance from the side

eyes, instead of the same distance. The row is straight instead of

procurved. The lip is broader than long instead of same breadth.

The superior spinnerets are only two-thirds the length of meta-

tarsus iv. stead of (sec. Koch) the same length.

The measurements (in millimetres) are :—

Long. Broad.

Cephalothorax ... 102 f vee front.

‘ | 83

PNoelOraN EI 465 soooc 9 54

Mandibles....... 6

Pat. & Metat.

Coxa. Tr. & fem. tibia. & tars.

Wegsyeccc scat 1 5 9 9 Op MESO

2 5 9 9 Qh. 3 ABO

3. 4 8 OA Tle pie Sy

9-4

4, 44 10 oe ee oo,

IPAWOHE Lonacds oeGop 5 6 7 A ee

Superior spinnerets-..... )27 912) 227 G:

There are two females from Waiuku, Auckland, N. Z., sent by

the Rev. W. H. Webster, after whom I have named them.

The following are from a small collection of Spiders forwarded

to the British Museum, Nat. Hist. Dept., by the late Capt. Hutton

and Mr. C. H. Tripp from Pitt Island, one of the Chatham group,

some 500 miles east of New Zealand.

340 MR. H. R. HOGG ON AUSTRALASIAN SPIDERS. [PANpId,

This includes two species of Dolomedes Latr., both apparently

new, and I therefore describe them below. The only two species

recorded from New Zealand are nearest to them in the genus.

L. Koch (Die Arach. Austr. vol. ii. p. 858) makes the following

synopsis of his Australian and New Zealand species :—

A. Cephalothorax longer than tibia iv.

Che eaten Kae sce eee cen cater ee

b. Eyes of second row markedly larger than the middle

eyes of first row.

a. No scopula on legs.

61. The legs furnished with a scopula.

a*, Third pair of legs shorter than the second.

a’, Scopula on front two pairs only (sec. Koch).

(L willadd) Eyes of 2nd and 3rd rows of equal size ... D. minor, L. K.

b3. Scopula on all legs. Eyes of 2nd row clearly (from New Zealand).

larger than those of 3rd.

a+, Scopula on tarsi and metatarsi. Mandibles

as long as front patella. Hyes dingy

brown or black. Hair on abdomen

upstanding, rough and coarse ............ D. huttoni, sp. n.

6+, Scopula on tarsi only. Mandibles longer

than front patella. Eyes bright topaz-

yellow. Hair on abdomen downlying,

smooth and fine ............ soncsagcuosen IDs UMD DOy IDs Vale

62. Second and third pairs of legs of equal length.

Scopula on all tarsi and metatarsi (sec. Koch). D. imperiosus L. K.

(from New Zealand).

DOLOMEDES HUTTONI, sp. n.

The cephalothorax is dark red-brown with short, smooth, yellow-

brown hair, paler and more bristly round the margin and on the

eye-space. The mandibles, hp, and maxille black-brown, with

rather long upstanding brown hair and light red fringes.

Sternum and coxe somewhat lighter brown, with thick yellow-

brown hair and some upstanding dark brown hair mingled there-

with.

Legs and palpi red-brown, thickly covered with yellow-brown

hair. Abdomen above brown with dark yellowish-brown hair.

A paler median longitudinal stripe on anterior half and two pairs

of dark muscle-spots, followed by five transverse corrugations, the

last close to posterior end. On under side thick rough yellow-

brown hair, with a depressed shield pattern on which are two

darker narrow longitudinal lines.

The cephalothorax is rounded at the sides, the cephalic part

only slightly raised up; a long deep longitudinal fovea and well-

marked side stripes on the thoracic part. The eyes of the second

row are not quite twice the diameter of the front median, from

which and from one another they are the latter’s diameter distant.

The front laterals are three-quarters the diameter of the median

which are that distance apart, side eyes from middle rather less.

The oval eyes of the third row are smaller than those of the

second, but larger than the front median.

The margin of the clypeus is three diameters of the front

median eyes distant from them.

1908. ] MR. H. R. HOGG ON AUSTRALASIAN SPIDERS. 34]

The mandibles are kneed at the base and as long as the patella

of the front pair of legs; on the inner margin of the falx-sheath

are four large teeth, and on the outer margin one large between

two small teeth and a thick fringe.

The lip is as broad as long, straight in front, and rounded at

the sides, about half the length of the maxille, which are well

rounded on the outer side and upper half.

Dolomedes huttoni, sp.n. Female.

a. Hyes*. 6. Epigyne. | c. Male palp.

The abdomen is straight in front, broadest two-thirds of its

length down and rounded at posterior end; the hair coarse and

rough.

* The figures are tenths of a millimetre; the figure 26 between the oval eyes of

the third row should be 30.

342 MR. H. R. HOGG ON AUSTRALASIAN SPIDERS. [Apr. 7,

The legs are stout with scopula on all the tarsi and metatarsi,

on the under side of which latter are long powerful spines. There

iS one spine on tibia iv. above, none on the others. The

rather long superior claws have about 7 short pectinations, The

inferior claw is bare.

The palpi have tibial joint slightly longer than the patella and

the anterior end of the distal joint is rather club-shaped.

The measurements of the female (in millimetres) are as follows.

The male is rather smaller.

Long. Broad.

Cephalothorax ... 134 1 1 soe rom

INndomlen! seer eee 143 10

Mandibles ......... 6

Pat. & Metat.

Coxa. Tr. & fem. tib. & tars.

(53 73

NOES Rane. ee it 44 11 { 3 12 = 404

2 4s 103 123 ils = BY

3 4 10 1 ill = 37

4 43 13 14 14 = 454

Pallipl eee secs aes 24 6 5 5 = 184

There are nine females and seven males (nearly all immature)

from Pitt Island (Chatham Islands).

I have named the species after the late Capt. Hutton of Christ-

church, who forwarded these specimens to England.

DOLOMEDES TRIPPI, sp. n.

The cephalothorax is dark yellow-brown with pale yellow down-

lying hair. In the female there are lateral longitudinal stripes

between the median line and the margin but none at the margin.

The eyes are bright topaz-yellow.

The mandibles are black-brown with thin upstanding brown

hair and yellow-brown fringes. The lip and maxille are lighter

brown with brown upstanding hairs; the sternum paler with

yellowish-brown hair.

The legs and palpi are lighter brown, with upstanding brown

and downlying pale yellow-brown hairs.

The abdomen above is thickly covered with smooth downlying

pale yellow-brown hair mottled with dark grey. The sides are

darker, and a long narrow median space underneath is darker

still.

The head part of the cephalothorax is rather prominent.

The clypeus is not quite so broad as the two front median eyes

and the space between them, which latter is rather less than their

diameter. The side eyes are smaller and nearly close up to the

former. The eyes of the middle row are half as wide againas the

front median, being rather farther from them than the diameter

of the latter and the same distance from one another.

1908. | MR. H, R. HOGG ON AUSTRALASIAN SPIDERS. 343

The oval eyes of the third row are in their long diameter

smaller than those of the second row but larger than the front

median,

The mandibles are longer than patella of front pair of legs, they

are stout and kneed at the upper half. There are four large teeth

on the inner margin of the falx-sheath, one large between two

small teeth and a thick fringe on the outer margin.

The lip is broader than long, rounded in front, and half the

length of the maxille.

Text-fig. 75.

22 #,

Dolomedes trippi, sp. n.

a. Male. | 6. Female. | cc. Eyes. | d. Male palp.

The abdomen is somewhat cylindrical, straight at the sides and

rounded at front and rear, The spinnerets small and quite

terminal.

The legs are stout ; the first and second pairs are equally long in

the female, but in the male the first pair is the longer of the two.

344 MR. H. R. HOGG ON AUSTRALASIAN SPIDERS. [Apr. 7,.

There are scopula on the tarsi of all legs. One spine above on

tibia iv.

Besides being larger this species differs from D. minor L. Koch,

to which it is rather close, in having scopula on all the tarsi,

instead of on the front two pairs only, in its mandibles longer than

instead of equal to the front patella. The patellar joint of palpi

equal to instead of shorter than the tibial; and the clypeus more

than twice as broad as the distance between the front median and

second row eyes, instead of only slightly more.

The measurements (in millimetres) are as follows :—

Female.

Long. Broad.

Cephalothorax ... 9 : in front.

AN OCOMNEM 2 5505500- 13 7

Mandibles ...... 5

Pat. & Metat.

Coxa. Tr. & fem. tib. & tars.

j 34 63

Nees racine: 1 3 9 I 10 9 = 31

2 3 9 10 9 = Sil

3 25 9 9 8i = 29

4 3 10 10 10 = 333

Erol eae: « cau eae 2 5 4 33) eae

Male.

Long. Broad.

935 a

Cephalothorax ... 9 { a in front.

Abdomen ...... .. 9 ba

Mandibles ...... 4 longer than front patella.

Pat. & Metat.

Coxa. Tr &fem. tib. & tars.

hesspe Pee 1 3 92 if Wit = 34:

2, 3 9 103 104 = Be)

3 24 9 i i = 302

Le Pg 10 daa 12 = Ah

Palo yes. ye 2 6 4. 4 = 16

There are one male and one female from Pitt Island. I have

named them after C. H. Tripp, Esq., by whom they were collected,

1908. } MR, C, TATE REGAN ON AN AUSTRALIAN CAT-FISH. 345

April 28, 1908.

Dr. Henry Woopwarp, F.R.S., Vice-President,

in the Chair.

The Secretary read the following report on the additions made

to the Society’s Menagerie during the month of March 1908 :—

The number of registered additions to the Society’s Menagerie

during the month of March was 147. Of these 103 were acquired

by presentation, and 17 purchased, 9 were received on deposit,

8 by exchange, and 10 were born in the Gardens.

The number of departures during the same period, by death

and removals, was 178.

Among the additions special attention may be directed to:—

A pair of Jaguars (felis onca), from Northern Paraguay,

purchased on March 21st.

Two Bay Lynxes (felis rufa), from North America, purchased

on March 28th.

Two Himalayan Ibexes (Capra sibirica), from Panginear Chamba,

presented by H.H. The Maharajah of Chamba on March 27th.

A Red Brocket (Mazama rufa) and a Savannah Deer (Odocoileus

americanus savannarum), from Venezuela, presented by A. Pam,

Ksq., F.Z.8., on March 25th.

A Collection of thirty-one birds, including two Red-tailed

Guans (Ortalis ruficawda), an Ochre-winged Dove (Leptoptila

ochroptera), and two White-necked Saltators (Saltator albicollis),

new to the Collection, from Venezuela, presented by A. Pam, Esq.,

F.Z.S.,0on March 25th.

Mr. C. Tate Regan, M.A., F.Z.S., exhibited a specimen of an

Australian Cat-fish (Cnidoglanis megastoma Richards.), dissected

from one side to show the supports of the vertical fins, and made

the following remarks :—

“Tn Teleostean Fishes the rays of the caudal fin are inserted

directly on the more or less expanded heemal spines of the posterior

vertebrae, which are generally fused to form a hypural bone;

sometimes a few procurrent rays above and below are supported

by the neural and hemal spines of the vertebre preceding the

hypural. The dorsal and anal fins differ from the caudal in that

each ray is inserted on one of a series of basal supports, the so-

called interneural and interhzemal spines.

“The Siluroid Fishes of the sub-family Plotosine have been

generally defined as having two dorsal fins, a short anterior one

and a longer posterior fin continuous with the caudal, similar to

the long anal. The so-called second dorsal fin proves to be a pro-

current portion of the caudal fin, which has extended forward

along the back and in the species exhibited has a base of more

than 3 the total length of the fish and is formed of about 130 rays;

346 ON THE SHAPES OF EGGS. [Apr. 28,

these are supported directly by the neural spines, in striking

contrast to the externally similar anal fin, the rays of which are

attached to a series of interhzemals.”

Mr. Henry Scherren, F.Z.S., exhibited on behalf of Mr. Walter

Burton, F.Z.S., some melanisticand black leopard skins, presented

to Mr. Clarkson Williams in Abyssinia, where they were obtained.

The melanistic skins showed a deepening of ground-colour, espe-

cially along and on each side of the median dorsal line. In the

black skins, of which two were shown, traces of markings were

visible in certain lights, and the melanism appeared to be due to

w deepening of the ground, not to a multiplication of the spots.

Mr. Scherren stated that black skins from the same locality had

been received by Mr. Rowland Ward, and, like these, made up

into rugs. Myr. Ward, however, had preferred not to exhibit till

he could procure skins obtained by a Huropean sportsman.

Prof. D’Arey Wentworth Thompson, C.B., F.Z.S., read a

paper entitled “On the Shapes of Egegs”*, which contained a

discussion of the physical causes to which may be ascribed the

various forms assumed by the eggs of Birds and other animals.

The egg, prior to the formation of the hard shell, is a fluid body

enclosed in a somewhat elastic membrane. Its form tends to be

spherical, but is hable to distortion by the peristaltic pressure of

the oviduct. From the nature and direction of the peristaltic

wave, compression will be likely to occur in the posterior part of

the egg, rendering it broader in front than behind; and it may

be shown mathematically that the form of the envelope or surface

of the egg will correspond to the formula

1 Il

py + Ty(= at ~)=P,

where p” is the normal component of external pressure at a point

where 7 and ¢’ are the radii of curvature, Tis the tension of the

envelope, and P the internal fluid-pressure. It may further be

shown mathematically that, after the egg is formed, its unsym-

metrical shape, with a blunt anterior end and a tapering posterior

one, is well adapted for its easy transference through the oviduct,

blunt end foremost. The yolk, surrounded by its own proper

membrane, remains approximately spherical whatever be the form

of the egg-shell: because there lies between the two a fluid (viz.

the white of the egg) which makes the pressure (p”) upon the

yolk practically constant.

* Published in ‘ Nature,’ vol. 78. p. 111 (1908).

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‘dup weutmayy say WL? T8P PAeMpoonA FAL)

TX Wel BOGL SZ al

PAS; LOS JEL XO.

West, Newman. imp.

G.M. Woodward delet lith.

ORECTOLOBUS TENTACULATUS.

CHILOSCYLLIUM PLAGIOSUM.

“NQOIGNI -.WNITTIAOSOTIHDO “3 TUG ESE Reo Ss ORM iO)

dum weurma yy sayy ; : “YT 1819p Pxempoom “WH

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EEN RIE

Wx id 306) SZa

1908. | ON SHARKS OF THE FAMILY ORECTOLOBID. 347

The following papers were read :—

1. A Revision of the Sharks of the Family Orectolobide.

By C. Tate Reeay, M.A., F.Z.8.

[Received April 1, 1908. |

(Plates XI.—XTII.*)

This revision of the Orectolobide was prepared some time ago

as part of a descriptive catalogue of the Selachians in the British

Museum, a work which I have had no opportunity of continuing

for more than two years. I have therefore thought it best to

publish some of those parts which are ready.

The suborder Galeoidei includes Sharks with an anal and two

dorsal fins, without fin-spines and with five gill-openings on each

side. There are five families, viz. Odontaspidide, Lamnide,

Orectolobide, Scyliorhinide, and Carchariide. The Orectolobide

are distinguished by the presence of oro-nasal grooves T, by having

the last two to four gill-openings above the base of the pectoral,

and by the posterior position of the dorsal fins, the first of which

is above or behind the pelvics. Anatomically they differ notably

from the other Galeoidei in having the mesopterygium expanded

distally and bearing nearly as many radials as the metapterygium

and in the reduction or absence of the triradiate cartilagimous

rostrum.

In the present revision twenty-one species are recognised and

are referred to eight genera; most of the species are from the

Indo-Pacific.

The considerable range of variation in physiological characters

* For explanation of the Plates, see p. 364.

+ In most Selachians the nasal cavities are separate from the mouth. Jn three

species of Scyliorhinus (S. canicula, S. edwardsii, and S. marmoratum) the nasal

cavities are so near the mouth that the large anterior nasal valves overlie the edge

of the upper lip, but there are no oro-nasal grooves. In the Randz oro-nasal grooves |

are present, but run to the corner of the mouth and do not divide the upper lip.

In the Cestraciontide and Orectolobide the oro-nasal grooves divide the upper lip |

into a median and two lateral portions; they thus correspond in position to the

embryonic oro-nasal grooves of the Amniote Vertebrates.

Most text-books of embryology lay some stress on the presence in Amniote embryos

of these grooves, which are supposed to represent an ancestral condition found in

the adults of a lower group, the Selachians. There can be little doubt, however,

that in the Selachians oro-nasal grooves are specialised structures which have arisen

independently in different families, none of which can be regarded as in any way

approximating to the ancestral type of the higher vertebrates.

It is now generally accepted that the Amniote Vertebrates are derived from the

Batrachians and the latter from the Crossopterygian fishes. In all these groups

the premaxillary and maxillary bones form the upper border of the mouth and

separate the external apertures of the oral and nasal cavities. In the Crossoptery-

gians there are no internal nares; in the Batrachians internal nares are present and

develop as perforations of the palate, and in the Amniota they are the persistent

inner ends of the embryonic oro-nasal grooves. It seems more likely that in this

case ontogeny repeats phylogeny in the Batrachians rather than in the Amniota ;

if communication between the oral and nasal cavities internal to the premaxillaries

and maxillaries originated as open grooves, such grooves must have been present

before the development of the premaxillaries and maxillaries, but this is improbable,

as the Crossopterygians have the bones of the upper jaw fully developed, but no

trace of internal nares or oro-nasal grooves.

348 MR. C, TATE REGAN ON SHARKS [ Apr. 28,

in the sharks of this family is of some interest, and may be

exemplified by comparison of two extreme types.

Rhinodon typicus is a large pelagic shark, with numerous small

teeth, long gill-rakers, minute spiracles, and wide gill-openings ;

the pectoral fins are acutely pointed, and the axis of the caudal

fin is so strongly turned upwards and the lower lobe is so much

produced anteriorly as to give the appearance of a deeply forked

symmetrical fin. The resemblances of this shark to the Basking

Shark (Cetorhinus maximus), which belongs to the family

Lamnide, are very striking.

Eucrossorhinus dasypogon is a ground-shark with strong

dentition, large oblique spiracles, and small gill-openings; the

head and anterior part of the body is strongly depressed and

fringed with dermal flaps; the pectoral fins are broad and obtuse

and the axis of the caudal fin is not directed upwards nor is the

lower lobe enlarged. This form shows many points of similarity

to the Angel-fishes (Sguatina), which belong to another suborder.

In some species (e. g. Chiloscyllium punctatum, C. grisewm, and

Stegostoma tigrinum) there is considerable variation in colour and

markings, partly due to changes which take place during growth.

The young of these species have dark cross-bars, which may

become replaced by spots or may disappear, giving rise to a

uniform coloration.

As a rule the pelagic forms (e. g. Ginglymostoma, Khinodon)

have no conspicuous markings; the littoral sharks (Parascylliwm,

Brachelurus, Chiloscyilium, Stegostoma) are spotted, barred, or

variously ornamented, the coloration beimg most brilliant in the

tropical species; whilst the ground-sharks (Orectolobus, Hucros-

sorhinus) have markings which probably resemble the rocks and

weeds among which they lurk.

In the Orectolobide, as in other sharks, the dorsal and anal

fins appear to be comparatively larger in the young than in the

adult.

Synopsis of the Genera.

I. Spiracles very small ; anal quite distinct from the caudal,

wholly or partly opposed to the second dorsal.

A. Origin of second dorsal behind that of the anal ... 1. Parascylliwm.

B. Origin of second dorsal above or in advance of that

of the anal.

Teeth tricuspid or multicuspid ............. cies 2. Ginglymostoma.

feat WOMODISDUEL — Snoscoovoacbocnsecae seoesconeopss cecsecossqanoseseca Gh JeOApOgloyD.

II. Spiracles well-developed ; anal either continuous with

the caudal or terminating directly in front of it.

A. Lower lip divided into two by a symphysial groove.

1. Sides of head and snout without dermal flaps ;

spiracle surrounded by a circular raised rim... 4. Brachelwrus.

9. Sides of head and snout more or less strongly

fringed with dermal flaps; spiracles wide

cblique slits.

Last two gill-openings closer together than the rest .........

Gill-openings equidistant ..............ssceee eee

B. Lower lip not divided by a symphysial groove.

Caudal fin of moderate length ..........-.ceceeeee eee 7. Chiloscyllium.

Gaudalitmliveryaelongabemcnunnceeereses tee meee settee tee mCeN S CEO OSLOML Ce:

. Orectolobus.

. Hucrossorhinus.

fore!

1908. | OF THE FAMILY ORECTOLOBID#. 349

1. PARASCYLLIUM.

Parascylliwm Gill, Ann. Lye. N. York, 1861, p. 412; Giinth.

Cat. Fish. viii. p. 410 (1870).

Head rather elongate; body very elongate, subcylindrical.

Mouth slightly arched, near the end of the snout; teeth small,

pointed, with or without accessory cusps; lower lip not divided

by a symphysial groove. Nasal valves separate, each with an

obtuse cirrus. Eye small; a longitudinal fold below the eye.

Spivacles very small, below the level of the eyes and posterior to

them. First four gill-openings of moderate width; last con-

siderably wider; last two close together and above the base of

pectoral. Dorsal fins subequal, the first behind the pelvics, the

second in part posterior to the anal, which terminates at a con-

siderable distance from the caudal. Caudal of moderate length ;

axis scarcely directed upwards; lower lobe notched posteriorly.

Pectorals broad, rounded.

Two species from Australia.

1. PARASCYLLIUM COLLARE.

Parascyllium variolatum (non Duméril) Giinth. Cat. Fish. viii.

p. 410 (1870).

Parascyllium collare Ramsay & Ogilby, Proc. Linn. Soc. N.S.

Wales, (2) ui. 1889, p. 1310; Waite, Mem. Austral. Mus. iv.

1899, p. 32, pl. i. fig. 2.

No dorsal keel. Mouth near the end of snout; fold of lower

lip broadly interrupted ; nasal cirrus short. First dorsal origi-

nating above the posterior edge of pelvics; free edge of the fin

straight or convex; length of base 2 the distance from second

dorsal, which originates above the posterior part of anal. Anal

longer than deep, its base rather longer than that of either

dorsal, but shorter than its distance from the caudal. Brownish

above, yellow below ; back with some broad dark transverse bars ;

upper parts of body and fins with round dark spots.

Coasts of New South Wales, Victoria, and Tasmania.

(860 mm.). Tasmania. M. Allport, Esq.

(750 mm.). Tasmania.

2. PARASCYLLIUM VARIOLATUM.

Henuseyllium variolatum Dumeéril, Rev. et Mag. Zool. 1853,

p. 121, pl. i. fig. 1, and Elasmobr. p. 327 (1865).

Parascyllium nuchale MacCoy, Ann. Mag. N. H. (4) xiii. 1874,

p. U5; pl. a.

A broad blackish transverse band extending from behind the

eye to the root of the pectoral, covered with small white spots ;

numerous white spots on the body ; along the side a series of six

white semicircles, which may unite to form an undulating stripe,

extending from pectoral to caudal; each fin with a pair of dark

spots at its free edge and one at its base. In structural characters

apparently very similar to P. collare, but evidently distinct.

Coasts of Victoria and Tasmania.

350 MR. C. TATE REGAN ON SHARKS [ Apr. 28,

2. GINGLYMOSTOMA.

Ginglymostoma Mill. & Henle, Arch. f. Naturgesch. 1837, i.

p- 395, and Plagiost. p. 22 (1841); Giinth. Cat. Fish. viii. p. 407

(1870).

Nebrius Riipp. Neue Wirbelth., Fische, p. 62 (1840).

Head broad, obtuse; body elongate, subcylindrical. Mouth

transverse, near the end of the snout; teeth small, tricuspid or

multicuspid ; lower lip not divided by a symphysial groove.

Nasal valves separate, each with a cirrus. Hye small; no longi-

tudinal fold below the eye. Spiracle very small, behind the eye.

Gill-openings of moderate width; last two close together; last

two or three above the base of the pectoral. First dorsal above

or partly behind the pelvics; second dorsal above or partly in

advance of the anal, which is free from the caudal. Caudal of

moderate length; axis directed upwards; lower lobe notched

posteriorly.

Four species.

Synopsis of the Species.

TI. Each dorsal and anal fin with the anterior angle broadly

rounded ; lower caudal lobe not produced anteriorly ;

teeth tricuspid or pentacuspid, the middle cusp the

strongest.

Origin of first dorsal opposite to that of the pelvics ... 1. cirratum.

Origin of first dorsal slightly in advance of posterior

GG OH? HA] GHP ANOS soe cesses gouavseoo ove 90ner nes0a9a00

II. Each dorsal and anal fin with the anterior angle

pointed ; lower caudal lobe somewhat produced

anteriorly.

Teeth with a strong median cusp and 3 or 4 small

CBFOS OM, GROIN SOS) 55 no soocne oe o59005. 208 0090001200 sananaoer

Teeth with the cusps graduated to the strongest, which

TE) TALE WASNT 26 aps onosuoocqonocbodeasedaneccentopcboooscocan 4b GOMCHIGIA,

2. brevicaudatum.

3. ferrugineum.

1. GINGLYMOSTOMA CIRRATUM.

Squalus cirratus Gmelin, Linn. Syst. Nat. p. 1492 (1788) ;

Schneid. Bloch’s Syst. Ichth. p. 128 (1801).

Squalus punctatus Schneid. t. c. p. 134.

Squalus punctulatus Schneid. t. c. p. 549.

Squalus argus Bancroft, Zool. Journ. v. 1834, p. 82.

Ginglymostoma cirratum Mill. & Henle, Plagiost. p. 13 (1841);

Duméril, Elasmobr. p. 334 (1865); Gunth. Cat. Fish. vin. p. 408.

(1870); Jord. & Everm. Bull. U.S. Nat. Mus. xlvui. 1896, p. 26,

and 1900, pl. iv. fig. 13.

Ginglymostoma fulvum Poey, Mem. u. p. 342 (1861).

Ginglymostoma caboverdianus Capello, Jorn. Sci. Phys. Lisbon,

1867, p. 167.

Nasal cirrus extending to or beyond the oral edge of the nasal

valve. Lower labial folds separated by a considerable interspace.

Teeth bicuspid or pentacuspid, the middle cusp much the strongest.

First dorsal a little larger than the second and considerably larger

1908. } OF THE FAMILY ORECTOLOBID. Sey):

than the anal ; origin of first dorsal opposite to that of the pelvics ;

free edge of the fin straight or convex, anterior angle broadly

rounded, posterior angle nearly rectangular; length of base

greater than the distance from second dorsal. Anal beginning

below the middle of second dorsal; free edge of the fin evenly

convex. Caudal fin 2 the length of the rest of the fish or rather

more ; lower lobe moderately deep and with the edge forming an

obtuse angle anteriorly. Pectoral with nearly straight free edge

and rounded angles, a little longer than broad, its length nearly

equal to its distance from the mouth or 2 of the distance from its

origin to that of the pelvics. Brownish; young with small,

scattered, round, blackish spots.

Tropical Atlantic ; Pacific Coast of Mexico.

1-2. (530 and 400 mm.) St. Croix.

3. (390 mm.) Cuba. Zool. Soc.

4, (280 mm.) Jalisco, Mexico. Dr. A. C. Buller.

5. (230 mm.) S. America. Sir R. Schomburek.

6-8. (740-2400 mm.) stuffed. W. Indies.

2. GINGLYMOSTOMA BREVICAUDATUM.

Ginglymostoma brevicaudatum Ginth. & Playfair, Fish.

Zanzibar, p. 141, pl. xxi. (1866); Giinth. Cat. Fish. viii. p. 408

(1870).

Nasal cirrus short, not nearly reaching the oral edge of the

nasal valve. Lower labial folds separated by a considerable inter-

space. Teeth tricuspid or pentacuspid, the middle cusp much the

strongest. Dorsal and ana! fins subequal, each with broadly

rounded anterior angle, straight or slightly convex free edge, and

obtuse posterior angle. First dorsal originating a little in

advance of end of base of pelvics; length of base less than the

distance from second dorsal. Anal beginning and ending respec-

tively a little behind the origin and end of the base of second

dorsal. Caudal fin rather less than 2 of the length of the rest of

the fish; lower lobe deep and with the edge forming an obtuse

angle anteriorly. Pectoral with nearly straight free edge and

rounded angles, as broad as long, its length equal to its distance

from the eye or 3 the distance from its origin to that of the

pelviecs.

Zanzibar : Seychelles.

1. (580 mm.), stuffed, Zanzibar. Lieut.-Col. Playfair.

type of the species.

2. Skull. Seychelles. Swinburne Ward, Hsq.

3. Jaws.

3. GINGLYMOSTOMA FERRUGINEUM.

Scyllium ferrugineum Less. Voy. Coquille, Zool. ii. p. 95 (1830).

Ginglymostoma concolor Mill. & Henle, Plagiost. p. 22, pl. vi.

(1841).

Ginglymostoma muelleri Giinth. Cat. Fish. viii. p. 408 (1870) ;

Proc, Zoou. Soc.—1908, No. XXIII. 23

352 MR, C. TATE REGAN ON SHARKS [ Apr. 28,

Klunz. Fisch, Roth, Meer. p. 230 (1870); Day, Fish. India,

p. 725 (1878).

Nasal cirrus extending to or nearly to the oral edge of the

nasal valve. Lower labial folds separated by a considerable inter-

space, Teeth with a strong median cusp and with 3 or 4 smaller

accessory cusps on each side. First dorsal a little larger than the

second or than the anal; all three with slightly concave free edge

and acutely pointed anterior angle; base of first dorsal opposite

to that of the pelvics, its length a little more than the distance

from the second. Anal beginning below the anterior part of

second dorsal. Caudal fin nearly 4 the length of the rest of the

fish ; lower lobe produced and acutely pointed anteriorly. Pectoral

with slightly concave free edge and acutely pointed anterior

angle, nearly twice as long as broad, its length nearly equal to

its distance from the end of snout or 2 of the distance from its

origin to that of the pelvics. Brownish.

Tropical Indo-Pacific.

Attains a length of 2400 mm.

4, GINGLYMOSTOMA CONCOLOR.

Nebrius concolor Riipp. Neue Wirbelth., Fische, p. 62, pl. xvii.

fig. 2 (1840).

Ginglymostoma concolor Cant. Cat. Mal. Fish. p. 395 (1850) ;

Ginth. Cat. Fish. viii. p. 409 (1870) ; Klunz. Fisch. Roth. Meer.

p. 232 (1870); Day, Fish. India, p. 811 (1878).

Ginglymostoma ruppellii Bleek. Verh. Bat. Gen. xxiv. 1852,

Plagiost. p. 91, and Nat. Tijds. Ned. Ind. iii. 1852, p. 83;

Dumeril, Elasmobr. p. 334 (1865).

Nasal cirrus extending to the oral edge of the nasal valve.

Lower labial folds separated by a considerable interspace. Teeth

multicuspid, the cusps subequal or graduated to the strongest,

which is not the median one. First dorsal a little larger than

the second or than the anal; all three with straight or slightly

concave free edge and acutely pointed anterior angle ; base of first

dorsal opposite to that of the pelvics, its length greater than the

distance from the second. Anal beginning below the anterior

part of second dorsal. Caudal fin 4 the length of the rest of the

fish ; lower lobe produced anteriorly. Pectoral with slightly

concave free edge and pointed anterior angle, 13 as long as broad,

its length equal to its distance from the mouth or 2 of the

distance from its origin to that of the pelvics. Brownish.

Indian Ocean; Malay Archipelago.

1. (660 mm.) Java. Dr. P. Bleeker,

2. (700 mm.) stuffed. Pinang. Dr. Cantor.

3. RHINODON.

Rhineodon (Smith) Mill. & Henle, Arch. f. Naturgesch, 1838,

1. p. 84.

1908. ] OF THE FAMILY ORECTOLOBID. 353

Rhinodon (Smith) Mill. & Henle, Plagiost. p. 77 (1841);

Giinth. Cat. Fish. vii. p. 396 (1870).

Micristodus Gill, Proc. Acad. Philad. 1865, p. 177.

Head broad, obtuse; body elongate, subcylindrical. Mouth

transverse, subterminal; teeth very small, unicuspid, pointed,

subconical, recurved; lower lip not divided by a symphysial

groove. Nasal valves separate; no nasal cirri. Eye small; no

longitudinal fold below the eye. Spiracle very small, behind the

eye. Gill-openings wide; last two above the base of pectoral.

First dorsal above the pelvics; second dorsal above the anal,

which is free from the caudal. Caudal of moderate length; axis

strongly directed upwards; lewer lobe considerably produced

anteriorly and without posterior notch.

This genus comprises a single species, which has generally been

placed in more or less close proximity to Cetorhinus maximus, a

shark which it appears to resemble in habits and in its large

size, small teeth, long gill-rakers, and wide gill-openings, as well

as in the form of the caudal fin and the keeled tail. There can

be little doubt, however, that /hinodon is allied to Ginglymostoma,

from which it differs in a few features of specialisation. The

‘curious dermal keels are like those met with in other Sharks of

this family, i.e. Stegostoma, Chiloscyllium.

1. RHINODON TYPICUS.

Rhinodon typicus Smith, Mill. & Henle, Plagiost. p. 77,

pl. xxxv. fig. 2 (1841); Smith, Ill. Zool. 8. Afr. Fish. pl. xxvi.

(1845); Dumeéril, Elasmobr. p. 428 (1865); Giinth. Cat. Fish.

vil. p. 396 (1870); Haly, Ann. Mag. N. H. (5) xii. 1883, p. 48;

Thurston, Bull. Madras Mus. 1884, No. 1, pl. iii. A; Gill,

Science, (2) xv. 1902, p. 824.

Micristodus punctatus Gill, Proc. Ac. Philad. 1865, p. 177;

Ginth. Cat. Fish. vii. p. 396 (1870); Jord. & Everm. Bull. U.S.

Nat. Mus. xlvii. 1896, p. 52.

Rhinodon pentalineatus Kishinouye, Zool. Anz. xxiv. 1901,

p. 694, fig.

Lower labial folds separated by a wide interspace. A median

dorsal keel and on each side 2 or 3 lateral keels, the lowest of

which is continued along the middle of the tail to the caudal fin.

First dorsal considerably larger than the second or than the anal,

its base opposite to that of the pelvics; free edge of first dorsal

slightly concave, anterior angle rounded, posterior angle pointed.

Second dorsal and anal equal and opposite. Lower caudal lobe

strongly produced and acutely pointed. Pectoral much longer

than broad, with slightly concave free edge and acutely pointed

anterior angle. Head and body with round whitish spots.

This large pelagic shark has been recorded from Florida, the

Cape of Good Hope, the Seychelles, Ceylon, Madras, Japan,

California, Panama, and Peru.

1. Large stuffed specimen. Ceylon. Colombo Mus.

‘2. Upper jaw and snout. Seychelles. Prof. E. P. Wright.

23*

304 MR. C, TATE REGAN ON SHARKS [ Apr. 28,

4, BRACHALURUS.

Brachelurus Ogilby, Proc. Roy. Soc. Queensland, xx. 1906, p. 27.

Head broad, depressed; body elongate, somewhat depressed

anteriorly, subcylindrical posteriorly. Dermal denticles small,

imbricated, more or less distinctly keeled. Mouth transverse ;

teeth small, tricuspid ; lower lip divided by a symphysial groove.

Nasal valves separate, each with a cirrus. Hye small; a longi-

tudinal fold below the eye. Spiracles moderately large, below the

level of the eyes and posterior to them; each is surrounded by a

raised circular rim. Gull-openings of moderate width ; last three

above the base of pectoral; last two a little closer together than

the others. Dorsal fins subequal, the first above or behind the

pelvics, the second in advance of the anal, which terminates

directly in front of the caudal. Caudal of moderate length; axis

not directed upwards ; lower lobe notched posteriorly. Pectorals

broad, with straight or convex edges and rounded angles.

A single species from Australia.

1. BRACH2ZLURUS MODESTUS.

Chiloscyllium modestum Ginth. Proc. Zool. Soc. 1871, p. 654,

1G ion

Chiloscylliium furvum Macleay, Proc. Linn. Soc. N. 8. Wales,

vi. 1881, p. 364.

Hemiscyllium modesitum Waite, Rec. Austral. Mus. iv. 1901,

_p. 88, fig. 9, and pl. iv. fig. 1.

No dorsal keel. Mouth nearer to level of eyes than to end of

snout; fold of lower lip not continuous; nasal cirrus long,

extending to posterior edge of lower lip. First dorsal originating

above posterior part of base of pelvics; free edge of the fin

straight; length of base greater than the distance from second

dorsal. Origin of anal below the end of base of second dorsal ;

length of base of anal less than 3 that of the caudal. Pectoral

extending a little more than of the distance from its origin to

that of the pelvics. Brownish, with more or less distinct darker

cross-bars and, in the young, with light spots.

Coasts of Queensland and New South Wales.

1. (550 mm.) stuffed, type Queensland.

of the species.

2. (580 mm.). Port Jackson. Imperial Inst.

5. ORECTOLOBUS.

Orectolobus Bonap. Faun. Ital. Pesc. 7 fase. (1834); Gill, Proc.

U.S. Nat. Mus. xviii. 1895, p. 212.

Crossorhinus Mull. & Henle, Arch. f. Naturgesch. 1837, 1.

p. 396, and Plagiost. p. 21 (1841).

Crossorhinus (part.) Gunth. Cat. Fish. viii. p. 413 (1870).

Head broad, depressed ; body elongate, more or less depressed

anteriorly, subcylindrical posteriorly. Dermal denticles small,

imbricated or juxtaposed. Mouth slightly arched, wide, sub-

1908. | OF THE FAMILY ORECTOLOBID¥, 355

terminal; teeth slender, pointed, the median ones enlarged and

unicuspid, the lateral teeth smaller, with small accessory cusps;

lower lip divided by a symphysial groove. Nasal valves separate,

each with a cirrus. Eye small; a more or less distinct longi-

tudinal fold below the eyes. Spiracles large, oblique, extending

forward to below the eyes. Gill-openings of moderate width, the

last a little wider than the rest ; last three or four above the base

of pectoral ; last two closer together than the others. Dorsal fins

subequal, the first above or behind the pelvics, the second in

advance of the anal, which terminates directly in front of the

lower caudal lobe or is continuous at the base with the latter.

Caudal of moderate length; axis not directed upwards; lower

lobe notched posteriorly. Pectorals broad, with straight or

convex edges and rounded angles.

Synopsis of the Species.

T. Nasal cirrus with a short branch at about the middle of

its length.

A. Branch of nasal cirrus bifid; on each side 3 to 5

dermal lobes above the upper lip, followed by 4 or 5

near the angle of the mouth and these Nee 2 ee

notched distally) at the side of the head . eeee: ) Lebanbatus:

B. Branch of nasal cirrus simple.

On each side 2 or 3 dermal lobes above the upper

lip, followed by 3 or 4 (the first and last of

which are bifid) near the angle of the mouth

and these by 2 pee notched ly at the

side of the head ........ Wesesssse. | 2a JQPOnICUs.

Dermal lobes all simple; ¢ on each side 2 above the

upper lip, 2 near the uncle of the mont, and 2

at the side of the head . He Hhootnesadedodpabes By) UGA

II. Nasal cirrus simple; dermai lobes Bale on each side

one above the upper lip, another near the angle of the

mouth, and a third at the side of the head.................. 4. tentaculatus.

1. ORECTOLOBUS BARBATUS.

Squalus barbatus Gmelin, Linn. Syst. Nat. p. 1493 (1788);

Schneid. Bloch’s Syst. Ichth. p. 128 (1801).

Squalus lobatus Schneid. t.c. p. 137.

Squalus appendiculatus Shaw, Nat. Mise. pl. pccxxyvtt.

Crossorhinus barbatus (part.) Dumér il, Elasmobr. p. 338 (1865) ;

Giinth. Cat. Fish. vii. p. 414 (1870).

Crossorhinus barbatus McCoy, Prodr. Zool. Vict. v. 1880,

pl. xliii. fig. 1; Macleay, Proc. Linn. Soc. N. 8. Wales, vi.

188151365; %.

A papilliform projection above the posterior part of each eye

(and in the adult another above the anterior part), Nasal cirrus

long, with a short bifid branch at about the middle of its length.

On “each side a series of 3 to 5 simple tentacle-like dermal lobes

above the upper lip, followed by 4 or 5 near the angle of the

mouth, the first and last of which are ramose and by 2, short,

broad, and distally notched, at the side of the head. First dorsal

€

396 MR. C. TATE REGAN ON SHARKS [Apr. 28,.

originating above the posterior part of base of pelvics; free edge

of the fin straight, posterior angle acute or rectangular; length

of base more than the distance from second dorsal, which ter-

minates a HEthils in advance of the origin of anal. Pectoral

extending 4 to 2 of the distance from its origin to that of the

pelvies. Bowman) with numerous white spots and markings,

many of which form circles or enclose irr egular areas.

Eastern and Southern Coasts of Australia.

1. (750 mm.) New South Wales. G. Krefft, Esq.

2. (600 mm.) Sydney. Imperial Inst.

3. (460 mm.) Tasmania. Haslar Coll.

4, (2000 mm.) stuffed. South Australia.

2. ORECTOLOBUS JAPONICUS.

Crossorhinus barbatus (non Gmelin) Miill. & Henle, Plagiost..

p. 21, pl. v. (1841); Schleg. Faun. Japon., Poiss. p. 301 (1850).

Crossorhinus barbatus (part.) Duméril, Elasmobr. p. 338 (1865);

Giinth. Cat. Fish. vin. p. 414 (1870).

Orectolobus barbatus Jord. & Fowler, Proc. U.S. Nat. Mus.

xxvi. 1903, p. 606.

Orectolobus japonicus Regan, Ann. Mag. Nat. Hist. (7) xviii.

1906, p. 435.

Closely allied to O. barbatus, but differing in the following

characters:—No papilliform projections above the eye. Nasal

cirrus with a simple branch. On each side 2 or 3 simple dermal

lobes above the upper lip, followed by 3 or 4 near the angle of

the mouth, the first and last of which are bifid, and by 2, short,

broad and distally notched, at the side of the head. Free edge of

dorsal fins straight. or slightly concave. Pectoral extending at

least 3 of the distance from its origin to that of the pelvics.

Yellowish, upper surface with brownish vermiculations or reticu-

lations; back with broad dark-brown cross-bars, with yellow

vermiculations.

Coasts of Japan and China.

1,2. (1000 and 780 mm.) types of the species. Japan.

3. ORECTOLOBUS ORNATUS. (Plate XI. fig. 2.)

Crossorhinus ornatus De Vis, Proc. Linn. Soc. N.S. Wales

vill. 1883, p. 289.

Differs from O. barbatus as follows :—Nasal cirrus with a short.

simple branch. Dermal lobes all simple, on each side 2 above

the upper lip, 2 at the angle of the mouth, and 2 at the side of

the head. Dorsal fins each with slightly convex free edge and

rounded posterior angle. Greyish; back with dark brown cross-~

bars ; a bar at the level of the pectorals and one in front of the

first dorsal have irregular edges and each encloses a pair of

ocelli ; the posterior bars nearly meet in the mid-ventral line ;

fins with large dark spots.

Queensland.

1. (180 mm.) Australia.

1908. | OF THE FAMILY ORECTOLOBIDA. 357

4, ORECTOLOBUS TENTACULATUS. (Plate XII. fig. 2.)

Crossorhinus tentaculatus Peters, Monatsb. Ak. Berl. 1864,

p. 123; Giinth. Cat. Fish. viii. p. 414 (1870); Macleay, Proc.

Linn. Soc. N. 5. Wales, vi. 1881, p. 365.

A papilliform projection above the posterior part of each eye.

Nasal cirrus long, simple. On each side a small simple dermal

lobe above the upper lip, a larger one at the angle of the mouth,

and a third at the side of the head. First dorsal originating a

little in advance of the posterior end of base of pelvies; free edge

of the fin straight or convex, posterior angle rounded or obtusely

pointed; length of base considerably more than the distance

from second dorsal, which terminates nearly above the origin of

anal. Pectoral extending 3-3 of the distance from its origin

to that of the pelvics. Yellowish, back greyish; on the back,

in front of the first dorsal fin, three large dark areas edged with

white, continuous or subecontinuous with dark vertical bars on

the sides; tail completely encircled by 3 dark vertical bands,

corresponding to the two dorsal and the anal fins; fins with large

dark spots.

Queensland.

1-2. (430 and 240 mm.) Cape York.

3. (220 mm.)

Haslar Coll..

6. HUCROSSORHINUS, gen. nov.

Crossorhinus (part.y Giinth. Cat. Fish. viii. p. 413 (1870).

Very closely allied to Orectolobus, but with broader and more

depressed head, smaller eyes, and wider spiracles than in any

species of that genus. No longitudinal fold below the eye.

Gill-openings rather small and of equal width, equidistant ; last

four above the base of pectoral.

A single species from Waigiou.

EUCROSSORHINUS DASYPOGON.

Crossorhinus dasypogon Bleek. Arch. Néerland. 1867, p. 400,

pl. xxi. fig. 1; Giinth. Cat. Fish. viii. p. 414 (1870).

Nasal cirrus ramose. Head margined by a nearly continuous

series of ramose dermal flaps, extending on each side from the

nasal opening to the pectoral fin; a transverse series of similar

flaps behind the mouth. First dorsal originating slightly in

advance of the posterior end of base of pelvics; free edge of the

fin straight or slightly convex, angles rounded ; length of base

scarcely more than the distance from second dorsal, which ter-

minates above the origin of anal. Pectoral extending + of the

distance from its origin to that of the pelvics. Upper parts of

head, body, and fins covered with a brownish network enclosing

small round whitish spots; a few small dark spots on the tail,

2 or 3 on each of the pectoral and ventral fins, one on each side

358 MR. C, TATE REGAN ON SHARKS ‘Apr. 28

at the end of the base of both dorsals and the anal, one on the

anterior part of the lower caudal lobe.

Waigiou.

1. (210 mm.) type of the species. Waigiou. Dr. P. Bleeker.

7, CHILOSCYLLIUM.

Chiloscyllium Mill. & Henle, Arch. f. Naturgesch. 1837, 1.

p. 395, and Plagiost. p. 17 (1841); Giinth. Cat. Fish. viii. p. 410

1870

ey yllium Mull. & Henle, Arch. f. Naturgesch. 1838, 1.

p. 83, and Plagiost. p. 16.

Synchismus Gill, Aun. Lye. N. York, 1861, p. 413.

Head obtuse; body elongate, subeylindrical. Mouth trans-

verse; teeth small, pointed, usually with one or two pairs of

small accessory cusps; lower lip not divided by a symphysial

groove. Nasal valves separate, each with a cirrus. Eye small;

no longitudinal. fold below the eye. Spiracles moderately large,

below the level of the eyes, and usually extending forward beneath

them. Gill-openings of moderate width; last three above the

base of pectoral; last two very close together. Dorsal fins sub-

equal, the first above or behind the pelvics, the second in advance

of the anal, which is continuous at the base with the lower caudal

lobe. Caudal of moderate length; axis not directed upwards;

lower lobe notched posteriorly. Pectorals broad, with straight or

convex edges and rounded angles.

Seven species from the Indo-Pacific,

Synopsis of the Species.

I. Mouth nearer to the end of the snout than to the vertical

from the eyes; origin of first dorsal behind the base of

pelvics; dorsal fins with concave free edges.

A. A large ocellus above the pectoral fin.

Base of first dorsal 2 to # its distance from the second; body

with scattered rounded dark spots ..............seeeee eee eee eee 1. ocellatum.

Base of first dorsal # its distance from the second ; body with

numerous close-set dark EWOWS. coccancdervonbespnasesecsceasunneana — Mb UPUS/DCCOMUGNRA

B. No ocellus above the pectoral fin ............. esses 8 freycineti.

II. Mouth nearer to the vertical from the eyes than to the end

of the snout; fold of the lower lip continuous.

A. Origin of first dorsal above the anterior part of base of

pelvics; dorsal fins with concave free edges ............ 4. punctatum.

B. Origin of first dorsal above the posterior end of base of

pelvics; dorsal fins with straight or convex free edges.

1, A single more or less distinct median dorsal keel.

Anal as deep as lower caudal lobe, its length } to 2 that of the

latter in front of the notch ; no white or pale spots ...... 5. griseum.

Anal usually deeper than lower caudal lobe, its length from

less than 2 to nearly # that of the latter im front of the

notch; body with round or oval white or pale spots......... 6. plagiosum.

Ah, Dimes Glare TRESS: Soo jacdoscuscsecasscconuaadouosauoacanscevcos Yin CCMICIE.

1908. ] OF THE FAMILY ORECTOLOBID®. 359

1, CHILOSCYLLIUM OCELLATUM.

Squalus ocellatus Gmelin, Linn. Syst. Nat. p. 1494 (1788) ;

Schneid. Bloch’s Syst. Ichth. p. 129 (1801).

Hemiscyllium ocellatum Mill. & Henle, Plagiost. p. 16 (1841);

Dumeéril, Elasmobr. p. 326 (1865).

Scyllium ocellatwm Blyth, Journ. As. Soc. Bengal, 1847, p. 726,

pl. xxvi. fig. 2.

Chiloscylliwm ocellatum Ginth. Cat. Fish. vii. p. 410 (1870),

’ No dorsal keel, except between the dorsal fins. Mouth nearer

to the end of snout than to the vertical from the eyes; fold of |

lower lip not continuous ; nasal cirrus extending to edge of nasal

valve. Nearly the whole of the spiracle below the eye. First

dorsal originating well behind the base of pelvics; free edge of

the fin concave ; length of base 2 to ? the distance from second

dorsal. Origin ‘of anal at a distance from the vertical from the

end of base of second dorsal greater than the length of the latter ;

length of base of anal about 2 of that of lower caudal lobe in

front of the notch. Pectoral extending 2 to ? of the distance

from its origin to that of the pelvics. Scattered round dark

spots on the body; a large ocellus above the pectoral fin; indis-

tinet cross-bars on the back; two large spots on the anterior edge

of each dorsal fin.

New Guinea and Northern Australia to the Kermadec Islands.

1. (630 mm.).

2. (590 mm.). Cape York. Herr Damel.

3-4. (410 and 420 mm.) Sunday Is. J. B. Jukes, Esq.

5. (400 mm.) Solomon Isds.

6. (360 mm.) N.W. Australia. Haslar Coll.

2. CHILOSCYLLIUM TRISPECULARE,

Hemiscyllium trispeculare Richards. Ic. Pisce. p. 5, pl. 1. fig. 2

(1843), and Erebus & Terror, Fish. p. 43, pl. xxviii. (1844).

Chiloscyllium trispeculare Giinth. Cat. Fish. viii. p. 411 (1870),

Scarcely specifically distinct from C. ocellatum, but with some-

what larger fins and with a different coloration. Length of base

of first dorsal 4 its distance from the second ; pectoral extending

4+ of the distance from its origin to that of the pelvies. Body

densely covered with brownish spots, the larger of which are

studded with several small darker spots; a large ocellus above

the pectoral fin, followed by one or two smaller ones; indi-

stinct cross-bars on the back; two large spots on the anterior

edge of each dorsal fin.

North-west Australia.

1. (550 mm.) type of N.W. Australia. Sir J. Richardson.

the species.

3. CHILOSCYLLIUM FREYCINETI.

Scyllium freycinetti Quoy & Gaim. Voy. Uran., Poiss. p. 192

(1824),

360 MK. C. TATE REGAN ON SHARKS [ Apr. 28,.

Scyllium malaisianwm Less. Voy. Coquille, Zool. 11. p. 94,

pl. vi. (1830); Giinth. Cat. Fish. viii. p. 411 (1870).

Chiloscyllium malaianum Mill. & Henle, Plagiost. p. 20 (1841);

Duméril, Elasmobr. p. 332 (1865).

Apparently closely allied to the two preceding species, but

without the ocellus above the pectoral fin; spots on the body

on the back and spots on the dorsal fins as in the two preceding

species.

Waigiou.

4, CHILOSCYLLIUM PUNCTATUM.

Chiloscyllium punctatum Mill. & Henle, Plagiost. p. 18, pl. in.

(1841); Bleek. Verh. Bat. Gen. xxiv. 1852, Plagiost. p. 22;

Duméril, Elasmobr. p. 330 (1865); Giimth. Cat. Fish. viii. p. 413

(1870).

Chiloscylliam grisewm Mill. & Henle, t. ¢. pl. iv.

No dorsal keel, except between the dorsal fins. Mouth much

nearer to the vertical from the eyes than to the end of snout;

fold of lower lip continuous ; nasal cirrus extending well beyond

the edge of nasal valve. Nearly the whole of the spiracle below

the eye. First dorsal originating above the anterior part of base

of pelvics ; free edge of the fin concave; length of base not much

less than the distance from second dorsal. Origin of anal at a

distance from the vertical from the end of base of second dorsal

equal to less than 3 the length of the latter; length of base of

anal from a little more than 2 to nearly ? of that of the lower

caudal lobe in front of the notch. Pectoral extending } to g of

the distance from its origin to that of the pelvics. Body with

or without small dark spots; young with broad dark cross-bars,

one across the head, one at the level of the pectorals, one just in

front of and another at the posterior end of each of the dorsal

fins, and three behind the second dorsal.

~ Malay Archipelago.

1. (640 mm.) 2, uniformly greyish. Thursday Is., Torres Str. Earl of Crawford.

2. (600 mm.) ¢, with scattered Singapore. Fisheries Exhib.

dark spots, without cross-bars.

3. (340 mm.) 9, with cross-bars, Java. Dr. P. Bleeker.

without spots.

5. CHILOSCYLLIUM GRIsEUM. (Plate XI. fig. 1, and Plate XIII.

fig. 1.)

Chiloscyllium plagioswm, var. 1, Mill. & Henle, Plagiost. p. 18

(1841).

Chiloscyllium griseum Mill. & Henle, t.c. p. 19.

Chiloscyllium plagioswm (part.) Cantor, Cat. Mal. Fish. p. 392

(1850).

Chiloscyllium plagiosum Bleek. Verh. Bat. Gen. xxiv. 1852,

Plagiost. p. 17; Duméril, Elasmobr. p, 328 (1865); Day, Fish.

Malabar, p. 267 (1865).

1908. ] OF THE FAMILY ORECTOLOBID. 361

Chiloscyllium indicum, vars. y and é (part.) Giinth. Cat. Fish.

vill. p. 412 (1870).

Chiloscyllium indicum Day, Fish. India, p. 726, pl. elxxxviil.

fig. 3 (1878).

A single more or less prominent dorsal keel. Mouth much

nearer to the vertical from the eyes than to the end of snout ;

fold of the lower lip continuous; nasal cirrus extending to the

edge of nasal valve or slightly beyond. Anterior part of spiracle

below the posterior part of eye. First dorsal originating above

the posterior end of base of pelvics; free edge of fin straight or

convex ; length of base from # to as long as the distance from

second dorsal. Origin of anal at a distance from the vertical

from end of base of second dorsal a little less than the length of

the latter ; anal as deep as lower caudal lobe, 3 to 2 the length

of the latter in front of the notch. Pectoral extending # to 4 of

the distance from its origin to that of the pelvics. Young with

very distinct, broad, dark cross-bars, which become somewhat

narrower on the sides and broaden out again and unite below,

the lower surface being uniformly dark; interspaces between the

bars in great part occupied by oblong or oval spots; bars about

12 in number, one across the snout, four more anterior to the

first dorsal; one at the end of the base of each dorsal and one

in front of the second dorsal; three or four behind the second

dorsal. Half-grown examples with bars less distinct and not

united below, sometimes edged with darker lines or series of

spots; interspaces and lower surface uniformly pale. Adults.

uniformly greyish or brownish.

Coasts of India to the Malay Archipelago.

I. Adults, without cross-bars.

1. (580 mm.) 6, type of Moluccas. Dr. P. Bleeker.

C. hasseltii.

2. (550mm.) 9°. Kurrachee. F. W. Townsend, Esq.

3. (460 mm.) ¢, C. ob- Moluccas.

scurum Gray.

II. Half-grown specimens, with more or less distinct cross-bars.

A. Cross-bars without darker edges.

1-38. (140-245 mm.) Vizagapatam. Capt. Mitchell.

4-6. (120-340 mm.) Madras. F. Day, Esq.

7-9. (120-270 mm.) Malabar. F. Day, Esq.

B. Cross-bars edged with darker lines or series of spots.

1. (245 mm.) Pinang. Dr. Cantor.

2-3. (170 and 190 mm.)

III. Young, with cross-bars very conspicuous and united below.

1. (122 mm.) Malay Peninsula. Mr. Evans.

Miiller and Henle’s typical specimen, from Malabar, evidently

belongs to this species. The discrepancies which they observe

between their specimens and the figure which they reproduce is

due, not to the inaccuracy of the latter, as they supposed, but to

its representing another species, viz. C’. punctatum.

362 MR, C. TATE REGAN ON SHARKS [ Apr. 28,

6. CHILOSCYLLIUM PLAGIOsUM. (Plate XII. fig. 1.)

Seyllium plagiosum Bennett, in Life of Rattles, p. 694 (1830).

Scyllium ornatum Gray, Il. Ind. Zool. i. pl. xeviii. fig. 2 (1832).

Chiloscyllium plagiosum, vars. 2, 3 and 4, Miill. & Henle,

Plagiost. p. 18 (1841).

Chiloscyllium plagioswm (part.) Cantor, Cat. Mal. Fish. p. 392

(1850).

oe margaritiferum Bleek. Ned. Tijds. Dierk. i. 1863,

p. 243; Dumeril, Elasmobr. p. 329 (1865).

Chiloseyllium indicum, vars. a and 3, Giinth. Cat. Fish. viii.

p. 412 (1870).

Chiloscyllium indicum Jord. & Fowler, Proc. U.S. Nat. Mus.

xxvi. 1903, p. 605, fig. 2.

Very closely allied to C. grisewm, but with quite a different

coloration. Dorsal fins usually shorter and more elevated, the

length of the base of the first 2 to + its distance from the second.

Anal usually shorter and deeper, its depth usually distinctly

greater than that of the lower caudal lobe, its length from less

than 2 to nearly # that of the latter in front of the notch.

Dark cross-bars arranged as in C. grisewm, but much narrower

and persistent in the adult; edges of the bars crenate; numerous

round or oval white spots on the cross-bars and at their edges, as

well as on the lower parts of the body; dark spots either few,

large, and definitely arranged in the mid-dorsal line and at the

edges of the bars, or small, numerous, and irregularly arranged on

head, body, and fins.

Coasts of China and Japan to the Malay Peninsula and

Archipelago.

1, 2-5, 6-7. (650mm., 370-540 mm., Japan.

125 and 245 mm.)

8. (580 mm.) Formosa. R. Swinhoe. Esq.

9. (450 mm.) Singapore. Fisheries Exhib.

10. (250 mm.) Manado. Dr. B. Meyer.

11. (220 mm.) Amoy.

12. (170 mm.) Manilla. ‘ Challenger.’

13. (150 mm.) China. J. R. Reeves, Esq.

7, CHILOSCYLLIUM INDICUM. (Plate XIII. fig. 2.)

Squalus indicus Gmelin, Linn. Syst. Nat. p. 1503 (1788);

Schneid. Bloch’s Syst. Ichth. p. 1387 (1801).

Squalus tuberculatus Schneid. 1. ec.

Chiloscyllium tuberculatwm Mill. & Henle, Plagiost. p. 19

(1841); Bleek. Verh. Bat. Gen. xxiv. 1852, Plagiost. p. 17;

Dumeril, Elasmobr. p. 331 (1865); Kner, ‘ Novara’ Fische,

p. 412 (1865).

Squalus caudatus Gronoy. Syst. p. 8 (1854).

Chiloscyllium phymaiodes Bleek. t. c. p. 21; Duméril, 1. c.

Synchismus tuberculatus Gill, Ann. Lye. N. York, 1861, p. 408.

Chiloscyllium indicum, vars. @, e, and ¢ (part.), Giinth. Cat.

Fish, vil. p. 412 (1870).

Three prominent dorsal keels. Mouth much nearer to the

1908. | OF THE FAMILY ORECTOLOBIDA. 363

vertical from the eyes than to the end of snout; fold of lower

lip continuous; nasal cirrus extending to the edge of nasal valve.

Anterior edge of spiracle vertically below the posterior edge of

eye. First dorsal originating above the posterior end of base

of pelvies; free edge of the fin straight or convex; length of

base 3 to 3 the distance from second dorsal. Origin of anal at a

distance from the vertical from end of base of second dorsal

considerably greater than the length of the latter ; length of base

of anal equal to or a little greater than that of the lower caudal

lobe in front of the notch. Pectoral extending about 2 of the

distance from its origin to that of the pelvics. Head and body

with dark reddish spots or vermiculations, some of which may

unite to form pairs of transverse stripes.

From the Cape of Good Hope to China.

1. (275 mm.) skin, type Gronow Coll.

of the species *.

2. (270 mm.) type of Java. Dr. P. Bleeker.

C. phymatodes.

3. (380 mm.) China. Sir J. Richardson.

4,5. (250 and 290 mm.)

6. (430 mm.) stuffed. Cape of Good Hope. Sir A. Smith.

8. STEGOSTOMA.

Stegostoma, Mill. & Henle, Arch. f. Naturgesch. 1837, i.

p- 395, and Plagiost. p. 24 (1841); Giinth. Cat. Fish. viii.

p. 409 (1870).

Head obtuse; body elongate, compressed posteriorly. Mouth

transverse ; teeth small, tricuspid, in many series, occupying in

each jaw a flat, four-sided area; lower lip not divided by a

symphysial groove. Nasal valves separate, each with a cirrus.

Hye small; no longitudinal fold below the eye. Spiracles mode-

rately large, nearly vertical, behind the eyes. Gill-openings of

moderate width, last three above the base of the pectoral ; last

two very close together. First dorsal partly above and partly

in advance of the pelvics, somewhat larger than the second,

which is partly above and partly in advance of the anal, which

terminates immediately in front of the caudal. Caudal very

long; axis not directed upwards; lower lobe notched posteriorly.

Pectorals broad, with straight or notched edges and rounded

angles.

A single species from the Indo-Pacific.

STEGOSTOMA TIGRINUM.

Squalus tigrinus Gmelin, Linn, Syst. Nat. p. 1493 (1788);

Forst. Zool. Ind. p. 24, pl. xiii. fig. 2 (1795).

Squalus longicaudus Gmelin, t.c. p. 1496.

* Tam unable to understand Dr. Giinther’s statement that the anal fin is cut

away in this specimen. It is true the species was described by Gronow as lacking

an anal fin, but this was because he made the very natural mistake of regarding the

anal fin as part of the caudal, as is evident from his description :—“ Lobis qui subtus

caudam occupant bint lineares, longi, equales, convext.”

364 ON SHARKS UF THE FAMILY ORECTOLOBIDE. [| Apr. 28,

Squalus fasciatus Bloch, Ausl. Fische, pl. exii. (1795);

Schneid. Bloch’s Syst. Ichth. p. 130 (1801).

Scyllium heptagonum Riipp. Neue Wirbelth., Fische, p. 61,

pl. xvi. fig. 1 (1840).

Stegostoma fasciaitum Mill. & Henle, Plagiost. p. 25, pl. xvi.

(1841); Cantor, Cat. Mal. Fish. p. 396 (1850); Bleek. Verh. Bat.

Gen. xxiv. 1852, Plagiost. p. 23; Duméril, Elasmobr. p. 336

(1865) ; Giinth. & Playfair, Fish. Zanzibar, p. 140 (1866); Klunz.

Fische Roth. Meer. p. 672 (1870).

Stegostoma carinatum Blyth, Journ. As. Soc. Bengal, xvi.

SA ap sad 2p mpl excxaye Oot Sele

Squalus cirrosus Gronov. Syst. p. 6 (1854).

Stegostoma tigrinum Giinth, Cat. Fish. viii. p. 409 (1870);

Day, Fish. India, p. 725, pl. clxxxvii. fig. 4 (1878).

Three dorsal keels, as in Chiloscyllium indicum, more prominent

in the adult than in the young; adults also with a pair of lateral

keels. Mouth a little nearer to the vertical from the eyes than

to the end of snout; fold of lower lip broadly interrupted; nasal

cirrus extending to the edge of nasal valve. First dorsal rather

elongate, gradually increasing in height posteriorly and ter-

minating above the end of base of pelvics; free edge convex,

angles rounded; second dorsal originating directly behind the

first, terminating above the middle of anal. Caudal as long as

the rest of the fish. Pectoral nearly reaching the origin of

pelvics. Young with broad dark brown cross-bars with blackish

margins, wider than the pale yellowish interspaces; during

growth the bars become paler and dark spots appear on them,

whilst the dark edges of the bars break up into spots; adults

without bars, with numerous vertical series of dark rounded

spots.

: Indian Ocean ; East Indian Archipelago ; China.

1-2. (1120 and 750 mm.) Zanzibar. Lieut-Col. Playfair.

3-4. (650 and 420 mm.) Formosa. R. Swinhoe, Esq.

5. (300 mm.) Rejang R., Sarawak. B. Low, Esq.

6. (200 mm.) N.W. Australia. Capt. D. Le Sauvage.

In addition to the specimens listed above, on which my de-

scription is based, there are some large stuffed examples from

India (Jerdon), Ceylon (Zool. Soc.), and Zanzibar (Playfair).

EXPLANATION OF THE PLATES.

Pruate XI.

Hig. 1. Chiloscyllium griseun.

2. Orectolobus ornatus ; 2a, head seen from below.

Prats XII.

Fig. 1. Chiloseyilium plagiosum.

2. Orectolobus tentaculatus ; 2a, head seen from below.

Prate XIII.

Fig. 1. Chiloscyllium grisewm.

2. C. indicum ; 2a, head seen from above; 24, from below.

1908. | ON A RARE OLIGOCH ATE WORM IN ENGLAND. 36D

2. A Note on the Occurrence of a Species of Phreatothrix

(Vejdovsky) in England, and on some Points in its

Structure. By Frank H. Bepparp, M.A., F.RS.

F.Z.8., Korresp. Mitgl. d. K. Bohm. Ges. Wiss., &e.

| Received April 28, 1908. ]

(Text-figures 76 & 77.)

A few days since Dr. Harmer, F.R.S., was so good as to

forward to me a number of worms which had been sent to the

Cambridge Museum for identification under the following

circumstances :—A gentleman resident near Cambridge has bored

a well upon his property, and in the water drawn from this well

appeared considerable numbers of an Oligochzete which proves to

belong to the genus described some thirty years since by Prof.

Vejdovsky, C.M.Z.S.*, from examples supplied to him from a

well in Prague. So far as I am aware, that memoir is the only

one of recent date which contains any information upon this

worm, which appears, however, to have been originally described

about one hundred years earlier from the same city. It is clearly

therefore a genus which is limited to underground waters; and

IT am not aware of any other record of its occurrence save those

referred to, and that to be communicated in the present report

to the Zoological Society of London.

I am therefore able to record the presence of this interesting

Lumbriculid in this country for the first time.

The account given of it by Vejdovsky + seems to me, after

having carefully examined a number of examples of the living

worm, to have included all the chief points in its structure.

There remain some minutie in which [ differ from Vejdovsky or

supplement him. The differences are, I am inclined to think,

due to a difference between the species.

In his account of the species given in his great work upon

the Oligocheta =, Vejdovsky figures the minute tactile processes

which stand stiffly out from the prostomium. JI find that these

processes are not limited to the prostomium—though doubtless

more numerous there than elsewhere,—but occur also even on the

seta-bearing segments for some distance back. As I shall bring

forward in the sequel some reasons for regarding this species as

being hitherto undescribed, this point may be possibly urged as

among those which prove this view.

The sete do not appear in their general arrangement or form

to differ from those of Phreatothrix pragensis. But in one of the

examples which I studied (text-fig. 76) I found a persistence of setze

on the first segment, which is at least extremely unusual among the

* Zeitschr. wiss. Zool. Bd. xxvii, p. 543 (1876), with preliminary notes in earlier

papers quoted by him.

+ Zeitschr. wiss. Zool. Bd. xxvii. p. 548.

Syst. u. Morph. d. Oligochzten, Prag, 1884, p, 54.

366 MR. F. E. BEDDARD ON A RARE [Apr. 28,

Oligocheta. On the first segment of one specimen the ventral

setzee were evident on both sides of the body, but showed an

inequality of development. On the right side there were two

setae, but both of them immature, the immaturity being unequal.

One of the sete was like the “‘Soies de remplacement” of other

segments. It was complete down to the node. The other seta

was merely a slender short rod of chitin, On the left side of the

body I only found one rudimentary seta, which was like the least

developed seta of the right side and merely a short slender rod

exactly of the same form and size.

Text-fig. 76.

Phreatothrix cantabrigiensis.

Ventral pair of sete of segment I.

The vascular system seems to present certain recognisable

differences from that of Phreatothrix pragensis described by

Vejdoysky, and it is really mainly by reason of this that I venture

to create a new species for this Phreatothrix from Cambridge.

Vejdovsky has figured * the way in which the dorsal vessel ends

anteriorly. It appears to bifurcate in the fourth segment and to

give off anteriorly to this point three pairs of lateral vessels

which join the ventral vessel.

T found in an example, studied with a special view of elucidating

the mode of bifurcation of the dorsal vessel, a different arrange-

ment anteriorly. The dorsal vessel (text-fig. 77) divided quite

at the anterior end of the body, and the first pair of transverse

vessels uniting it with the ventral vessel arose from it some

way behind this point of bifurcation, On the other hand, these

‘two lateral vessels joined each half of the ventral vessel formed

by the splitting of the dorsal vessel some way before the latter

united to form the single ventral blood-vessel. This very con-

siderable difference appears to me to be quite of specific value,

and there is hardly room for an error so very considerable, either

on my part or on that of the Bohemian Professor.

* Zeitschr. wiss. Zool. ¢. c. Taf. xxxix. fig. 2.

1908. } OLIGOCHZTE WORM IN ENGLAND. 367

There is another important point in the structure of the

vascular system in which I find that my species does not agree

with Phreatothrix pragensis. It is characteristic of the latter, as

of some other Lumbriculids, that the dorsal vessel in the intestinal

region gives off a series of blindly ending transverse appendages,

which are contractile. Such structures demand the living worm

for their successful demonstration. J examined several specimens

with great care in order to ascertain if these characteristic cecal

vascular trunks were to be seen; and I utterly failed to see them.

Text-fig. 77.

Phreatothrix cantabrigiensis,

Anterior end of dorsal and ventral vascular trunks.

D. Dorsal vessel. | H. “Hearts.” | V. Ventral vessel.

I cannot think that they were present in the greater part of the

intestinal tract, where I sought them in vain. I have the more

confidence in my ability to see these contractile appendages in

that I recognised the network of non-contractile vessels upon the

gut, which Vejdovsky does not record in Phreatothrix pragensis.

These vessels formed a close lattice-work upon the gut, quite

Proc. Zoou. Soc.—1908, No. XXIV. 24

368 MR. F. E. BEDDARD ON A RARE [Apr. 28,

similar to that which has been described, for instance, by Stole in

Lophocheta*, whose excellent figure of the same has been repro-

duced by myself in my ‘Monograph of the Oligocheta’ 7. This

system of capillaries was only to be clearly seen during the diastole

of the dorsal blood-vessel.

In certain other minutie of structure the present examples also

do not agree with the species P. pragensis; for instance, I found in

several examples of my species that the investment of chloragogen

cells of the alimentary tract commences in segment vill.

Vejdovsky mentions segment xil. as being that in which this

investment is first visible in Phreatothrix pragensis. Vejdovsky

has mentioned that in the living worm the mouth-aperture is

seen to be covered with ciliated cells, the cilia being in active

motion. It appeared to me that this ciliation in my species

extended also on to the prostomium, but certainly not far on to that

part of the body. It was in any case very clearly visible in the

mouth and just at its outside. This state of affairs was, however,

quite unrecognisable in one of the two specimens in which I

particularly studied it. It is not very easy to miss actively

vibratile cilia in this position, so that I am confident in my

assertion that there was no active ciliation in one specimen.

Whether this is a variable character in the species or was due to

temporary cessation of movement { I am unable to say.

The nephridia of Phreatothriz are in some ways remarkable.

Vejdovsky has pointed out that a single pair may occupy a

number of segments. I found precisely the same extension of

the nephridia through several segments in the examples which I

have studied. There is, however, some difference in the earlier

account given by Vejdovsky from the later account of the

nephridia of Phreatothrix pragensis which is important, in view

of the facts which I have noted in the position of the nephridia

of my own species of Phreatothrix. In the earlier paper on the

species (that in the ‘Zeitschrift fiir wissenschaftliche Zoologie’)

he figures § two pairs of nephridia, a pair in each of segments vii.

and vili., each pair only occupying its own segment, though the

funnel of course lies in the segment in front. In the ‘System

und Morphologie der Oligochzten’ || he figures a pair which open,

indeed, on to the vulith segment, but which extend back as far

as the xivth. These are followed by another pair which open on

to the xivth segment, but which extend back as far as the xxist.

Both individuals figured are represented as sexually mature. It

may be that Vejdovsky had before him examples, not only of

Phreatothrix pragensis but also of the species with which I deal

here, and which I believe to be distinct from that.

I have myself found a nephridium on each side in segment Vii. ;

* Abhandl. Bohm. Ges. Wiss. (2) vii. (1889).

+ Oxford, 1895, p. 241, fig. 41.

+ I kept the specimen under intermittent observation for most of a morning, and

supplied fresh water at times.

§ Tom. cit. pl. xxxix. fig. 2. || Pl. xi. fig. 18.

1908. ] OLIGOCHZTE WORM IN ENGLAND. 369

apparently these nephridia are confined to that segment, with

the exception, of course, of the funnels. In the following segment,

viz., viil., there was another pair of rather smaller excretory

glands. A third pair of the same size, or nearly so, were to be

found in segment ix. Then followed a gap and the next pair

were in segment xv., extending, however, through the two

following segments as well—that is to say, three in all, exclusive

of the segment which contains the funnel. Further back still

a pair of nephridia extended through four segments which I

have not mapped accurately. It is clear that in any case this

species shows some differences from both accounts given by

Vejdovsky. But the facts are not irreconcilable with his earlier

account, which, as I have suggested, may refer to the same species

which I consider here, or may merely lend evidence to the view

that the species varies in the number and position of its

nephridia.

The generative organs appear to be like those of Phreatothrix

_pragensis, save in one respect. I have, however, examined only a

single sexually mature individual. In Phreatothrix pragensis

there is a second pair of spermathece in segment xii. rather

smaller than those of segment xi. I looked carefully for, but was

unable in my species to find, the second pair. I noted the

sperm-sacs to lie in segments xi., xii., and xiii. The sperma-

thece and atria open behind the ventral pair of sete, which are

present and have not disappeared. This is quite in accord with

Vejdovsky’s statements concerning Phreatothria pragensis.

To resume: this species from Cambridge appears to differ from

that described by Vejdovsky from Prague in the following

points, viz.:—(1) The chloragogen investment of the gut begins

in segment vii.; (2) the dorsal and ventral vessels divide to

reunite much further forward in the body ; (3) there are no con-

tractile appendages of the dorsal vessel; (4) the number and

arrangement of the nephridia is somewhat different; (5) the

spermathece of segment xii. have totally disappeared.

It seems to me that we have here a considerable number of

differences which warrant the erection of a new species for the

worms from Cambridge, which I propose to term Phreatothriz

cantabrigiensis, or, in accordance with Michaelsen’s views *,

Trichodrilus cantabrigiensis. Possibly this second species with

nephridia extending through a considerable number of segments

may be regarded as furnishing an argument for reinstating the

genus Phreatothrix. On the other hand, inasmuch as this

character is to be found in Séylodrilust vejdouskyi—a species

belonging to a genus where there are no contractile appendages

to the dorsal vessel,—the question of the generic distinctness of

other Lumbriculids is perhaps raised by the facts which I am

here able, to bring to the notice of the Society.

* Ohgocheta in ‘ Das Tierreich,’ Berlin, Oct. 1900, p. 58.

+ Benham, Quart. Journ. Micr. Sci. vol. xxxiii. 1891, p. 42.

24%

370 MRS. E. W. SEXTON ON A [ Apr. 28,

3. On the Amphipod Genus Trisehizostoma.

By HE. W. Sexton.

[Received March 14, 1908. |

(Plates XIV.-XXI.*)

During the cruise of the ‘ Huxley’ in the Bay of Biscay,

August 1906, a small collection of Amphipoda was taken, which

was handed to me by Dr. Allen for examination.

Amongst them was a large female specimen of Z'rischizostoma

niceense. On looking into the literature of the group there

appeared to be so much confusion with regard to the two genera

Guerina and Trischizostoma, that a thorough revision has been

necessary.

To those who so kindly assisted me in this, my best thanks

are due: to Prof. G. O. Sars for his gift of a co-type of 7. raschii;

to Dott. A. Della Valle for sending me his series of preparations

of Guerina niceensis ; to the Rev. T. R. R. Stebbing for permitting

me to see his (as yet unpublished) manuscript on the $8. African

Lysiauasside, containing the description of a new species of

Trischizostomat; to Mons. EK. Chevreux for his information con-

cerning his specimen of Guerinella niceensis; to Dott. A. Brian

for his paper on a specimen from the Gulf of Genoa; to Mr. W.

M. Tattersall for his kindness in forwarding all his specimens

to me as soon as he heard I was engaged on this paper; and to

Dr. Calman for much assistance and for revising the manuscript.

I. Huisvroricat.

The first specimens of this genus were taken at Nice by the

Rev. F. W. Hope and sent by him to the Director of the Naples

Museum, Dott. Achille Costa, to be described and added to the list

of Crustacea then being prepared for publication in the ‘ Fauna

del Regno di Napoli’ (2).

The description was written by Costa in 1853,a new genus

Guerinia, Hope, being formed, and the type species, Guerinia

miceensis figured, and was published at once with an account

of two other crustaceans, under the title of “Tre nuovi

Crostacei discoperti dal Rev. Gugl. F. Hope” (1). The description

was published again, later in the same year, in the ‘ Fauna,’ but

the plate, although alluded to as plate vil. in the text, was

omitted.

Costa recorded the specimens as parasitic on various fish, ‘“ sopra

uno squalo e sul merluzzo ordinario,” to which they were clinging

* Wor explanation of the Plates, see p. 400.

+ Since this paper was sent to press, Mr. Stebbing’s article, cited below (20),

containing the description of this new species, T’vrischizostoma remipes, has been

published in the ‘Annals of the South African Museum,’ vol. vi. pt. 1: South

African Crustacea, pt.4, by the Rev. T. R. R. Stebbing, M.A., F.R.S., F.LS., F.Z.8,

With 14 plates. Cape Town: April 2,1908. Pp. 59-64, pl. xxxiv.

ZS. 1908; Pl XIV:

EW.Sexton del.

MP Parker lith. Parker b West imp.

TRISCHIZOSTOMA NICASENSE, Costa.

P ZS. 1908, PIX:

}

——

= W.Sexton del ;

* MP Parker lith Parker & West imp.

TRISCHIZOSTOMA WNICAENS#E, Costa.

P_Z.S. 1908, Pl XVI.

Parker & Westinsp.

EW Sexton del

MP Perker lith.

Costa.

NICAENSE,

TRISCHIZOS TOMA

My idea

SPI DIBA

LEE So

Azo EE IZ

Henne eres

——

TSF

P.Z.S. 1908, Pl XVII.

E W.Sexton del.

Parker & West imp.

MP Parker hth.

TRISCHIZOSTOMA NICASENSE, Costa, figs. l-la.

SCHII, Hsmark & Boeck, fig.18.

TRISCHIZOSTOMA RA

P.Z.S. 1908, Pl. XVII.

be LLE_E

VY, / —

7) Y

EW. Sexton del.

MP Parker hth. Parker k West imp.

TRISCHIZOSTOMA PASCHI, Esmark & Boeck.

IP Gos), WSYOS, IPI IIDC:

E W.Sexton del :

MP Parker hth. Parker & West imp.

TRISCHIZOSTOMA RASCHII, Esmerk & Boeck, figs. 2-1.

TRISCHIZOSTOMA NICASENSE, Costa, fig.1.

P.Z.S. 1908, Pl. XX.

E W.Sexton del.

MP Parker lith. Parker & West imp.

TRISCHIZOSTOMA RASCHII, Esmark & Boeck.

P.Z.S. 1908, Pl. XXxI.

E W.Sexton del.

MP Parker hth. Parker & West imp.

TRISCHIZOSTOMA RASCHIiI, Esmark & Boeck, figs 118; 15-18.

TRISCHIZOSTOMA NICAENSE, Costa, fig. 14.

1908. | GENUS OF AMPHIPOD CRUSTACEA. Bye

by the strong claws of the first gnathopods. The figure given is

that of a female, 25 mm. in length.

In 1860, Axel Boeck published his ‘Observations on the Nor-

wegian Amphipoda’ (3). In this work he added a new tribe to

the three already established primary divisions of the Amphipoda.

This tribe, Prostomate, was created for the reception of a “‘ new

and remarkable form,” 7rischizostoma, which Boeck regarded as

a transition between the Hyperide and Gammaride, resembling

the former in the structure of the head, the eyes, the antenne,

and the abdomen, and haying much in common with the Gam-

maride, especially with the family Orchestide, and the genera

Opis and Anonyx.

The Prostomate contain only the one genus Trischizostoma (so

named by Esmark, who first distinguished it, from the trifid tube

formed by its mouth organs) with the type species 7’. raschii, in

describing which Boeck specially notes the peculiar structure of

the first gnathopods.

The specimens described by Boeck, three large females, were

dredged by Prof. Rasch at ‘“‘ Havbroen,” a bank 20 miles off the

west coast of Norway, in 100 fathoms.

Tn 1862, Spence Bate in his British Museum Catalogue (4)

described and figured Guwerinia niceensis, placing the genus

Guerinia in the subfamily Phowides of the Gammaride, between

the genera Lafystius and Lepidactylis. He adds, “For the

‘description of this animal I am dependent upon the accuracy of

the Rev. Mr. Hope’s figure in the pamphlet quoted (Three New

Crustacea, Fauna of Naples)”; but both the description and the

figures, though evidently copied from Costa’s, are inaccurate and

misleading. For instance, in describing the second gnathopod,

Costa states that it has one joint less than the normal number,

being “entirely without a nail,” and further, that “the fifth and

last (article), which represents the hand, is the shortest of all, com-

pressed, narrow at the base, dilated inferiorly, where it is fringed

with stiff hairs which increase in length towards the anterior

angle, the longest being twice the length of the hand”*. Spence

Bate gives the normal number of joints, applying Costa’s descrip-

tion of the hand (propodos) to the nail (dactylos) thus— “ pro-

podos very small: dactylos triangular, dilated, compressed,

flattened at the apex, fringed with long hairs.” In order to bring

the figure into agreement with his description he has emphasised

the line by which Costa indicated the articulation of the last

joint, so making it appear as another very small joint and calling

it the ‘“ propodos.”

In 1865, Lilljeborg (5) published two papers on the Lysianassina,

the “ Lysianassa magellanica ete.” in English, and the “ Bidrag

till Kinnedomen” in Swedish. They contain practically the same

* “T] quinto ed ultimo (articolo) che rappresenta la mano é il pit corto di tutti,

compresso, stretto alla base, dilatato inferiormente, ove é ornato di peli rigidi e

crescenti in lunghezza verso V’angolo anteriore, i maggiori essendo lunghi il doppio

della lunghezza della mano.”

372 MRS. E, W. SEXTON ON A | Apr. 28,.

matter. In the table given of the Gammaride, he brackets to-

gether as closely related “Subfamilia 4. Phowina (Phoxides,

Spence Bate), and Subfamilia 5. Zvrischizostomatina (Prostomatee

A. Boeck)”—“ partes oris appendiculares non tubiformes ” in

Phowina, and “ tubiformes” in T'rischizostomatina.

In 1867, Costa again refers to Guerinia in the catalogue of the

Crustacea sent to the Paris Exposition (6) and gives the plate

promised but not included in the ‘ Fauna di Napoli.’

In 1870 Boeck published his ‘Crustacea Amphipoda Borealia’(7),

a synopsis of the larger treatise then being prepared for the press.

In this work he alters his previous classification of the group,

the former tribe Prostomate being reduced to a family of the

Gammaride. The diagnoses here given of the family Prosto-

matide with its one genus Tvrischizostoma and one species

T.. raschit are repeated in the later work, ‘De Skandinaviske og

Arktiske Amphipoder’ (8), where Boeck again emphasises the

resemblance to the Hyperina, and to the Orchestide and Lysianas-

sine among the Gammarina.

Boeck records the capture of several specimens, all females, by

Storm, in the Throndhjemsfjord, “‘ parasite on a shark ”—length

from rostrum to telson of the largest specimens 25-30 mm. ; and

of “‘one very young one’ ” taken by himself trawling in the

Christianiafjord, at the depth of 60 fathoms.

Sars in the ‘Oversigt af Norges Crustaceer,’ 1882 (9), follows

Boeck’s classification, giving coraclere Tribe II. Guinnanmina. Fam. I.

Trischizostomide, one genus, Trischizostona, one species, T. raschii.

In 1885, Carus in his ‘ Prodromus’ (10) places Guerinia in the

subfamily Phowine of the Gammaride between the genera.

Lilljeborgia Sp. Bate, and Gammarus (Fabr.) Sp. Bate. His

description appears to be a literal translation of Spence Bate’s

definitions (4). He notes G. miceensis as peculiar to the

Mediterranean fauna.

In 1886, Bovallius (11) established a new tribe Synopidea,

intermediate between Gammaridea and Hyperiidea, in which he

placed the family Z’rischizostomatide. He gives full descriptions

and figures of the adult female and a young male (5 mm. in length)

of Trischizostoma raschii. The peculiar aspect of the first

enathopods he discovered to be due to torsion in the adult, being

wholly absent in the young form.

About 20 young specimens were taken by Bovallius in July

1871 at Tjgttd, Norway, 20 miles south of the Polar Circle, in 80

fathoms; in July 1880 he found some adult females, “ parasites

on, or in company with, an Asterias” in Hardangerfjord, S.W.

Norway, in 250 fathoms, the largest of these measuring 22 mm.

He mentions Capt. Collin as having obtained specimens from the

west coast of Novaya Zemblya.

Stebbing in 1888 (12), in his introduction to the ‘Challenger ’

Amphipoda, p. xix, writes: ‘ A connection between the Hyperina

and the Lysianasside has already been indicated by Boeck, who

placed the family Prostomatide at the head of the Gammarina,.

1908. ] GENUS OF AMPHIPOD CRUSTACEA. 313

in immediate sequence to the Hyperina because of the agree-

ment which he considered to exist between that family and the

Hyperide and Orchestide. The Prostomatide are in close

relationship with the Lysianasside and might, in my view, well

be included in the older family.” Stebbing was the first to per-

ceive the great resemblance between the Mediterranean and

Norwegian forms, and says (p. 272) of Guerinia nicwensis : ‘* It is

beyond doubt, generically, perhaps also specifically, identical with

the later Trischizostoma raschii Esmark & Boeck, 1860”; and

again (p. 321) of Trischizostoma, “the genus, at least, is assuredly

a synonym of Guerinia Hope & Costa.”

In 1890, Sars (13) demonstrated clearly that Trischizostoma is

a true Lysianassid, basing his conclusion on the structure of the

oral parts, anterior antenne, posterior gnathopods, and the bi-

articulate 3rd uropod, and pointing out that “the urosome, which

Boeck considered Hyperiidean in character, essentially differs by

being divided into three distinct segments.” He, therefore,

places the genus in the family Lysianasside, giving detailed

descriptions and figures of the adult female and the young form.

In the summers of 1890-91 Sars obtained several specimens, all

females, most of them ovigerous, in the Throndhjemsfjord, on the

common black dogfish (Spinaa niger). The largest measured

28 mm.

Bonnier in his discussion of the “ Lysianassides” (14) 1892,

refers to the difficulty of differentiating between the various

genera, and suggests ‘des diagnoses courtes basées sur les

véritables différences morphologiques de la structure des somites

et leurs appendices, ou des clefs dichotomiques permettant

d’arriver au genre et a l’espéce par l’examen d’un petit nombre de

caractéres, sans avoir & comparer une a une les descriptions et les

figures des nombreux types qui constituent la famille des Lysianas-

sides.” He first gives the characters common to the family, and

then proceeds to characterise the different genera according to the

structure of the 1st maxilla, the maxilliped, the Ist pereeopod,

the telson and the antennule. The Ist maxilla in particular he

considers useful in characterising most of the forms; for example,

the four genera, T’rischizostoma, Acidostoma, Acontiostoma, and

Amaryllis are at once distinguished from the rest by the modifi-

cation of the maxillary palp. He discusses at length the con-

clusions of Bovallius (11) and Sars (18), agreeing however entively

with Sars.

In 1893, Della Valle (15) in his ‘Sistematica’ gives 10 sub-

orders of the Gammarina, the tenth, Lysianasside, including the

two genera—Guerina and Trischizostoma. On p. 770 he points

out that the essential characters for distinguishing the Lysianassid

genera are—the peduncle of the anterior antenne ; the mandible ;

and the posterior gnathopod. In the table following, Guerima

and Trischizostoma (placed in close relation to Amaryllis and

Acidostoma) ave defined thus: “Nei gnathopodi anteriori dell’

adulto l’articolazione del 3°, 4°, e 5°, articolo é tale che il margine

374 MRS. E. W. SEXTON ON A [Apr. 28,

unguicolare della mano diventa anteriore. (La mano stessa é

enormemente ingrossata) ; nei piedi mascellari il 1° articolo del

palpo é pit lungo del 2° in Guerina,” this article being “ pit

breve del 2°” in T’vrischizostoma.

In answer to Stebbing’s words as to the “‘ generic and probably

specific identity of the two forms” he says:—‘ Senza dubbio fra la

Guerina del Golfo di Napoli e il Zrischizostoma della coste di

Norvegia ognuno a prima vista nota grande rassomiglanza, sopra

tutto per la forma generale del corpo, per la grandezza degli occhi

e per 1 gnathopodi anteriori. Nondimeno, considerando meglio

le varie appendici e piu di ogni altra cosa le parti boccali e 1 piedi

toracici del gruppo medio, la differenza dei due Gammarini riesce

evidente non solo specificamente, ma anche genericamente.” The

name Guerinia, being preoccupied in 1830, Della Valle changes to

Guerina. Figures are given of the male, the oral parts, gnatho-

pods, two perzeopods, and the urosome.

The three specimens described, all males, were taken on

“merluzzi” ; two on November 22, 1881, off Cuma in 250 metri ;

and one on December 10, 1881, “al largo d’Ischia ” at 150 metri.

Length 12-13 mm.

In 1895, Stebbing (16) in an article contributed to ‘ Natural

Science,’ again emphasises the close relationship of the two forms,

and considers the characters given by Della Valle too trivial to

warrant the placing them in different genera.

In 1903, Brian (17) published a note on the capture of a

Guerinia nicewensis in the Gulf of Genoa. The specimen, an ovi-

gerous female, 18 mm. in length, was found by Sig. Borgioli

in the mouth of a Chlorophthalmus agassizi. ‘The colour of the

living animal he describes as “ variante fra il giallo sporco e il

rosso mattone.”

Chevreux (18) in 1905 in his ‘ Liste des Gammarina’ taken by

the ‘ Princesse-Alice’ gives: 1. Guerinella niceeensis (Costa), and,

in an explanatory footnote, his reason for changing the name;

Guerinia and Guerina being so much alike as to lead to confusion,

and Guerinella conforming to the rules of nomenclature adopted

by the International Congress of Zoology.

The specimen, an ovigerous female, 19 mm. long, was taken

17/7/04 in the “ filet a grande ouverture” in the Bay of Biscay,

lat. 46° 15! N., long. 7° 09’ W., haul from 0-3000 metres. This

is the first record of this species out of the Mediterranean.

In 1906, Stebbing (19) in ‘ Das Tierreich’ combines the two

forms in one species 7'rischizostoma niceense, genus Trischizostoma,

family Lysianasside, taking 7. raschii as the female, and Guerina

niceensis as the male ; but in a manuscript as yet unpublished (20)

dated 24/7/07, which he has been kind enough to allow me to

make use of, he separates them specifically, still retaining, how-

ever, the one genus and including in it a third and new species

from South Africa. As he has pointed out (16) Boeck’s generic

name 7'rischizostoma, 1860, supersedes the Guerinia 1853 of Costa,

preoccupied in 1830.

1908. | GENUS OF AMPHIPOD CRUSTACEA. 375

The South African specimens are males, 10-13 mm. in length,

from Buffalo Bay—one taken in 32 fathoms 8.W. by W. ? W.,

34 m., and the others in 47 fathoms, lat. 33° 9’ 30’ S., long. 28° 3’

00” K.

In the following ‘Description of Species,”

the specimens

described are :—

One large ovigerous female, 7’. nicwense, 23 mm. long, taken

by Dr. Allen, 26.viii.06, on the ‘ Huxley’ in the Bay of Biscay, in

246 fathoms; lat. 48° 7’ N., long. 8° 13’ W.; Agassiz trawl.

Free-swimming.

And seven specimens taken by Mr. W. M. Tattersall on the

‘ Helga,’ as follows :—

7.vili.1904. Four specimens, 7’. nicwense: three females, re-

spectively 20, 20°5, and 22 mm. long, and one male, 21:5 mm.,

the first male recorded out of the Mediterranean. 50 miles

W.N.W. of Tearaght, Co. Kerry, Ireland, 396 fathoms. Net at

237 fathoms. Free-swimming.

5.xi.1905. One specimen, 7’. nicwense, a male, 20 mm. long,

50 miles W. 2 N. of Tearaght, Co. Kerry, Ireland, 411 fathoms.

Net at 75 fathoms. Free-swimming.

February 1906. One specimen, 7’. nicwense, a male, 21°5 mm.

Lat. 51° 54’ N., long. 11° 54’ W., 460 fathoms. Net at 350

fathoms. Free-swimming.

August 1906. One specimen, 7. raschii, immature female, 9 mm.

Lat. 50° 37' N., long. 11° 12’ W., 250-542 fathoms. ‘Taken in a

small net attached to the trawl. This is the first record of this

species out of Norwegian waters.

Il. DeEscRIPTion oF SPECIES.

Only three species of this genus are known so far: 7’. nicwense,

recorded from Naples, Genoa, Nice, the Bay of Biscay, and the

west coast of Ireland, of which the male and female have both

been taken; 7’. raschii, ranging from the Arctic Circle to

Christianiafjord, Norway, and from the west coast of Ireland,

only the female and young of this species are known; and

Mr. Stebbing’s new South African species, of which the male only

has been captured.

Detailed descriptions and figures of the first two species are

given here to prove the justice of their inclusion in the same

genus.

1. TRISCHIZOSTOMA NICHENSE (Costa). (Plates XIV., XV.,

XVI., XVII. figs. 1-12, Pl. XTX. fig. 1, and Pl. X XI. fig. 14.)

Syn. 1853. Guerinia niceensis Hope in litt., Costa (1).

1893. Guerina nicewensis Della Valle (15).

1905. Guerinella niceensis Chevreux (18).

1906. Trischizostoma nicwense, 3, Stebbimg (19).

Both the male and the female of this species are known. The

male has been figured by Della Valle, but no accurate figure of

376 MRS, E, W. SEXTON ON A [ Apr. 28,

the female has been published hitherto. The figure given by

Costa is evidently that of a female, but his specimens cannot be

traced. The last mention of them is in the ‘ Annuario’ (6),

where they are referred to as forming part of the Collection of

Crustacea sent to the Paris Exposition, 1867. The method of

preservation is described, the specimens being dried and mounted

in glass cells.

Though the geographical range of this species is wide, the

specimens taken have been few in number: Costa’s specimens ;

the three males described in Prof. Della Valle’s work (15); a

female from the Gulf of Genoa; two females from the Bay of

Biscay, one taken by the ‘ Princesse-Alice,’ and one by the

‘Huxley’; and Mr. Tattersall’s six specimens, all from the

west coast of Ireland, three females and three males. Thanks

to Mr. Tattersall’s kindness in permitting me to examine these

last and to dissect a male specimen, I have been able to satisfy

myself that the oral parts taken by Della Valle as characters

to differentiate the genera Gwuerina and Trischizostoma are

really identical in structure in both forms. The first maxilla,

in particular, which he describes as lacking the inner plate,

and with the palp reduced to a small and simple tubercle,

will be seen to possess not only the inner plate, but a minute,

distinetly bi-articulate palp (Pl. XV. figs. 1 & 2). These struc-

tures, however, are so exceedingly fragile and pellucid as to

render dissection very difficult. The little leaf-like palp arises

in a small hollow inside the margin of the outer plate, and sets

out at right angles to it; when mounted for the microscope the

weight of the cover-glass is quite sufficient to depress it into

the hollow, thus giving the effect of a little tubercle.

The description of the male is taken principally from the Irish

specimen, 21°5 mm. in length; that of the female from the

‘Huxley’ specimen, 23 mm.in length. All the measurements are

taken in the same way, from the tip of the rostrum to the tip of

the telson, along the medio-dorsal line.

There is little difference between the sexes, the principal dis-

tinguishing characters being found in the antenne.

The Jntegument is very characteristic, having the appearance of

“pitting”; under a high power each little pit is seen to be irre-

gularly six-sided and fringed with sharp spines (Pl. XIV. fig. 2).

The Head is much deeper than the perzeon, about as long as

deep; rostrum broad, apically rounded, curving right over the

bases of the superior antenne. The head is longer than the

first segment of the perzon (2°5mm. to 1-5 mm.),about as long

as the first segment and half the second. yes large, dark brown

in colour, the pigmented masses on each side numbering not less

than 60 ommatidia, arranged in eight or nine transverse rows,

with an irregular row of smaller unpigmented ommatidia entirely

surrounding the pigmented masses and meeting in the medio-

dorsal line.

- Prr#on.—The Ist segment is the longest; the 2nd, 3rd, and

1908. } GENUS OF AMPHIPOD CRUSTACEA. 377

4th are shorter and subequal; the 5th is smaller again; and the

6th and 7th are the shortest of all and subequal. All, except the

first, are produced at the posterior angles and rounded.

Side-plates.—Ahbout half as deep as the body (see perzeopod figs.

for the correct proportions. They are not well represented in

the figure of the whole animal owing to the immense distension

of the ovisac forcing them out of the normal position). The Ist

is small, triangular, almost entirely covered by the large 2nd.

This side-plate is the largest of all, greatly dilated inferiorly and

produced forward, posterior margin straight. The 3rd is almost.

as deep as long, with the anterior distal angle produced forward,

posterior margin straight, inferior margin rounded; the 4th is

similar, but smaller; the 5th and 6th successively smaller,

inferiorly bilobed; the 7th is the smallest, subquadrate.

PuiEon.—First three pleon segments subequal, large, equalling

the first pereeonal segment in length. Epimeral plates large and

rounded, the 2nd and 3rd with a lateral carina which in the 2nd

terminates in a denticle at the postero-lateral angle. The 4th

segment is deeply depressed dorsally ; the 5th is shorter than the

4th or 6th; the 6th is depressed dorsally and emarginate for

the insertion of the telson.

SuPERIoR AnTENNA.—Jemale (Pl. XIV. fig. 3). The first joint

of the peduncle is as long as the two following taken together,

broader than long, with a fringe of about 24 of the so-called

“auditory sete” around the distal posterior angle, and 8 smaller

similar sete: in a cluster on the proximal posterior margin.

The primary flagellum consists of one long broad joint and

eight small. The Ist joint nearly equals the peduncle in length

and is as long as the eight small joints taken together. It carries

on its inner surface two longitudinal bands of laminar hyaline

filaments (see fig.), about 38 transverse rows in each band. The

2nd joint widens distally and is furnished at the inner posterior

angle with a long, rigid, slightly curved spine reaching to the tip

of the flagellum; the 3rd has a similar but shorter spine inserted

at the outer anterior angle. The 2nd, 3rd, 4th, 5th, and 6th

joints are all fringed on the inner margin with a row of small,

stiff, curved sete, deeply inset ; the apical joint is tipped with two

long stiff sete.

The accessory flagellum is inserted anteriorly in a deep emargi-

nation of the peduncle and is composed of one long laminar joint.

and two small ones.

The number and proportions of the joints appear to vary with

the age of the specimen. The ‘Huxley’ specimen, just described,

and one of the Irish specimens, 20°5 mm. in length, have each

nine joints in the primary flagellum, with the first joint cylindrical

and swollen. The other two Ivish specimens, 20 mm. and 22 mm.

respectively, have eight joints, the first joint much more slender,

and incurved like that of the male, and the accessory flagellum

much longer in proportion. These two females appear to be

younger than the others—the claws and spines are much less worn.

378 MRS. E. W. SEXTON ON A [Apr. 28,

Male (Pl. XIV. figs. 4 & 5).—This antenna is much longer in

the male than in the female, measuring 5 mm. in a specimen

21:5 mm. in length, as compared with 3 mm. in a female specimen

of 23 mm.

The peduncle is not so long as in the female. First joint

nearly twice as long as the 2nd and 3rd taken together (measured

along the inner surface) with the ‘auditory setz ” as in female.

The primary flagellum consists of nine joints. The Ist is half as

long again as the others taken together; it is incurved and

‘covered on its inner surface with dense masses of the long sensory

filaments. The six following joits are narrow proximally,

widening distally, and carrying on their inner margins fringes of

the small, stiff, curved sete, the 2nd and 3rd having the long

rigid, outstanding spines as in female, and the 5th, 6th, and 7th

each with a calceolus. The 8th and 9th are slender and cylin-

drical, the 9th tipped with one long stiff bristle, one long seta and

‘one small one.

The accessory flagellum is much smaller than in the female, not

reaching to half the length of the first jot of the primary. It

is composed of one long laminar joint, one small joint slightly -

constricted in the middle, and a minute apical joint. The Ist jomt

has four small sete inset on the distal margin; the 2nd one

“auditory ” and one simple seta; and the terminal joint has two

of the “ auditory” and two or three simple sete.

Inrertor Antenna.—Vemale (Pl. XIV. fig. 6). The Ist joint of

the peduncle is produced posteriorly downwards in a thick laminar

lobe ; it is hollowed behind, and in this hollow lies the small

2nd joint with its large antennal cone; the 3rd joint is small, as

wide as long; the 4th is more than twice the length of the 3rd ;

the 5th is not as broad as the 4th and very slightly longer.

Sete.—The first three joints carry no sete. The 4th is fur-

nished on the posterior margin with seven long auditory sete

proximally and one at the distal angle; with eight tufts of small

setee on the anterior margin. The 5th has twelve of these tufts

along the anterior margin, and a fringe around the anterior angle ;

with one small seta at the posterior angle.

The flagellum consists of 29 joints, the first the largest ; 26 of

these are short, wider than long, decreasing gradually in size,

each with a row of sete inset anteriorly above the distal angle ;

the three terminal joints are very slender, cylindrical, the apical

joint having one long stiff bristle and one auditory seta.

In the Irish specimens the proportions of the joints of the

peduncle are the same as described above ; the number of joints in

the flagella vary—the 20°5 mm. specimen having 27; the 20 mm.

having 20; and the 3rd specimen of 22 mm. with 22.

Male (Pl. XIV. figs. 7,8, 9). The first two joints of the peduncle

as in female; the 3rd a little longer ; the 4th twice the length of

the 3rd; the 5th as long as the 3rd and 4th taken together, more

slender than the 4th, narrow proximally, slightly curved, with

the anterior margin concave.

1908. ] GENUS OF AMPHIPOD CRUSTACHA. 379

Sete.—The first three joints are without sete. The 4th has

twelve auditory sete on the posterior margin (fig. 9), eleven prox-

imally and one at the distal angle; anteriorly it is provided with

nine groups or rows of small setze set transversely on the margin,

with one auditory and one small seta at the distal angle. The 5th

joint has one auditory seta at the posterior distal angle; and, on

the anterior margin, eleven of the small transverse rows of sete,

with a fringe of sete and one auditory seta at the distal angle.

The flagellum is half as long again as the peduncle, filiform,

much more slender than that of the female. It consists of 38

small joints; the first with the posterior margin shorter than the

anterior, and the others with the anterior angle a little produced

downwards, giving an oblique look to the articulations. The first

25, subequal in length, gradually decrease in width, the following

joints being longer and narrower. Hach joint bears anteriorly a

little bunch of sete at the distal angle. There is a calceolus on

each of the first three joints, then one on alternate joints to the

35th, the calceolus (fig. 8) being set on a little protuberance above

the bunch of sete. The first joint also bears a large “ Rheich-

zapten.” The three terminal joints are exceedingly slender, the

apical one tipped with two sete.

In the second specimen, 21:5 mm. long, the flagellum is twice

the length of the peduncle and consists of 40 joints, the terminal

four very minute and without calceoli. The first four have each

a calceolus, after which they occur on alternate joints tothe 36th.

The third specimen, 20 mm. in length, has lost the tips of both

antennee ; 33 joints still remaining on each.

Ora Parrs.— Upper and Lower Lips (Pl. XIV. figs. 10 & 11).

The upper lip is elongate, deeply hollowed, apex entire and pro-

duced forward. The lower lip is divided at the tip into two

lobes, with a minute pellucid structure between the apices.

Manprpie.—Lemale (Pl. XIV. figs. 12 & 13). The large basal

portion carrying the palp is produced forwards in a laminar

process, with truncate cutting-edge, feebly spatulate, no molar.

On the inner surface distally are three spinules.

The Ist jomt of the palp is very small; the 2nd large, about

six times as long as the Ist, broad, rounded anteriorly, carrying

along the distal half of the anterior surface inside the margin a

row of 25 long, curved bristles, with plain shafts and minutely

serrate tips. The 3rd joint is lanceolate, as long as the 2nd,

but only half the width, with 17 long bristles on the anterior

margin of the right mandible, 18 on the left mandible. These

bristles are plumose for half their length and curved, the distal

three being more widely spaced, smaller and more curved than the

others. At the apex of the joint, set at a different angle from

the others and from each other, are two bristles, the apical one

half the length of the other, which is the longest on the palp;

the shafts are dentate on either side, and the tips are long, stiff

and finely plumose. The greater part of the joint is thickly

covered with fine transparent flat spines.

380 MRS. E. W. SEXTON ON A [Apr. 28,

Male (Pl. XIV. fig. 14). The apex of the front part of the trunk

is more spatulate than that of the female. The proportions of

the joints of the palp vary a little also, the 5rd joint being slightly

longer than the 2nd, which is more elongate in form than that

of the female. The row of bristles on the 2nd joint commences

lower down the margin; in the right mandible they number 34

on the 2nd joint, and 27 on the 3rd; the left mandible having

31 on the 2nd, and 26 on the 3rd. The numbers appear very

variable. In the Mediterranean specimen examined they were :

right mandible 37 and 26 respectively; and on the left 30

and 25.

Frrst Maxiuua (Pl. XV. figs. 1 & 2).—The outer plate is elon-

gate, with five strong claws apically, four much curved, and

serrated (see fig. 1, detail). The plate is contracted below the

apex and again at the insertion of the palp. The small pellucid

inner plate is of very delicate structure, unarmed. The palp is

bi-articulate, minute, set in a hollow and united along its length

to the inner surface of this hollow, the laminar leaf-like second

joint being twice the length and breadth of the first.

There is practically no difference between the sexes, except

that the first maxilla of the male is longer than that of the female

and the apical claws are longer.

SeconpD Maxitia (Pl. XV. fig. 3).—Slightly larger in male

than in female. Jnner plate small, tipped with one setule; outer

-plate broad and rounded distally, hollowed underneath, with three

setules on the apical margin and one on the inner side.

Maxiuuipeps (Pl. XV. figs. 4 & 5).— Female. The basal joints

deeply curved; the 2nd joint of the one maxilliped fused with

that of the other for nearly three quarters of its length. Jnner

plates fairly large, narrowing apically, unarmed. Outer plates

extending beyond the distal margin of the first joint of the palp ;

large, unarmed except for one or two microscopic setules.

The lst jomt of the palp is large and broad; the 2nd short,

widening a little distally, with one seta at each angle; the 3rd is

the largest of all (measured along the outer margin), lightly

curved, with seven transverse rows of strong bristles on its under

surface (see detail, fig. 5), and two sete on the inner distal

margin. Distally the under surface of the joint is covered with

a spinose armature similar to thet of the mandibular palp. The

Ath joint is subequal in length to the 2nd, narrow and unarmed.

Male. The basal portion and plates are the same as in the

female, but the proportions of the joints of the palp vary.

The Ist joint is the longest; the 2nd the shortest, with one

seta on its inner angle and two clusters of the long bristles distally

on the outer margin; the 3rd joint is slightly shorter than the

1st, curved, densely setose on its under surface, with about ten

transverse rows of long bristles; the 4th joint is subequal to it

in length, but only half its width and unarmed.

First GNATHOPOD (Pl. XV. figs. 6, 7, & 8) very powerful, with

the chitinous margins of extraordinary thickness, and longi-

1908. | GENUS OF AMPHIPOD CRUSTACEA. 381

tudinal ridges of chitin on all the free joints. The 2nd joint is

very long and curved; the 3rd, 4th, and 5th are subequal, the

srd and 4th continuing the curve of the 2nd; the 5th with a

longitudinal ridge produced to a point on each side of the articu-

lation with the 6th (see fig. 6). The 6th or “hand” is usually

carried as in fig. 6, immensely swollen, with the inner side (fig. 8)

rounded and more swollen than the outer (fig. 7). The palm

margin is bordered with strong, incurved, mobile teeth, 15 in

number, each with a tubercle behind (see fig. 8, detail), and with

five long, slender, mobile spines above the margin. The hand

has the deep groove or channelling described by Costa, “una

scanalatura nella faccia esterna,” into which the acuminate tip of

the long claw fits. At the prehensile angle, on the outer edge of

this groove is a very long, mobile, curved spine, with a small one

beside it, each with a tubercle posteriorly, while on the under

edge are two medium-size spines, one larger than the other; all

four spines are provided with small flagella. The powerful curved

claw or “ finger” is very long, longer than the palm margin, with

12 or 13 minute sensory setules along its inner surface.

The only difference between the sexes is—in the male the hand,

while equally broad, is slightly longer than that of the female, and

the palm margin is provided with more spines.

females. In the ‘ Huxley’ specimen, 23 mm., the width of the

hand is 2°5 mm., with 5 long spines and 15 small teeth on the

palm margin; hand figured (fig. 7). Of the Irish specimens,

the first 20°5 mm. long measures 3 mm. across the hand, with 5

long and 16 small teeth, hand as fig. 6; in the second specimen,

20 mm., the hand measures 3 mm., 5 long and 18 small teeth,

hand inverted; in the third female, 22 mm., the hand measures

2°75 mm., 5 long and 18 small teeth, hand as fig. 7.

Males. In the specimen figured, 21°5 mm. in length, the hand

measures 3 mm., 6 long and 19 small teeth (fig. 6). The second

specimen, 21'°5 mm., is exactly similar; the third of 20 mm.

length measures barely 3 mm. across the hand, 6 long and 18

small teeth, hand carried as in fig. 6.

SeconD GnatHopop (PI. XVI. figs. 1, 2, 3, 4, & 5),—The 2nd

joint is very long and slender, curved, widening distally; the

3rd is rather more than two thirds the length of the Ist, longer

than the two following taken together, laminar, the distal half of

the joint wider than the proximal; the 4th joint is small, narrow

proximally, twice as wide at the distal end, rounded posteriorly,

with the posterior margin twice the length of the anterior; the

5th is longer than the 4th, ovate elongate, narrow at both

extremities, considerably expanded posteriorly; the 6th bends

backward towards the 5th, almost discoidal in shape, as wide as

long, very narrow proximally, but expanding both anteriorly and

posteriorly. The 7th joint or claw is very minute, set trans-

versely in the middle of the distal margin of the 6th, so that the

point of the claw impinges against the under surface of the margin,

It is completely concealed by the dense masses of setze, which is,

389 MRS. E. W. SEXTON ON A | Apr. 28,

no doubt, the reason why it escaped detection by Costa (2) (‘‘ privo-

affatto di unghia”).

Sete.—The 2nd, 3rd, and 4th joints each carry a seta at the

posterior distal angle; the 5th is covered on the anterior and

posterior surfaces with dense masses of delicate, hyaline, sensory

hairs (fig. 4), with a fringe of long jointed setx at the anterior

angle, and four clusters of the same along the posterior margin

(for detail see fig. 5). The 6th joint is thickly covered with the

hyaline hairs, with a large number of the jointed set anteriorly,

increasing in length to the anterior angle, the longest being

twice the length of the joint; the posterior angle also bears a

cluster of shorter, jointed sete. These “hyaline hairs” and

“jointed sete” are peculiar to the 5th and 6th joints; the 6th

jot having yet another kind, stiff, curved, and serrate, similar

to that figured for 7’. raschwi (Pl. XX. fig. 3). A cluster of about

four to six of these is to be found on each side of the claw, and

three just beyond its tip. The claw itself is denticulated on its

under surface, and the portion of the margin against which it

impinges is thickly dentate (fig. 2).

PrR£OPODA.—Branchial vesicles occur on all the perzeopoda, long

and much pleated on the first and second, shorter and more

divided on the three posterior pairs. Incubatory lamelle are

attached to the first three, as well as to the second gnathopod.

The sixth joint of each perzeopod is produced over the base of the

claw in two delicate transparent plates, or ‘‘ dactyloptera” (see

Spence Bate, 4. p. 317), with pectinate margins (Pl. XVI. fig. 6).

The tactile spines of the perzeopoda and uropoda are of similar con-

struction. Hach consists of a stout shaft, blunt-tipped, carrying

subapically a slender flagellum (cf. Pl. XVI. fig. 3).

Frrst Perorop.—Ffemale (Pl. XVI. fig. 7). 2nd joint long,.

a little longer than the two following taken together, expanded,

rather narrowed proximally; 3rd very small; 4th long, a little

dilated anteriorly, 5th subequal to the 4th in length ; 6th longer

than the 5th, narrow; the 7th or claw moderately curved, about

half the length of the 6th.

Sete.—The 2nd and 3rd joints each carry one seta at the

posterior angle. The 4th has, anteriorly, one small seta and one:

large spine at the distal angle, with three sete along the poste-

rior margin and one at the angle. The 5th has two sete at the

anterior angle ; one at the posterior, and four along the margin.

The 6th has one small seta at the anterior angle; the posterior

margin is dentate, with seven strong sete inserted at intervals.

along it. The claw is provided with eight denticles on the

proximal half of the inner margin.

Male. 2nd and 3rd joints as in the female; 4th and 5th slightly

longer, the 6th and 7th distinctly longer, than in the female.

Setee.—The posterior margins of the 4th and 5th have each one

seta more, and the 6th two sete more, than the female; the

posterior margins of both the 5th and the 6th are dentate; the

claw with ten denticles.

1908. | GENUS OF AMPHIPOD CRUSTACEA. 383

SECOND PER&oPoD.— Female (Pl. XVI. fig. 8). Very similar to

the first pereeopod, but with the 2nd, 3rd, 4th, and 5th joints

shorter ; the claw is equally as long; the 2nd joint a little more

expanded than in the first pereeopod ; the setze on the joints much

less in number; claw without denticles.

Male. 2nd, 3rd, 4th, and Sth joints as in the female; 6th and

7th longer.

Setce as in the female.

The THirp PER#opop is the shortest of all.

Female (Pl. XVI. fig. 9). The 2nd joint long, broadly oval,

anteriorly rounded, posterior margin laminarly expanded and

produced a little downwards, widest proximally. The 3rd small;

the 4th and 5th practically subequal, the 4th a little dilated

posteriorly. The 6th is nearly as long as the 2nd, narrow. Claw

moderately curved, nearly two thirds the length. of the 6th.

Sete.—The 2nd joint has one minute sensory setule indented

in the middle of the posterior margin and three in the ante-

rior margin, with one large spine at the anterior angle. ‘The

3rd carries one spine on the anterior angle. The 4th has one

spine and two or three of the sensory setules posteriorly ; with

four spines on the anterior margin and two at the distal angle.

The 5th has three on the anterior margin and two at the angle.

The 6th is provided posteriorly with three minute setules and

anteriorly with five spines. The claw is apparently without

denticles.

The branchial vesicle extends to the distal margin of the 5th

joint.

Male. The 2nd joint is as long as the 2nd and 3rd taken

together in the female, with the anterior margin lightly concave ;

the following joints a little longer, the 6th decidedly so.

Setew.---As in the female, except that the 6th is furnished with

a few scale-like spines on the anterior distal angle, and the claw

bears four denticles.

Fourth Perxopop.—female (Pl. XVI. fig. 6, Pl. XVII. fig. 1).

2nd joint long, lightly concave anteriorly, laminarly expanded

posteriorly, and a little produced downwards; the 3rd small;

the 4th and 6th subequal to each other in length; the 5th shorter

than the 4th or 6th; 7th half the length of the 6th.

Setee.—The 2nd joint has one minute sensory setule inserted mid-

way on the posterior margin and one spine at the anterior angle.

The 3rd carries one at the anterior angle. The 4th has four on

the margin, and one large one at the angle posteriorly ; with four

at the anterior angle and six on the margin, two of these being

inserted in the same indentation, one a little behind and below

the other, an arrangement characteristic of the hinder perzeopoda.

The 5th carries two of these pairs and two single spines on the

anterior margin, and three large spines on the angle, which is

produced underneath in a pectinate fringe (cf. Pl. XVII. fig. 3).

The 6th bears five spines on the anterior margin, the distal half

of which is dentate. The claw is provided with two denticles,

Proc. Zoou. Soc.—1908, No. XXV. 25

384 MRS. E, W. SEXTON ON A [Apr. 28,

Male. The proportions of the joimts to each other are the same

as in the female, but all are longer, the 2nd, for example,

equalling the 2nd and 3rd of the female taken together.

Sete practically the same as in the female. The 5th has three

pairs and two single spines; the 6th one pair and five single,

with the distal half of the margin dentate; the claw has four

denticles.

Firrn Persopop.—Female (Pl. X VII. figs. 2 & 3). 2nd joint

long, a little concave anteriorly with an oblique groove mid-

way along the margin in which the branchial vesicle rests, less

expanded posteriorly than the preceding pereeopod ; 3rd small ;

Ath as long as the 2nd and longer and broader than the 5th or

6th ; 5th and 6th subequal ; 7th half the length of the 6th.

Setcee—This perzeopod is armed anteriorly with a formidable

array of spines. The 2nd joint has the one minute setule

posteriorly ; one large spine on the anterior angle. The 3rd has one

large spine. The Ath carries six on the margin and one at the

angle posteriorly; one pair and six single on the margin and

three at the angle anteriorly. The 5th has three of the sensory

setules and one spine posteriorly; the anterior margin dentate

with six pairs and one single spine inset and three at the angle

(fig. 3). The anterior margin of the 6th is also dentate with

one pair and five single spines inset; four sensory setules poste-

riorly. The claw is provided with five denticles (seven in another

specimen).

Male. Proportions of the joints to each other as in the female,

but all longer.

Sete.—Nine spines on the anterior margin of the 4th joint ;

three pairs and four single spines on the 6th, the rest as in the

female ; five denticles on the claw (three on the Mediterranean

specimen).

Puropops (Pl. XVII. figs. 4, 5, 6, 7).—-The peduncles are

large and stout, with two little Dane spies (fig. 5). The

rami are about twice the length of the peduncles. The cuter

ramus has 22 articulations, the long first joint carrying on both

margins proximally a tuft of fine hyaline sensory hairs with

flattened tips; the inner ramus is provided with similar tufts

and has six cleft spines (fig. 7) on the inner side of the first

joint, with an uncinate plumose seta (fig. 6) on each of the

following thirteen joints on the inner side.

Uropops (Pl. XVIT. figs. 8, 9, 10).—The peduncles are large

and stout; that of the Ist uropod ‘longer than the rami; that

of the 2nd subequal to the rami in length ; while in the 3rd the

rami are slightly longer than the peduncle. The apices of the

lst uropods reach considerably beyond the peduncles of the 3rd ;

those of the 2nd and 3rd are almost on a level (fig. 8). The

rami of the Ist and 2nd pairs are greatly curved.

The first wropod has seven small spines on the inner margin

of the peduncle which is serrated for half its length, with small

serrations along the distal margin also. ‘The outer ramus

1908. |] GENUS OF AMPHIPOD CRUSTACEA. 385

is shorter than the inner, with the outer margin armed with

seale-like spines and the inner with strong teeth (fig. 10). The

inner ramus has the margins serrated (fig. 9), the under margin

beset with two spines in the female and four in the male.

In the second wropod also, the outer ramus 1s shorter than the

inner. The peduncle is serrated along the distal margin and

around the inner angle, with two spines on the angle. The

rami as in the first uropod but without spmes. The puter ramus

of the third uropod is bi-articulate, with a spinule inserted at the

outer distal angle of the first joint ; all the margins serrate.

Texson (Pl. XVII. figs. 11 & 12) as bro ad as long, margin

entire. It carries four oblique rows of microscopic spiules, two

on each side of the medio-dorsal line, the distal rows longer than

the proximal. It is also provided with two rows of tufted setules,

four on one side and three on the other (fig. 12), each set in a

little pocket (cf. 7. raschii, fig. 13). These tufted setules are

peculiar to the telson.

The principal points of difference between this species and the

fellowing, are :—The shape of the rostrum, curvedin 7’. nicwense,

horizontal in 7’. raschii; the size and “veal er of the ommatidia ;

the shape of the 2nd, 3rd, and 4th side-plates; the 2nd mexilla,

the inner lobe small in 7. nicwense, subequal to the outer in

T. raschii; the maxilliped palp, the first joint long, the second

short in 7. niccwense, the reverse in 7’. raschii; the shape of the

hand of the first gnathopod ; the Ist and 2nd perzeopods, the

2nd joint being dilated in 7’. nicwense, the 4th in 7. raschii; the

shape of the Qnd joint of the 5rd pers weopods : ; and lastly the shape

of the telson, the margin entire in 7’. nicwense, excavated in

T. raschit.

TRISCHIZ0STOMA rascHit Ksmark & Boeck, 1860. (Plate X VII.

fie a Piss NOVEM SOXULX es 20) iss XOXC PONG iescel

13, 15-18.)

Syn. 1860. Zrischizostoma raschii Boeck (8).

1865. x ,, Lilljeborg (5).

1870. ue », Boeck (7).

1872. os », Boeck (8).

1886. = » Bovallius (11).

1890. re a poTasi(QLS))

1893. te », Della Valle (15).

1906. oe nicewense 2 Stebbing (19).

1907. 5 raschii Stebbing (20).

The measurements quoted in the following description are

taken from three specimens: an adult female from Norway,

measuring 26 mm. from the tip of the rostrum to the tip of

the telson; the immature specimen from the West of Ireland,

9 mm.; and a young one taken from the incubatory pouch of the

first specimen, 7mm. long. ‘The female had seven young still re-

maining in the ovisac ; the measurements of their appendages ete.

25*

386 MRS. E, W. SEXTON ON A [ Apr. 28,

do not vary by the fraction of a millimetre. The intermedite

specimen appears to be an immature female; no incubatory

lamellee are developed, but the proportions of the antennal joints

agree with the adult form.

Head.—Much deeper than the pereeon, as long as deep; ros-

trum horizontal, apically rounded. In the adulé the head is

slightly longer than the first segment of the pereeon (3 mm. to

2-5 mm.); in the immature it is much longer; and in the young

it is nearly twice as long (as 9:5). The huge Hyes cover nearly

the whole surface of the head; they are composed of a great

number of small, dark brown ommatidia, arranged in rows, with

one row of unpigmented ommatidia following the contour of the

pigmented mass and meeting in the medio-dorsal line. There

are not less than 154 of the dark ommatidia, each side, but they

are too closely crowded together to permit of ascertaining the

exact number.

In the immature the eyes are reniform, almost but not quite

touching dorsally, and not extending so far forward or downward

as in the adult; not less than 120 pigmented ommatidia. In

the young a large number of unpigmented ommatidia.

Prrxzon.—The Ist segment of the perzeon in the adult is the

largest, the four following subequal, the 6th and 7th the smallest,

all, except the first. produced at the posterior angle and rounded.

Side-plates.—Adult (see pereopoda figures). The Ist is very

small, triangular, almost completely hidden by the 2nd, which

is greatly dilated inferiorly, forming a large triangular lobe,

reaching anteriorly to the infero-lateral margin of the head,

and produced to a great length downward posteriorly, slightly

emarginate behind. The 3rd is in shape obliquely oval, anterior

margin convex, posterior emarginate, less than half as wide

and slightly shorter than the 2nd. The 4th is shorter again,

rounded anteriorly and deeply emarginate posteriorly, inferior

margin truncate. The Sth is bilobed, posterior lobe deeper

and wider than the anterior ; 6th of similar construction, much

smaller. The 7th is the smallest, wider than deep, with rounded

corners. In the tmmatwre specimen the proportions of the

perzeon segments and side-plates are the same as in the adult.

In young. The first pereeonal segment is the largest, all the

others subequal ; body deep; side-plates not so large in propor-

tion as in adult, those of the second segment differing a little in

shape.

Prieon.—Adult (Pl. XVIII. fig. 1). The first three segments

large and subequal, the Ist pleon segment equalling the first

pereeonal segment in length (measured along the medio-dorsal

line). Epimeral plates large and rounded; those of the 2nd

segment with a diagonal carina terminating in a denticle at the

postero-lateral angle, and those of the 3rd segment almost

rectangular; 4th segment with a deep depression dorsally ;

5th and 6th smaller, the 5th being slightly shorter than the 6th,

which is emarginate dorsally, for the insertion of the telson.

1908. | GENUS OF AMPHIPOD CRUSTACEA. 387

Immature. The proportions are the same as in the adult.

Young. First three pleon segments a little longer propor-

tionately than in adult, the plates rounded inferiorly, no trace of

lateral carina in second.

Superion Antenna (Pl. XVIII. figs. 2 & 3)—In the adult

female, the lst joint of the peduncle is twice as long as the other

two taken together, broader than long, posterior margin concave,

carrying a series of auditory setz distally; the 2nd joint also

has three or four of these sete distally ; the 3rd jomt is deeply

excavated anteriorly for the insertion of the accessory flagellum.

The primary flagellum consists of ten joints; the 1st equalling

the first joint of the peduncle in length, and so broad as to appear

a continuation of it. It carries, on its anterior surface, a great

number of broad, hyaline, sensory filaments, arranged in two

longitudinal rows, about 22 groups of the filaments in each row.

The 2nd joint is short, widening distally, with a fringe of short

stiff setee at the anterior distal angle; the 5rd joint is produced

at the anterior distal angle for the insertion of a long curved

spine which reaches nearly to the tip of the flagellum; the

Ath joint 1s cylindrical with a much shorter similar spine, and

with a transverse row of small sete in the middle of the anterior

margin ; these three joints have each a row of set inset midway

along the posterior margin. The following four joints decrease

gradually, each carrying at the distal anterior angle a fringe of

small sete. The apical joint is very small, with one long stiff

bristle and two auditory sete.

The accessory flagellum consists of one long laminar joint

and three small jomts. The Ist equals in actual length the

first joint of the primary flagellum; it carries three groups of

long sete on its upper margin, with a group of six sete at the

distal angle, two long, two short, and two “auditory.” The

2nd joimt is very small, cylindrical, rounded, with two long

setee and one auditory distally ; the 3rd is longer and more

slender than the second, slightly constricted in the middle, with

one long seta and one auditory; the apical joint is minute,

furnished with a long stiff bristle and three sete.

As stated before, the number of joints in the flagella of the

antenne appears to vary with the age of the specimen. In the

primary flagellum, the adult has 10, the immature 7, and the

young 4; in the accessory flagellum, the young form has only one

small joint, the immature specimen has two, and Boeck, Bovallius,

and Sars record the number as two for the adult, but in the

specimen here described there were three (fig. 3). The inferior

antenne also show this variation, the adult having 25, the others

10 and 4 respectively.

In the immature specimen (Pl. XVIII. fig. 4) the 1st joint

of the peduncle is not so long in proportion as in the adult; the

2nd and 3rd are subequal, taken together not so long as the

first. The primary flagellum has 7 joints; the first joint large

and stout, furnished with two rows of the sensory filaments,

388 MRS. E. W. SEXTON ON A (Apr. 28,

about 10 groups in each row; the remaining six joints taken

together are twice the length of the first. The accessory flagellum

is tri-articulate.

In the young form (Pl. XVIII. fig. 5) the 2nd and 3rd joints

of the peduncle taken together are equal in length to the Ist.

The primary flagellum is very little longer than “the peduncle,

and is composed ‘of four joints, the Ist the longest ; the 2nd and

3rd are each armed with a stiff spine, similar to those of the

adult; the apical jot is tipped with a long bristle, four simple

sete, and one auditory. The accessory fla gellam i is biarticulate,

and much longer than in the adult, equalling the first and

second jomts of the primary flagellum ‘taken together. There is

an auditory seta on each joint.

INFERIOR ANTENNA.—ddult (P]. XVIII. fig. 6). About twice

the length of superior antenna. The Ist joint of the pedwnele is

produced posteriorly in a thick lobe, a little hollowed behind ;

the 2nd joint is small, the antennal cone large in proportion ;

the 3rd is as wide as long, broadening distally ; the 4th exceeds

the 5th in length and is much broader. The first three carry no

sete ; the 4th has twelve long auditory sete along the posterior

margin, nine of which are inset on the inner surface and three on

the outer; the anterior margin carries seven groups of long fine

setee proximally, and seven clusters of small sete arranged in

transverse rows across the margin ; six of these little groups are

continued down the 5th joint. At its distal posterior angle there

is one long auditory and one simple seta.

The flagellum, which is about a third longer than the peduncle,

has 25 joints, the first the largest, decreasing gradually to the

tip which carries one long stiff bristle, two fine sete, and one

small one; each jot has a row of small sete anteriorly, inset a

little above the distal angle.

In the immature form the proportions of the peduncle joints

are the same. The 4th joint carries six of the auditory sete ;

the 5th has four sete at intervals on the anterior margin and

one auditory one distally, and one auditory and one fine seta

posteriorly as in adult. The flagellum has only 10 joints, the

first the largest; all the joints are provided with the little rows

of sete as in adult.

In the young (Pl. XVIII. fig. 7) the flagellum is shorter than

peduncle and consists of only four joimts, the first of which much

exceeds the other three taken together in length, each furnished

distally with a cluster of rather long seten, the apical joimt

carrying one long stiff bristle and four small sete.

OraL Parts (Pl. XVIII. fig. 8)—Upper and Lower Lips

(figs. 9, 10, & 11). The upper lip is elongate, apex emarginate ;

the lower lip i is of a more delicate structure, bifid, with the lobes

lanceolate; both lips so hollowed as to appear very narrow, but

when flattened out, of considerable breadth.

ManoprzBtss (Pl. XVIII. fig. 12).—Strong basal portion carry-

ing the large palp, produced forward as a long narrow process ;

1908. | GENUS OF AMPHIPOD CRUSTACEA. 389

cutting-edge obliquely truncate; no molar. The Ist joint of the

palp is very short, twice as broad as long; 2nd joint long and

broad, with a series of long stiff bristles commencing at the

posterior distal angle and crossing the joimt to the anterior

margin, where it is continued downwards in a double line for

half the length of the margin. These bristles, numbering 50,

are closely crowded, and are inserted inside the margin on the

outer face; on the under side there are 5 large strong ones inset

at intervals on the margin. The 35rd joint equals the second in

length. This jomt is covered with a dense mass of fine trans-

parent spines giving it a furred appearance. It is furnished

with 15 long, curved, plumose bristles along the anterior margin,

inset on the under side, the distal three forming a distinct group,

shorter, more curved, and at wider intervals than the others.

At the apex are two bristles set at a different angle from the

others and from each other, the upper one the shorter of the two,

both with dentate shafts and plumose tips.

In the young (Pl. XVIII. fig. 13) the 1st joint of the palp is

longer in proportion, the second joimt being only three times its

length, instead of five times as in the adult. The 2nd joint is

shorter than the third, with no bristles developed; the 3rd has

the two apical bristles, the distal one of the small group, and the

distal one of the long series.

First Maxinua.—ddult (Pl. XVIII. figs. 14 & 15; cf. also

Pl. XIX. fig. 1). With outer plate elongate, contracted below the

apex, and divided at the tip into five large claw-like teeth, four

of them much curved ; the fifth is not at the same level but set a

little farther down on the inner side, with a small spine at its

base on the inner margin, and a tuft of fine sete just below. The

palp is small, bi-articulate, set broadside on, the first joint very

minute, the second twice as long and tipped with two setules,

one longer than the other. The inner plate is very small, not

a quarter the length of the outer portion, unarmed, of delicate

structure.

In the young (Pl. XIX. figs. 2 & 3) the inner plate and the

palp ave much longer in proportion. The owter plate is rounded

at the apex, the five teeth not being separated as in adult. The

palp is bi-articulate, the two joints subequal, the apex tipped with

two setules.

SeconD Maxinia.—dAdult (Pl. XIX. fig. 4). The two plates

almost equal, the znner slightly the shorter, both hollowed under-

neath, giving the appearance of much less than their actual

breadth. The owter plate carries two minute setules at the tip,

one inset at the outer surface and one on the inner, and a few

hyaline, sensory sete on the inner margin. The inner plate has

the apex obliquely truncate, beset with three setules, the distal

one the shortest. This plate also has a few of the hyaline setee.

In the young (fig. 5) the proportions are the same, the plates

being rounded at their apices.

Maxiuuipeps.—dAdulé (Pl. XIX. fig. 6). The maxillipeds are

390 MRS. E. W. SEXTON ON A [Apr. 28,

deeply curved and hollowed, forming the lower portion of the

oral tube; the second joint of the one maxilliped is fused with

that of the other for more than two-thirds the length. Jnner

plates narrow, linguiform, tipped with two spinules ; ; outer plates

large, hollowed, so close together as to conceal the inner plates,

armed on the inner edge w ih ten uncinate spinules.

The palp is geniculate at the middle. The Ist joint is the

shortest, broad distally ; the 2nd and 3rd successively longer ; the

4th considerably the longest of all, being twice the length of

the first, lanceolate, one setule apically, finely serrated on both

margins, the serrations being stronger on the inner edge.

Sete.—The Ist joint has one small seta at the inner angle ;

the 2nd carries three long sete on its outer angle; the 3rd has

one proximally inside the margin on the outer surface, two

clusters on the margin, and a femmes of six around the outer

angle, with three on the inner angle.

In the young (fig. 7) the 3rd joint i is the longest; the 2nd and

Ath subequal to each other in length.

The Ist joint has one small seta; the 3rd carries one at each

distal angle, while the 4th has the apical setule and the margins

not serrated.

First GnarHorop.— Adult (Pl. XIX. fig. 8). The 2nd joint

is very long, slightly curved, widening a little at the distal end ;

the 3rd and Sth are subequal in length; the 4th is the

smallest, greatly curved, the outer margin of the joint being

five times as long as the inner. The 6th joimt or “ hand” is bent

over to such an extent that the articulation of the finger,

normally situated at the infero-anterior angle, appears to originate

from the infero-posterior corner instead, and what would

normally be the under surface thus becomes the upper. The

curve backwards is so extreme that this articulation almost

touches the third joint, and the fourth and fifth are nearly hidden

on the outer side. The hand is enormously developed, rounded-

triangular, greatly inflated, more so on the upper side. The

palm margin is straight with 15 strong recurved teeth, gradually

decreasing in size posteriorly. At the extreme anterior angle is

a small groove into which the tip of the “ finger” fits. On the

upper edge of this groove are two strong spines; on the under

edge three spines, with large tubercles at the bases of two of

them. One of these spines is minute, one broad and strong,

similar to the upper ones ; and the third, strong, tapering, mobile,

is situated at the anterior end of the row of teeth, reaching, when

close against the palm, to the base of the sixth tooth. The

“finger” or claw is large and curved, not extending beyond the

palm margin. There ave no sete on any of the joints.

Immature (Pl. XIX. fig. 9).--The second joint is much more

curved than in adult. The hand sets out more, at right angles

to the other joints. The small teeth number 15, the posterior

five being barely visible, and there are two very large mobile spines

at the anterior angle and two smaller just appearing.

1908. | GENUS OF AMPHIPOD CRUSTACEA. 391

Young (Pl. XIX. fig. 10).—The proportions of the first five

joints are the same as in adult, but the hand differs somewhat.

It is held in the normal position, not inverted, though showing a

tendency to curvature. It is more ovoid in shape, produced

posteriorly beyond the palm limit, which is defined by a spine

opposed to the tip of the claw. Close to this spine. one small

tooth can be seen, the distal one of the small series. None of the

joints is furnished with sete except only the claw, which has

two minute ones at the tip.

The hand is much larger in the young in proportion to the

size of the animal than in the adult; measured across the widest

part, in the young it is 1:5 mm. to 7 mm. of total length ; in the

immature 2 mm. to 9mm. total length ; and in theadult 3°75 mm.

to 26 mm.

Suconp Gnaruopop. Adult (Pl. XIX. fig. 11; Pl. XX. figs. 1,

2,3,4,&5). The 2nd joint is long, broadest distally ; the 3rd nearly

as long as the second ; the 4th short, narrow proximally, rounded

posteriorly with the posterior margin twice as long as the

anterior; the 5th is nearly as long as the third; the 6th is

broadly ovate, only half the length of the fifth, very narrow at

the base, but expanding laminarly on both sides ; the 7th or claw

is minute, nearly hidden under the long sete, and situated in the

middle of the distal margin of the sixth.

Sete.—There are no sete on the 2nd jomt; one small one

distally on the 3rd; a little fringe of fine sete on the posterior

margin of the 4th, with three small spinesacross the distal angle ;

the 5th is furnisked with two dense masses of very delicate,

hyaline hairs (fig. 5), one mass covering the anterior surface and

the other the posterior. At the anterior angle is a fringe of

12 strong, flexible sete, reaching to the distal margin of the

succeeding joint. Each seta consists of a stout shaft, finely and

transversely serrated for half its length, with a slender flagellum

(see fig. 4). Another cluster of these setee, shorter in length,

appears at the posterior distal angle and extends partway under

the joint. These two kinds of sensory sete, the “ hyaline” and

the “jointed,” are peculiar to the 5th and 6th joints of this

gnathopod. The 6th joint is nearly covered with the hyaline hairs,

with two thick tufts on the distal angles; the posterior angle

has a small group of the jointed sete; while on the anterior

angle is a cluster of about 50, the longest being twice the length

of the joint. On the distal margin are two smal] clusters of the

stiff, slightly curved, serrated bristles peculiar to this joint

(fig. 3). They are placed one on either side of the claw, the

group on the upper surface of the joint containing three bristles

and that of the under side four; between them is the denticu-

lated groove into which the claw fits. Bordering this groove are

6 stout sensory spines, three on each side, with their thick shafts

embedded for two thirds their length, and their large flagella

reaching beyond the apices (fig. 1). The curved under surface

of the claw is also covered with minute denticles. It is provided

392 MRS. E. W. SEXTON ON A [ Apr. 28,

with a distinct nail and bears on its distal margin one small

bristle, similar to those of the lateral clusters.

The large branchial vesicle reaches to the distal margin of the

3rd joint.

The tncubatory lamella is lanceolate in shape, and extends to

the distal margin of the 2nd joint. It is bordered with fine,

delicate sete of great length, the tips of which are slightly

expanded (fig. 2).

In the young (Pl. XX. figs. 6 & 7) the 3rd joint is only

half the length of the 2nd; the 4th is small, half as long as the

5th, which equals the 3rd. The 6th is much larger than in the

adult, rounded, with the two masses of hyaline hairs, and four of

the long, jointed sete on the anterior angle. The claw also is

much larger, with a distinct nail, three strong sete at the tip, and

one on the distal margin.

Sete.—The 2nd, 3rd, and 4th joints each carry one small seta

near the posterior angle; the 5th has two of the long jointed

setee posteriorly, one small seta anteriorly, no hyaline hairs

developed.

Prermopopa.—The lst pereopod is the longest, the 3rd the

shortest ; the three posterior pairs successively increase in length.

Branchial vesicles are attached to all; incubatory lamelle to the

first three.

Frrst Persxorpop.—dAdult (Pl. XX. fig. 8). The 2nd joint is

very long, broad; 3rd short; the 4th long, with a laminar ex-

pansion anteriorly ; the 5th ovate, shorter than the 4th; the

6th narrow, subequal to the 4th; the 7th half the length of the

6th, with a distinct falciform nail, the posterior margin with a

slight excavation proximally.

Setw.—The 4th joint carries five spines anteriorly ; these spines

are similar in construction to those of 7’. niewense, but with the

flagellum much shorter and nearer the apex. There is one spme

at the posterior angle, and, indented in the posterior margin, are

two small sensory setules. The 5th joint carries three of these

sete anteriorly, two posteriorly, and six spines around the poste-

rior angle, one considerably larger than the others. The 6th and

7th have their posterior margins thickly dentate, the anterior

margins serrate, the sixth joint with three spinules inset in the

posterior fringe, two in the anterior, and three at the anterior

angle.

The branchial vesicle at its widest is only half the width of the

one attached to the second gnathopod.

The immature has the 4th jot expanded.

In the young (Pl. XX. fig. 9) the proportions of the joints

are very different, the 4th, 5th, 6th, and 7th being practically

subequal. The 4th joint is not expanded.

Sete.—The 2nd, 3rd, 4th, and 5th are each provided with a

small seta at the posterior angle, the 6th with one at the anterior

angle.

Srconp Permorpop.—Aduwlt (Pl. XX. figs. 10 & 11). 2nd joint

1908. ] GENUS OF AMPHIPOD CRUSTACEA. 393

broad, and as long as the two following taken together ; 4th long

and very broad, laminarly expanded on both sides; 5th half the

length of the 4th, ovate; 6th narrow, much shorter than the 4th ;

7th half the length of the 6th.

Sete.—The 31d jot has one small seta on the posterior angle.

The 4th has six spines anteriorly inset along the chitinous margin

on the upper surface; and, on the posterior margin, 14 or more

microscopic sensory setules set in little indentations. The 5th

carries one spine anteriorly, and four (2 large and 2 small) on the

posterior angle; the posterior margin has four of the minute

setules, and is thickly dentate along its distal half. The 6th and

7th as in the first pereeopod, the 7th having several minnte spines

on its upper surface.

In the immature the 4th joint is expanded, about twice the

width of the succeeding joint.

In the young (Pl. X XI. fig. 1) the 4th joint is hardly longer

than the 5th, and not expanded; the 6th is the second longest

joint of the perzeopod.

Setz as in the first peraeopod.

Toirp Prermorop.—Adult (Pl. X XI. fig. 2). Second jomt as

long as the three following taken together, laminarly expanded,

anterior margin rounded, posterior straight ; the posterior part

is produced downwards with its lower margin truncate. 3rd joint

very small; 4th long, a little dilated posteriorly ; 5th and 6th

shorter than the 4th and much narrower; 7th about two thirds

the length of the 6th.

Sete.—The 2nd, 3rd, and 4th joints have each, at the anterior

angle, one strong spine; the 4th also bears at intervals along the

anterior margin six small spines, and four larger ones posteriorly.

The 5th has one large one midway on the posterior margin, and

two at the anterior angle; the whole of the anterior margin as

well as that of the 6th and 7th joints is strongly dentate, the 5th

and 6th each having two small spines inset.

In the young (Pl. X XI. fig. 3), the 2nd joint is not so long

nor so expanded as in the adult; the 4th and 5th are subequal ;

the 6th and 7th are subequal and a little longer than the

preceding.

Sete.—Vhe 2nd, 3rd, 4th, and 5th joints each carry one seta

at the anterior angle ; the 6th has two at the posterior angle.

FourtH Per#opop.—dAdult (Pl. X XI. fig. 4). The 2nd joint is

long, laminarly expanded, with both margins rounded ; the poste-

rior margin is produced downwards, lobate. The 4th is a little

dilated posteriorly, a little longer than the 5th and 6th which are

subequal; the 7th is about two thirds the length of the 6th; the

nail small and straight.

Sete.—The 2nd joint is furnished at its anterior angle with

one large spine and seven long fine sete extending beyond the

succeeding joint. These sete have each a slender shaft, with a

fine hair-like tip. Indented on the posterior margin are five

very minute sensory setules. The 3rd jomt has two spines; the

394 MRS. E. W. SEXTON ON A [Apr. 28,

Ath, five large ones posteriorly, five small on the anterior margin,

and three large on the anterior angle; the 5th carries one on the

posterior margin, two on the anterior, and two at the distal angle ;

and the 6th has two anteriorly. The distal half of the anterior

margin of the 5th and the whole of the anterior margins of the

6th and 7th are dentate. The 7th joint is serrate posteriorly.

In the young (Pl. XXI. fig. 5) the joints are slightly longer

than in the preceding pereopod, but the proportions are the

same.

Firru Perorop.—dAdulé (Pl. XXI. fig. 6). The 2nd joint is

longer and narrower than that of the preceding pereopod, con-

cave anteriorly, rounded posteriorly and produced downwards in

a deep lobe; the 4th, 5th, and 6th successively shorter and nar-

rower, the 4th slightly dilated posteriorly; the 7th small and

straight, much shorter than in the other pereopods, not quite

half the length of the 6th.

Setce.—The 2nd joint has five minute sensory setules posteriorly

so deeply inset as to give a crenulated appearance to the margin.

The 2nd and 3rd joints at their anterior angles each have five

large spines. The 4th has nine large strong spines set in deep

indentations of the posterior margin, and seven smaller along the

anterior margin, with four at the anterior angle—one large and

three small. The 5th carries one small one on the margin, and one

at the angle posteriorly ; its anterior margin is strongly dentate

with five spines inset along the proximal half, the two distal ones

being placed together in the same indentation; at the anterior

angle are four spines, one large and three small. The 6th has the

anterior spinose fringe with two small spines inset ; the anterior

angle carries three,and the posterior two. The 7th joint is also

dentate anteriorly ; the nail minute, not faleate. The branchial

vesicle attached to this pereeopod is very small.

In the young (Pl. X XI. fig. 7) the 2nd joint is expanded; the

4th, 5th, and 6th are successively longer, not shorter as in the

adult ; the 7th is curved, eaualling the 6th in length.

Sete.— The 2nd joint has one setule on the posterior margin; the

anterior angles of the 2nd, 3rd, and 4th have each one, that of

the 5th carries three, while the 6th has one at each angle.

PiEopopa.— Adult (Pl. X XI. fig. 8). The pleopoda are large and

powerful. The peduncle is stout, with two little coupling-spines

on the inner side, of the same construction as those figured for

T. niceense; the rami twice the length of the peduncle, and

subequal. The outer ramus of the 1st pleopod consists of 21 joints,

the first much the largest, carrying six plumose setee on each side

in addition to the distal two; the remaining 20 joints successively

decrease in size, each furnished with two long plumose sete. The

Ist joint of the inner ramus bears on its inner side a tuft of fine

hyaline hairs and eight cleft spines; the seta at the mner angle

of each of the 13 following joints is uncinate. The inner ramus

of the 3rd pleopod has seven cleft spines and fifteen uncinate

sete.

1908. | GENUS OF AMPHTPOD CRUSTACEA, 395

In the young the peduncle is much longer in proyortion, with

one seta on the outer side and two coupling-spines on the inner.

The rami are very little longer than the peduncle; the outer

ramus is composed of 4 joints, the Ist nearly three times as long as

the other three together, each joint provided with two long plumose

sete; the ener ramus has 3 joints, the Ist four times the length

of the other two together, two plumose sete to each joint, and one

large cleft spine on the first.

Uroropa.—A dult (Pl. XXI. figs. 9, 10, 11,13; <¢f also fig. 14),

The apices of the Ist uropods reach considerably beyond the

peduncles of the 3rd; those of the 2nd and 3rd pairs are on a

level,

The peduncle of the lsé wropod is broad and long, equal to the

outer ramus in length, with one small seta on its outer margin,

and ten large, strong spines on the inner edge. The rami are

broadly lanceolate, curved, the outer being the shorter of the two.

The inner ramus has both margins serrated, with three spines on

the inner one ; the outer ramus bears six minute sensory setules

(fig. 11) deeply indented in its upper margin, the distal half of

which is serrated, as is also the under margin.

The rami of the 2nd wropod are subequal, longer than the

peduncle and slightly curved. he peduncle bears one small

spine on the inner angle. The upper margin of the outer ramus

is beset with five minute setules, no serrations, all the other

margins finely serrated.

The peduncle of the 3rd wropod is short, wider than long, the

rami twice its length, the outer one bi-articulate. The outer

margins of both rami are plain, the inner edges serrated; the

small 2nd jomt of the outer ramus having 12 or 13 fine

serrations.

For the young, see Pl. X XI. figs. 12, 15,16. The outer margin

of the outer ramus of the Ist and 2ud uropods is deeply notched

near the apex. In this notch, set as it were in a little pocket, is

a large sensory seta, similar io the sensory setule of the adult, but

on a much larger scale (fig. 12).

Trtson.— Adult (Pl. XVIT. fig. 13, Pl. X XI. fig. 17). Rounded

anteriorly, narrowing posteriorly, longer than the peduncle of the

3rd uropod, and almost as broad as long. The apex is obtusely

truncated, distinctly emarginate, tipped with two setules. It

has four transverse rows of spinules dorsally, as described for

T. micwense, and six of the tufted setules. (For arrangement and

detail, see Pl. XVII. fig. 13; cf. fig. 11.)

In the immature (Pl. XXI. fig. 18) and young forms (fig. 15)

the telson is incised for about one fifth of its length, with

the apices rounded.

IIT. Generat Remarks.

The Norwegian form of this animal when first discovered was

considered of sufficient importance to rank as one of the principal

396 MRS. E. W. SEXTON ON A [ Apr. 28,

divisions of the Amphipoda, the tribe Prostomate being established

by Boeck for its reception.

Boeck noted many points of resemblance to the Lysianassidee

the structure of the upper antenne, the powerful hand of the first

gnathopod, and the slender characteristic second gnathopod ; but

it remained for Sars to prove conclusively its relationship to that

family, in which it now rests.

Boeck considered, and Bovallius and Sars are in agreement

with his views, that the peculiar tubiform structure of the mouth

indicates a parasitic habit. Most of the specimens have been

taken on fish. The first were caught by Prof. Rasch by lowering

in the trawl a freshly-killed and skinned bird, to the body of which

they clung, sucking the blood. Herr Storm found it in the

Throndhjemsfjord “ parasite on a shark” (Boeck), as well as on

the common black dogfish (Sars). Bovallius captured several

specimens in the Hardangersfjord, “ parasites on or in company

with an Asterias.” Prof. Sars has taken it in the Throndhjems-

fjord, in all eases clinging to the skin of the black dogfish (Spinaa

niger). Mr. Tattersall’s specimen, the only one yet recorded out

of Norway, was taken in a small tow-net attached to the trawl.

The Mediterranean form also has always been recorded as taken

on fish. The first specimens were found, as Costa states, on

“squalo” and “ merluzzi” ; the three examined by Della Valle

were taken on “ merluzzi,” and the one recorded by Brian was

discovered in the mouth of a “ Chlorophthalmus agassizi.” On

the other hand, Chevreux’s specimen from the Bay of Biscay, the

‘Huxley’ specimen from the north of the Bay of Biscay, and

Mr. Tattersall’s six specimens from the West of Ireland, were all

free-swimming. Myr. Stebbing’s 8.-African species also appears

to have been captured free-swimming.

It would seem better, all things considered, to describe this

genus as predatory rather than parasitic. All the Lysianasside

are carnivorous : 1n many of the genera they move in vast hordes

devouring any carrion they find, and not hesitating to attack

anything living unable to escape them, such as fish caught in a

net. Many instances of this could be cited ; to take an example

from my own observation—in the Nassa-pots placed in Plymouth

Sound by the Marine Laboratory 1 have found the common little

Orchomenella nanus swarming in thousands, not only eating the

dead crabs used for bait, but devouring the living polyps of the

Perigonimus on the Nassa shells, and attacking the Nassa them-

selves whenever extruded. And again, in a dredging taken off

the Eddystone, over 18,000 Scopelocheirus hopei were found in two

Echinus-shells. A statement of Prof. Della Valle’s supports this

view in regard to 7’. nicwense; in the ‘Fauna’ (15), p. 287, he

says, referring to his three specimens :—‘* Uno di questi individui

portava ancora imprigionato nel formidabile organo di presa dei

suoi gnatopodi posteriori un pezzo di carne del pesce su cui era

attaccato.” (Cf. also Chevreux, 21.)

Two features render this genus noteworthy

the inflation of one

1908. | GENUS OF AMPHIPOD CRUSTACEA. 397

or more joints of the pereeopods and the peculiar torsion of the

first gnathopod. This torsion is caused by the curvature of the

fourth and fifth joints, by which the normal position of the hand

and finger is reversed. Even in the young 7’. raschii taken from

the incubatory pouch a slight curvature can be seen, and the

immature specimen, 9 mm. in length, shows it already completed.

All the adult specimens of 7. raschii yet discovered are females,

and all have this torsion. In 7’. nicwense some specimens show

it and others not. Costa’s specimens are figured with the hand

normal; Della Valle records his three specimens, all males, with

the torsion; Brian’s specimen, ovigerous female, normal ; Chev-

reux’s, ovigerous female, with the hand twisted; while the

‘ Huxley’ specimen is an ovigerous female, and normal. In the

six taken by Mr. Tattersall, three males and three females, the

way the hand is held can be plainly seen. The second joint

of the first gnathopod is long and much curved; the third or

elbow makes an acute bend forward; the fourth bends back

towards the second; and the fifth turns a little forward and

outwards in such a way as to hold the sixth joint or “ hand ” with

its inner side uppermost, so that the claw and curved spines

are on top, and the articulation of the claw appears to be in

the infero-posterior angle instead of the infero-anterior angle as

in the normal position. The “ hand ” is enormously inflated, and

subquadrate. It is carried usually with the claw articulation

outwards and a little raised, the tips of the claws meeting under

the animal. All the joints are strengthened with ridges of chitin,

and all their margins are remarkably strong and thick. The

animal is evidently able to turn the hands in any direction at

will; this ean be plainly seen in Mr. Tattersall’s specimens. In

one or two both hands are normal; im one specimen one hand is

normal and the other twisted, in the others the hands are in the

position just described—these differences being evidently due to

the animal’s contortions when being preserved.

Another peculiarity of this genus is the inflation of one or more

joints of the pereeopoda: in 7. raschzi the fourth joint of the first

and second pereopods ; in 7’. nicwense the second joint of the first

and second pereopods; and in Mr. Stebbing’s new species the

sixth joint of the fifth perzeopod.

The sensory equipment of these animals is remarkable, there

being no less than twenty-five specialised forms of sete, four of

which are peculiar to the antenne, five to the oral parts, four to

the second gnathopoda, one to the incubatory lamell, four to the

pleopoda, and one to the telson. As the number and arrange-

ment of the sete appear constant, I have thought it well to give

them in detail.

The chief differences in the three species hitherto known lie in

the shape of the 2nd, 3rd, and 4th side-plates, the antenne, the

2nd and 4th joints of the anterior pereopods, the 2nd joints of

the hinder pereeopods, and in the telson. Intheadult 7. nicwense

the telson is entire; in 7’. raschii deeply incised in the young and

398 MRS. E. W. SEXTON ON A [Apr. 28,

immature, and lightly excavated, almost entire, in the adult; in

Mr. Stebbing’s species, ‘ perhaps not fully adult,” deeply incised.

'The incision appears to be a character of the young, but this point

cannot be settled till the young ot 7’. nicwense and the adult of

the last species are known.

This genus is a deep-water form, not being recorded from less

than 60 fathoms in northern seas, 30-40 fathoms in southern

waters.

TV. D&Frrnrrion oF GENUS.

Family Lysianasside.

Genus TriscHizostoMA Boeck & Esmark, 1860.

1853. Guerinia (preoccupied, 1830) A. Costa (1).

1860. Trischizostoma Boeck (8).

1865. 4 Lilljeborg (5).

1870. ns Boeck (7).

1872 # Boeck (8).

1886. p Bovallius (11).

1888. ie Stebbing (12).

1890. ; G. O. Sars (18).

1893. Trischizostoma & Guerina Della Valle (15).

1895. Trischizostoma Stebbing (6).

1905. Guerinella Chevreux (18).

1906. Trischizostoma Stebbing (19).

Body thick, slightly compressed, rounded dorsally.

Head longer than first segment of person, with a short,

anteriorly-rounded rostrum, produced over the bases of the

superior antenne.

Eyes very large, contiguous dorsally, occupying the whole sur-

face of the head, except the rostrum and anfero-lateral margin.

Superior Gniene pedunele very stout, with first joint larger

than the other two ; first jot of primary flagellum so large as to

appear a continuation of the peduncle, thickly fringed with sensory

filaments; accessory flagellum 3- or 4- articulate, with the first

joint long and laminar.

Inferror anteana much longer than superior antenna ; flagellum

in male longer than in female.

Oral parts greatly projecting inferiorly.

Mandibles with large body carrying very large 3-articulate,

densely setose palp ; cutting-edge truncate, unarmed ; no molar.

Mawxilla 1. outer plate elongate, shghtly contracted just below

the apex, which is divided into 5 claw-like teeth; inner plate

small, unarmed ; palp small, bi-articulate.

Mawillipeds large, outer plates partly encompassing the oral

parts; inner plates narrow; palp 4-articulate.

Perceon: last two segments smaller than preceding ; infero-

posterior angles of segments produced backwards and rounded.

Side -_plates J ilist small, nearly triangular, almost completely

1908. | GENUS OF AMPHIPOD CRUSTACEA, 399

covered by the large second pair, which are dilated inferiorly ;

5th and 6th inferiorly bilobed ; 7th small, subquadrate.

Branchial lamelle very large, much pleated.

ls¢ gnathopod enormously developed, subchelate, prehensile.

Pleon: first 3 segments very large, rounded; the 4th with a

deep depression dorsally ; the 5th the shortest.

Uropoda with broadly lanceolate, serrate rami; outer ramus of

3rd uropod bi-articulate.

Telson small, broad, entire or apically incised.

LITERATURE.

(1) 1853. Cosra, AcuiLLE.—“ Descrizione di tre nuovi Crostacei

del Mediterraneo, discoperti dal Rev. G. F. Hope.”

Kstratto dal fascicolo 83° della Fauna del Regno di

Napoli, pp. 3-6, pl. i.

(2) 1853. Cosra, AcuitLE.—Fauna del Regno di Napoli.

(3) 1860. Bock, AxEn.—‘ Bemaerkninger angaaende de ved

de Norske Kyster forekommende Amphipoder.” For-

handlinger ved de Skandinaviske Naturforskeres :

Ottende Mgde i Kjobenhavn, 8-14 de Juli, 1860,

pp. 631-677. Translated by W.S. Dallas, F.L.S., in

‘Annals and Magazine of Natural History,’ ser. 4,

no. 17, May 1869.

(4) 1862. Spence Barn, CHARLES.—Catalogue of the Specimens

of Amphipodous Crustacea in the Collection of the

British Museum. London, 1862, pp. 110-111,

pl. xiv a. figs. 5 &e.

(5) 1865. Litusepore, Wiii1am.— Bidrag till Kinnedomen

om underfamiljen Lysianassina inom underordningen

Amphipoda bland Kraftdjuren,’ p. 9, in Upsala Uni-

versitets Arsskrift, 1865, Mathematik och Natur-

vetenskap, 1.—‘“On the Lysianassa magellanica

H. Milne-Edwards, and on the Crustacea of the

Suborder Amphipoda and Subfamily Lysianassina:

found on the coast of Sweden and Norway,” p. 18,

Nova Acta Societ. Scient. Upsaliensis, ser. 3, vol. vi.

(6) 1867. Cosra, AcHinLE.—Annuario del Museo Zoologico della

R. Universita di Napoli. Anno iv., 1864. Napoli,

1867,

(7) 1870. Bozck, Axen. “Crustacea Amphipoda borealia et

arctica” in Christ. Vidensk. Selsk. Forhandlinger,

ISO; joe We):

(8) 1872. Bonck, AxeL.— De Skandinaviske og Arktiske Am-

phipoder.” Forste Hefte: med 7 kobberstukne

Tavler. Christiania, 1872, pp. 95-97, pl. i. fig. 1,

pl. i. fig. 2.

(9) 1882. Sars, GEorce Osstan.— Oversigt af Norges Crus-

taceer,” in Christ. Vidensk. Selsk. Forhandlinger,

1882, No. 18, p. 20.

Proc. Zoou. Soc.—1908, No. XX VI, 26

400

(10) 1885.

(11) 1886.

(12) 1888.

(13) 1890.

(14) 1892.

(15) 1893.

(16) 1895.

(17) 1903.

(18) 1905.

(19) 1906.

(20) 1907.

(21) 1895.

. The whole animal magnified,

showing actual leng th

MRS. E. W. SEXTON ON A [Apr. 28,

Carus, JuLtius Vicror.—Prodromus Faune Mediter-

ranee, Vol. I. pars .: Arthropoda. Stuttgart, 1885,

p. 411.

Boyatuius, Cart.— Amphipoda Synopidea, with 3

plates (presented to the Royal Society of Sciences of

Upsala, 10th May, 1886), pp. 22-31, pl. 101.

Sreppine, Rev. T. R. R. Report on the Scientific

Results of the Voyage of H.M.S. ‘Challenger’

Amphipoda, 3 Parts, Vol. xxix., pp. xix, 272, 323, 576.

Sars, G. O.—Crustacea of Norway. Vol. I. Am-

phipoda. Christiania and Copenhagen, 1895, pp. 29-

32 & 673, pl. xil.

Bonnter, JuLES. — “ Les Amphipodes du Boulon-

nais (1).” Bull. Sci. de la France et de la Belgique,

tome xxiv. pp. 165, 173.

De.LiA VALLE, ANToNIO.—Fauna und Flora des Golfes

von Neapel: Gammarina, 2 vols., Monog. xx. pp. 770,

775-780, pl. 61. figs. 10-22.

Sreppine, T. R. R. ‘A Passage-at-Arms over the

Amphipoda.” Natural Science, vol. vi. No. 38,

pp. 265-266. (April 1895, London.)

Brian, ALEssANDRO. — Bollettino del Naturalista.

Siena. Hstratto dall’ anno xxiii. n. 4, 1903.

CuEvreux, Epovarp.—Bulletin du Musée Océano-

graphique de Monaco, No. 35, 5 Mai 1905.

STEBBING, T. R. R. Ios Tierreich, eine Zusammen-

stellung und Kennzeichnung der Tene ion Tierformen.

21 Lieferung. Amphipoda: it Gammaridea, pp. 12-14,

fig. 3. Berlin, 1906.

Sreppine, T. R. R. MSS. on the South-African

Lysianassidee. 24.vii.1907.

Cuevreux, E. Mém. Soc. Zool. France, vol. viii. p. 425.

EXPLANATION OF THE PLATES.

PLATE XIV.

with line above

T. niceense.

2. Cuticle, taken from underneath the rostru un, 5 2 Bi a X 265.

3. Superior antenna, 2 ‘ Huxley’ specimen (with, tip

of sensory filament X 265) . Sots ep x 17.

4, Superior antenna, ¢ Irish specimen . PBR ac theese ee S< to

5. Accessory flagellum, superior antenna, d ............ % x 42.

6. Inferior antenna, 2 ‘ Huxley’ specimen ............... 3 Xx 17.

7. » 6 Irish specimen .. 3 x<Lie

8. Calceolus from inferior antenna 3, the third from

the tip ..... 35 X 265

9. Two an ditoe ‘sete from ‘the 4th joint. ‘of the

peduncle, inferior antenna, ¢ Ivish specimen... % X 265.

10. Upper lip, ¢ Ivish specimen ......... Rene Ae Bs x 42.

11. Lower lip flattened out, ¢ Irish specimen Seeaeorseene a xX 42.

12. Right mandible, 2 ‘ Huxley’ SpeCimMeM vee eceeee eee eS x 42.

18. Cutting process, @ ‘ Huxley’ specimen ............... 5 XxX 75.

14. Left mandible, g Ivish specimen ..................... 00 0 x 42.

1908.}

(re)

ig.

CoN pe

OU om

for)

GENUS OF AMPHIPOD CRUSTACEA.

PLATE XV.

First maxilla, g Irish specimen (with 4th spine

X 265, to show the serration) .... :

First maxilla, ¢, magnified to show palp ‘and inner

plates eee Sana eis

. Second maxilla, ES Trish specimen Ss

. Maxilliped, ¢ Ivish specimen Be NGC REGRET ea ret

. Palp of maxilliped, inner surface, 2 ‘Huxley’

specimen

. First gnathopod, g Irish specimen .......0..0......

ae 55 OTE s Gn bsd C=y'aaaat) 02163000) |

inner surface, 2 ‘ Huxley’ spe-

cimen (with three of the small teeth

22 22

Prate XVI.

. Second gnathopod, 2 ‘ Huxley’ specimen ms

. Claw of second gnathopod, ¢ Irish specimen _......

Terminal joints of second gnathopod, 2 ‘ Huxley ’

SPECIMEN Use eon cnemeainen eos ces uote

Tip of “ hyaline hair,’ > 6th joint, second gnathopod,

OT Huxley ‘ specimen

. Tip of “jointed seta,” 6th joint, ‘second gnathopod,

‘Huxley’ specimen

. One of the “ dacty ‘loptera” “of the 4th pereopod,

7. First pereopod, 2 ‘ Huxley’ specimen. ache iite

8. Second ,, ONSHnaxl ey 2 specimen!” 5.520... .aee:

o> ae aes

“I

co oO

10.

iil,

12.

13.

SOMNANE whore

Prate XVII.

. Fourth pereopod, ‘Huxley’ specimen ............

Fifth 3 9 ‘Huxley’ specimen..

Fringe of spines | on anterior angle of 5th joint,

5th pereopod, ‘ Huxley’ specimen a

. Second pleopod, * Huxley’ specimen

Coupling spines, second pleopod, ‘Huxley? " spe-

cimen

> doo OE uncinate + seta, “second “pleopod, |“ Husley’

specimen

. Tip of 5th cleft spine, “second pleopod, ‘Huxley’

specimen

. Uropoda and telson, o Mediterranean specimen. sodede

. Serrations, inner margin of inner ramus, first

uropod, ubanveiguld eciaa telah wena Bk taraeaeenereirestte

Serrations, inner margin of outer ramus, first

uropod, @ .. BEE OsEee

Telson, 3 Trish specimen. : ees

Sensory setule from telson, Ir ish specimen. icin ae dees

Telson, 2 BUSH athe

Prats XVIII.

First three segments of pleon, adult 2

. Superior antenna, adult 2 .

Terminal joints of accessory ‘flagellum, “superior

PRAUNeT TOE. 7 il pga aeB ee Soe aG eae ae AS Gre Ane Res Bi ie

Superior antenna, immature specimen

* young specimen

: Inferior antenna (seen from the under side), adult 2.

oy AVOUUTE SIDES HIME, gnc dagooa ong done 085 008

Oral. parts in position, 9 end

. Upper lip, 2

Lower lip with one lobe flattened, oe Rhee NOR

T. nieeense.

T. niceense.

T. niceense:

» Xabou

T. raschit.

T. raschii.

XXX XXX

XxX

MX KXKXKXKK XX

401

42.

Fig.

oS wo bo be

Sem ND

Ie, dl

meee sve

Sco cst Oe PO

. First maxilla, young specimen

. Apex of the outer plates first maxilla, young

. Second maxilla, adult § Q

: Palp of maxilliped, right side, adult 2

. Maxillipeds, slightly flattened, seen from the under

ON A GENUS OF AMPHIPOD CRUSTACEA.

. Upper and lower lips, young ie

. Mandible, adult 2

young? specimen

: First, maxilla, adult Oe ass

. Apex of the outer plate, first mavilla, adult OF

PuatEe XIX.

Apex of the outer plate, first anes adult a

For comparison ( !

specimen

FOWAS SOACMINED poosasenscooonncesv0s see

side, young specimen

. First onathopod, showing the malo surface, 9

ay 5 of immature specimen ..............-

4 5 of a young specimen, taken from

the incubatory pouch

. Second gnathopod, adult 9

PLATE XX.

Claw of second gnathopod, adult 9

. Tip of seta from the ineubatory lamella, second

enathopod, 2

. One of the stiff setee, ¢ distal margin, 6th joint, second

gnathopod, Oe Saeed

Tip of “jointed seta,” “anterior. angle, “6th. joint,

second gnathopod, 2 A aia ci Si toacte nag ara neeepacnd eer

Tip of “hyaline hair,” anterior angle, 6th joint,

SQGOMG! GATT HO GCL, SY ooscsovanaan soe oan raccovconnssdasode

Second gnathopod, young specimen Bvetan

. Claw of the second gnathopod, young specimen

vesicle and the incubatory lamella i ae icaneeie sees

First pereeopod, young specimen

Second pereopod, adult Q .

. “Nail” of the 7th joint, 2nd peropod, adult. Q

Prate XXI.

. Second perzeopod, young specimen

. Third perxopod, adult 2

young specimen

b Fourth peropod, adult @

young specimen

. Fifth pereeopod, adult 2

young specimen oog06

: Tip of distal cleft spine, second pleopod, adult o.

. First uropod, adult 2 ....

. Proximal spine, mner ramus, first ‘uropod, ‘adult. .

. Sensory setule, outer ramus, first uropod, adult @ ..

” ” » » second uropod, young

specimen SEAM zai tS

5 For compar ison

; Telson and uropoda, young specimen ..................

. Second joint, third mEGu ed) young ay

. Telson, adult 2 x ST slags

>, Immature specimen -

T. raschii.

T. nic@enseé.

T. raschit.

[ Apr. 28,

xxx XX

DRwW eS

BLO OHO FO

xx XXX XXKXXX XX

Lo or

X 265.

x 435.

> %about500.

> *aboutd500.

by)

1p: niccwense.

T. raschii.~

«KK KX KK KKKKK

x 42.

X 265.

x I

x 42.

xe ie

X 265.

OSE

PAP AWA

hp repre ee be

loner in f

SS eon)

1908. ] ON GEORGE THE FOURTH’S GIRAFFE. 403

4, On certain Errors with reference to George the Fourth’s

Giraffe. By Henry Scuzrrey, F.Z.S.

[Received April 27, 1908. |

For more than seventy years the duration of life in captivity

of the Giraffe presented to George IV. by the Pacha of Egypt

has been considerably understated. The error undoubtedly

originated in one of the Society’s publications, and consequently

has been widely copied. The following appears to be the earliest

of the erroneous statements, and the source of all the rest :—

“In the year 1827 a female specimen in bad condition arrived at Windsor as

a present to H.M. George IV. from the Pacha of Egypt. This individual, after

lingering afew months, died, and its mounted skin and skeleton adorn the museum

of the Society 7 *.

This was repeated in the next List, the last of the series,

published in 1844, whence it was copied into D. W. Mitchell’s

Guide, which appeared in 1852, with this addition :—‘“ The

animal died partly from bad management and partly from

infirmity of constitution.” In the edition of 1858 the supposed

contributory causes are omitted, the statement being :—‘ It

lived, however, only a few months at Windsor.” This was

repeated in the first edition of Dr. Sclater’s Guide, published

in 1859, and onwards in each successive publication down to

the fifth edition of the Official Guide. It also occurs in a

paper by Mr. Lydekker, F.R.S.7, on ‘Old Pictures of Giraffes

and Zebras,” in which it is stated that “the animal survived

but a short time at Windsor,” Dr. Graham’s Renshaw being

quoted as the authority.

As a matter of fact, the animal lived at Windsor for two years

and two months; and the evidence as to the dates of its arrival

and death is here set out. At the end of 1826 the Giraffe was

sent from Cairo to Malta, where it was kept over the winter. In

May 1827 it was shipped in the ‘ Penelope’ for London, with two

Egyptian cows, in charge of two Arab keepers and an interpreter.

These were landed at the Duchy of Lancaster Wharf, Waterloo

Bridge, on the evening of Saturday, August 11th, 1827; and on

Monday morning Cross took the Giraffe to Windsor in one of

Richardson’s caravans. The King “hastened to inspect his

extraordinary acquisition, and was greatly pleased with the care

which had been taken to bring it to his presence in fine order” §.

The animal was referred to by the author of ‘The Menageries’ ||

(i. p. 343) as still living in June 1829; and in an article published

in 1833, under the editorial supervision of Owen—if, indeed, it

was not written by him,—it was said to ‘‘ have increased eighteen

* ‘List of Animals in the Gardens,’ p. 35 (1837).

+ P.Z.S. 1904, ii. p. 339.

t ‘Natural History Essays’ (p. 105, 1904).

§ ‘ Literary Gazette,’ Aug. 25, 1827, p. 554.

|| ‘ Library of Entertainmg Knowledge.’

404 ON GEORGE THE FOURTH’S GIRAFFE. [Apr. 28,

inches in less than two vears”*. In 1854 the true date was

again indicated :

“The giraffe sent to the French menagerie did well; that presented at the same

time to George IV. was the shortest and weakest. [The Consuls of each nation drew

lots for the choice.| She was never in good health, and had been roughly treated,

and though she had grown eighteen inches up to June 1829, she sank gradually and

died in the autumn of that year” fF.

By 1879 the correct date was given by Dr. Wilhelm Stricker :

“Hin zweites nach London bestimmtes Exemplar traf im August 1827 anderthalb

Jahre alt, daselbst ein, verendete aber schon im Oktober 1829 ebenfalls an Gelenk-

krankheit wahrscheinlich, weil es in Afrika auf weite Strecken geknebelt auf dem

Riicken von Kameelen transportiert worden war ” f.

Major Fortuné Nott stated that the Giraffe ‘“‘ died in 1829” § ;

and while collecting material for my book on the history of the

Society I was so fortunate as to meet with evidence that enabled

me to fix the exact date of the animal’s death.

A paragraph in the ‘ Times’ of Oct. 19, 1829, taken from the

‘Windsor Express’ of Oct. 17, puts the actual date beyond doubt.

It runs thus :—

“Messrs. Gould and Tomkins, of the Zoological Gardens, are now dissecting the

giraffe which expired on Sunday last [Oct. 11]. We understand that when the skin

is stuffed His Majesty intends making it a present to the Zoological Society.”

So that the Giraffe lived exactly two years and two months in

this country, having been landed in London on August 11, 1827.

At the meeting of Council held on August 28, 1830, Lord

Auckland, President of the Society, submitted “an offer by

Sir William Freemantle of the skin and skeleton of the girafte

that lately died at Windsor.” It was resolved that the Secretary

be requested to inform Sir W. Freemantle that they thankfully

accept the same, and will also bear the charges of preserving and

setting up the animal. In the second edition (1838) of the

‘Catalogue of Mammalia in the Society’s Museum,’ the animal is

entered as being “ presented by His Majesty William IV.”

Another error with regard to the animal is that it was

well known to the people of the metropolis—a popular favourite.

The writer of the article in the ‘ Zoological Magazine,’ previously

referred to, stated that, owing to the distance from town at

which this animal was kept and the state of confinement which

its weakly condition rendered indispensable during the latter part.

of its existence, the living Giraffe was seen in this country by

comparatively few individuals.

In the paper by Mr. Lydekker already cited, the author, in

describing a painting by R. B. Davis, then in the Royal Collection

at Windsor, having on the back the note “ Portrait of Giraffe

belonging to His Majesty” |, suggested that the artist took “a.

+ ‘English Cyclopedia’ (Nat. Hist.), ii. col. 1028.

t ‘Geschichte der Menagerien,’ S. 25.

§ ‘ Wild Animals Photographed and Described,’ p. 225.

|| P. ZS. 1904, ii. p. 342 (text-figure).

1908. | ON THE BREEDTNG-HABITS OF A CICHLID FISH. 405

portrait of the Cape Giraffe for his model, and he may have

copied Paterson’s specimen in the British Museum.” It should,

I think, be remembered that Richard Bennett Davis was an

artist of good reputation. He was afterwards animal-painter to

William IV., and the Court Officials of that day must have been

aware of his presence for the express purpose of painting the

Giraffe, and indeed of his whole history. He owed his education

as an artist to royal favour. His father had been huntsman to

the royal harriers; George III. took favourable notice of some of

the boy’s sketches, and placed him under Sir William Beechey.

In 1806 he sent to the Academy a painting, “‘ Mares and Foals

from the Royal Stud at Windsor.” His appointment as animal-

painter to the King was made in 1831, and he painted the

cavaleade which formed the coronation procession of that

monarch *,

There is, moreover, incontestable evidence that he did paint the

Giraffe, and practically all that we know about the habits of the

animal in captivity came from his pen. He contributed a very

interesting account of the Giraffe to the ‘ Literary Gazette’t; and

in the following passage occurs the first mention of the bilobed

teeth known to me :—

“The upper lip is longer than the lower one, which assists the tongue in drawing

in the boughs; but when grinding its food it is contracted. It has no teeth or

nippers in the upper jaw, and the outside ones are divided to the socket. It is

a ruminating animal, and lies down when it chews the cud.”

The fact that Davis painted a picture of the Giraffe was stated

by the editor of the ‘ Literary Gazette,’ who expressed “ our

obligation for this paper to Mr. R. B. Davis, who, while painting

the picture of the animal for His Majesty, had many oppor-

tunities of observing its peculiarities.” In ‘ Menageries’ (p. 348)

Davis's account of the bilobed teeth is quoted, but this important

part of his narrative is omitted from the ‘ Zoological Magazine’

and the ‘ English Cyclopeedia.’

5. On the Breeding-Habits of a Cichlid Fish ( Tilapia nilotica) .

By Caries L. Boutencer, B.A., F.Z.8.

[ Received March 23, 1908. |

During a short expedition in the spring of last year to

Lake Qurun, in the Faytm province of Egypt, I was able to

make some observations on the breeding-habits of Tilapia

nilotica, the ‘‘ Bolti sultani” of the native fishermen.

These interesting Cichlids abound in the brackish waters of

the lake and are extensively fished for by the natives, the Birket

el Qurun supplying most of the markets of Lower Egypt with

this excellent food-fish.

Tsolated observations have been recorded on the habits of the

* Dict. Nat. Biogr. xiv. p. 171. + Dec. 1, 1829.

406 ON THE BREEDING-HABITS OF A CICHLID FisH. | Apr. 28,

Bolti; but a complete account of the nesting-processes has never

been published.

On visiting the fish-markets on the shores of the lake, and

inspecting the baskets full of Boltis, the first thing to strike one

is the fact that some individuals are more brilliantly coloured than

others: these the natives declared to be males, the females being

less vividly coloured. I accordingly dissected and sexed a few

examples, and found this statement to be correct.

The males, which grow to a larger size than the females, have

the whole of the ventral surface of the head and beily, almost as

far back as the commencement of the anal fin, of a bright red

colour; the throat is of a particularly brilliant hue. The pectoral

and pelvic fins are bright red, and the top of the head also bears

red pigment—not, however, so vivid as on the ventral side, and

inclining rather to violet.

In the females, on the other hand, the red colour is absent from

the top of the head and from the pectoral fins; the throat and

belly are pink.

The brighter colours and the large size of the males render

these easily distmguishable from the females, even when viewed

in the water.

In common with many other Cichlid fishes, Zilapia nilotica

excavates holes for breeding-purposes ; these were first observed

by Loat in Lake Menzaleh. In Lake Qurun these nests or,

as the natives call them, ‘ houses” of the Bolti are to be found

close to the shore in two to three feet of water. I found them

to occur almost exclusively on the north or desert side of the

lake. The reason for this is, I think, quite evident to anyone

familiar with the Birket el Qurun; for on this side the water

becomes rapidly deeper offshore, enabling the fish when disturbed

to dart off into deep water, where they are safe from their

numerous enemies. On the south side of the lake the conditions

are very different, there being often fifty yards of muddy shallows

to cut off their retreat.

The nests are merely basin-shaped holes scooped out in the

sand, usually among reeds or tamarisk bushes. The largest one

I saw measured just over a yard in diameter and one and a half

feet deep. I had the good fortune to be able to watch a Bolti at

work on one of these nests. The latter was nearly completed

when I commenced my observations at the beginning of May

and was occupied by a large male with brilliantly coloured head

and fins; there were no signs of a female in the neighbourhood.

The male remained by himself in the nest during the two days

that I observed him, and was occupied chiefly in smoothing down

the sides of the excavation ; this he did by revolving round and

round with his tail in the centre, brushing away dirt from the

sides of the nest with his fins. On the third day I returned

early to the nest and found it unoccupied. A short distance

away, however, thé same male was to be seen nowaccompanied by

a female; courtship was evidently in progress, the male swimming

Pt GS oO Sar Le Oaie

Jd.Green del.et lith.

MEGALOPHRYS NASUTA.

Ieee iS), USOle) IPL DOSTIU.

J. Green del et lth.

MEGALOPHRYS MAJOR.

Wed Gus) NOS), Pl. JOU.

27 See ee

J.Green del et lth.

MEGALOPHRYS ROBUSTA.

lean SOS). PLL On,

d.Green del.et lth.

1. MEGALOPHRYS GRACILIS. 2 M MEITBROPUS.

S. IMC IGUAS SIT,

1908. | ON A GENUS OF ORIENTAL BATRACHIANS. 407

about in front of the female, exhibiting his brilliantly coloured

throat and fins to their best advantage.

During a short absence from my post of observation, the pair

was disturbed by fishermen with casting-nets and never returned

to the nest.

We see, however, that it is the male only who constructs the

nest, and it is not until the latter is completely finished that he

goes out in search of a mate. This is by no means unusual; and

a similar state of things has been described in the American

Cichlid, Geophagus brasiliensis, and in many Centrarchids. /

The native fishermen confirmed my observations and informed

me that they made use of their knowledge by catching the large

males over the holes, the fish being too occupied by their nest-

making to be easily disturbed.

I was unfortunately unable to witness the oviposition; but an

intelligent native whom I cross-examined informed me that male

and female repair together to the nest where the eggs are

deposited, the female later taking them into her mouth. He

ridiculed the idea of a reverse process of parturition, which Loat

asserts to be the belief of some of the Nile fishermen.

A number of fish were brought to me containing ova or young

in the mouth; these, as was to be expected*, all proved to be

females. One specimen, carrying young measuring about 10 mm.

in length, had the oviducts full of large ripe eggs, and was evi-

dently prepared to mate again as soon as the first brood was

disposed of, as already ascertained by Pellegrint in Tilapia

galilea.

6. A Revision of the Oriental Pelobatid Batrachians (Genus

Megalophrys). By G. A. Boutencsr, F.R.S., V.P.Z.8.

[Received April 6, 1908. ]

(Plates XXII.-XXV.t and Text-figure 78.)

My attention has been drawn to the generic characters of the

Pelobatide of the Oriental Region through the notes published

by Mr. Beddard a year ago in these ‘ Proceedings’ § on the

anatomy of Megalophrys nasuta, in which he stated that the

single specimen examined by him had proccelous vertebra, whereas

the geuns Megalophrys had been defined by Cope and myself

as having the vertebre opisthoceelous. A skeleton of an adult

Megalophrys nasuta which I had prepared confirmed Mr. Beddard’s

statement; but at the same time I find the vertebre to be pro-

ceelous also in some specimens of IZ. montana, the type of the

genus, and of MM. longipes, of which species other specimens

* Cf. G. A. Boulenger, Tr. Zool. Soc. xvii. 1906, p. 538

+ Mém. Soc. Zool. France, xvi. 1904, p. 311.

{ For explanation of the Plates, see p. 430.

§ P.Z.S. 1907, p. 324.

408 MR. G. A, BOULENGER ON ORIENTAL [ Apr. 28,

showed them to be opisthoceelous. It is therefore clear that the

character, however important it may appear at first, is worthless

even as a specific character in these Batrachians. I had to a

certain extent foreseen this result in 1882*, when [ refused to

divide these frogs into larger groups according to the mode of

articulation of the vertebrae, as had been done by Cope? and

Lataste {, remarking that: “In this case, this character does not

seem to me to be of such importance as was believed by some

authors. It has been noticed that, in some individuals of various

species, the intervertebral spheres do not become firmly attached

to either centrum at maturity, and one adult specimen of

Xenophrys monticola in the British Museum exhibits a very

anomalous mode of articulation, the fourth vertebra being

biconcave, the fifth biconvex, the others proceelian.” Curiously,

I was soon after censured by R. Blanchard § for not having

divided the Tailless Batrachians into two primary groups: the

Opisthoccelous and the Proccelous.

Still I persisted in using the character of the opisthoccelous

vertebre for separating Megalophr ys from Leptobrachium, with

which I had been obliged to unite Giinther’s Yenophrys w hen it

was shown that the presence or absence of vomerine teeth could

not be used as a generic character in this group ||. Now an

unexpected discovery shows the last genus to be still more closely

related tc the first.

Some time ago, whilst collecting in Darjeeling, Dr. Annandale

found some curious tadpoles agreeing so closely with those of

Megalophrys montana, first described from Java by Prof. Max

Weber and since found in the Malay Peninsula. However,

owing to the fact that M. montana has never been recorded

from ithe Himalay as, whilst Xenophrys monticola (now ealled

Megalophrys parva) is common there, doubts arose in his mind

as to the correctness of the identification, and he sent me a large

series of specimens, adult and young, of the latter species,

together with several specimens of the problematic larva, one of

which has the limbs fully developed. <A study of this material

has convinced me that the so-called Xenophrys monticola has

the same sort of tadpole as Megalophrys montana, so closely

resembling it that I can only distinguish it by the whitish

colour of the belly, which in the Malay species is dark brown.

I have therefore no hesitation in abandoning the genera

Xenophrys and Leptobrachium and uniting them with the first-

described Megalophrys. And as the species are in want of

revision, I have seized this opportunity for submitting them all to

a renewed study, the results of which appear in this paper.

Before proceeding with the descriptions of the species, I wish

to reply to two criticisms of Beddard’s in the above-quoted paper.

* Cat. Batr. Ecaud. p. 432. + J. Acad. Philad. (2) vi. 1866, p. 67.

t Actes Soc. Linn. Bord. xxx. 1879, p. 330.

§ Bull. Soc. Zool. France, 1885, p. 584.—Reply by Boulenger, op. cit. 1886,

p. 320. || Ann. Mus. Genova, (2) vil. 1889, p. 750.

1908. | BATRACHIANS OF THE GENUS MEGALOPHRYS. 409:

First, he remarks that Iam mistaken in describing J/egalophrys

as with ‘outer metatarsals united,” in opposition to Pelobates

with “outer metatarsals separated by web.” He finds that

“both frogs agree in the separation by web, only that the web is

more extended towards the tip of the toes in Pelobates.” The

figure given by him of MJegalophrys (fig. 92, p. 326) shows, how-

ever, that the web does not extend into the metatarsal part of

the foot, the outer metatarsal segments being completely bound

together by the integument; whilst in Pelobates, when spread

out, they are seen to be separated by a deep groove into which

the web penetrates, as in the true Frogs.

Secondly, Mr. Beddard still thinks the fusion of the coccyx

with the sacral vertebra to be an important character in

Megalophrys nasuta, although I have shown it to be variable

in Pelobates. I find it to be likewise variable in WVegalophrys,

as may be seen from the following notes on various vertebral

columns examined by me :—

In a dry skeleton of JZ. montana and in two smaller specimens

in spirit of the same species, which I examined when preparing

the ‘Catalogue of Batrachians’ published in 1882, I found the

presacral vertebre, 8 in number, opisthoccelous, and the coceyx

articulating with the sacral vertebre by one condyle, as stated by

Cope. In another skeleton, which has since been prepared, the

vertebree are proceelous, and the coccyx is likewise distinct from

the sacral vertebra.

Ina skeleton of a large IW. nasuta, from Kina Balu, in which the

dermal ossification forms a complete roof over the

Text-fig. 78. skull, similar to but thinner than that of Pelobates

cultripes, and not extending over the temples,

the vertebral column is as in the second spe-

cimen of M. montanu, except that the coccyx is

immovably fixed to the sacrum, although the

line of separation is visible on the lower aspect.

Exactly the same condition obtains in a large

M. major. In a smaller specimen of JZ. nasuta

from the same locality the coceyx is quite free

from the sacral vertebra and the vertebra are

opisthoccelous.

The vertebre are proccelous or opisthoceelous

in M. longipes, and the coceyx is completely

fused with the sacrum.

Vertebral column In several specimens of JZ. parvus the ver-

one tebre are proccelous and the coccyx is free and

ventral aspect. movable.

A skeleton of I/. pelodytoides is interesting as

showing an anomaly somewhat similar to one described by Howes

in Bombinator*. Anomalies, we know, are frequent in the:

Discoglosside and also in Pelobates t. There are two triangular

* Proc. Anat. Soc. 1890, p. xvi, fig.

+ Of. Adolphi, Morphol. “ein xx. 1895, p. 449, pl. xix.

410 MR. G. A, BOULENGER ON ORIENTAL [ Apr. 28,

sacral diapophyses on the left side—the first on the ninth vertebra,

as normal, the second being part of the coccyx; on the right

side the ninth vertebra has a slender diapophysis, as on that

preceding it, and the triangular process forms part of the

Coccyx.

T have alluded above to irregularities in the mode of arti-

culation of the vertebre, which lose all systematic importance

from the fact that, in these lowly forms, the intervertebral

sphere of cartilage which is at first distinct from the so-called

centra, as they ossify, become attached either to the vertebra

anterior or posterior to each, if not remaining independent. We

may thus find, on the same vertebral column, proccelous, opistho-

celous, or amphiceelous vertebre ; and it is now quite clear that

such a character is worthless in the Pelobatidz even as a specific

character. I may add that the same will probably be found to

be the case in the Hemiphractide. Brocchi has described the

vertebre of Hemiphractus as opisthocelous. JI have not been

able to verify the statement, but I find them procelous in a

skeleton of the closely-allied Oeratohyla bubalus.

Genus MEGALopHrys Kuhl.

Pupil erect. Tongue circular or pyriform, entire or nicked

and free behind. Vomerine teeth in two small groups, if pre-

sent. Tympanum distinct or hidden under the skin. Fingers

free, toes free or shortly webbed; outer metatarsals united.

Sternum with a bony style. Coccyx, if distinct from the sacral

vertebra, with simple articulation.

South-eastern Asia.

Synopsis of the Species.

I. Profile of snout obliquely truncate, strongly projecting beyond lower jaw;

canthus rostralis angular and loreal region vertical or shghtly oblique.

A. Head at least once and a half as broad as long (to occiput) ; tibio-tarsal

articulation not reaching eye; vomerine teeth usually present ; tympanum

hidden or feebly distinct.

End of snout without or with a mere indication of a

dermal appendage; outer border of upper eyelid pro-

duced in the adult into a pointed appendage, which

does not, as a rule, measure more than half diameter

DEOVE eiciees con ere costal leaden Menon eae seer Rae Maa See oer

Snout terminating in a pointed dermal appendage; outer

border of upper eyelid produced into a pointed append-

age, which is at least $ diameter of eye in the adult ... 2. WM. nasuta Schleg.

1. M. montana Kuhl.

B. Head not more than once and two-fifths as broad as long; tympanum

distinct.

1. Tibio-tarsal articulation reaching beyond eye ; vomerine teeth present.

Upper eyelid with a horn-like tubercle; tibia 13 to

12 times in length from snout to vent; toes with a

slicht mudumentlof web) sscscsdessscesssese ee see se

Upper eyelid without appendage; tibia 12 to 2 times

in length from snout to vent ; toes + to 3 webbed ...... 4, M. major Blgr.

3. W. longipes Blgy.

1908. | BATRACHIANS OF THE GENUS MEGALOPHRYS. 4\1

2, Tibio-tarsal articulation not reaching beyond eye (rarely slightly beyond) ;

tibia at least twice in length from snout to vent; toes with a slight

rudiment of web.

Vomerine teeth present ; tympanum about + diameter of

eye, which equals the distance between the two ......... 5. WM. robusta Blev.

Vomerine teeth present; tympanum 2 to = diameter of

eye, which is less than the distance between the two... 6. IZ. parva Bley.

Vomerine teeth none; tympanum 2 to } diameter of eye,

which is less than the distance between the two......... 7. MM. bettgeri Bley.

II. Snout rounded or vertically truncate, not or but feebly projecting beyond lower

jaw.

A, Tibio-tarsal articulation reaching the eye, or beyond ; head as long as broad

or slightly broader than long; tympanum perfectly distinct ; no vomerine

teeth.

Tibio-tarsal articulation reaching tip of snout; tibia a

little more than 3 length from snout to vent; snout

rounded sre ono cad avngpAceonmaRBorBenaduansoeaBAScagsevaHecooasvess Sh JUL Giracolas Gillie.

Tibio-tarsal articulation reaching eye; tibia } length

from snout to vent; snout truncate ........ . 9. MW. heteropus Bigy.

Tibio-tarsal articulation reaching eye or between eye and

nostril; tibia 3, or a little less than 3, length from

HUT, THO) WEG S SHMONNE WOLVES 4. conaavesomssnoudasonanscoonaan. WOK) JE, pelodytoides Blev.

B, Tibio-tarsal articulation not reaching the eye; tympanum feebly distinct or

hidden.

1. Head not more than once and } as broad as long; no vomerine teeth :

toes + to + webbed.

Foot longer than head ..............-..ses ceereereeerssesesesses, LL. UM, boulengeri Bedr.

Foot as long as head . 12. MW. hasseltii Tsch.

2. Head at least once and $ as broad as long; one or several conical tubercles

on upper eyelid ; vomerine teeth usually present.

Loreal region very oblique; toes $ to webbed ............ 13. DM. carinensis Blgr.

Loreal region very oblique; toes nearly free or with a

Mele LUCIMENGIOR WED Ee ae watpenyteeneqta-weesesea-aanns aay An, 0 fee Blow

Loreal region vertical; toes with a mere rudiment of

VC) 0) stiaisadeacis Sadan aeemopiec atone ee a scan cer Gosmnarense acct eee tel Ustad RLU TITS Bler.

1. MEGALOPHRYS MONTANA,

Megophrys monticola Kuhl, Isis, 1822, p. 475.

Megalophrys montana Kuhl, in Férussac, Bull. Sc. Nat. ii.

1824, p.83; Wagler, Syst. Amph. p. 204 (1830); Tschudi, Class.

Batr. p. 82 (1838); Dum. & Bibr. Erp. Gén. viii. p. 458 (1841);

Giinth, Cat. Batr. Sal. p. 36 (1858), and Ann. & Mag. N. H. (4)

xi, 1873, p. 419; Bouleng. Cat. Batr. Ecaud. p. 442 (1882) ;

M. Weber, Ann. Jard. Bot. Buitenzorg, xy. Suppl. ii. 1898, Ton) Ove

Laidlaw, Proc. Zool. Soc. 1900, p. 889; Annandale, Fasc. Mal.,

Zool. 1. p. 275 (1903).

Ceratophrys montana Gravenh. Delic. Mus. Zool. Vratisl. p. 47

(1829); Schleg. Abbild. p. 29, pl. x. fig. 3 (1837),

Ceratophryne montana Schleg. Handl. Dierk. ii. p. 57 (1858).

Megalophrys montana, part., Giinth. Rept. Brit. Ind. p. 413

1864).

Megalophrys montana, var. aceras, Bouleng. in Annandale &

Robins. Fase. Mal., Zool. p. 131, pl. v. fig. 1 (1903),

Tongue entire or feebly nicked behind. Vomerine teeth

usually present, in two widely separated small groups just

behind the level of the choane. Head large, 14 to 1 7? times as

412 MR. G. A, BOULENGER ON ORIENTAL [Apr. 28,

broad as long, usually defined behind by a more or less distinct

transverse fold; snout truncate or obtusely pointed, projecting

beyond lower jaw, as long as or a little shorter than eye ; canthus

rostralis sharp; loreal region vertical or a little oblique, concave ;

nostril equally distant from eye and from end of snout; inter-

orbital space concave, its width 14 to 2 times that of upper

eyelid (narrower in the very young); tympanum usually feebly

distinct, rarely hidden, its diameter 4 to # that of eye, from

which it is separated by a distance equal to the diameter of the

latter. Fingers obtuse or feebly swollen at the end, first as long

as or a little longer than second, which measures 2 to } length of

third; no subarticular tubercles; no distinct metacarpal tubercles.

Toes rather short, obtuse or feebly swollen at the end, with a mere

rudiment of web or, at most, + webbed ; no subarticular tubercles ;

a flat, very indistinct inner metatarsal tubercle. Tibio-tarsal arti-

culation reaching the shoulder, the commissure of the jaws, or the

temple; tibia 2 to + length from snout to vent; foot as long as or

shorter than tibia. Skin of upper parts smooth or with scattered

conical warts, old specimens with bony deposits on the head and

anterior part of the back; a strong glandular fold from eye to

shoulder, usually another on each side of the back; upper eyelid

with a sharp, raised edge, which is produced into a more or less

distinct point or “horn”; this point may be very indistinct (var.

aceras Blgr.),or very much developed, measuring nearly # diameter

of eye; as a rule it does not measure more than 4 diameter of

eye; an indication of a similar appendage on the tip of the snout

rarely present *; a more or less developed pointed tubercle usually

present behind the commissure of the jaws; limbs usually with

oblique transverse glandular ridges; throat smooth, belly with small

tubercles. Olive-brown above, uniform or variously marked with

darker or lighter; a more or less distinct large triangular dark

spot between the eyes, the base forwards, and a dark oblique bar

below the eye; limbs with more or less distinct dark cross-bars ;

lower parts pale brown, spotted or marbled with darker ; a white

tubercle on each side of the breast. Male without vocal sac.

Measurements of specimens from Java :—

3. Q.

TDIHOOR, SaVOVOIG WO) MEM Ye edovauoeaoeeudooedeus 5D 88 millim.

Length of head (to occiput) ............... 19 Or ge aes

Widthvotdheads jis peow mere echoes seen 29 ae)

ene uhvor smoutie rece ne. ere ecece ee ebeer ee ete 6 Soe

Diameter of eye ..... MOMS ee ec oe eee 6 Sh hoy

Iranierore orien yale Sadecceroesceossaesuouscdsec 10 AS vn

iDiameber (oh bympam UME emer eeectrer 4 An pee

Distance between eye and tympanum ... 6 Dy tee

iBLore sani e ee eens cee ee Cet ec ee 35 Dan ee

1 SE ave emg Sana ett Ame tee oe arts SonGoas deat oN 16 2 ee

ie have Well era) op oGnene Manton ds Seo ddec osuob accesses LOMO es.

ARIE sea eGanG agate a aaa AE Wan sk enindnasnannods 23 Big

HOGG Rash oes cone ae Cee ene eee nen eee 21 SOM

* In a specimen from Java, where the species is most abundant.

1908. | BATRACHIANS OF THE GENUS MEGALOPHRYS. 413

Originally described from Java, this curious-looking frog has

since been found in other parts of the Malay Archipelago and in

the Malay Peninsula. The numerous specimens in the British

Museum are from Java, Borneo, Palawan, Balabac, Samar and

Dinagat Ids. (Philippines), Sumatra, and Bukit Besar in Jalor.

It is a sluggish and thoroughly nocturnal animal. Nothing

has been observed concerning its breeding-habits ; but I find the

eges to be large, those in the oviduct of a specimen 83 millim.

long measuring 3 millim. in diameter. The larvee were first

described by Max Weber* from Java, and Laidlaw? and

Annandale ¢ have published observations on specimens obtained

in the Malay Peninsula. ‘These tadpoles are found in mountain-

streams with gravelly beds and are remarkable for the funnel-like

float formed by the lips, which are beset with minute horny teeth ;

these are not connected in any way with definite ridges or lamelle,

but radiate along the anterior surface of thefunnel. According to

Annandale, the funnel-shaped lip is capable of assuming two very

distinct forms, according to the position of the tadpole :—(1) When

the animal is hanging from the surface-film, as it frequently does,

this structure becomes a translucent rhomboidal or lozenge-shaped

float, depressed in the centre towards the mouth, but otherwise

nearly flat; (2) when, on the other hand, the animal is resting on

the bottom, the float takes on the appearance of a pair of slender

processes, continued upwards on the sides, like a pair of horns.

As in other Pelobatidee, the spiraculum is sinistral. The tail is

more than twice as long as the body, the total length of the

largest tadpole being about 40 millim. The coloration is of a

very dark brown, even on the belly.

2. MrecALopHrys NasuTA. (Plate XXII.)

Ceratophrys montana, var., Schleg. Abbild. p. 30 (1837).

Megalophrys montana (non Kuhl), Cantor, Cat. Mal. Rept.

p. 140 (1847).

Ceratophryne nasuta Schleg. Handl. Dierk. ii. p. 57, pl. iv.

fig. 72 (1858); Giinth. Cat. Batr. Sal. p. 36 (1858).

Megalophrys montana, part., Ginth. Rept. Brit. Ind. p. 413

(1864).

‘ eae chystt Edeling, Tijdschr. Nederl. Dierk. 1864,

p. 205, and Nat. Tijdschr. Nederl. Ind. xxvii. 1864, p. 265, pl. —.

Ceratophrys nasuta Schleg. Dierentuin, Rept. p. 58, fig. (1872).

Megalophrys nasuta Giinth. Ann. & Mag. N. H. (4) xi. 1873,

p. 419; Bouleng. Cat. Batr. Ecaud. p. 443 (1882); 8. Flower,

Proc. Zool. Soc. 1899, p. 913; Werner, Zool. Jahrb., Syst. xiii.

1900, p. 498; Isenschmid, Mitth. Nat. Ges. Bern, 1903, p. 21;

Beddard, Proc. Zool. Soc. 1907, p. 324, figs.

Tongue entire or feebly nicked behind. Vomerine teeth pre-

sent, in two widely separated small groups on a level with the

posterior border of the choanz. Head large, 14 to 2 times as broad

* Ann. Jard. Botan. Buitenzorg, 1898, Suppl. ii. p. 5.

+ Proc. Zool. Soe. 1900, p. 889.

{ Fascic. Malay., Zool. p. 275 (1908).

414 MR. G. A. BOULENGER ON ORIENTAL [Apr. 28,

as long, usually defined behind by a more or less distinct trans-

verse fold; snout truncate in profile, pointed from above, and

projecting beyond lower jaw, as long as or a little shorter than

eye; canthus rostralis sharp; loreal region vertical or a little

oblique, concave; nostril equally distant from eye and from end

of snout; interorbital space concave, its width 13 to 2 times that

of upper ‘eyelid (narrower in the very young ‘); tympanum often

hidden ; if distinct, its diameter 3 to § that of eye, from which it

is widely separated. Fingers obtuse or feebly swollen at the end,

first as long as or a little longer than second, which measures $ 2 to

2 length of third; no subarticular tubercles ; no distinct meta-

carpal tubercles. Toes rather short, obtuse or feebly swollen at

the end, with a mere rudiment or web or, at most, + webbed * ;

no subarticular tubercles; a flat, very indistinct inner metatarsal

tubercle. Tuibio-taisal anealeition reaching the shoulder or the

commissure of the jaws tT; tibia 3 to 2 length from snout to vent ;

foot a little shorter than tibia. “Sls of. upper parts smooth or

with a few scattered warts; adult with bony deposits on the head

and anterior part of the back, which may completely fuse with the

skull, and form a bony shield on the presacral part of the body ;

a glandular fold from eye to shoulder, sometimes continued on the

side of the body; another fold usually present higher up on the

back, from behind the head to the sacral region ; frequently, but

not constantly, a subconical tubercle on the scapular region and

another in the middle of the sacral region; similar tubercles,

exceptionally, on the head; upper eyelid with a sharp, raised

edge, produced into a long point or “ horn,’ which is at least 2,

and often quite, as long as the eye in the adult; a similar, but

shorter appendage on the end of the snout, and a still smaller one

at the angle of the jaws ; limbs with or without oblique transverse

glandular ridges ; lower parts smooth, or belly with small tubercles.

Olive-brown above, uniform or variously marked with darker and

lighter; a more or less distinct Y-shaped dark marking between

the eyes and on the occiput; a dark oblique bar below the eye ;

lower parts dark brown, or spotted or marbled with dark brown.

Male with an internal vocal sac.

Measurements of two specimens from Sumatra, found pairing

by Dr. H. O. Forbes :—

3. ;

Mom SNa mb bO, Web ci admcebeet ye ace eeee - 82 125 millim.

Length of head (to occiput) ............... 24 SO wer

IWadibligor headle€ 285 ie. 4 Reps ie ok Pe 4] GOR ae

Menethiotisnout leh... seme these een a OUgS ANS

DiamSter Ofjeye nie h. hee die Rin deat af eae?

Interorbital width ..... Rasa cier cre Aa Seti a 13 20), (ae

Diameter of tympanum ..........:.-..--.--: 4 6. ape

Distance between eye and tympanum... 8 UG oe

* Tsenschmid (7. c.) describes a specimen from Sumatra in which the toes are

nearly half webbed.

+ The eye in a specimen from Sumatra, according to Isenschmid (J. ¢.).

1908. ] BATRACHIANS OF THE GENUS MEGALOPHRYS. 4J5

Horeslimibseeseeee SH OREO SD OEE aR Eee enn aan 54 74 millim.

lS lehave daca s Sho cae pee ee ee eee ena 20 oly eae

JEtnAClllnin OM 2 iianasac stone ometen tee eee 96 NSyee

GR OY eyo es Mee Wis 2 On Race gee a 30 AD. ae

TO OG 8 ones Nee IEP ere Sati assde hs whe 29 40°,

Known from the Malay Peninsula (Penang, Perak), Sumatra,

Borneo, and the Natuna Islands.

The above description is almost a repetition of that of IZ. mon-

tana, so closely are these species allied to each other. As I

mentioned in 1882, the only important difference between the two

resides in the greater length of the palpebral horn-like appendage,

and the presence of a similar appendage on the tip of the snout in

M. nasuta. To this may be added the somewhat more anterior

position of the vomerine teeth in J/. nasuta.

Dr. H. O. Forbes has observed the frog to pair, in Sumatra *,

on the banks of streams, the male embracing the female round

the lumbar region. Recently transformed young, some with

remains of the larval tail, measuring 15 to 17 millim. from snout

to vent, were obtained in Penang by Capt. Flower; the palpebral

and rostral appendages are absent in three very young specimens,

and I do not see how they can be distinguished from J/. montana

at a similar stage of development.

3. MEGALOPHRYS LONGIPES.

Megalophrys longipes Bouleng. Proc. Zool. Soc. 1885, p. 850,

pl. lv.; Giinth. Ann. & Mag. N. H. (5) xx. 1887, p. 316; A. L.

Butler, Journ. Bombay N. H. Soc. xv. 1904, p. 400.

Tongue feebly nicked behind. Vomerine teeth in two small

groups just behind the level of the choane. Head moderate, much

depressed, once and 7 as broad as long; snout obliquely truncate

in profile, projecting beyond lower jaw, nearly as long as eye;

canthus rostralis sharp; loreal region vertical, concave; nostril

equally distant from eye and from end of snout; inter-

orbital space concave, as broad as or a little broader than upper

eyelid; tympanum distinct, 2 to 3 diameter of eye, its diameter

equal to or less than its distance from eye. Fingers slender,

feebly swollen at the end, first as long as or a little longer than

second, which measures about 2 length of third; no subarticular

tubercles; no distinct metacarpal tubercles. Toes long and

slender, feebly swollen at the end, with a slight rudiment of

web; no subarticular tubercles ; no distinct metatarsal tubercle.

Tibio-tarsal articulation reaching beyond tip of snout, knee reach-

ing axil or shoulder; tibia 2 to 2 length from snout to vent; foot

shorter than tibia. Skin smooth above, with small warts on the

flanks, and two pairs of delicate glandular folds originating on the

scapular region and converging posteriorly, the outer extending to

the sacral region ; a strong fold from the eye to the shoulder; a

* “AM Naturalist’s Wanderings in the Eastern Archipelago’ (London, 1885), p. 154,

Proc. Zoou. Soc.—1908, No. XX VII. 27

416 MR. G. A. BOULENGER ON ORIENTAL [ Apr. 28,

short horn-like tubercle on outer edge of upper eyelid; lower

parts smooth. Olive-brown above; sides of head with oblique

dark bars; a large triangular dark marking between the eyes;

limbs with dark ecross-bars; hinder side of thighs dark brown,

with one or two round whitish spots ; belly dotted or spotted with

brown; throat and breast dark brown, or with large dark brown

spots or symmetrical markings. Male with an internal vocal

sac.

@s OF

TNAOMAN, SNOW WO WEIN, 6404s sece0c0ses soe scase- AT 65 millim.

Length of head (to occiput) ............... 14 TDs psen

Widthiot Neagle. of es tens AW robe fae cer 7 BPO

emer no lelsmO UG aye) errr ee eae 4 Orcas

Diemeteroh eye wr si ic tan Geet re ceee ae 5 6

lamteror-loitall avait eee eee anes ee nee eer 5 cle was

Drameterton chyna aun eee Sheeran 3 34s,

Distance between eye andtympanum ... 4 On ue

ore mma Gen. ce 4 caer Aa ee re hee ol LS IN se

TDs C10 LR Adair ib ce ae Nee i Be ler ee aR 12 NG) gues

1a Grave Mla) oy Spee eee tee Rd aoe Near doe oray Tey 5

AST Tar terete tee tek ecg a eae ds cat see 27 Bo 55

NOX Rec dele Sea is te NT cmos oa orien ea chore 22 33

Only known from the mountains of Perak, in the Malay

Peninsula, between 3000 and 4500 feet. Mr. A. L. Butler has

observed it to be the commonest frog on the hills above 3000 ft.

and to be entirely nocturnal, being found in the daytime under

logs, rocks, or in holes in banks, and in densely shaded spots among

dead leaves. These frogs appear to be quite bewildered by the

sunlight, and when disturbed give one jump into the open and

make no other attempt to escape. When seized in the hand they

frequently open their mouths widely for some seconds. Mr. Butler

has never seen this frog enter water of its own accord, and he

suspects very large ova (4 inch in diameter), containing tadpoles

with the hind limbs and tail well developed, which he found

under damp moss on tree-trunks, to belong to it.

4, MuGALoPHRYs Mason, n.n. (Plate XXIII.)

Xenophrys gigas Jerdon, Proc. As. Soc. Beng. 1870, p. 85, and

in Anders. Proc. Zool. Soc. 1871, p. 200.

Xenophrys monticola, part., Anders. 1. c.; Bouleng. Cat. Batr.

Eceaud. p. 441 (1882).

2 Ixalus lateralis Anders. Journ. As. Soc. Beng. xl. 1878, p. 29,

and Anat. Zool. Res. Yunnan, p. 844, pl. Ixxviii. fig. 5 (1879).

Leptobrachium monticola, part., Bouleng. Ann. Mus. Genova, (2)

vil. 1889, p. 720, Faun. Ind., Rept. p. 510 (1890), and Ann. Mus.

Genova, (2) xii. 1893, p. 344.

Tongue entire or feebly nicked behind. Vomerine teeth in two

small groups on a line with the posterior border of the choane.

Head moderate, more strongly depressed than in the preceding

1908. ] BATRACHIANS OF THE GENUS MEGALOPHRYS. 417

species, 17 to 12 times as broad as long; snout obliquely truncate

in profile, projecting beyond lower jaw, nearly as long as the eye;

canthus rostralis sharp ; loreal region vertical or slightly oblique,

concave ; nostril equally distant from eye and from end of snout ;

interorbital space flat or slightly concave, as broad as or a little

broader than upper eyelid; tympanum more or less distinct,

about half diameter of eye, the distance between eye and

tympanum nearly equal to 3 or once the diameter of the former.

Fingers with feebly swollen tips, first as long as or a little shorter

than second, which measures about 2 length of third; no sub-

articular tubercles; no distinct metacarpal tubercles. Toes

moderate, with swollen tips, 7 to + webbed, the web extending as

a lateral fringe; no subarticular tubercles ; metatarsal tubercle

flat, indistinct. Tibio-tarsal articulation reaching the tip of the

snout or a little beyond, rarely only to between eye and nostril ;

tibia a little more than 4 or 2 (rarely only 3) length from snout to

vent ; foot considerably shorter than tibia. Skin smooth, or with

fine granular asperities on the back; fine glandular ridges may

be present on the body and across the limbs, one on each side of

the back being constant and a V-shaped one behind the head very

frequent; a glandular fold from the eye to the shoulder. Brown

above, with darker, light-edged symmetrical markings, the most

constant being a triangular spot between the eyes; a dark band

capping the end of the snout as far as the nostrils and extending

along the loreal and temporal regions to the shoulders; a light

streak along the upper lip, sometimes interrupted by a dark brown

bar below the eye ; limbs with irregular dark cross-bands ; hinder

side of thighs dark brown, with a round white spot, which is

nearer the leg than the vent; belly whitish, throat and breast

brown or marbled with brown, with darker, light-edged spots on

the lower lip, the last of which extends as a curved band to the

anterior face of the arm. Male with a subgular vocal sac and

fine brown nuptial asperities on the inner side of the first and

second fingers.

Measurements of type specimens from Darjeeling :—

Cre ese

IRON SINCWNG WO WEIN 4 seo soetoonosauesosoes se dete ouirnalline

Ikength of head (to occiput) (....027.2-.... 22 PASS aiuiher

Wadthrotebeadh 22) Bes 6 or SAN 29 BO cl gy

enothyote smote s\n. aes 9

Diameberomeyey iy...) wees: 9

littrerormbibalewacit nee ye ee eee 8 ORME

Diameterlof ty mpamum) 92-53) eee 4

Distance between eye andtympanum ... 7

Hore: ime get renee sen eee 48 OE

MED eat SY SA tere te amy Sree dah earn 21 eee

HE=Tina cli Innate eer re yn een ssh eres ee (eo G2 tae

MRTOTES ete eee teen er Anima ae Na 4] DN ater

HOO LG Peete eres rite a ecu in an Me ROR 37 AR tu

418 MR, G. A. BOULENGER ON ORIENTAL [Apr. 28,

This frog was indicated, rather than described, by Jerdon in

1870, from specimens collected by him at Darjeeling and in the

Khasi Hills, now preserved in the British and Indian Museums.

Further allusion to it was made soon after by J. Anderson,

who regarded it however as the adult of Giinther’s Yenophrys

monticola. After examining about 50 specimens, measuring

32-94 millim. from snout to vent, I am convinced that Jerdon was

right. Most of the specimens in the Indian Museum are from

Cherrapunji, Khasi Hills. I have examined specimens from the

Karin Hills, in Upper Burma, and from the Man Son Mts.,

Tonkin (3000-4000 feet). If I am right in regarding Anderson’s

Txalus lateralis as a young M. gigas, the range of the species

extends to Western Yunnan.

I regret the name chosen by Jerdon cannot be used for this

species, as there is an earlier Megalophrys gigas Blyth, 1854,

which is, however, a synonym of Rana liebigii Gthr. I have

therefore proposed the new name Megalophrys major.

5. MEGALOPHRYS ROBUSTA, sp.n. (Plate XXIV.)

Tongue feebly nicked behind. Vomerine teeth in two small

groups on a line with or just behind the posterior borders of the

choane. Head moderate, once and 2 as broad as long; snout

obliquely truncate in profile, projecting beyond lower jaw, as long

as eye; canthus rostralis sharp; loreal region slightly oblique,

concave ; nostril equally distant from eye and from end of snout,

or slightly nearer the former ; interorbital space concave, broader

than upper eyelid; tympanum distinct, about half diameter of

eye, which equals the distance between the two. Fingers with

feebly swollen tips, first as long as or a little longer than second,

which measures about 3 length of third ; no subarticular tubercles ;

no distinct metacarpal tubercles. Toes moderately long, with

feebly swollen tips, with a very slight rudiment of web; no sub-

articular tubercles; metatarsal tubercle flat, indistinct. Tibio-

tarsal articulation reaching the eye, or between eye and nostril ;

tibia 4 or a little less than 4 length from snout to vent; foot a

little shorter than tibia. Skin of upper parts smooth or finely

granulate ; a fine glandular ridge on each side of the back and a

V-shaped one behind the head ; a strong glandular fold from the

eye to the shoulder. Brown above; a triangular dark marking

between the eyes; a dark temporal band and a dark vertical bar

below the eye; other dark bars sometimes present on the upper

lip; hind limbs with indistinct dark cross-bars ; hinder side of .

thighs dark brown, with a round light spot nearer the leg than

the vent; belly white; throat and breast spotted or marbled with

brown.

Measurements of two specimens in the Indian Museum,

Calcutta :—

OF len

JNO, STOW WO WIA oc acungvoes sooacceas see 114 54 millim.

Length of head (to occiput) ............... 30 Sa aes.

1908. | BATRACHIANS OF THE GENUS MEGALOPHRYS. 419

Byte Elon |

IWatcthyoitneatcl te eee nee he eae 43 21 millim.

eng thyoitcmout ems sacs eee. ene 10 ee

Diameter of eye tre tc: des eee take 10 eH od

haterorbitalswidisierne te seee eee eee eee 13 Or Res,

Diameter Of tyanpamunny eens. eese eee 4 a ee

Distance between eye and tympanum ae ed) oa

IHOKewiimM peer eerer me ee eaten centne coe 66 36 Be

NG AVEC.) 22 sae aman co NO Uae ean A ata: 27 Ae at

(Pindeltmalys ocsce anc Noc 2. es aes s cole ones aeee ee 165 Soma

RN ORE Riss 8 sees arn ee eed ama eer aa yt 53 Dilfer ea

EL OO Gite eters Gan ira nea enamine serie AQ PAD) Jil he

I have examined five specimens, all from Darjeeling, collected by

Dr. J. Gammie, and preserved in the Caleutta Museum. I have

been allowed to retain one (here figured) for the British Museum.

6. MEGALOPHRYS PARVA.

Xenophrys monticola Giinth, Rept. Brit. Ind. p. 414, pl. xxvi.

fig. H (1864).

Xenophrys monticola, part., Anders. Proc. Zool. Soc. 1871,

p. 200; Bouleng. Cat. Batr. Hcaud. p. 441 (1882).

Leptobrachium monticola, part., Bouleng. Ann. Mus. Genova, (2)

vil. 1889, p. 720, and Faun. Ind., Rept. p. 510 (1890).

Leptobrachium parvum Bouleng, Ann. Mus. Genova, (2) xiii.

1893, p. 344, pl. xi. fig. 2.

Tongue entire or feebly nicked behind. Vomerine teeth in

two small groups on a line with or just behind the posterior

borders of the choanz. Head moderate, 17 to 13 times as broad as

long ; snout obliquely truncate in profile, projecting beyond lower

jaw, nearly as long as eye; canthus rostralis sharp; loreal region

vertical, concave ; nostril equally distant from eye and from end

of snout; interorbital space flat, as broad as upper eyelid ; tym-

panum more or less distinct, 2 to 2 + diameter of eye, the distance

between the two less than the diameter of the latter. Fingers

with feebly swollen tips, first as long as or a little shorter than

second, which measures about 2 length of third; no subarticular

tubercles : no distinct metacarpal ‘snare ls, Toes rather short,

with feebly swollen tips, with a slight rudiment of web, often

with a feeble dermal ridge on the lower surface, but without sub-

articular tubercles; metatarsal tubercle flat, indistinct. Tibio-

tarsal articulation reaching the eye or not quite so far; tibia 4 or

a little less than 4 length from snout to vent; foot shorter than

tibia. Skin of upper parts smooth or granular, with small

glandular warts which may form symmetrical ridges, very variable

in arrangement; a strong glandular fold from the eye to the

shoulder ; lower parts perfectly smooth. Brownish above, with

more or less distinct darker symmetrical markings, the most

constant of which forms a triangle or a Y between the eyes; some

of the warts edged with blackish; upper lip with vertical dark

420 MR. G. A. BOULENGER ON ORIENTAL | Apr. 28,

bars; limbs with more or less regular dark cross-bars ; hinder

side of thighs marbled with brown, usually with a round light

spot nearer the leg than the vent ; lower parts whitish, throat and

breast spotted or marbled with brown, or with brown symmetrical

marking; a round white spot often present on each side of the

breast. Male with a subgular vocal sac, and with fine brown

nuptial rugosities on the first and second fingers.

GE OF

IromesnoubbOuveltan i tessa ace eeeece 42 52 millim.

Length of head (to occiput) ............... ig AD) eee

INV Ghvok mead « .:cemmericacicons cee lance 14 5S aah

Gemethvol snout cesses e eeraeeee 4 Diets are

Damme ter, Of ieye Wi.,.\. bites Sy Baby te ol As Ox ais

Ifronerserelonivell Syl, Sodaeudaanodedodocsenesaccn 4 Oak Nos,

IDrevameiiere Ore whysa0yORWONUNIM, 66 oeescecoscsosncn0 3 Amite

Distance between eye and tympanum ... 3 A ee

Hours MMA Fsrararaic y wire etels/e saipiscarcere eisai eee 29 3) os

TAG Up Ra eere ee eR ARR Tee cee coes Gosh eee soscre I lowes

Te buaye sl inan) ope eens one t dann Guanes eon ee een 65 (fox Pie

RUG AeA SOA SA Ue een RIOR Cc eRe San ene 21 A

Lore teratoma dated goabeenesteoten sane onc ae 18 20m a

The type specimens of Xenophrys monticola*, in the British

Museum, are from the Sikkim Himalayas and the Khasi Hills.

Others, from Darjeeling (Jerdon) and Pegu (Theobald), are pre-

served in the same Institution. I have examined about 40

specimens from Darjeeling and Cherrapunji belonging to the

Calcutta Museum. Leptobrachium parvum was described from

specimens obtained by Fea in the Karin Hills, Upper Burma.

No direct observations have been made on the habits of this

species, but an inspection of the oviducts of females on the point

of spawning show the eggs to be large, 2 millim. in diameter.

Tadpoles obtained by Dr. Annandale at Kurseong and referred

by him provisionally to J/. montana of Kuhl? belong, I feel

convinced, to this species, in which the toes are nearly free. These

tadpoles, with funnel-shaped mouth, agree very closely with those

of the former species, as known from specimens obtained by

Messrs. Annandale and Robinson in the Malay Peninsula, differ-

ing only, so far as I can see, in the whitish belly.

7. MEGALOPHRYS BOETTGERI.

Leptobrachium boettgeri, Bouleng. Proc. Zool. Soc. 1899, p. 171,

pl. xix. fig. 3.

Tongue entire. Vomerine teeth absent. Head moderate, 14

to 13 times as broad as long ; snout obliquely truncate in profile,

projecting beyond lower jaw, nearly as long as eye; canthus

* The specific name of which must be changed, as being preoccupied in the genus

Megalophrys.

Proc. As. Soc. Beng. (2) ii. 1906, p. 290.

1908. ] BATRACHIANS OF THE GENUS MEGALOPHRYS. 421

rostralis sharp; loreal region vertical, concave; nostril equally

distant from eye and from end of snout; interorbital Space flat,

as broad as upper eyelid ; tympanum very distinct, 3 to = diameter

of eye, equal to or greater than its distance from latter. Fingers

with feebly swollen. tips, first and second equal, about 5 length of

third ; subarticular and carpal tubercles indistinct or atcome Toes

slender, with feebly swollen tips, with a slight rudiment of web,

often with a feeble dermal ridge on the lower surface, but without

subarticular tubercles ; a small, oval, flat inner agian! tubercle.

Tibio-tarsal articulation reaching the eye; tibia 4 ora little less

than 3 length from snout to vent ; foot shorter than tibia. Skin

smooth, itl small scattered warts on the head and back; a

elandular fold from the eye to the shoulder; two small white

warts close together on the chin and one on each side of the breast

near the insertion of the fore limb. Dark grey or brown above,

with symmetrical blackish markings; upper surface of snout and

scapular region light; a whitish blotch on the upper lp below

anterior half of eye; limbs with dark cross-bands; a small round

white spot on the back of the thigh; throat and breast brown or

brownish; three longitudinal, blackish, light-edged markings on

the throat ; large blackish spots on the side of the belly ; posterior

part of belly and lower surface of thighs dirty white. Male with

internal vocal sac.

Ge :

NRO, SIVOUIG WO WEN ) Goadoasdosuesnecoadanne: 30 46 millin.

Length of head (to occiput) ............... at Ue rer

Wiidthiot mead’ (cite: ccd sane Setadde nnn dte 13 FOR Res

ILGTASN, CH GMOWIG Soegeoosoose hboossocdoonsce sed 3 ANA.

Diameter Oneeveawars ne das een noah 4 DRA

ne rorwitalewaditlyeseeaesece secretes 33 Dyn

Diameter of tympanum ..................+.- Lass Dey

Distance between eye and tympanum ... 23 Sel iiss

ores tems t attet tasclada tats eared ae 24 DORA:

TEV Cy eee ciety unc de oa aaetcvans en 10 20a

TeTenmitel @ilairiatbo eats eet ates Nc ea Me 0 o4 Gu

UDNOTEas raricraaterets Me Si os eh oreeMne sacs 17 ae ae

HOO UEY Rae Aner orice temitras a circ s sarees 15 EO ees

Described from six specimens obtained by Mr. J. D. La Touche

at Kuatun, N.W. Fokien, China. <A specimen from Kiukiang has

been noticed by Boettger (Ber. Senck. Ges. 1894, p. 141) under

the name of Leptobrachiwm monticola.

8. MEGALOPHRYS GRACILIS. (Plate X XV. fig. 1.)

Leptobrachium gracile Giinth, Proc. Zool. Soc. 1872, p. 598 ;

Bouleng. Cat. Batr. Ecaud. p. 440 (1882).

Tongue nicked behind. Vomerine teeth absent. Head

moderate, as long as broad or slightly broader than long ; snout

rounded, not projecting beyond lower jaw, as long as or slightly

shorter than orbit; canthus rostralis distinct ; loreal region not

429 MR. G, A. BOULENGER ON ORIENTAL [ Apr. 28,

very oblique, concave ; nostril a little nearer end of snout than

eye; interorbital space a little narrower than upper eyelid;

tympanum distinct, its diameter + that of eye, from which it is

Separated by a space nearly equal to its own diameter. Fingers

slightly swollen at the end, first and second equal and about

§ length of third; no subarticular tubercles; two metacarpal

tubercles, inner very large, outer small. Toes rather long and

slender, with blunt or slightly swollen tips, with a very short

web at the base; no subarticular tubercles, but a more or less

distinct dermal ridge along the lower surface ; a feebly prominent,

elliptical inner metatarsal tubercle. Tibio-tarsal articulation

reaching tip of snout; tibia a little more than 1 length from

snout to vent; foot shorter than tibia, but longer than head.

Skin smooth ; a curved glandular foldabovetympanum. Greyish

olive to dark olive above, with more or less distinct darker spots

or symmetrical markings on the head and body and cross-bars on

the limbs ; the type specimen has a whitish spot below the eye,

and the upper arm and elbow whitish; lower parts dirty white,

with or without irregular brown spots. Male unknown.

TOIOTEN SHOVORUIG WO) WEIN 4454 be scac see easesnees 44 millim.

Length of head (to occiput)............... IRS eae

Widithrotihiead’ 2: mea see ee ee Ok ee

kenge thyor (snout) sere sane en aeee Dee.

Miamletertatreyeyyn! si. Neen Gen Vad. Sige ee

Ibmberord oninayl AGHE®, | 4 caganeesceacoonsbbon ose Ay gies

Diameterrot tyampannn eee yen en Ze as

Distance between eye and tympanum... Pa

More diam a pete eee SU A AOR DO deed tee SHS) Pi

FEL ATA NE MEARNS AMUN) EES Se UNE IN er clai le oe aie

fin ol limo eee eee ene han ele oe

gore Th eared Amati es, mre AE RP IES Didnt

ORG Pie. Set eee ae aaa eet ee Sear ed amcor

Young specimens with remains of the larval tail measure as

much as 30 millim., whilst a female ready to spawn measures only

41. The eggs in the oviduct are very large and comparatively

few, the diameter of the vitellus being 2 millim.

Founded on a single specimen from Matang, Borneo, this

species has since been found on Mount Kina Balu, Borneo, and

at Gunong Tahan, Pahang, in the Malay Peninsula, at an altitude

of 5200 feet.

~ 9. MEGALOPHRYS HETEROPUS. (Plate XXV. fig. 2.)

Leptobrachium heteropus Bouleng. Ann. & Mag. N. H. (7) vi.

1900, p. 186.

Tongue nicked behind. Vomerine teeth absent. Head

moderate, as long as broad; snout truncate at the end, not

projecting beyond lower jaw, a little shorter than orbit ; canthus

1908. | BATRACHIANS OF THE GENUS MEGALOPHRYS. 423

rostralis strong; loreal region not very oblique, concave ; nostril

a little nearer end of snout than eye ; interorbital space as broad

as upper eyelid; tympanum distinct, its diameter 4 that of eye,

from which it is separated by a space equal to its own diameter.

Fingers blunt, first and second equal and about # length of third ;

no subarticular tubercles; two metacarpal tubercles, inner very

large, outer small. Toes moderately long, blunt, webbed at the

base only, the web continued as a slight fringe along each side ;

a strong dermal ridge or keel along the lower surface of the third

and fourth toes ; a small, feebly prominent, oval inner metatarsal

tubercle. Tibio-tarsal articulation reaching eye; tibia 4 length

from snout to vent ; foot shorter than tibia, but longer than head.

Skin smooth, with small tubercles on the upper eyelids. Grey

above, with darker light-edged symmetrical markings, the largest

occupying the middle of the back ; a black lumbar spot; a black

canthal and temporal streak; black spots on the sides; dark

cross-bars on the limbs; lower parts grey, speckled with black ;

a round whitish spot on each side of the breast, at the base of the

arm, another on the back of each thigh.

Measurements of the single ( 9 ) specimen :—

IRON SNOUNG HO) AVES. snoacagaupacmodnesesaase 33 millim.

Length of head (to occiput) ............... ieee:

sWalditng@ielneardien once sannccccene ee. seston: Ices

iene thvoismoubess steers cae oseseece i leew

Diameter Ob Cy jens snnce ee eres Meee: Ay gs

Ibatwarcore ortHall Wal) Ns coancondaapcsoscdehactans 3) ease

Diameter of tympanum .................. PAN

Distance between eye and tympanum... 2 _,,

(Boner linn bpenaeae cece reeci ec ceee sive ah AMA DO ae

5 LNT aN Le wR NS On Lane Mae a Sluice

eelciogol ena teen caacie ante cienrmadiewern ena. s tees AS ee

MILD Te Ayre Pp eRe Natta eet mrerincter Mier arama Gi ae

1 RYOKO U MSC cc nor men en Riley A lies Se Met er ray eS ae

A single specimen from the Larut Hills, Perak, 3500 ft.,

formerly in the Selangor Museum, is now in the British

Museum.

10. MEGALOPHRYS PELODYTOIDES.

Leptobrachium pelodytoides Bouleng. Ann. Mus. Genova, (2)

xill. 1893, p. 345, pl. xi. fig. 3.

Tongue notched behind. Vomerine teeth absent. Head

moderate, as long as broad or a little broader than long; snout

rounded, searcely projecting beyond lower jaw, a little shorter

than orbit; canthus rostralis distinct; loreal region oblique,

concave ; nostril equally distant from eye and from end of snout ;

interorbital space as broad as upper eyelid ; tympanum distinct,

its diameter 3 or 2 that of eye, from which it is separated by a

space equal or inferior to its own diameter. Fingers with slightly

swollen tips, first and second equal and about # length of third ;

424 MR. G. A, BOULENGER ON ORIENTAL [Apr. 28,

no subarticular tubercles ; two metacarpal tubercles, inner large,

outer small. Toes moderately long, with slightly swollen tips,

one-third webbed or less, the web extending as a fringe along

each side, and with a more or less distinct dermal ridge along the

lower surface ; a small, feebly prominent, oval inner metatarsal

tubercle. Tibio-tarsal articulation reaching eye, or between eye

and nostril; tibia 3, or a little less than 3, length from snout to

vent, as long as or slightly longer than foot. Upper parts with

small smooth warts, lower smooth. Olive above, with dark spots,

marblings, or symmetrical markings, which may be edged with

lighter ; upper lip with dark vertical bars ; limbs with dark cross-

bars; sometimes a round, white, dark-edged spot on back of

thighs, nearer tibia than vent; white beneath, throat of male

brown. Male with an internal subgular vocal sac.

The largest type specimen measures 37 millim. from snout to

vent. The following are the measurements of an adult male

from Burma (one of the types) and of a female from Tonkin in

the British Museum :—

IM ROL, SOKGIIG WO) WEG 25 asogncpncnacusconcece 30 42 millim.

Length of head (to occiput) ..... ey eae 10 TAs pe 9

Wrhaidthvottheadhs sat .retace jee Scene ater 10 oe ere.

IDeyavenl de Oil SINOWIG aosonaciasooossanssgcscedcbante 34 Ona pias

Diameter toiteyeren ct: see. en. neuen eee: DS Oi gine

Ibmoe rove otal WWACIOIN! Gocn4sctcas2ccqnocobeosuods 5) AN ale

Wiamleterg ol auymnpoamunn ee eneep eee ee sence 2 OE Alias

Distance between eye and tympanum ... 14 PRN

Ione dlimallopererctcs scot eae es ore comer nee 19) DO's aie

| GEE OL ae Mire, ae Steere RR Realy Sa ont 8 is

IEGirva Satria et. ete ak eee ein nerr er A6 Dillon ee

BNO TES gecer sea iia stn eee oe ene eran tte: 15 aS Peer

IOOEF wireline sic soe trake ab ROk ee mec hele eetomatan 15 Lops

This species was discovered in the Karin or Karennee Hills, east

of Toungoo, between Burma and Siam, by the late L. Fea. ‘The

types were obtained at Thao (1300-1400 m.) and in the district

of the Karin Bia-po. I refer to this species a number of speci-

mens from a collection made in the Man-Son Mountains (3000—

4000 feet) in Tonkin, on the Kwangsi frontier. These specimens

differ from the types only in having the web between the toes

less developed, being reduced to a rudiment in females, whilst

none of the male specimens can be said to have them more than

one-fourth webbed. Yet these specimens were procured during

the breeding-season, the abdomen of the female being distended

with very large ova (diameter of vitelline sphere 2 millim.). In

all other respects, however, the resemblance with the typical

M. pelodytoides is so great that I cannot adopt any other course

than to regard them as specifically identical, a course which is

fully justified by the amount of variation in this character

exemplified by J. hasseltii.

1908. | BATRACHIANS OF THE GENUS MEGALOPHRYS. 425

11. MEGALOPHRYS BOULENGERI.

Leptobrachium boulengert Bedriaga, Przewalski Exped., Amph.

p. 638, pl. i. fig. 7 (1898).

Tongue entire behind. Vomerine teeth absent. Head mode-

rate, little broader than long; snout rounded, scarcely projecting

beyond lower jaw, about as long as orbit; canthus rostralis dis-

tinct; loreal region oblique, feebly concave; nostril nearly equally

distant from end of snout and from eye; interorbital space a

little broader than upper eyelid; tympanum hidden. Fingers

obtuse, not swollen at the end, first and second equal; no distinct

subarticular tubercles; two moderately large carpal tubercles.

Toes rather short, blunt, half-webbed, the web existing as a fringe

to the tips; no distinct subarticular tubercles ; inner metatarsal

tubercles small, oval, feebly prominent. ‘Tibio-tarsal articulation

reaching the temple; tibia about + length from snout to vent ;

foot longer than head. Upper parts warty, the warts of unequal

size; a strong glandular fold from eye to shoulder; lower parts

smooth, except posterior part of belly, which is granulate. Light

greenish grey above, with an ill-defined olive-brown vertebral

band which expands into two branches between the eyes; some

of the larger warts also olive-brown; a dark canthal and temporal

streak ; lower parts yellowish.

IDTROWN SUVOWIH WO) EINE “Goncdeuadsagcseceenecace A9 millim.

emeislnok ead syne 23 see pans ee

\WIG hela Gre UGE VOIP Ais Ree DUM RM Rare asta ata Loss 3

ILperaerHly Olt SIMOTWI GE pasha cnesdeoasoosecbacoapsen Gres

Duame ber OM CVC yc scnwge vets eee ae A 13

linberorinitalayidiGloeereeeree eee canes Ay hates

Loins) Jubom| os “aba aeesterm anon aoace enemeenacaeten 29)es bas

da rec Glal broil oy pee Maar nape nants earn ran Ans tea ne Oya

GUTSY, ARR os MANS He cea ae TN coe LG tages

H EXC (OLR Weam enacts ey Te OO ORES eerie re NOT 5 oat

Dy-Chu River, Upper Yangtse Kiang, China.—Types in St.

Petersburg Museum.

12. MEGALOPHRYS HASSELTIH. (Plate XXV. fig. 3.)

Leptobrachium hasseltii Tschudi, Class. Batr. p. 81 (1838);

Giinth. Cat. Batr. Sal. p. 36 (1858); Bouleng. Cat. Batr. Hcaud.

p- 441 (1882), Zool. Rec. 1885, Rept. p. 24, Proc. Zool. Soc.

1890, p. 37, and Faun. Ind., Rept. p. 511 (1890); Isenschmid,

Mitth. Nat. Ges. Bern, 1903, p. 20; Van Kampen, Zool. Jahrb.,

Syst. xxii. 1905, p. 712.

Rana hasseltti Schleg. Handl. Dierk. 11. p. 56, pl. iv. fig. 71

(1858).

Leptobrachium montanum Fischer, Arch. f. Nat. li. 1885, p. 44.

Tongue nicked behind. Vomerine teeth absent. Head large,

about once and 7 as broad as long; snout rounded, not projecting

beyond lower jaw, about as long as orbit; canthus rostralis

426 MR. G. A. BOULENGER ON ORIENTAL [ Apr. 28,

distinct ; loreal region very oblique, concave ; nostril a little nearer

end of snout than eye; interorbital space a little broader than

upper eyelid; tympanum hidden or feebly distinct, its diameter

3 to § that of eye, from which it is separated by a space less than

its own diameter. Fingers obtuse, not swollen at the end, first

and second equal or first the longer, third nearly twice as long as

second ; subarticular tubercles, if distinct, irregular in their dis-

position ; two moderately large carpal tubercles, inner a little

larger than outer. Toes short, obtuse, like the fingers, webbed

at the base in females, + to 4 webbed in males; third toe not

reaching beyond base of antepenultimate phalanx of fourth ; sub-

articular tubercles sometimes distinct, sometimes more or less

confluent into an obtuse ridge; inner metatarsal tubercle small,

oval, feebly prominent. ‘Tuibio-tarsal articulation reaching the

shoulder; tibia 4 to 2 length from snout to vent ; foot as long as

head. Skin smooth or with small tubercles above, granular on

belly ; a glandular fold from eye to shoulder. Brown, grey, or

pale olive above, with small or large dark brown spots or marblings,

which may be irregular or form a symmetrical pattern; a more

or less distinct dark canthal and temporal streak; sides of snout

with dark vertical bars ; limbs with dark cross-bars; throat and

belly dirty white, or brown speckled with white. Male with an

internal vocal sac.

IDTROTN SICOUNS WO AYSIU sagoaanonadas sdovansodac AT 74 millim.

ens thvor headin tovocenput)aeeeree eee IO sb 7a) A. as

IWaicthvotihieadl: ten vee cee een etic rns 20 Sere

MenothvolicnoutieGnieshepire eon te 6 10

Diameter Ol CVE hee oe sehen eee eee ) On

JbnerRorelomal Wahid) soeacgaossdsaoeoodoosusuces 6 OR Ree.

IDMERITNE Ce Gt wT OKADA A nonacoaenoosenensec

Distance between eye and tympanum ... aie

(Hore slinma bees ese enna betelentt erie ee 33 50a

TELE NOVGLs wake abodes tolde SB CUSEE Ce Lest cee Enna ic 11 LG 4 fae

2 LIWE\C ba Misa Ohi seen dees eeeasere Nercmmeoeuncreera doa 53 To

MIEN AED, Cantera colnet asien teen sem at oot 16 24 Cy,

INO O.bs oe ok. Hiaabbss bla oe oege ees We eee eae 3 15 23

Burma, Malay Peninsula, Sumatra, Borneo, Palawan, Java.

The larva has been first noticed by me, from specimens from

Sumatra and Perak, and others have since been obtained in

Selangor by Mr. Butler, and in Perak by Dr. Hanitsch. These

tadpoles are of the same type as the typical Pelobatids of Europe,

but remarkable in being marked all over with numerous deep black

dots or round spots. No observations have been made on the

breeding-habits, but it is probable that the eggs are laid in the

water, being similar to those of Pelobates. Eggs from the oviducts

of a female 65 millim. long measure 2 millim. in diameter.

Mr. A. L. Butler observes (Journ. Bombay N. H. Soe. xv. 1904,

p- 397) that the larval period of existence is very prolonged, and

1908. | BATRACHIANS OF THE GENUS MEGALOPHRYS. 427

that the tail does not disappear until the size of about 40 millim.

(from snout to vent) is attained.

13. MEGALOPHRYS CARINENSIS.

Leptobrachium carinense Bouleng. Ann. Mus. Genova, (2) vii.

1889, p. 748, and Faun. Ind., Rept. p. 511 (1890); W. Sclater,

Proc. Zool. Soc. 1892, p. 8347; Bouleng. Ann. Mus. Genova, (2)

xill. 1893, p. 345, pl. xii.

Tongue very indistinctly nicked behind. Vomerine teeth

absent or in two widely separated small groups between the large

choane *. Head very large and externally depressed, once and

7 to twice as broad as long; snout rounded, as long as eye or

orbit, not projecting beyond lower jaw ; canthus rostralis distinct ;

loreal region very oblique, slightly concave ; nostril equally dis-

tant from eye and from end of snout; interorbital space nearly

twice as broad as upper eyelid; tympanum completely hidden.

Fingers short, blunt or with slightly swollen tips, first and second

equal, about } length of third; no subarticular tubercles ; meta-

carpal tubercles indistinct. ‘Toes short, blunt or with slightly

swollen tips, 4 to 3 webbed, the web extending as a slight fringe

on each side; no subarticular tubercles; a very large, oval, flat

inner metatarsal tubercle. Tibio-tarsal articulation reaching

axil or shoulder in female, commissure of jaws in males; tibia

4+ to 2 length from snout to vent; foot longer than head. Skin

with bony deposits on head and anterior part of body; a strong

transverse fold defines the head behind; a strong glandular

fold from eye to shoulder; upper eyelid with conical tubercles,

two to four of which may be enlarged and form very short

“horns”; body with small scattered smooth warts, which are

Jarger and may be conical on the sides ; an oblique glandular fold,

parallel with the supratemporal fold, on each side of back, as far

as sacral region; more or less distinct oblique glandular folds

across the limbs; throat finely granulate, belly nearly smooth.

Upper surface of head and sides of body yellowish: back between

the dermal ridges purplish grey (brown in spirit); eyes, dermal

ridges, and larger tubercles bordered with black; some of the

lateral tubercles pure white; limbs purplish grey; gular region

brown or purplish black ; belly brown or whitish. Male with an

internal vocal sac.

oie Qe

IDAOUN SAO OG WOVE. capacsesvesedossndeoboue 123 = 150 millim.

Length of heal (toroceipuit) a-. aes Be Bie Res

IWidithvot ead een cae De comet 58 (0) takes

Pemethvott Mout ater. tect. ack 12 Italy

DiMebeLIOMeVe = tretmen eee eter tran 10 oe

* The palate was toothless in the specimen originally described. Vomerine

teeth were subsequently found, by Mr. W. Sclater and by myself, in other

specimens. That the character is not of specific importance in this case is further

evidenced by the fact that a large female specimen from Fea’s collection has a group

of vomerine teeth on the left side and no trace of them on the right side.

428 MR. G. A. BOULENGER ON ORIENTAL | Apr. 28,

Gia OF

itnterorbitalewdtihpereneeeeeneee eee eemeer 17 22 muillim.

Hore dimiby, ic: ee. seats va cope ote ees ante 66 (24

gS eho ts Pepe oa are irene ns WA AOR 6 or Sb cect 28 Se

MDa: Litianllopys Seach 1. ee ee, ee Soe L3i OOM

LID Si ee Aart ORR IN een aba Se eens US 45 Sle yaa

RO Ot Aceh eas eR as See ts cae oe 44 53

Several specimens were obtained by the late L. Fea on the

western slope of the Karin or Karennee Hills, east of Toungoo,

Burma, at an altitude of 2500 feet or upwards. The species has

been recorded from Mergui, Tenasserim, by W. L. Sclater.

According to Fea, the males show themselves very irritable when

disturbed, opening their wide gape, ready to bite at the slightest

provocation, emitting at the same time a shrill cry. The female

is of more gentle disposition. The strong jaws of this frog enable

it to overpower small mammals, and a large female contained, in

addition to insects, a small squirrel.

14. MEGALOPHRYS FE.

Megalophrys fee Bouleng. Ann. Mus. Genova, (2) iv. 1887,

p. 512, and v. 1887, p. 423, pl. v.

Leptobrachium fee Bouleng. op. cit. vii. 1889, p. 750, and Faun.

Ind., Rept. p. 512 (1890).

Tongue feebly nicked behind. Vomerine teeth usually present,

in two widely separated small groups just behind level of choanz.

Head very large and extremely depressed, once and # to twice as

broad aslong; snout rounded, hardly as long as eye, not projecting

beyond lower jaw; canthus rostralis distinct ; loreal region very

oblique, slightly concave ; nostril equally distant from eye and from

end of snout; interorbital space slightly concave, twice as broad

as upper eyelid; tympanum completely hidden. Fingers short,

blunt, first and second equal, # length of third ; no subarticular

tubercles ; metacarpal tubercles indistinct. Toes short, blunt,

nearly free or with a mere rudiment of web at the base; no sub-

articular tubercles; a very large, oval, flat inner metatarsal

tubercle. Tibio-tarsal articulation reaching axil, shoulder, or

commissure of jaws; tibia + to # length from snout to vent; foot

longer than head. Skin with bony deposits on head and anterior

part of body; a strong transverse fold defines the head behind ;

a strong glandular fold from eye to shoulder; upper eyelid with

tubercles, one of which is larger and conical and may be developed

into a rather long horn-like appendage in the adult. Body and

limbs above with scattered small smooth warts; no longitudinal

glandular folds; lower parts smooth. Olive-brown above; a

T- or Y-shaped darker marking on the head, the transverse

branch between the eyes, often dividing the head into a lighter

anterior and a darker posterior portion ; a dark temporal band,

the lip below it yellowish ; lips, eyes, and some of the larger

warts on the body edged with blackish ; some of the warts whitish ;

1908. | BATRACHIANS OF THE GENUS MEGALOPHRYS. 429

lower parts dirty white to dark brown; throat sometimes spotted

with dark brown. Male with an internal vocal sac.

The type specimen, a female, measures 110 millim. from snout

to vent.

Measurements of specimens in the British Museum :—

Cir ;

INAGLTN AOU WO WWW gooodenvoosononcooonscne 82 106 millim.

Length of head (to occiput) ............... 25 lam

IWiaelthiote lieaclinpecrerc cache epee pats. 46 Mis | oo

ID@ravedslo Oi SSMOWNG oJocvocssdecooroosce hes Sialereel 8 Or eee

Diameteriotreyer sje. seve. v.cerctccest ee 9 ‘LOWS, 2

Ibaynercor oeayl ARAIClG cossonnnoconooncooosebanend 13 GRA NEe.

Hore: Tmmilosmenanee pe yeete. Manteca. cee, 45 DOM.

UNsTieaYa CU yee ener art Netes a eg ctl hl MUSSER Bs 24 2Ore) ee

HFetirnavclialiraat lope siea es eae eae AnD She aN he eh M1 Ne USO),

il itl Terenas Meyeine oe eerie te Slo oR Nea Hs eal ae, 3 SO eee

JOO eae ke ences matali ecocre ue ae ce aR Ce cae D1 4]

M. few was discovered by the late L. Fea in the Kakhyen Hills,

east of Bhamo, Upper Burma. But the species extends further

east, as the British Museum possesses a specimen obtained by

Hr. Fruhstorfer in the Man-Son Mountains, Tonkin (altitude

3000-4000 feet) ; this specimen, in which the supraocular “horn”

is very feebly developed and vomerine teeth are absent, I had

first referred to LZ. carinense (Ann. & Mag. N. H. (7) xii. 1903,

p. 186).

15. MEGALOPHRYS BALUENSIS.

Leptobrachium baluense Bouleng. Ann. & Mag. N. H. (7) iv.

1899, p. 453, and Journ. Str. Br. As. Soc. xxxiv. 1900, pl. ii.

figs 1

Tongue entire. | Vomerine teeth in two widely separated small

groups just behind level of choanz. Head moderately large, much

depressed, nearly twice as broad as long; skin adherent to the

rugose skull; snout rounded, much shorter than orbit, not pro-

jecting beyond lower jaw; canthus rostralis strong; loreal region

vertical, concave; nostril equally distant from eye and from end

of snout ; interorbital space slightly concave, nearly twice as broad

as upper eyelid; tympanum feebly distinct, 2 diameter of eye,

about 4 its distance from eye. Fingers rather elongate, slender,

blunt, first a little longer than second, which is 2 length of third ;

no subarticular tubercles; no metacarpal tubercles. Toes mode-

rately long, slender, blunt, with a mere rudiment of web; no

subarticular tubercles, but a feeble median ridge or keel under the

toes; a very indistinct oval inner metatarsal tubercle. Tibio-

tarsal articulation reaching shoulder; tibia 2 length from snout

to vent ; foot much longer than head. Skin perfectly smooth ; a

very small conical tubercle near the border of the upper eyelid,

above the pupil. Back and upper surface of snout dark grey;

430

posterior half of upper surface and sides of head blackish brown ;

a curved light streak, the concavity turned forwards, across upper

eyelids and interorbital region, followed by a Y-shaped blackish

marking ; two light spots on upper lip, below eye ; large blackish-

brown partly confluent spots on the back; sides dark brown,

light-edged above; limbs dark brown, with rather indistinct

ON A GENUS OF ORIENTAL BATRACHIANS.

darker cross-bars; throat brown, belly brownish white.

From

SAOUNG WO) WEIN scencsooacacccocauceaces

Length of head (to occiput)...............

Wadthvotvhieddy je. 15 eee Pec eer tere ecr se

IDES OE STON “egandnnodaceacedsAonaneso:

DiamMme borhokte yeu as Whee cee eee ken ee

lnxtexorioutallgwidtl ences: sae eee

Diameter of tympanum ..................

Distance between eye and tympanum...

(Horevliriibs jebte) aeaeeeice, chet. eee ace as

Tibia

Known from a single female specimen, full of ripe eggs 3 millim.

in diameter, discovered by Dr. Hanitsch on Mount Kina Balu,

ee we ewww ee we wee wee een were te eee eee ene cce

Borneo, at an altitude of 4200 feet.

EXPLANATION OF THE PLATES.

PratEe XXII.

65 millim.

a. Megalophrys nasuta, p. 411. Brunei, Borneo.

}. Side view of head of a.

c. Head of specimen from Penang.

Pravre XXIII.

Megalophrys major, p. 416. Type.

With side view of head.

PraTE XXIV.

Megalophrys robusta, p. 418. Type.

With side view of head.

PLATE XXV.

Fig. 1. Wegalophrys gracilis, p. 421. Type.

la. Side view of head.

10. Lower view of hand, <2.

Ie. 55 5 foot, X 2.

Fig. 2. Megalophrys heteropus, p. 422. Type.

2a. Side view of head.

Fig. 3. Megalophrys hasseltii, p. 425. Java.

3a. Side view of head.

No. 54.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON®

March 3rd, 1908.

G. A. BoutencEr, Esq., F.R.S., Vice-President, in the Chair.

The Secrurary read a letter from Mr. Roland Trimen, F.R.S.,

F.L.S., F.Z.S., in which he described the presence of a cuticular

claw-like growth, about seven lines in length, on the tip of the tail

of a domestic cat in his possession. The Secretary exhibited, for

comparison, the tail of a young lion on which such a structure is

well known to occur.

The Secrerary exhibited some skins of the Coypu (Myocastor

coypu) lent to him for the purpose by Mr. C. Hawkins, and called

attention to the dorso-lateral position of the nipples of the mam-

mary glands. The peculiar position of these organs in the Coypu

and some of its allies had long ago been discussed by Sir Richard

Owen, but apparently was not known to writers of recent text-

books.

Mr, F. E. Brepparp, M.A., F.R.S., Prosector to the Society,

exhibited a preparation of the colon and rectum of the Badger

(Meles meles), and directed attention to the unusual size of the

Payer’s patches.

Dr. P. Cuaumers Mircuent, M.A., F.R.S., Secretary to the

Society, read a paper entitled “On a Young Female Kordofan

Giraffe,” and illustrated his remarks by lantern-slides. He com-

pared the coloration of this specimen, born at the Gardens, with

that of its parents and with that of a young female Giraffe from

Nigeria, now living in the Society’s Collection, and stated that

* This Abstract is published by the Society at 3 Hanover Square, London,

W., on the Tuesday following the date of Meeting to which it refers. It will

be issued, free of extra charge, to all Fellows who subscribe to the Publications,

along with the ‘ Proceedings’; but it may be obtained on the day of publieation

at the price of Sixpence, or, if desired, sent post-free for the sum of Siz

Shillings per annum, payable in advance.

the evidence to be derived from the study of this specimen

strengthened the case for the distinctness of the Giraffes from

Kordofan and Nigeria.

Mr. F. E. Bepparp, M.A., F.R.S., Prosector to the Society,

gave an account of his communication entitled ‘“* A Comparison of

the Neotropical Species of Corallus, C. cookii with C. madagas-

carvensis; and on some Points in the Anatomy of Corallus

caninus.”

Mr. R. I. Pococs, F.L.S., Superintendent of the Gardens, gave

a description of a new species of Monkey of the genus Cercopi-

thecus, which differs from C. neglectus principally in the absence

of the black band across the head, in the reddish tinge of the

hairs beneath the callosities, and in the similarity in colouring

between the tail and the body. He proposed to name this new

Monkey Cercopithecus ezre.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, the 17th March, 1908, at half-past Hight

o'clock P.m., when the following communications will be made :—

1. Watrer A. Kipp, M.D., M.R.C.S., F.Z.S.—Some Obser-

vations on the Effects of Pressure upon the Direction of Hair in

Mammals.

2. O. Tuomas, F.R.S., F.Z.S., and R. C. Wroveutron, F.Z.S.—

The Rudd Exploration of S. Africa.—IX. List of Mammals

obtained by Mr. Grant on the Gorongoza Mountains, Portuguese

8.E. Africa.

3. R. I. Pocock, F.L.8., Superintendent of the Gardens.—

Notes upon some Species and Geographical Races of Serows

(Capricornis) and Gorals (Nemorhedus) based upon Specimens

exhibited in the Society’s Gardens.

The following communication has been received :—

Dr. Kxup ANpDERsEN.—A Monograph of the Chiropteran

Genera Uroderma, Enchisthenes, and Artibeus.

Communications intended for the Scientific Meetings of the

ZOOLOGICAL Society oF Lonvon should be addressed to

P. CHALMERS MITCHELL,

Secretary.

3 Hanover Square, Lonpox, W.

March 10th, 1908

No. 55.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

March 17th, 1908.

‘Dr. Henry Woopwarp, I'.R.S., Vice-President,

in the Chair.

The Secrerary read a Report on the additions that had been

made to the Society's Menagerie during the month of February

1908.

Mr. G. A. Bounencer, F.R.S8., V.P.Z.8., exhibited a remarkably

malformed Plaice (Plewronectes platessa) from the London market,

which had been given him by the Secretary of the Fishmongers’

Company. One of the most striking peculiarities of the specimen

was that the colouring and scaling of the two sides of the tail

were exactly alike and similar to the normal condition of the

coloured side. Mr. Boulenger suggested that the tail might be

a regeneration in which an ancestral condition had reappeared.

Dr. Waurer Kipp, F.Z.S., F.R.S.E., read a paper on “Some

Observations on the Effects of Pressure upon the Direction of

Hair in Mammals.” This paper was a sequel to other com-

munications on the subject of the direction of hair, and consisted

chiefly of the observed effects of the pressure of harness on

certain regions of the coats of domestic horses. This pressure

was shown to produce reversed areas of hair, and it was held

that these results supported the view put forward in other papers

that changes in the arrangement of hair are due to mechanical

* This Abstract is published by the Society at 3 Hanover Square, London,

W., on the Tuesday following the date of Meeting to which it refers. It will

be issued, free of extra charge, to all Fellows who subscribe to the Publications,

along with the ‘ Proceedings’; but it may be obtained on the day of publication

at the price of Sixpence, or, if desired, sent post-free for the sum of Sie

Shillings per annum, payable in advance.

eauses. Fifty-three cases were brought together and eight

different regions of the coats of the Horse were shown in which

the effects a pressure were found.

A paper was read by Messrs. OLDFIELD Tuomas, F.RB.S., F.Z.8.,

and R. C. Wroveuton, F.Z.8., on “ Mammals obtained by

Mr. C. H. B. Grant in the Gorongoza Mountains, Portuguese

S.E. Africa,” being the ninth of the series of papers on the

Mammals of the Rudd Exploration of South Africa. 150 speci-

mens were dealt with, belonging to 31 species and subspecies, of

which three were described as new.

Mr. R. I. Pocock, F.L.S., the Superintendent of the Gardens

read a paper entitl ed “ Notes upon some Species and Geogra-

phical Races of Serows (Capr icornis) and Gorals (Nemorhedus).

based upon Specimens exhibited in the Society’s Gardens,” and

pointed out that the “grey” Goral of the Himalayas was origi-

nally described by Hardwicke as Antilope goral, and that the

“brown” Goral, to which the specific title goral has been applied

in recent literature, required a new name. He proposed to call

it Vemorhedus hodgsont. Concerning the genus Capricornis, he

stated that although only one form had ‘been hitherto distin-

guished from the “Himalayas, the available material pointed to

the existence of at least four subspecies in that mountain-range.

The follows ing were characterized as new :—Capricornis swma-

traensis oman, with a rufous-brown head, from Kashmir ;

C. s. rodoni, with a white interramal area and throat-patch,

white underside and lower legs, and a long thick coat with under-

fur, from Chamba; C. s. jamrachi, with short black coat, blackish-

brown underside, and partially rufous lower legs, from Darjiling.

A black form from Selangor, with a scarcely appreciable quantity

of red in the main, was also described as new under the name

C. s. robinson.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, the 7th April, 1908, at half-past Hight

o'clock p.m., when the following communications will be

made ;—

1. Dr. Kyup Anprrsen.—A Monograph of the Chiropteran

Genera Uroderma, Huchisthenes, and Artibeus.

2. Sir Ray Lanxester, K.C.B., F.R.S., F.Z.S.—On certain

Points in the Structure of the Cervical Vertebree of the Okapi

and the Giraffe.

8. H. BR. Hoge, M.A., F.LS., F.Z.8.>Some Austra lian

Spiders.

The following communication has been received :—

Mrs. E. W. Sexton.—On the Amphiped Genus Zrischizostoma.

Communications intended for the Scientific Meetings of the

FooLOGICAL Socrery oF LONDON should be addressed to

P. CHALMERS MITCHELL,

Secretary.

3 Hanover Square, Lonpon, W.

March 24th, 1908.

No. 56.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

April 7th, 1908.

Dr. Henry WoopwarD, F.R.S., Vice-President,

in the Chair.

On behalf of Mr. Tuomas Coprinetoy, Dr. A. Smita Woop-

WARD, F.R.S., F.Z.S., exhibited a collection of 168 stones, weighing

altogether 7 Ibs. 13 oz., taken from the stomach of an Klephant

shot by Mr. H. Thornicroft in Northern Rhodesia. The animal

was a large male, with tusks weighing 45 lbs. each. The stones

showed no signs of attrition.

Dr. OC. W. Anprews, F.R.S., F.Z.S., exhibited a restored model

of the skull and mandible of Prozeuglodon atrox And. ‘This

animal is one of the links uniting the true Zeuglodonts with the

land Creodonts. It is found in the Middle Eocene of Hgypt,

where also the earlier type, Protocetws, was discovered by Fraas

at a somewhat lower horizon. The model was constructed by

Mr. F. O. Barlow, for the British Museum of Natural History.

The Szcrerary exhibited a photograph of two young living

examples of the Forest-Pig of Central Africa (Hylocherus

meinertzhagent Thomas). The photograph had been taken by

Mr. T. J. Morson, of Limoru, who had obtained the pigs in the

Limoru and escarpment forest about 353 miles from Mombasa, at

an elevation of between seven and eight thousand feet.

The Szcrerary stated that he had been informed by the High

* This Abstract is published by the Society at 3 Hanover Square, London,

W., on the Tuesday following the date of Meeting to which it refers. It will

be issued, free of extra charge, to all Fellows who subscribe to the Publications,

along with the ‘ Proceedings’; but it may be obtained on the day of publication

at the price of Sixpence, or, if desired, sent post-free for the sum of Siz

Shillings per annum, payable in advance.

Commissioner for New Zealand, that the Chamois presented by

the Emperor of Austria to New Zealand, and which had been

successfully taken to New Zealand by one of the Society’s staff

in the beginning of 1907, had been seen in the locality in which

they were liberated, one of the females being accompanied by a

strong, healthy-looking kid.

The SEcRETARY presented, on behalf of Dr. Knup ANDERSEN, a

communication entitled ‘A Monograph of the Chiropteran Genera

Uroderma, Hnchisthenes, and Artibews.” The work was based on

an examination of the material in the British and United States

National Museums, and contained a discussion of the homologies

of the teeth and molar cusps in Stenodermatous Bats, a full

description of the genera mentioned in the title, their species and

subspecies, with a discussion of their probable inter-relations, and

finally remarks on the bearing of the present geographical distri-

bution of the species and sub-species on a former connection of

the West Indian Islands with continental America.

The SECRETARY read a communication from Sir H. Ray Lan-

KEsTER, K.C.B., F.R.S., F.Z.8., entitled “Qn certain Points in

the Structure of the Cervical Vertebre of the Okapi and Giraffe.”

The paper dealt chiefly with the posterior cervical and anterior

dorsal vertebre, the Author concluding that where the Okapi

differed in these respects from the Girafie, it resembled other and

particularly bovine’ Artiodactyles. It also included a discussion

of the zygapophysial articulations of the cervical and dorsal

vertebre in the Giraffe, Okapi, and some other May “als.

Mr. H. R. Hoee, M.A., ¥.Z.S., gave an account of a paper on

“Some Australian Spiders,” in which he gave further notes on

the type species of the genus Jssulena, hitherto known only by

two specimens, a synopsis of the New Zealand genus Hexathele,

with description of two new species, and a description of two new

species of Dolomodes Latyeille, from Pitt Island of the Chatham

Group, showing affinities with the only two species recorded from

New Zealand.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, the 28th April, 1908, at halt-past Hight

o'clock P.M., when the following communications will be

made :—

1. Mrs. E. W. Sexton.—On the Amphipod Genus Z'rischizo-

stoma.

2. C. L. Boutencer, B.A., F.4.8.—On the Breeding-Habits of

a Cichlid Fish (Tilapia milotrea).

Wii

3. C. Tarn Ruaan, M.A., F.Z.S.—A Revision of the Sharks of

the Family Orectolobide.

4. G. A. Boutenesr, F.R.S., V.P.Z.S.—A Revision of the

Oriental Pelobatid Batrachians (Genus Megalophrys).

The following communications have been received :—

1. C. F. Jennxin.—The C. Crossland Collection of Calearea from

Zanzibar and Wasin (British Hast Africa),

2. R. HE. Turner, F.Z.8.—Notes on the Australian Fossorial

Wasps of the Family Sphegide, with Descriptions of new Species.

Communications intended for the Scientific Meetings of the

ZOOLOGICAL Society or Lonpon should be addressed to

P. CHALMERS MITCHELL,

Secretary.

3 HANovER Square, Lonpon, W.

April 14th, 1908.

ContTENTS (continued).

April 7, 1908 (continwed).

Page

e Secretary. Exhibition of a photograph of two young living examples of the Forest-

Pig of Central Africa (Hylocherus meinertzhagent Thomas) ..++eesseessseeeerrece 203

e Secretary. Notice of the successful acclimatization of Chamois in New Zealand .... 204

A Monograph of the Chiropteran Genera Uvoderma, Enchisthenes, and Artibeus. By

GHD PANDERSEN eictuiete islets! sectia rails cidleseie nitiate orn areia'e'« sy veie/tlm) cievniei ove ei at'elaia sy ait/«im/a'elersiaie 204

On certain Points in the Structure of the Cervical Vertebre of the Okapi and the

Giraffe. By Sir Ray Lannusrmr, K.C.B., F.B.S., F.Z.S. 0.0.2.6. eee cece eee eee es 320

Some Australasian Spiders. By H. R. Hoae, M.A., BOA ee Recs Ainae t a ee ene BO

April 28, 1908.

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

Waele l O08) oie ea siece sce eee ealee «i Grd mio 5) SRI es er a ete Sbsienee po 345

hy. C. Tate Regan, M.A., F.Z.S. Exhibition of, and remarks upon, an Australian Cat-

fish (Cnidoglanis megastoma Richards.) 1+ ..++++.. cece cece cee cee e ence eee e eee 345

Mr. Walter Burton, F.Z.8. Exhibition of some melanistic and black Leopard skins ...... 346

Le Beare smite pI cy orem File mhae sr Gt ccshenc ey Cid Hens colada: OOint Olane GE metn minnie 346

I, :. Revision of the Sharks ofthe Family Orectolobide. By C. Tarr Ruean, M.A., F.Z.8.

® (Plates XU-XIID) ..... Re Amen Nash rhe rene eae etntage SON itunes Saha Werte ak 347

. A Note on the Occurrence of a Species of Phreatothrix (Vejdovsky) in England, and on

some Points in its Structure. By Frank HE. Brepparp, M.A., E.R.S., F.Z.S., Korresp.

Mitgl. d. K. Bohm. Ges. Wiss., &e. eee ce eee cece ee eee teen eect teen e ees 365

8. On the Amphipod Genus Trischizostoma. By E. W. Sexton. (Plates XDV. = OX ee ond,

£ On certain Errors with reference to George the Fourth’s Giraffe. By Henry Scuurren,

PEER ea es Ree eterna NTs Meare ine taker slls, ¢ a arviotc saan ots, ciauacacees wePeieroae ea Siekoner et aan 403

. On the Broeding-Habits of a Cichlid Fish (Tilapia nilotica), By Cuaruzs L, BOULENGEE, ,

a Revision of the Oriental Pelobatid Batrachians (Genus Megalophrys). By G. A.

Bovutencrr, F.R.S., V.P.Z.8. (Plates XXIT-XXV.) .... eee ee eee pois gen es 407

EL ist of Texti-figures ee ce ce eee cece eee tence ere eet teen teen eet nett nees Xvil

List ak ea Grade Denins Sebunrs 6 sopoecmncboocbooaD ourocGurcopoouoocT eh eae ee XX

Ende Bey eta oy SR e eM Fat Oph UAL) reece, «Slat auc 6 vluosok OR MU aaTS EST OU aint eco nub hecet at aheeemey ar See pel cum

LIST OF PLATES,

1908, pp. 127-430.

€

Plate ’ Pag

X. ‘New Species of Cercopithectis .... ...cecccecsscsccscece 158

XI. 1. Chiloscyllium griseum. 2. Orectolobus ornatus ......

XII. 1. Chiloscyllium plagiosum. 2. Orectolobus tentaculatus } 347

XIII. 1. Chiloscyllimm griseum. D2 CoOL aornsaluiete's epee

XIV. ~

xy. TYASCRUZOSLOMM MLCLENSE! Ie sie a vic oye ayea en a n)e/n1~ seoin|aie ie) eee

XVI. ,

XVII. 1-12. Trischizostoma niceense. 13. T. raschii, ........

DVL. LoaschicostomaTQschit.xe.nrc nsession ehia toes Se eRe 370

XIX. 1. Trischizostoma niceense. ZU PA SCHUU \eretcetreteraere |

MEKG | Wrischizostom@maschit, «2/0 «002s te sole laste = aces) oie tclenees

XXI. 1-13; 15-18. Trischizostoma raschit. 14. T. niceense .... )

AXIE, Megalophrys nasuta .. csv... . ceeess cece REI Macc bc )

XXIIE. Megalophrys major ..-... 0... ve oh wt oe es sec ee emer ‘ 407

ORD Vin Meg nloplrySPGUUSHG. . oe cs 1% ce cielo cate clen = ania ee tanene |

XXV. 1. Megalophrys gracilis. 2. M. heteropus. 3, M. hasseitii .)

NOTICE.

The ‘ Proceedings’ for the year are issued in four parts, paged consecutively,

so that the complete reference is now P. Z. 8. 1908, p... . The Distribution

is as follows:—

Papers read in January and February, issued in June.

_ » March and April, » », August.

i » May and June, 1» 3) October.

A » November and December,,, ,, April.

‘ Proceedings,’ 1908, pp. 1-126, were published on July 8rd, 1908.

The Abstracts of the papers read at the Scientific Meetings in |

March and April are contained in this Part.

4p 9700