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PROCEEDINGS
OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS LLE 2
OF THE
ZVOLOGICAL SOCIETY
OF LONDON.
1906, pp. 1-462.
(JANUARY—APRIL.)
PRINTED FOR THE SOCIETY, AND SOLD AT THEIR HOUSE IN HANOVER-SQUARE, LONDON:
MESSRS. LONGMANS, GREEN, AND CoO, PATERNOSTER ROW.
eel
4
| gas bes te B
OF THE
QV! £3
COUNCIE AND “OFFICERS
OF THE
ZOOLOGICAL
SOCIETY
OF LONDON.
1906,
COUNCIL.
His Grace THe DuKe or Beprorp, K.G., President.
Srr ALEXANDER BatirD, Br.
JoHN Rose Braprorp, Esq., M.D., “D.8c., > F.RS., Vice- President.
Major The Hon. Wruuiam E. CAVENDISH.
F, Dawtrey Drewirtr, Esq., | M.A., M.D.
CHARLES DrumMmonn, EsqQ., Treasurer.
Sir Epwarp DuraAnp, Br., C.B.
FREDERICK GILLETT, Esq., Vice- President.
W. R. Ociivie-Grant, Esq.
Major Tur Marquis oF Hamitron, M.P.
| JosepH Jackson Lister, EsqQ.,
M.A, ERS. Sir Epmunb Gives Loper, Br., Vice-President.
_ E.G. B. Mrapre-Watpo, Esq.
P. Cuatmers MircuHetty, Ksq., M.A., D.Sc., F.R.S., Secretary.
E. Lort Puiturrs, Esa.
Howarp SAunpers, Esq., Vice- President.
Davin Seru-Smiru, Esq.
| OLDFIELD Tuomas, Esq., F.R.S.
CHARLES 8. Tomes, Esq., M.A., F.R.S., Vice-President.
Aveustus F, WIENER, Esq.
Henry Woopwarp, Esq.,LL.D., F.R.S., Vice-President.
PRINCIPAL OFFICERS.
P, CHaLMErs MircueEtt, Esq., M.A.,D.8c., F.R.S., Secretary. FrANK E. BeppArp, Esq., M.A., F.R.S., Prosector.
R. I. Pococr, Esq., F.LS., Superintendent of the Gardens. CHARLES GABRIEL SELIGMANN, Esq., M.R.C.S., L.R.C.P.,
Pathologist.
Mer. F. H. Wavernouse, Librarian.
Mr. Joun Barrow, Accountant.
Mr. W. H. Coxz, Chief Clerk.
Mr. Georce ArtuHur Douptepay, Clerk of Publications.
Mr.
Gardens.
Artuur THOMSON, Assistant Superintendent of the
| ate
fy [i
A
mH Af
A, say /
LIST OF CONTENTS.
January 16, 1906.
The Secretary. Report on the Additions to the Society’s Menagerie during the month of December 1905 .........
Prof. E. A. Minchin, F.Z.8. © Exhibition of a living specimen of a Lemur (Galago) from Uganda ............ cece
Dr. F. G. D. Drewitt, F.Z.S. Exhibition of a white variety Ge TN Olorananoray IWIN: Sor seeeuabatodneson sbecoaonsarueceocpe dud
Mr. Oldfield Thomas, F.R.S. Exhibition of, and remarks upon, the skull of a new Forest-Pig ......-++-1seeeeseee es
1. On Mammals from South Johore and Singapore collected by Mr. C. B. Kloss. By J. Lewis Bonuore, M.A.,,
B.L.S., F.Z.S. (Plate LD.) 2... eee cee eee ee cee ee teen eee nees
2. Contributions to the Anatomy of the Ophidia. By Frank EB. Bepparp, M.A., F.R.S., Prosector to the Society
3 On the Minute Structure of the Teeth of Creodonts, with especial reference to their suggested resemblance to Marsupials, By Cuarues 8. Tomes, M.A., F.R.S., Vice- [DISS WAAS | senddablbrbausadcdocrane sogbeeec psdnapameaucosbaneneaesn
Page 1
bo
bo
bo
iv Page 4, Synopsis of the Toads of the Genus Vectophryne B. & P., with special Remarks on some known Species and Description of a new Species from German East Africa. By Dr. Jean Roux, Curator in the Basle Museum of Natural History. (Plate IT.) ............:.eeseeeeeeeeeneeeees 58
5. On some Bones of the Lynx from Cales Dale, Derbyshire. By W. Storrs Fox, M.A., F.Z.8. .......0.0-seeeesnereteonee 65
February 6, 1906.
Mr. Frederick Gillett, F.Z.S. Exhibition of a case of mounted cubs ot the “Timber Wolfie ce tackess or cdaneaceer eee eee 73
Dr. C. W. Andrews, F.Z.S. Exhibition of some models of the skulls and mandibles of Meritheriwm and Paleo- PUDSEOMONG © sides Bae CaP TOT ONS Se RT ee 73
Dr. Walter Kidd, F.Z.8. Exhibition of a series of lantern- slides of sections of skin from the palmar and plantar surfaces of Mammals:amnd) Birdsis.cucac<nn- eo aaeccwdesoeeenen 73
1. Notes on the Histology and Physiology of the Placenta in Ungulata. By J. W. Jenxryson, M.A., D.Sc., Assistant to the Linacre Professor of Comparative Anatomy, Oxford. (Plate III.) 73
i a a)
2. Note on the Cavies of the Genus Dolichotis, and on Living Specimens of D. salinicola. By Sir Epmunp Loprr,
Bt; EZ. CP Leelee) neces dec eet anidcc SduGeioe ganenen: ae 96 3. Description of a new Fly of the Family Zabanide. By
GERTRUDE RICARDO%.3.stce tes sinme deeepiies « seeheiees. See eee 97 4, On Trichorliza, a new Hydroid Genus. By E.S. Russext.
(RUBE Win) evn as cian eos oe: sedaneintewaseweegmeehai anaes eee eee 99
1
. A List of the Mammals obtained by Messrs. R. B. Woosnam and R. E. Dent in Bechuanaland. By Haroip NCHWANN, 2:8. (Plate: Vil.) i. ti. P ees eee eee 101
W
6. On a Central African Ratel and Water-Chevrotain. By ie dbyaniaicdtaisy A(eTeNes TTS) cadendoposos checcepaumerreaano:
7. The Articulation of the Vertebrate Jaw. By H. Gzorcr IAS PUR RETR eal yen ees 2 Mice ly Rs kd Sal Sb ale klk i,
February 20, 1906.
The Secretary. Report on the Additions to the Society’s Menagerie during the month of January 1906
Sir Reginald Talbot, K.C.B. Extract from a letter from, concerning the supposed breeding of a Mule...............
My. R. I. Pocock, F.Z.8. Exhibition of a photograph of a
Ring-tailed Lemur carrying its young on its back ......
Dr. A. Smith Woodward, F.R.S., F.Z.S. Exhibition of a drawing of the skeleton of Rhynchosaurus articeps ......
1. On Breeding Experiments with Lepidoptera. By L. Don- casTER, M.A., F.Z.S., Mackinnon Student of the Royal Society, and the Rev. G. H. Raynor, M.A., F.E.S. (ETN reo Yi GEIL) Se ered Seren eae Ante einen Ae
2. Contributions to the Osteology of Birds.—Part VIII. The ‘““'Tracheophone ” Passeres; with Remarks on Families allied thereto. By W. P. Pycrart, F.Z.8., A.LS., &e..
3. The Rudd Exploration of South Africa.—IV. List of Mammals obtained by Mr. Grant at Knysna. By Oxp- FIELD THomAsS, F.R.S., and HAarotp ScHWANN, F.Z.S. ...
4, Notes on the Living Specimens of the Australian Lung- fish, Ceratodus forstert, in the Zoological Society’s Collection, By BasHrorp Dean, Ph.D, (Plate IX.)...
March 6, 1906. Mr. G. A. Boulenger, F.R.S8., V.P.Z.S. Exhibition of a
specimen of, and remarks upon, a giant Frog from Cameroon
Se eee eee sees eee rere saeeaeeeeseaseesessereeroeserseosseoes
114
168
vi Tage Mr. R. T. Giinther. Exhibition of, and remarks upon, Meduse from Lake Tanganyika o......cccsccsceseetencnceees ies)
Mr. G. A. Boulenger, F.R.S., V.P.Z.S. Notice of a Memoir entitled “‘ Report on the Collection of Fishes made by Dr. W, A. Cunnington during the Third Tanganyika Hxpedition, 1904-05” ........secsecreceerseecececsevererenscees 180
1. Zoological Results of the Third Tanganyika Expedition, conducted by Dr. W. A. Cunnington, 1904—-1905.— Report on the Mollusca. By Epear A. Sirs, 1.8.0., BEZE Ss (Plate X.)i 5. sdosner seerenmanenene rahe rere ee teem 180
2. Zoological Results of the Third Tanganyika Expedition, conducted by Dr. W. A. Cunnington, 1904-1905.— Report on the Macrurous Crustacea. By W. T. Carman, D.Sc., British Museum (Natural History). (Plates 08 =X PV) aise Foe Saeco ate do tae cts tite raed ae eee 187
3. Zoological Results of the Third Tanganyika Expedition, conducted by Dr. W. A. Cunnington, 1904—1905,— Report on the Oligocheta. By Frank EK. Brpparp, K.RS., Prosector to the S0eeby a. ...1 pecs. wocccinmesee eoeeee 206
4. Zoological Results of the Third Tanganyika Expedition, conducted by Dr. W. A. Cunnington, 1904-1905.— Report on the Porifera, with Notes on Species from the Nile and Zambesi. By R. Kirxparricn, F.Z8. (Plates XV Vp rset rae ce oe eer paar 218
5. A Note on “ Flying” Snakes. By R. SHetrorp, M.A., CIMLASS. us ctecotet picts o2 sop nesetaphehacGeateceavsune cs eee 227
March 20, 1906.
The Secretary. Report on the Additions to the Society’s Menagerie during the month of February 1906 ......... 230
The Secretary. Exhibition of a paper cutting representing the print of the foot of an Indian Elephant ............... 230
Mr. John Bowes, F.Z.S. Exhibition of a tooth of the Mammoth from near Herne Bay
Dr.
Myr.
vil
Walter Kidd, F.Z.S. Exhibition of lantern-slides of sections of skin from the palmar and plantar surfaces of Mammals
<} ah Scale] eis} ekehele)6iais)(e) ela iajalel™ielaleialelelaie|alsleisielalels\e/ujslelulaisyai vis) afefele)ete)s
r. Oldfield Thomas, F.R.S. Description of a new sub-
species of Bear, Ursus arctos shanorwm
soe ees ees see eeess ene
R. E. Holding. Exhibition of, and remarks upon,
specimens illustrating anomalies and variations in teeth. ‘
1. Note on Deaths occurring in the Society’s Gardens during
1905. By C. G. Szniemann, M.B., M.R.C.P.
ee ce eer ec ooe
2. A Monograph of the Coleoptera of the Genus Sciobius
Schh. (Curculionide). By Guy A. K. Marsuwatt, F.Z.S. (GE TGs a NG V/AISTOIE she ONUIONG: i aU eal a tee aaeene se raei tf ae a
3. A Contribution to the Study of Evolution based upon the
Mr.
Mr.
Mexican Species of Cnemidophorus. By Hans Ganow, F.RS., F.Z.8. (Plate XX.)
Soe eee ees e reese esse ee eeeosaseecns
April 10, 1906.
F. KE. Beddard, F.R.S. Exhibition of, and remarks upon, a dissected specimen of the Lizard Trachysawrus rugosus showing abdominal ribs
Peer eceee cores se erases cesseees
R. I. Pocock, F.Z.S. Exhibition of the skull of a Horse showing preorbital pits
feces eeeseceeesosoesoesoeesere
1. On the Fresh-water Fishes of the Island of Trinidad, based
on the collection, notes, and sketches made by Mr. Lech- mere Guppy, Junr. By C. Tare Reean, B.A., F.Z.S. (OPT HHES) 2. 0-1 =P, GYD Men OA nea cactopMeenGoane sh qubn Anne Dag
2. The Marine Fauna of Zanzibar and British East Africa,
from Collections made by Cyril Crossland, M.A., B.Sc., F.Z.8., in the Years 1901 and 1902. — Alcyonaria. By Prof. J. ArrHur Tuomson, M.A., University of Aberdeen, and W. A. Henprrson, M.A., B.Sc., Car- negie Fellow, University of Aberdeen. (Plates XX VI.- OX aes BPP so uBE SOS COs abt Ont de 6 .Ue meu ana e ne HA GaIP na:
236
376
307
378
393.
. vill Page 3. On Cyclopia in Osseous Fishes, By James F, Gemma, A AED PP ae PORTE, NV et ee ae ae. 443
4. Notes on Supernumerary Eyes, and Local Deficiency and Reduplication of the Notochord in Trout Embryos. By James E. Gemuitzt, M.A., M.D. (Plate XXXIII.). 449
5. On Three New Forms of Butterfly of the Genus Heli- conius. By Prrcy I. Larny, F.Z.8., F.E.S. (Plate SET 2) RL @ 452
PIG IP Tel A IB INI OWA I WAL Syl OF THE
CONTRIBUTORS,
With References to the several Articles contributed by each.
Page Anprews, CHARLES WiLi1AM, D.Sc., F.R.S., F.Z.8., of the British Museum (Natural History). Exhibition of some models of the skulls and mandibles of Meritheriwm and Palccomastodon .............eeceeee seen (33 Bepparp, FRANK E., M.A., F.R.S., Prosector to the Society. Contributions to the Anatomy of the Ophidia ......... 12 Zoological Results of the Third Tanganyika Expedition, conducted by Dr. W. A. Cunnington, 1904-1905.—Report on the Oligocheeta ...........ccccceeeceeeee ee et eee ec ete eneneenens 206
Exhibition of, and remarks upon, a dissected specimen of the Lizard Trachysaurus rugosus showing abdominal
x Bonuote, J. Lewis, M.A., F.LS., F.Z.S.
On Mammals from South Johore and Singapore
collected by Mr. °C. B. Kloss. (Plate £.)...c0.s2sc:cn.s.senes 4
Bou.Eencer, Grorcr ALBERT, F.R.S., V.P.ZS.
Exhibition of a specimen of, and remarks upon, a giant Brom Troms CAMOLOON a 4.)a..5-<e~ orn an: Ween = eae eee 179
Notice of a Memoir entitled ‘‘ Report on the Collection of Fishes made by Dr. W. A. Cunnington during the Third Tanganyika Expedition, 1904-05” .............0.... 180
Bowss, Joun, F.Z.S8.
Exhibition of a tooth of the Mammoth from near erie Hay i. (2 fais. cis « ve ane ae ne eaplep eee Reee ae eee ene: eee 231
Catman, WituiAm TuHomas, D.Sc., F.Z.8S., of the British Museum (Natural History).
Zoological Results of the Third Tanganyika Expedition, conducted by Dr. W. A. Cunnington, 1904—-1905.— Report on the Macrurous Crustacea. (Plates XI—XIV.)......... 187
DEAN, BasHrorp, Ph.D., of Columbia University, New York.
Notes on, the Living Specimens of the Australian Lung-fish, Ceratodus forsteri, in the Zoological Society’s Collection, > (Plate Lee). he SUA aes. Peas Weasenees : see ehiene 168
Doncaster, Lronarp, M.A., F.Z.8., Mackinnon Student of
the Royal Society, and Raynor, Rev. G. H., M.A.,
On Breeding Experiments with Lepidoptera. MP la bemVabInl 2), i515. sich tian e'aese states s seh ep aes peek eee eet Ree 125
X1
Page Drewitt, Freperic Gxrorce Dawtrey, M.A., M.D., F.R.C.P., F.Z.8. Exhibition of a white variety of the Common Mole ... 2
Fox, Wiutram Storrs, M.A., F.Z.8,
On some Bones of the Lynx from Cales Dale, Derbyshire. 65
Gapow, Hans, M.A., Ph.D., F.R.S., F.Z.8.
A Contribution to the Study of Evolution based upon
the Mexican Species of Cnemidophorus. (Plate XX.) ... 277 GemMiLL, JAMES F., M.A., M.D., Embryological Laboratory, University, Glasgow. On Cyclopia in Osseous Fishes. (Plate XX XIT.)...... 443 Notes on Supernumerary Eyes, and Local Deficiency and Reduplication of the Notochord in Trout Embryos. GET Gp NO NONCTNIET tes eed a re rae canes Sian oisarad ap dalccrts aie ours 449 GiLuerT, FreDErRIcK, F.Z.S. Exhibition of a case of mounted cubs of the Timber- INO liver ieaca te tice caer t Cr Aan SM eth ae Se hace ations 73 Gtntuer, R. T., of Magdalen College, Oxford. Exhibition of, and remarks upon, Medusze from Lake pibeinne crmnivallicaye sy tee snc store caine wiacaietcal Caden sales ecient caine su uelenas 79
Henperson, W. A., M.A., B.Sc., Carnegie Fellow, Uni- versity of Aberdeen, and Tuomson, Prof. J. ARTHUR, M.A., University of Aberdeen.
The Marine Fauna of Zanzibar and British East Africa, from Collections made by Cyril Crossland, M.A., B.Sc., F.Z.8., in the Years 1901 and 1902.—Alcyonaria. (Plaites ; NEKCVi LRN IRET eRe aie WON he et eda ck 393
Ho.pine, R. E.
Exhibition of, and remarks upon, specimens illustrating
anomalies ang variatlons ane teeull wes. serene eeeeenenetets
JENKINSON, J. W., M.A., D.Sc., Assistant to the Linacre Professor of Comparative Anatomy, Oxford.
Notes on the Histology and Physiology of the Placenta miUnoulate. (Plate: VEE.) 0). o.cacsasece ces scoeaenea eens
Kipp, Watter, M.D., M.R.C.S., F.Z.S.
Exhibition of a series of lantern-slides of sections of skin from the palmar and plantar surfaces of Mammals Bnd: Birds. es fico Pokacecosne tle acok oe RE Ee eee
Exhibition of lantern-slides of sections of skin from the
palmar and plantar surfaces of Mammals ..................
Kirkpatrick, Ranpoten, F.Z.S., of the British Museum (Natural History).
Zoological Results of the Third Tanganyika Expedition, conducted by Dr. W. A. Cunnington, 1904—1905.— Report on the Porifera, with Notes on Species from the Nile and Zambesi. (Plates XV.-XVIT.) .............0.08.
Larry, Percy I., F.Z.S., F.E.S.
On Three New Forms of Butterfly of the Genus Heliconius.’. (Plate XXORTV,.) fig. wet ad 8, Mieke
Lover, Sir Epmunp Gites, Bt., V.P.Z.S.
Note on the Cavies of the Genus Dolichotis, and on Living Specimens of D. salinicola. (Plate IV.)..........4.
Page
233
73
73
231
218
452
96
X1li
Page LyprkKeEr, Ricuarp, B.A., F.R.S., F.Z.S. On a Central African evel and Water-Chevrotain. GE TeIbe RAV NIE) i. Sits k era eaten teh anrat ae ei acre MEM at ch hana rt le 112 MarsHatt, Guy A. K., F.Z.8. A Monograph of the Coleoptera of the Genus Sciobius Schh. (Curculionide). (Plates XVIII. & XIX.) ......... 236 Mincuin, Prof. E. A., F.Z.8. Exhibition of a living specimen of a Lemur (Galago) HOT MU SATAC AN cco cic sete aa oe sermon yy hee Sa nn yee 2 Mircuet, P. Cuautmers, M.A., D.Sc., F.R.S., Secretary to the Society. Report on the Additions to the Society’s Menagerie during the month of December 1905 ....................000. 1 Report on the Additions to the Society’s Menagerie durine; the month of January 1906 235342022. ae.. we see atin e un 123 Report on the Additions to the Society’s Menagerie during the month of February 1906........................005 230 Exhibition of a paper cutting representing the print of ihe oot omanpe indian Mlephamteneneeene ee kaccsnecnaee 230 Pocock, Recinaup Innes, F.L.8., Superintendent of the Society’s Gardens. Exhibition of a photograph of a Ring-tailed Lemur GRAMMING THES, OMT? OO. TUES IORVOLS! Soon ronedeboncudecdsodscchoonono 124 Exhibition of the skull of a Horse showing preorbital POUUSE face cece sage a serene sath cei eee seen He a cweM aI EI aaa 377
PycraFrr, WitLiAM Puane, F.Z.8., A.L.S., of the British Museum (Natural History). Contributions to the Osteology of Birds.—Part VIII.
The ‘‘ Tracheophone” Passeres ; with Remarks on Families Cullbvevd le Maver He ROME BAM RCH ACHSS GaSe SBS aaron Tee tet Sere era iaaers NBS
XIV Page Raynor, The Rev. G. H., M.A., F.E.S., and Doncaster,
Leonarp, M.A., F.Z.S., Mackinnon Student of the Royal Society.
On Breeding Experiments with Lepidoptera.
(Plate VIII.)
Recan, C. Tate, B.A., F.ZS., of the British Museum (Natural History).
On the Fresh-water Fishes of the Island of Trinidad, based on the collection, notes, and sketches made by
Mr. Lechmere Guppy, Junr. (Plates XXI.-XXV.)...... 378
Ricarpo, Miss GERTRUDE,
Description of a new Fly of the Family Tabanide...... 97
toux, Dr. Jean, Curator in the Basle Museum of Natural History.
Synopsis of the Toads of the Genus Vectophryne B. & P., with special Remarks on some known Species and
Description of a new Species from German East Africa.
(Binte Te)! 1). Se aera oe oe eee 58
RusseEtL, E.S8., of the Embryological Laboratory, University of Glasgow.
On Trichorhiza, a new Hydroid Genus. (Plate V.)... 99 Scuwann, Haroun, F.Z.8.
A List of the Mammals obtained by Messrs. R. B. Woosnam and R. E. Dent in Bechuanaland. (Plate VI.) 101
ScHWANN, Haroxp, F.Z.S., and Tuomas, O_priexp, F.R.S., F.Z.8.
The Rudd Exploration of South Africa—IV. List of Mammals obtained by Mr. Grant at Knysna
XV
Senigmann, C. G., M.B., M.R.C.P., Pathologist to the Society.
Note on Deaths occurring in the Society’s Gardens Gunmen rT OO OAS ae Hata tales sl aehiceen Men iows ciate te alvaln ats crolsaualdeee ole ;
SuELFoRD, Ropert, M.A., F.L.8., C.M.Z.S,
AWNotesome cr Rilivamoy aS nallcess ies tyee jaar es He leraclle eres :
Smira, Epear A., 1.8.0., F.Z.8.
Zoological Results of the Third Tanganyika Expedition, conducted by Dr. W. A. Cunnington, 1904—-1905.—Re- monureny thes Miolimsceale = CE labewNe)) Mace acteetlrtctnsetete eects
SPURRELL, H. GuorGE F.
The Articulation of the Vertebrate Jaw ...............--.
Tasor, Maj.-Gen. Sir Rueryatp, K.C.B., F.Z.S8., Governor of Victoria.
Extract from a letter from, concerning the supposed oreedimonotra MVNULG Suse aaclacom cepacia sweats e woe ceeiaceule
THOMAS, OLDFIELD, F.R.S., F.Z.S.
Exhibition of, and remarks upon, the skull of a new INESUAPUS | wadancecscsnodar suo sboe UbouseocrEnDes auconpeycotocuconu:
Description of a new subspecies of Bear, Ursus arctos
STOTT ee ee get AES ELTON MARGIE AST Rania
Tomas, OLDFIELD, F.R.S., F.Z.S., and Scuwann, Haroxp, 1a Ags),
The Rudd Exploration of South Africa.—IV. List of Mammals obtained by Mr. Grant at Knysna ...............
Page
234
bo bo ~]
180
114
bo
XV1
Tuomson, Prof. J. ArtHur, M.A., University of Aberdeen, and HeEnpEerson, W. A., M.A., B.Sc., Carnegie Fellow, University of Aberdeen.
The Marine Fauna of Zanzibar and British East Africa, from Collections made by Cyril Crossland, M.A., B.Sc., F.Z.8., in the Years 1901 and 1902.—Alcyonaria. (ares ee VL OKC) Gs sees ace odes telee cea ee
Tomes, CHARLES S., M.A., F.R.S., V.P.Z.S.
On the Minute Structure of the Teeth of Creodonts, with especial reference to their suggested resemblance to Marsupials cists: stitid.). -ics seep sieeehoks-n die watt nc eds ea Re
Woopwarp, ArtHuR Situ, LL.D., F.R.S., F.Z.8.
Exhibition of a drawing of the skeleton of Rhyncho-
SUUPUS GLUCADS: «.. a cichs' wearer ake tee te edee eee rere
Page
393
45
LIST OF PLATES.
1906, pp. 1-462.
Plate Page Il, Dg Wis cis, Bs HIS INGBSO ado Go ob am oaedo Ks eoddue c 4 ll. 1. Nectophryne hosti. 2. N. everetti. 3. N. macrotis. ASW INE CORI Ue Te PNA al 6 ROE ua LSA eR 58 Ill. Histology of the Placenta of the Cow and Sheep ...... 73 IW, JDOUCeORS SHULTWOOUD, osbcobobvoncosondoocoeuDeoodnOEs 96 Wo Spear OMI cosh ecood ec osupoceoonaoodueDuGaNS 99 WIL) WWE WOOSMGHRD occoo sb oBmgDooCoOURS Op Or Hededisy aeons snaps 101 VII. The Black Ituri Ratel (Welhvora cottont) ..........+. 112 VIII. Figs. 1-8. Angerona prunaria and var. sordiata. Figs. 4,5. Abraxas grossulariata and var. lacticolor .......+.04. 125 IDK, | CORTON ORE ooooosebooddccgnaycace aun gooodoUdE 168 X. Mollusca from Lakes Tanganyika and Victoria ........ 180 XI. 1-1a. Palemon mooret. 2-8. Limnocaridina retiarius. } QL JL, WU saccsccdbceseaouHcooadsbenaopuOK XII. 15-22. Limnocaridina similis, 23-29. L. latipes. 30-37. | PS SOCTUS Has elee Cy eierst serene eehey ovskel cued Me eiiekehstey ish eto 187 XI. 38-44. Limnocaridina spinipes. 45-52. Caridella cunning- ois SBS; Ch OMI cchoonedodondpangoevobocec
XIV. 57-64. Atyella brevirostris, 65-72. A. longirostris .... )
XV. XVI. | African Freshwater Sponges ......00..:-5.05-s.0500- 218
XVII. ae Coleoptera of the Genus Sctobius ............-.+00- 236 XX. Distribution of Cremidophorus in Mexico..........++.- 277
Proc. Zoou. Soc.—1906, Vou. I. b
XVill Plate Page XXI. Tetragonopterus guppyt. 2. Haplochilus harti. 3. Curi-)\ matus argenteus....... Be sm fost ool atin =i nlisodee to RetelloRe le XXII. Girardinus guppyi, d. 1a. Q. 2. Chirodon pulcher. 3. Corynopoma risiit, 6. 3a. 2. 4. Letragonopterus (378
LENTUTUS AOS ENUM ALLS, eicea a ee KOM .cPanducheniplerus PUSEE 20%. hase ate, hen keke eee MXIUV., Peeudauchentpterus Guppy? <...02200% osaeen oe ene | XXV. 1. Acara pulchra. 2. Polycentrus schomburghkii ...... y) XXVI. ) XXVII. ae >Alcyonarians from Zanzibar ............0...00eeee 393 D.OO.F XXXII. XXX. Cyclopia in Osseons Wishes .0)52.). Mase caseeee eee 443
XXXII. Supernumerary Myes and absence of Notochord in Trout 449 MAATY..” New forms of Hehcontas.. Fe 0: «swe oon eee 452
LS) J
bo bo bo bo b&b bb
LIST OF TEXT-FIGURES.
1906, pp. 1-462
Page
a, upper, and 4, lower right molars of Hylocherus rimator .... 3
Region of umbilicus in newly-born Anaconda (Eunectes noteus) 13
Region of umbilicus in a young Anaconda (unectes noteus).. 14 Kidney and adjacent organs in newly-born Anaconda (Eunectes
TOECUS Ae eM A eran ie esaqetisl oleeUmar ata scaysts Covet ene tes sta ce Mea eee lay Renal afferent vein of left side of body and its connections in
newly-born Anaconda (HLumectes not@us)........-......... 17 Kidneys of newly-born Anaconda (Lwnectes noteus) .......... 23 Liver and portal veins of young Anaconda (Hunectes noteus) .. 23 Region of umbilicus in newly-born Bitis nastcornis ......,... 35
we) f)
Veins of neck of newly-born Bites nastcornis ......0..+0..005 OF Veins in region of kidney of newly-born Bitis nasicornis ...... 38 Portal veins of newly-born Brtis nasicornis.................. 40
Hypsiprymnus. Longitudinal section of dentine and ener! 48 Thylacinus. enpicuainal section of dentine and enamel...... 49 . Dasyurus. Longitudinal section of dentine and enamel ...... 49 5. Hyena. Longitudinal section near apex of cusp ...... Eo elie eke) Hyzena. Transverse SINC BICIN AE ies A RARE 6 GUO BO aa eo 6 51 Ocelot. The enamel-prisms are not shown, Pom some dle fine! tubes pass a little way into the enamel .................. 51 ishyenodoD, MV ORORECHON Sedan abodobbosabadsocseasnes OF 9. Mesonyx. Longitudinal section ........ yest aan HE Ah oo 53 0, nanos Deming ISIN Gass donde sie tooo ccs id the Rane .. 64 le O rycen Wentinoswellamresenye dur gee tear irri et 54 2. Sinopa. Dentine perished wexcepian: wlacesuin tne me nario) 3. Borhyena. Dentine perished, enamel well preserved ........ 55 4. Borhyena. WDentine well preserved in places................ 56 Dy Cy Modiclis.) PeOneitudinal SeCrOl et ene ining a ony. ors Salo 6. Remains of Felis lynx from Cales Dale, Derbyshire .......... 67
xx Page 27. Details of the formation of an accessory cotyledon in a Cow’s placenta of the 6th month. .........--. sees cece eee eeee 77 28. Sheep.—Detail of cotyledonary crypt ..........++ Uinatans Chews 79 29, Sheep.—Formation of new crypt-cavities by the downgrowth
Wteris of SHEGp ©... /tabmuels stele wie tie siecle ste : . Rhomboidal and lanceolate crystals of bilirubin obtained from
of cell-masses from patches of unmodified—not flattened— GPIGHOl ia Ke sis ai. Wp selec sikle Siaiare dys ote vp ole ne Meas ek iet 81
. Section through one side of the chorionic ring and base of the
diverticulum allantoidis of the Sheep .............- arnt 82 aand 6. Amniotic epithelial thickening, c. Glycogenic epithelium
of the allantoic stalk, and d, e. Glycogenic connective-tissue
cells, of Cow of 4 months ...... el Col UAT es ede tere fete eens 84 eda ciate Ree 91
a chloroform solution of the dried allantoic bodies of the
PET aia igiia’s alee yore eye ote toupee erate ete wisn Serna we tigate povelefenaetee 93 BGS It tICOl WW Gly 2 cig tity epsteearetr tere wiaiarnyey cveretefe lel encl cre oret vc ener 115 Bb PARLE EM MDATG), oe 1 2fe taxa eterene fotse mighelnt ovareywielotm eierelayaze\'oca teeth eee 115 SG aSTAM OL {AWS.) CYC.) Wesuee cio steintee eee ieeiake creme ote vie rele 115 37. Skull of Dog. A. Glenoid fossa pepe by a process of the
squamosal bone. B. Cylindrical condyle of lower jaw .... 116 38. Diagram of jaws. Type2..... Berets cater! tote i eee tn v LLG 39. Diagram of jaws. Type 2. Showing fen crowns of he eth
set in a plane at right angles to the greatest pressure ...... Ly. 40, Diagram of jaws. Type 2. Showinp the emenentia articularis . 117 41, Diagram of jaws. Type 2. Showing lengthening of jaws in
order that incisors may be widely separated, &c. .......... 118
. Diagram of jaws. Type2. Showing the introduction of a second
angle to procure wide separation of the incisors, &e. ...... 119 AS, lull ‘of Draguls Javanvens’ versie seve a'eleighs si0s) Hot ieee pila genaheck gi) 44, Diagram of Snake’s jaws...........- majeteai Vien Siselols bales teaotcea tate 120 45, Skull of Iguanodon bernissartensis ....... sah tavbog uae eve Rae 121 46, Diagram of jaws. Type2R..... se nee als dices *\ «are ste 47. Skull of Batagur Tortoise ...... SEAES Soret erico ety ee CLE Reith dec 122 48. Ring-tailed Lemur and young .......... sia unalimeaads shomenate’ 124 49. Lateral aspect of the skulls of :-—a. Pseudocoluptes boissineauti.
b. Hylactes megapodius. c. Batara cmerea. d. Dendroco-
laptes picumnuUs voc cccrecseos Tease pecbsle tuys Cal ua ob eae eae ae 136
. Ventral aspects of the skulls of :—a. Hylactes megapodius.
b. Philepitta jala. ¢. Pseudocolaptes boissineaut?. d. Den- drocolaptes picumnus. e. Batara cinerea. f. Xiphorhynchus trochilirostris. g. Dorsal aspect of skull of Psewdocolaptes .. 140
. Form of the posterior border of the sternum and the relations of
the articulations of the bones of the Shoulder-girdle at the FOFAMIGN GOSSELIN. 6.2 wicvalae sie yer a erijione amt talet aetna 148
. Phylogenetic tree indicating the probable relationships of the
soTracheoplione” Passeres’.':... ai. s . ccsrieie ena eeeene aes
. Ceratodus forsteri in various positions of rest and movement .. 173 . Ceratodus forstert in various positions of rest and movement .. 175
72.
75 76, ilies
79.
82. 85. 84. 80.
. a, skull, and 0, last lower molar of Ursus arctos Shera OHO. > Se
XX1
Ceratodus forstert in various positions at the time of its coming (RO) Hae) SATE TKO) LONNIE Ge Seon as dblons cou 6 Mea snaveln hoa MHid A ventral scale of Chrysopelea ornata Shaw ..........+..... 228 Diagrammatic transverse sections of the body of Chry aeoreiee OPMWHEE soboavdookooongbo6os soodoooe s0do000Db OAD OO DOD Skull of Ursus arctos shanorum, lateral view ............0.08
bo bo Co bo me ©
bo (SG) i)
Front and nde views of skull of Cercopithecus patas bearing
SUPCRMMNIMenany prem Olarsw sew esa a eerste coatsay ate hag Lepidosis of the front of the left forearm of Ooamidanion us
MELUCANUS 5 ss) -lt Ser aeeh sued ttebaley ofedeoctoyerot aencieree uate) ialeiitary ts ZOO) Lepidosis of the under surface of the left forearm of Cnemido-
PBLORUSLUIUICUCROU IS WC CoE Valve inveneh yyy ota tac ere aie 55) kei Lepidosis of the under surface of the forearm of Cnemedophorus
TIA WOT EDN Mob E REG bin's 4 ONO OG Bb 8 bs SMB OIE h dace Hebi olia neter Bepidosks of the collar and throat of Sa ae ie tessellatus, Lepidosi of the collar and throat of Oncaninionl on us pommel, Bye linigion of Pattern of Coamiioutar us deppet, sien 6 to 11 ale
StLIpess ... Hn OIG BROS OE SIGS I he ety enue a Wg met oes RSMO YZ 0) Evolution of Pattern of Cnemidophorus immutabilis A to 1, and
of C. guttatus from youth F to old agel ............:... 291 Evolution of Pattern of ne scalaris from youth to
co ize)
EXOUUN Bary sa joys) ener el eee Ie le Sapetialndar aneraeie) ohana eter Evolution of Pattern of Cnemidophorus communis from A to D anGUOriC wer tears fromm AGtO PW Elune rei: Were ee Woks eh eae Hvolution of Pattern of Cnemedophorus tessellatus A to D and of CRD ides hl asie clertineh nese tee Hea shes ot ee Pew niia Materayos Cnemidophorus sexlineatus and C. denne RAR Mapai etenetc ol saretKoO Cnenudophorus deppet ......... BSEsTeR Ci Re MIRIAM Bas eS Ca Td Cnemidophorus sexlineatus aud C. ile sane ANA SN metab Shoo uD Cnemidophorus immutabilis 1... ce. ee ee TON Pera ebotl 2.3 Oe
Cnemudophorus immutabilis and C. guttatus........0+eee ee ees S24 GCremidop ors SCalanismraiers sade erate hs ele niee LOOO Cnemidophorus communis occidentalis ..c.ccrceccceeercceess OA Cnemidophorus communis ..........005 Pe fear eI voce IIE lie ee Cnemidophorus communis occidentalis and C. c. australis...... 353 Cnemidophorus bocourt: from Oaxaca .............1..++..+- SOU Cnemidophorus mexicanus typicus, C. m. irae, and C. tmmuta-
(BLOT GOS AOR Cidan OES BOVE On BU Te Re DANS sooeclabouDovecgbes, Boe) Cnennidophorus mexicans Cypicusy Ws he seen as SO Cnemidophorus mericanus var. balsas ......... OER Scio NIG Pane Se: Portion of ventral surface of Trachysaurus rugosus .......... 3876 STOUR RIAIOHUES BUSY Soo ge sobawoeooneee mreberapae a ateyey ere sar Aelia)
Proc. Zoou. Soc.—1906, Vou I. C
LIST OF NEW GENERIC TERMS
PROPOSED IN THE PRESENT VOLUME (pp. 1-462).
Atyella (Crustacea) ......... 187, 201 Melissomorpha (Insecta) ......... (
Caridella (Crustacea) ...... 187, 198
fe PROCEEDINGS |.
1
OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
OF THE
ZOOLOGICAL SOCLETY OF LONDON. 7, (0G!
‘Pagus .f—17.8. CONTAINING PAPERS READ IN
JANUARY ann FEBRUARY.
JUNE 1906.
PRINTED FOR THE SOCIETY, SOLD AT THEIR HOUSE IN HANOVER SQUARE. ene LONDON : | ' MESSRS. LONGMANS, GREEN, AND Co., PATERNOSTER-ROW.
[Price Twelve Shillings. | Gites \ \, &e K be a S
LIST OF CONTENTS.
1906, pp. 1-178.
January 16, 1906.
The Seeretary. Report on the Additions to the Society’s Menagerie during the month of December, LOOM se leisayes! tetera cvs cr wictogaiate ls ele ovavevescestateie ts miate ar tate ee
Pee eenre eceee
Prof. E. A. Minchin, F.Z.S. Exhibition of a living specimen of a Lemur (Gadago) from Uganda 1
Ce i ee a ay
Dr. F. G. D. Drewitt, F.Z.8. Exhibition of a white variety of the Common Mole
Mr. Oldfield Thomas, F.R.S. Exhibition of, and remarks upon, the skull of a new Forest-Pig
ee i er ran y
1. On Mammals from South Johore and Singapore collected by Mr. C. B. Kloss. By J. Lewis Bonnorn, M.A., F.L.S., F.Z.8. (Plate I.)
i ey
to
. Contributions to the Anatomy of the Ophidia. By Franx E. Bupparp, M.A., F.R.S., Prosector £0 the Socteo yrs cin eleyeiaielereledn eiiece eel elm icurtela) ait |x'steh a ae elevalela aivie: detieherere regi
i)
. On the Minute Structure of the Teeth of Oreodonts, with especial reference to their
suggested resemblance to Marsupials. By Cuartes 8. Tomuus, M.A., F.RS., Vice-Pres.Z.8.
4. Synopsis of the Toads of the Genus Nectophryne B. & P., with special Remarks on some known Species and Description of a new Species from German East Africa, By Dr. Jean Rovx, Curator in the Basle Museum of Natural History, (Plate II.) ..
5
. On some Bones of the Lynx from Cales Dale, Derbyshire. By W. Srorrs Fox, M.A., BZA 55 Yoiais alo aiayStuccha a eoaiatogecetesertpeua leva sey eletak olen tant atts fala cr sie relia neRt taint agra een
February 6, 1906. Mr. Frederick Gillett, F.Z.8. Exhibition of a case of mounted cubs of the Timber-Wolf .
Dr. C. W. Andrews, F.Z.S. Exhibition of some models of the skulls and mandibles of Meritheriunt and Paleomastodon
Cr ee eee ner ae i cer re ea
Dr. Walter Kidd, F.Z.S. Exhibition of a series of lantern-slides of sections of skin from the palmar and plantar surfaces of Mammals and Birds............-.e0eceecucees
Page : 1
bo
12
45
58
65
Contents continued on page 3 of Wrapper.
PROCEEDINGS
OF TIIE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
OF TIIE
ZOOLOGICAL SOCIETY OF LONDON.
1906, Vol. I. (January to April).
January 16, 1906.
Howarp SAunvErRS, Esq., Vice-President, in the Chair.
The Secretary read the following report on the additions that had been made is the Society’s Menagerie in December 1905 :—
The registered additions to the Society’s Menagerie during the month of December were 203 innumber. Of these 26 were acquired by presentation and 25 by purchase, 108 were received on deposit, 42 by exchange, and 2 were born in the Gardens. The total number of departures during the same period, by death and removals, was 182.
Amongst the additions special attention may be directed to :—
An Agile Gibbon (Hylobates agilis), from Sumatra, deposited on Dec. 16th.
A Cross’s Guenon (Cercopithecus crossi), from the Cameroons, deposited on Dec. 29th. New to the Collection.
A Grysbok (Vototragus melanotis), from South Africa, deposited on Dec. 18th.
A Snowy Owl (Vyctea seandiaca), from Arctic Europe, purchased on Dec. 16th.
A Madagascar Tree-Boa (Corallus madagascariensis), deposited on Dec. 19th.
The Secretary also exhibited a series of photographs of the Red Proc. Zoou. Soc.—1906, Vor. I. No. I. 1
2 MR. OLDFIELD THOMAS ON [Jan. 16, Deer illustrating the growth of the antlers, which had been pre- sented to the Society by Mr. Walter Winans, F.Z.S.
Prof. E. A. Minchin, F.Z.S., exhibited a living specimen of a Lemur (Galago) which he had brought home with him from Entebbe, Uganda.
Dr. F. G. D. Drewitt, F.Z.S., exhibited, and made remarks upon, a white variety of the Common Mole.
Mr. Oldfield Thomas, F.R.S., exhibited the skull of a female Forest-Pig (Hylocharus), which had been received from Mr. G. L. Bates, who had obtained it from natives of the Ja River, Cameroons, thus confirming ¢ the reports already published * as to the occurrence of H: ylocherus on the west side of Africa. It was to be noticed, however, that the Ja River was not on the West Coast watershed, but rose some 150 miles inland and flowed eastwards to join the general Congo system by way of the Nyoko and Sanga tributaries ; so that this place was the furthest westward point of the true Congo basin, slightly further west in lat, 3° N. than even the main mouth in 6° 8. It was therefore quite likely to be the western limit of the genus Hylocherus.
On comparing the ‘skull of the Ja pig with those from Central and East Africa alr eady in the Museum, now numbering five of different ages and sexes t, Mr. Thomas had come to the conclusion that it could not be assigned to the same species, on account of its much smaller teeth, and therefore proposed to call it Hylocherus rimator { (Thomas, Abstr. P. Z. 8. No. 25, p. 1, Jan. 23, 1906). For while the general shape of the skull appeal to be similar, the teeth thr oughout were very mar kedly narrower, the difference being especially noticeable in the large posterior tooth, m°* (text- fig. T, a). This tooth was not only narrower at its broadest, anterior, point, but tapered poster iorly much more strongly and ended in a point, while in the Eastern form the tooth was broad, tapered but shghtly, and had a broadly rounded end. Throughout, the enamel of the teeth appeared to be much less heavily coated with cement, so that they had a markedly lighter and more delicate appearance. In the lower jaw the same characters were observable, m, (text-fig. 1, 6) being especially long and narrow, its length more than three times fe greatest breadth. while in ‘the allied forms the length was barely 22 times the Pode
There was also a portenn difference in the detailed structure of
* P.Z.S. 1904, ii. p. 196.
+ The Museum was indebted to Mr. C. W. Haywood for one young skull and two skeletons, and to Col. Harrison for a fine pair of adult skulls.
{ [The complete account of the new species described in this communication
appears here; but since the name and preliminary diagnosis were published in the ‘Abstract,’ the species is distinguished by the name being underlined.—Ep1Tor. |
1906. | A NEW FOREST-PIG. 3
the teeth, as might be seen by a comparison of the figures (text- fig. 1) now given with those formerly published.
The skull itself was comparatively light and delicate, with a low muzzle and slender zygomata. Its dimensions were as follows :—
Basal length (approximate) 325 mm. ; zygomatic breadth 176 ; nasals, length 191, breadth 42; interorbital breadth 88; height of muzzle in front of premolars 57; least breadth of maxillary zygomatic processes, below orbit, 42; least vertical breadth of zygomata, behind true orbit, 36; palatal length 232; breadth across sockets of canines 98; breadth between tips of canines 181; greatest diameter of canines 24; length of upper tooth-row, from front of p* 97; m’, length 42°3, breadth 17:5; length of lower tooth-row, from front of p,, 99; m, 48°2 x 16.
Text-fig. 1.
a, wpper, and 4, lower right molars of Hylocherus rimator.
Type. Old female skull. B.M. No. 6.2.21.1. Collected by Mr. G. L. Bates.
Native name ‘“ Engak” (G. L. Bates).
The remarkable character of the suppression of p* and persistence of mp’, described in the typical specimens of HH. meinertzhageni, was present in this specimen, as in other examples of the genus since received, and was evidently normal.
Another interesting point to be noticed was that the prenasal bone in all fully adult specimens of Hylocherus was firmly united to an ossified mesethmoid.
The following papers were read :—
1*
4 MR. J. LEWIS BONHOTE ON MAMMALS [Jan. 16,
1. On Mammals from South Johore and Singapore collected by Mr. C. B. Kloss. By 8. Lewis Bonyorz, M.A., PvE Shes:
[Received December 7, 1905. | (Plate I.*)
The following is an account of a small collection of mammals collected by Mr. C. B. Kloss in the southern part of Johore and the island of Singapore. From the latter place, which has been for long the emporium of that portion of the East, and from which many new species have been described, genuine wild collected
ecimens are most welcome, and up to the present but few
ecimens have been received from Johore. During the last few years several collections from the Malay Region have been received in both London and Washington, so that the number of novelties in the present collection is not so large as it might otherwise have been, but nevertheless, of the 17 species sent home, two are new and represent J/us ochraceiventer and M. whiteheadi of Borneo. In my former paper on Messrs. Annandale and Robinson’s collection, the synonymy of the various species was brought up to date, and in the present paper reference is given only to those papers that have been published since.
TUPAIA FERRUGINEA Rafiles.
Tupaa ferruginea Rafiles, Trans. Linn, Soe. xiii. p. 256 (1822) ; Flower, P. Z.8. 1900, p. 336; Miller, Proc. Biol. Soc. Wash. xiii. p. 193 (1900); id. Proc. U.S. Nat. Mus. vol. xxvi. p. 472 (1903).
a-e. $. Pelepak, Johore, March 1905,
S-h. 2. Pulai, S.W. Johore, Nov. & Dec. 1905.
i-l. 2. Johore Bahru, Feb. & March 1904.
m. 2. Woodlands, Singapore, 2nd Aug., 1904.
This species occurs as far north as Tenasserim, where it meets
with 7. f. belangeri, a paler form.
TUPAIA MALACCANA Anders.
Tupaia malaccana Anders. Zool. Res. Yunnan, p. 134 (1879); Bonhote, Fase. Malay., Zool. vol. i. p. 13 (1903); Miller, Proc. Wash. Acad. Sci. il. p. 230 (1900) ; id. Proc. Acad. Nat. Sci. Philad. 1902, p. 157; id. Proc. U.S. Nat. Mus. vol. xxvi. p. 472 (1908).
a. S$. Kangka Aming, 3rd June, 1905.
by Oe Kaneka Kuli, 25th Nov., 1904.
c. 2. S&S. Malaya, 24th June, 1904,
A series of very typical specimens. RATUFA AFFINIS (Raffl.).
Seiurus affinis Rati. (nec Horsf.) Trans. Linn, Soc. xiii. p. 259 (1822).
* For explanation of the Plate, see p. 11.
dort AT LOSS USIG »® Be"
Titel WO" SOGI “SZ al
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1906. ] FROM JOHORE AND SINGAPORE. 5
Ratufa affinis typica (Ratiles) Bonh. Ann. & Mag. N. H. (7) vol. v. p. 495 (1900).
Ratufa affinis (Rafiles) Miller, Proc. Wash. Acad. Sci. voi. ii. pp. 73 & 77 (1900).
a,b. 3 2. Pelepak, Johore, March 1905.
With reference to a note by Mr. Miller (Joc. cit. supra), I believe that his &. affinis is undoubtedly the same as my R?. affinis typica. In my description of this species I noted that the hairs “when newly-grown” were annulated; but this annulation dis- appears extremely quickly, so that itis very seldom that a specimen in annulated pelage is procured.
With regard to Miller's 2. pyrsonota described in the same paper, this is the Malayan form of 2. ephippiwm from Borneo and is, as Mr. Miller notes, quite distinct from 2. afinis. It should really be known as &. ephippium pyrsonota, which saves any possibility of confusion. Mr. Miller has since 1900 described many new species representing island forms or local races of F. affinis or ephippium, but, from his strict adherence to binomial nomenclature, which has prevented him from recognising the true specific difference between affinis and ephippiwm, it is impossible to tell to which of these species his new races belong, and in consequence to determine the true specific range. Jam, however, inclined to believe that they are all races of 2. ephippiwm, and that the true R. affinis will be found to have a very restricted range, confined to the Malay Peninsula and possibly Java.
Sciurus TENuIS Horsf.
Seiurus tenuis Horsf. Zool. Res. (1824); Thos. P.Z. 8. 1886, p. (6; Flower, P.Z.S. 1900, p.357; Mill. Proc. Wash. Acad. Sci. ego 2 ele (900) iids Proc) eatcads Nat. Sci Philad) 1902) p: 5; id. Proc. U.S. Nat. Mus. xxvi. p. 452 (1903).
a-b. 2. Gunong Pulai, 1500’, Nov. & Dec. 1904.
ce. Qimm. Mt. Austin, 8. Johore, 3lst Aug., 1904.
d. 2. SS. Malaya, 8S. Johore, 24th June, 1904.
e, f. 2. Woodlands, Singapore, June & Aug. 1904.
These specimens, which are practically topotypes, are therefore of the typical form. This species, however, appears to be a remarkably stable one, Mr. Miller stating that specimens from Sumatra were indistinguishable from topotypes. At the same time S. tenwis surdus Mill., from the north of the Peninsula, is distinctly paler and greyer.
SCIURUS VITTATUS Raffles.
Sciurus vittatus Rafiles, Trans. Linn. Soc. xiii. p. 259 (1822) ; Bonh. Fasc. Malay., Zool. vol. 1. p. 22 (1903).
Sciurus notatus miniatus Miller, Proc. Wash. Acad. Sei. ii. p. 79 (1909).
Sciurus penirsularis Miller, Smithsonian Miscell. Coll. vol. xlv.
p. 10 (1903).
6 MR. J. LEWIS BONHOTE ON MAMMALS [ Jan. 16
a,b. § @. Pelepak, Johore, 27th March, 1905. ce. 9. Kangka Aming, Johore, 26th May, 1905. d. $. Mount Austin, Johore, 3lst Aug., 1905.
e. 9. Johore Bahru, 10th May, 1904.
ft. 2. Pasir Gudang, Johore, 26th Feb., 1904.
g. 2. Pasir Gudang, Johore, 11th May, 1905.
h. 2. Woodlands, Singapore I., 15th Aug., 1904.
i,k. 6 2. Singapore I., June & Aug. 1904.
Mr. Miller has made two new species of the Peninsula forms of Se. vittatus. He first separated the N. Malayan form from those in the south under the name miniatus, and more recently the Southern Peninsula form has been named peninsularis, to distinguish it from the typical Sumatran race.
The series sent home by Mr. Kloss has enabled me to re- consider the matter very thoroughly, and has still further confirmed my opinion that this very variable species cannot be split up into the geographical races proposed by Mr. Miller.
To take the case of S. v. miniatus first of all. Itis characterised, compared with specimens from Singapore Island, by having the ved of the under parts much darker and the terminal pencil or sometimes the entire distal half of the tail uniform red beneath. The type locality of this subspecies is given as Trong in Lower Siam, but its exact distribution is not stated.
In the British Museum there are specimens answering to this description from various localities along the whole length of the Peninsula including Singapore Island, so that its distribution would appear to be identical with that of the paler form, as in many localities they are found side by side.
S. miniatus therefore can on no account be considered as a geographical race or true subspecies, and it must be either a distinct species or a mere colour-variety. The fact that inter- mediates in all stages occur renders its status as a true species impossible; and there are specimens showing on the under parts the deep red of miniatus as well as the lighter orange of true vittatus. ‘These, then, are the reasons for regarding miniatus merely as a colour-variety of S. vittatus.
Now as regards S. peninsularis. The characters distinguishing it from the true S. vittatus of Sumatra are said to be as follows: “more yellowish under parts, less yellowish cheeks,” ¢.e. a paler variety.
The types of SN. vittatus, which are in the British Museum, are absolutely indistinguishable from specimens taken in the Penin- sula. It should, of course, be noted that Tapanuli Bay, whence Mr. Miller procured his series, which he regarded as typical of S. vittatus, is a considerable distance from Bencoolen, the type locality ; and the Tapanuli Bay specimens may represent a distinct race, but this is most improbable, as this species also varies con- siderably in Sumatra, since darker specimens with the red tail are also to be found there.
The fact therefore emains that Malay specimens may be
1906. | FROM JOHORE AND SINGAPORE. 0
identical in all respects with the types of vittatus, and since this is the case Mr. Miller’s peninsaularis cannot stand.
‘It should always be remembered that among truly variable Species, of which there are not many, but of canal S. finlaysont is another good example, geographical forms are very unlikely to be found: for the fact that it is variable means (i) that natural selection does not restrict it to a particular type; (ii) that in consequence it almost certainly does not breed true. So that, although the majority of individuals in any one place may show a tendency to a particular form of coloration, other varieties will be so frequently appearing as to prevent the fixin g of any particular character. Now a true geographic race, to which forms alone trinomials should be restricted, should be the only race found in its type locality, intermediates being found only in the country approaching that of another race. It is therefore evident that S. vittatus is, as regards colour, a very variable species, and as such it behoves all systematists to be extremely cautious before naming any races thereof.
It is only fair to add that the type of miniatus came from the hills at an altitude of about 1000 feet, and may represent a hill form; but at present there is no evidence of this, and such evidence as there is, namely the dark and light under parts being found on the same individual, tends to disprove it.
SCIURUS NIGROVITTATUS BILIMITATUS Mill.
Sceiurus biimitatus Mill. Smithsonian Mise. Coll. vol. xlv. p. 8 (1903).
Sciurus nigrovittatus Horsf., Bonh. Ann. & Mag. Nat. Hist. (7) vii. p. 452 (1901); id. Fase. Malay., Zool. vol. i. p. 23 (1903).
a. 2. Pelepak, Johore, 24th March, 1905.
b,c. 3. Kangka Aming, Johore, 26th May, 1905.
d. 3. Johore Bahru, 20th Feb., 1905.
es 9. - oF 18th June, 1904.
Mr. Miller has separated the Peninsula and Javan forms chiefly on account of the almost entire absence of the light stripe in the latter. I have been able to examine only one specimen from Java, in which the stripe is certainly almost obliterated. This is, however, by no means the case with the Sumatran form, in which the stripe is very well marked.
FUNAMBULUS INSIGNIS PENINSULE Miller.
Funambulus peninsule Mill. Smithsonian Mise. Coll. vol. xlv. p- 25 (1903). a,b. 62. Kangka Aming, Johore, 29th May, 1905.
I have placed these examples under Mr. Miller’s name, al- though his description does not exactly agree with specimens from the Peninsula, and it is probable that his single specimen is hardly typical of the Malay form. I was, however, in error
§ MR. J. LEWIS BONHOTE ON MAMMALS [Jan. 16,
when I stated (Fasc. Malay., Zool. vol. i. p. 26) that the Malay and Sumatran animals were indistinguishable, since they are undoubtedly distinct, the former being of a much yellower and warmer tint. On the other hand, Messrs. Stone and Rehn (Proc. Acad. Nat. Sci. Philad. 1902, p. 133) mention a Sumatran specimen as approaching /’. 7. diversws Thos., in which the colours generally are much brighter and the sides of the body nearly pure chestnut. Further Sumatran material is, however, necessary before this matter can be satisfactorily settled.
The following are the races of Funambulus insignis hitherto described :—
F. insignis typicus F. Cuv. Mamm. 1821, pl. 233. Sumatra. F. i. diversus Thos. Ann. Mag. Nat. Hist. (7) vol. ii. p. 248 (1898). Baram, Sarawak.
», niobe Thos. ibid. p. 249 (1898). Pajo, Sumatra. ,, castaneus Mill. Proc. Wash. Acad. Sci. vol. ii. p. 217 (1900). Anambas I. ,, jalorensis Bonh. Fasc. Malay., Zool. vel. i. p. 25 (1908). Bukit Besar, Jalor. » peninsule Mill. Smithson. Mise. Coll. vol. xlv. p. 25 (1903). Trong, L. Siam. » rostratus Mill. ibid. p. 24 (1903). Tina Balu, Batu I. », obscurus Mill. ibid. p. 23 (1903). S. Pagi I., Sumatra.
Mus vocirerans, Miller.
Mus vociferans Miller, Proc. Biol. Soc. Wash. vol. xii. p. 138 (1900); Bonhote, Fasc. Malay., Zool. vol. i. p. 33.
a-k.73,492. Mt. Pulai S. Johore, Nov. & Dec. 1904 and Jan. 1905.
As stated by Mr. Miller in his original description, this species is similar to Jus sabanus Thos., but brighter and with a very long tail. The extreme length of the tail is sufficient to distinguish it from all other members of the jerdoni group. It would appear to be somewhat local, as the specimens in this collection are all from Mount Pulai. The following is a short description that may enable this species to be recognised :—
Colour of upper and under parts sharply divided. Above rufous buff, thickly grizzled with black, the black being almost absent on. the flanks. Under parts white. Feet brownish with white margins. ‘Tail bicolor and with its terminal portion white.
Skull of typical jerdoni group, 7. e. rather elongated, small for size of animal, bullee very small.
Dimensions. Head and body 229 mm.; tail 382; hind foot 45; ear 24.
Skull—greatest length 56 mm.; zygomatic breadth 26.
Since the publication of my paper quoted above, Mr. Miller has described several island forms of this species in Smithsonian Misc. Coll. vol. xlv. pp. 28 et seq. (1903).
1906. | FROM JOHORE AND SINGAPORE. 9
Mus surirer Mili,
Mus surifer Mill., Proc. Biol. Soc. Wash. xiii. p. 148 (1900) ; Bonh. Fasc. Malay., Zool. vol. i. p. 26 (1903). 236,79. Pelepak, Johore, March 1905. 129,18 2. Pulai, Johore, Dec. 1904 to March 1905. 39. Kangka Kuh, Johore, 28th Nov., 1904. . Kangka Aming, Johore, 29th May, 1905. Mt. Lun Chu, Johore, 13th May, 1905. Johore Bahru, 21st May, 1905. Kangka Keteho, Tebraun.
This large series of Mus surifer calls for. little comment; the examples show a slight variation in size, and the young are much duller in colour, being of a uniform brown (hair-brown, Ridgw.). The adult pelage first appears along the flanks. Mr. Miller’s distinction of the white of the foot being cut off from the white of the inside of the thighs owing to the brown colour encircling the ankle, appears to be a perfectly constant feature.
OS
Mts INAS, sp. nov. (Plate I. fig. 1.)
Mus whiteheadi Thos., Bonh. P. Z.8. 1900, p. 880.
The Malayan representative of Mus ochraceiventer Thos. Fur long and thickly interspersed with spines. General colour above uniform grizzled rufous (ochraceous rufous, Ridgw.), fading to pale ochraceous (pinkish buff, Ridgw.) on the under parts. Feet and hands brownish white. Tail rather shorter than the head and body, almost naked and bicolor.
Skull. The only skull available is in such a bad state of preservation that a description is impossible.
Dimensions of type (in flesh). Head and body 162 mm.; tail 5 2es hindstoob sl.
Habitat. Gunong Inas, Perak. Also found in Johore.
Type. B.M. 2.11.15.2. Adult 9. Gunong Inas, December 1899.
The type specimen of this species was erroneously referred by me some years ago to MW. whiteheadi. The advent of another specimen in the present collection proves it to be sufficiently distinct from the Bornean form to merit description. It may be distinguished from Jus ochraceiventer by its more rufous colour above and paler colour beneath, while the tail 1s very much more markedly bicolor. It is distinguishable from J/. whiteheadi by its larger size.
Mus Kuossi, sp. nov. (Plate I. fig. 2.)
The Malayan representative of Jus whiteheadi Thos. Fur short, thickly interspersed with spines. General colour above rufous-buff (orange-buff, Ridgw.), grizzled with dark brown, the buff colour becoming purer on the flanks and shading to pale ochraceous (cream-buft, Ridgw.) on the under parts. Feet and hands white. Tail shorter than the head and body, almost naked and bicoler.
10 MR. J. LEWIS BONHOTE ON MAMMALS [ Jan. 16,
Skull of the Mus jerdont type with small bulle, very similar to that of Mus whiteheadi but smaller.
Dimensions (of type in flesh), Head and body 120 mm. ; tail 105; hind foot 26; ear 17.
Skull. Greatest length 32 mm.; basilar length 24; palatal length 7; diastema 8; length of incisive foramina 5; length of nasals 11 ; ; zygomatic breadth 9 ; interorbital breadth 5:5 ; breadth of brain- eS at roots of zy gomata 13; length of molar series 5.
Habitat. Johore, Malay Peninsula.
Type. Coll. C. B. Kloss, No. 218. Adult 2. Collected on Mount Pulai, 8. Johore, 1600’, on the 31st December, 1904.
This species may-be easily distinguished from Mus whiteheadi, the only species with which it might be confused, by its paler and yellowish coloration and slightly smaller size. Mr. Kloss has sent over 5 specimens altogether, viz.
abs a. Pelepak, etious) Mandl 1905.
ce. 6. Mt. Lun Chu, Johore, May 1905.
d,e. 2 (one the type). Mount Pulai, Johore, Dec. 1904.
Mus vaurpus Mill.
Mus validus Mill. Proce. Biol. Soc. Wash. vol. xii. p. 141 (1900); Bonh. Fasc. Malay., Zool. vol. i. pp. 34 & 37 (1903).
a,b. 6 2. Kangka Kuli, Johore, 24th Nov., 1905.
This species is probably the Malay form of Jus muelleri. Mr. Miller in his original description states that this species bears a great external resemblance to J/us bowersi; but this is hardly the case, for the general colour of Mus bowersi is greyish and its fur is fairly soft, whereas in the species under consideration the general colour is dark brown, each hair having lighter annulations, Sa the fur is very harsh. Mus bowerst has in Seon the ter site| portion of the tail white, whilst in Mus validus the tail is uni- colorous and dark.
There are no skulls with these specimens, but the cranial characters have been carefully given by Mr. Miller.
Mus gArAk Bonh.
Mus jarak* Bonh. Journ. Fed. Mal. States, vol. 1. no. 3 (1905),
a. 9. Pelepak, Johore, 6th Jan. 1905.
b,c. 6 2. Kangka, Senibong, 8.W. Johore, 10th July, 1905, and 12th Sept., 1904.
d-h. 3. Pasir Gudang, 8.W. Johore, 10th May, 1905.
A series of eight skins sent home by Mr. Kloss proves this species, which I had supposed to be an island form, to occur in
* Mus jarak nearly allied to Mus jalorensis, but darker on the upper parts, the fulvous tips being browner and greatly diminished in size. Type locality, Pulau, Jarak, Str. of Malacca. A full description of this species is published by the Selangor Museum.
1906. | FROM JOHORS AND SINGAPORE. Tal
the south of the Peninsula. Individuals vary somewhat amongst themselves, some being rather lighter in their general coloration than others, but the lightest havea considerably darker appearance than Jus jalorensis.
They belong, as I pointed out in my former paper, to the Pyctoris sub-group of Mus rattus, and may be distinguished by their soft fur, medium size, short tail, and pure white under parts, the hairs being white to their bases.
MUS GRISEIVENTER Bonh.
Mus grisewenter Bonh. Fasc. Malay., Zool. vol. i. p. 30 (1903).
a-c.2 36,1 2. Pelepak, Johore, March 1905.
d-g.3 6,1 9. Pulai, S.W. Johore, Nov. 1904 and Jan. 1905.
h. 3. Johore Bahru, 3rd April, 1905.
in. 3 ¢,2 9. Kangka Senibong, Johore, Sept. 1905.
o. 3. Pasir Gudang, Johore, llth May, 1905.
This is the really common House-rat of the Peninsula; its greyish or yellowish under parts enable it to be easily recognised from M., jalorensis or M. jarak, the hill forms of Mus rattus. In size, too, it is rather larger than the preceding species.
Mus norvecicus Erxl.
Mus norvegicus Erxleben.
Mus decumanus Pall., Flower, P. Z.S8. 1900, p. 362.
73,9 2. Johore Bahru, April, June, and August, 1904.
So far as can be judged, this Rat is apparently found only on the Peninsula in the neighbourhood of shipping.
Mus concotor Blyth.
Mus concolor Blyth, J. A. 8S. B. xxviii. p. 295 (1859); Bonh. Fasc. Malay., Zool. vol. i. p. 38 (1903).
a-c. 6. Pelepak, Johore, March 1905.
d. $. Tebraun, Johore, Sept. 1905.
e. d. Johore Bahru, July 1904.
fig. 62. Kangka Senibong, Sept. 1905.
h. 2. Pasir Gudang, May 1905.
Mus muscuuus Linn.
Mus musculus Linn. Syst. Nat. xu. p. 83 (1776); Flower, IP Assi OOS Fos BOM,
a. 6. Johore Bahru, Feb. 1904.
EXPLANATION OF PLATE I.
Mus inas, p. 9. Mus klossi, p. 9.
12 MR. F. E. BEDDARD ON THE [Jan. 16,
2. Contributions to the Anatomy of the Ophidia. By Frank E. Bepparp, M.A. F.R.S., Prosector to the
Society. [Received Decembez 7th, 1905. |
(Text-figures 2-11.)
CONTENTS.
(1) On the Vascular System of the Anaconda, on the Characters of the newly-born Young of Hunectes noteus, and on the Differences between the two Species of the Genus Hunectes, viz. FE. murinus and EF. noteus: p. 12.
(2) Some Notes upon the Venous System of Python sebe: p. 27.
(3) Some Notes upon the Anatomy of I/ysia scytale, bearing upon its systematic position: p. 31.
(4) The Structure of the Young Bitis nasicornis, with Notes on other Vipers: p. 34.
(5) Considerations respecting the Primitive Structure of the Lungs in the Sqnamata: p. 41.
(1) Notes on the Vascular System of the Anaconda, on the Characters of the newly-born Young of Kunectes noteeus, and on the Differences between the two Species of the Genus Kunectes, viz. KE. murinus and E. noteus.
The opportunity of examining two newly-born* examples of a little-known species of Anaconda, viz. Hunectes noteus, is so unlikely to be of frequent occurrence, that I have thought it desirable to make as detailed a study as possible of the veins, while the fact that comparatively little is known of the venous system in the genus Hunectes seemed to me to render this study still more desirable. So far as [ am aware, some notes by myself f upon the intercostal arteries and the anterior abdominal vein are all that has been published upon this genus of Boine snakes so far as concerns the vascular system.
On the other hand, there is abundant material for comparison with the anatomy of the vascular system of Python, and less with that of Hry«wS and Boa
External characters of Young.—The newly-born young of this snake retain some of the embryonic veins and the yolk-sac is not completely absorbed. This woulda ppear to be a great disadvantage until we reflect that the Anaconda is so very aquatic an animal that the young are possi bly born in the water 4]. The navel ‘is a very large aperture in the ventral median line, measuring 14 mm. x9 mm. orl2mm.x7mm. It lies 44 or 54 mm. from ‘the vent.
* These young Anacondas were born on Sept. 27, 1905, and one of them was just living when it came into my hands.
+ “Notes upon the Anatomy of certain Snakes of the Family Boide,” P. Z.S. 1904, vol. 11. p. 107.
a6 ailopiingon & Pancoat, Trans. Amer. Phil. Soc. v. 1837, p. 121; Jacquart, Ann. Sci. D Nat. ® iv. 1855, p. 321; Retzius, K. Vet.-Ak. Handl. 1830; Beddard, Paves: 1904, vol. i. p. 362; Panizza, ‘Sopra il systema linfatico dei Rettili,’ Pavia, 1833 (I ave hi seen this memoir).
§ Beddard, P. Z. 8. 1904, vol. ii. p. 107. i Gadow, incorporated i in Bronn’s Thier. vi. Abth. iii. §{ Lam indebted to Mr. Pocock for reminding me of this.
1906. ANATOMY OF THE OPHIDIA. 13
The whole lengths of the two specimens which I have examined were as follows :—Specimen A, 21] inches; Specimen B, 192 inches. The distance between umbilicus and vent was naturally greater in the larger specimen, and the umbilicus itself larger.
In the above given measurements I regard as the navel not merely the actual aperture in the skin through which the yolk-plug* pro- trudes, but the whole area which is devoid of scaling. The yolk- plug in the interior of the body extends from the gall-bladder anteriorly to the end of the kidneys posteriorly, and lies above the fat-body. It is a dense solid plug. Meckel’s diverticulum arises from it just behind the umbilicus, and enters the small intestine about an inch behind the pancreas.
The umbilicus itself is so exactly median in position (text-fig. 2 that it has divided the epigastric vein, which, instead of lying to one side or the other, forms a loop surrounding it. As elsewhere this vein is single, the position of the umbilicus could, I imagine, be detected in a more fully adult snake by this splitting and rejoining of the epigastric vein.
Text-fig. 2.
Region of umbilicus in newly-born Anaconda (Eunectes noteus).
umb, Umbilical area; 7.S. Ventral scales anterior to umbilicus, which are still divided into two; V.S’. Similar ventral scales behind the umbilicus.
In the two newly-born Anacondas the area of the navel em- braced nine of the ventral scales, which are split into halves, each half lying on either side of the soft median area. Anteriorly and posteriorly two scales showed a median groove, indicating, it is to
** Messrs. Mole & Urich (P. Z. S. 1894, p. 505) mention the existence of ‘traces of the umbilical cord” in newly-born Hunectes murinus.
14 MR F. E. BEDDARD ON THE [Jan. 16,
be presumed, that they had been originally split, but that the two halves had come together in the course of the gradual obliteration of the area of the navel. Both specimens were practically identical in the disposition of the scales. It is remarkable, however, that in an individual of the same brood measuring 22+ inches long, and older by three weeks (it died on Oct. 17), the median furrowing of the ventral scales was continued for a long way behind the navel (text-fig. 3). The actual area of the navel, much narrower in this specimen, corresponds to seven scales. ‘This older individual is a male, as shown by the shape of the cloacal claws; I did not dissect it.
Text-fig. 3.
Region of umbilicus in a young Anaconda (Hunectes noteus). Lettering as in text-fig. 2.
Current treatises on Zoology have largely ignored the fact that among the Boide the rudiments of hind limbs offer sexual characters which are unmistakable. They are obvious, for instance, in the genus Eryx, and in the species with which I am concerned, viz. Hunectes noteus. That this fact is obviously not generally known is perhaps due to Duméril and Bibron. These
1906. | ANATOMY OF THE OPHIDIA. 15
authors, in their classical ‘ Erpétologie Générale, vol. vi. 1844, remark, in a general sketch of the Pythons and Boas, upon the investigations of Mayer on the rudimentary hind limb, but say nothing as to sexual differences in form of the ‘ ergots,” 7. e¢., claws in which these limbs terminate. Later on they write of Hunectes murinus (p. 531) :—‘* Ergots coniques, courts, recourbés et pointus,” stating also that they are “d'une trés petite dimension chez des femelles ayant plus d’un metre de long.” In Sow again (p. 503) it is said that these claws are present, but ‘“ néanmoins plus développés chez les males chez les femelles.”. The inference from these observations would surely be that while sexual differ- ences exist between individuals in respect of those claws, they are merely a matter of varying magnitude. It was possibly for this reason that text-books, at least those which I have referred to, have not noticed the matter. In the adult male Humectes notewus there is a very conspicuous pair of claws, which are sharply pointed, compressed, and curved, the lower surface being ridged. In the adult female there is not a claw at all, but a bluntly conical straight process, quite unlike the claw of the male. In both cases, however, this terminal process is ensheathed in two bract-like scales. In the young individuals the differences are as well marked as in the adult.
The mental groove in the young Anacondas differs from that of theadult male. In both adults the mental groove was bordered by five scales on each side. ‘The two individuals, a maleand a female, appeared to be identical in the disposition of these scales.
In the two youngest individuals, which died immediately on birth, the conditions were as follows :—In one snake, on the right side the mental groove was bordered by only three scales; but the middle one was very large, and indentations appeared to show that it was in reality composed of three. On the left side there were only fom: scales bordering the mental groove, but the anterior one of these was partly divided into two; thus the total on both sides is really five as in the adult. The second specimen was identical on the right side, but on the left the second scale, and not the first, was obviously composed of two. In the young Anaconda which lived for three weeks* there is a still closer approximation to the conditions observable in the adult. On the left side of the mental groove all five scales are separated ; on the right there were four scales, the second being much the largest and obviously representing two.
Kidneys.—Vhe kidneys of these young Hunectes have a form which is remarkable. It is illustrated in the accompanying figure (text-fig. 4,p.17). The part of the kidney which apparently corre- sponds to the whole kidney of other Snakes has the usual form which is characteristic of Serpents and needs no particular remark. The kidney does not, however, end off in front without much diminution of diameter. It suddenly narrows to form a slender
* This specimen shed its skin at any rate once, which doubtless accounts for the change, not therefore referable to individual variation.
16 MR. F. E. BEDDARD ON THE [Jan. 16,
forward prolongation, which is as long as, or longer than, the posterior region of the kidney.. There is no break whatever between these two sections; and their appearance as regards texture and colour is identical. The thinner anterior part of the kidney may pehaps be a mesonephros, persistent in these young forms. On the two sides of the body the two kidneys differed very considerably in dimensions. The right kidney is, as in other Snakes, more advanced than the left kidney, and its anterior end actually passes a trifle beyond the gall-bladder and all but reaches theliver. This kidney is altogether 108 mm. long, of which 44 mm. belong to the posterior bro vad region of the gland, the slender anterior portion being thus much the longer. The right-hand kidney is altogether only 92 mm. long, and the broader posterior region is here the longer of the two sections, measuring as it does 48 mm.
The slender anterior prolongation of each kidney is not, of course, to be confounded with the adrenal body. This latter is plainly distinguishable from the kidney-tissue by its yellow colour and different texture. It les in the middle section of the. anterior region of the kidney.
The gonads were not visible in either specimen. But I believe them both to be females. This conclusion was arrived at owing to the nature of the gonad-ducts. These ducts were of com- paratively large calibre and without the close windings of the male ducts Moreover they were prolonged forward in the case of the right-hand one to a point anterior to the gall-bladder where the duct appeared to end freely. In the region of the anterior portion of the kidney the duct was attached by an evident though narrow mesentery to the substance of the kidney. The actual course of the gonad- duct of the right side is shown in the figure annexed (text-fig. 4, p. 17). Anter iorly it lies to the outside of the kidney. At the junction between the anterior more slender and the posterior stouter region of the kidney it crosses over and lies to the inside of the kidney.
The diameter of this tube appears to me to be too great to allow of its being identified with the sperm-duct. Another and, as [ believe, very strong reason forbids this identification. To the inside of each kidney, along the anterior thinner region of that organ only, is a more slender duct than the gonad-duct, which, however, presents the same general appearance. This duct com- mences some way in front of each kidney, but the exact mode of its commencement [I have been unable to ascertain. It follows the kidney fairly closely, lying on the opposite side to that occupied by the gonad-duct, to a point some little way in front of the junction between the anterior and posterior sections of the gland, and there gradually disappears. It seems to me that this str ucture must be unquestionably homologised with the mesonephric duct ; and if so, the gonad-duct can hardly be the sperm-duct. That it is the mesonephric duct seems to be necessary from the impossi- bility of identifying it with anything else; and if so, then the
1906. ] ANATOMY OF THE OPHIDIA. 17
section of kidney along which it lies is possibly to be looked upon as mesonephros, the incorporation of which with the metanephros has perhaps caused the withering of the end of the tube, no longer needed as a secretory conduit.
Text-fig. 5.
ue
—y se ~~ OS
= Ss
SBE SS
Text-fig. 4.— Kidney and adjacent organs in newly-born Anaconda (Eunectes noteus).
gob. Gall-bladder ; . Kidney ; WM. Mesentery attaching oviduct to mn, mesonephros ; Od. Oviduct ; P.c. Posterior cardinal vein arising on kidney (?) ; Sr. Suprarenal portal veins: U. Ureter; v.c.i. Vena cava posterior or renal efferent vein ; Wd. Wolffian duct. :
Text-fig. 5.—Renal afferent vein of left side of body and its connections in newly-born Anaconda (Hunectes noteus). Ant. Abd. Anterior abdominal veins; F'.B. Fat-bodies from one of which the right anterior abdominal seems to arise; Int. Intestine; P. Parictal branches of R.aff., renal afferent, which is seen to reach kidney anteriorly.
Proc. Zoou. Soc.—1906, Vou. I. No. II. 2
18 MR. F. E. BEDDARD ON THE (Jan. 16,
I have had the opportunity of comparing the structure of the kidneys in the two young Hunectes noteus with the structure of those of an adult male of the same species. There is in the adult no forward prolongation of a narrower region of the kidney. The organs are of the normal Ophidian form, of equal diameter throughout, and ending anteriorly in a blunt rounded extremity. That there may have been some microscopic traces in the tissues surrounding the sperm-duct is of course possible ; but there was nothing obvious to the naked eye. This state of affairs in the adult snake confirms, as I think, my opinion that the narrow anterior region of the renal organ in the newly-born young is to be looked upon as mesonephros. I may observe that the testes are very long bodies, and that that of the right side extends as far forward as the gall-bladder. The growth of the testis may account for the disappearance of the mesonephric portion of the renal organ. With the disappearance of the supposed mesonephros there is correlated, perhaps in this species but not in some other Boids, the disappearance of the posterior cardinal vein, which, as I describe in this paper, accompanies that gland in the young snakes.
Umbilical Vein.—The umbilical vein was not absorbed in either specimen. I was able to trace it along its whole course in the body from the navel onwards to the anterior end. The conditions which obtain in this snake show that Prof. Hochstetter’s discovery of the independence of the umbilical vein from the anterior abdo- minal of the adult applies to Hwnectes as well as to the reptiles (Lacerta and Tropidonotus) whose development he studied. There were two veins to be seen running from the umbilical aperture. The right-hand vein had the longest course, and is, I take it, the representative of the right umbilical vein. The second vein was traced along Meckel’s diverticulum to the alimentary canal, where it joined the portal system. This vein is, as I think, the omphalo- meseraic. The umbilical vein pursues a straight course between the two fat-bodies and over the gall-bladder. Arrived at the liver, it passes beneath this organ, between it and the ventral body-wall. There were no branches to be detected anywhere, and there was certainly no connection between the vein and the two anterior abdominal veins at any point that I could ascertain. Indeed it was easy to observe the umbilical vein running across the anterior abdominals. The vein is fairly closely attached to the ventral parietes in the region of the liver. It gives off no branches to the liver, with which it has no relations save those of superposition. At the extreme anterior end of the liver the umbilical vein joins the vena cava posterior just where the latter emerges from the liver. The two then run as one vessel to the heart. It is for the reason that the umbilical vein joins the vena cava that I regard it as the right-hand of the two primitive umbilicals. For in Hochstetter’s figure* illustrating the relation of these various veins
* Morph. Jahrb. xix. 1892, pl. xvii. fig. 15, V.u.d. and V.u.s.
1906. ] ANATOMY OF THE OPHIDIA. 19
in the feetal snake, the right-hand vein is depicted as joining the vena cava, while the left-hand vein is lost in the liver plexus. The umbilical vein was turgid with blood and of equal calibre with the efferent renals or other principal blood-vessels of the reptile. It is noteworthy that the umbilical vein until it reaches the region of the liver appears to run in an accurately median course. This clearly suggests that only one umbilical vein is present. In any case only this vein is obvious, unless the vein identified by me above with the ophalomeseraic be really the left umbilical.
In any case it is clear that this vein belongs to the fetal circu- lation, inasmuch as it passes through the navel to the fetal membranes, and that it has nothing to do with what are usually held to be the equivalents in the Ophidia of the anterior abdominal vein or veins in other reptiles. A remarkable fact about this vein is not merely its presence in the young when born and able to feed for themselves, but its persistence in the fully adult snake. In an Anaconda* (Hunectes murinus 3) dissected in May 1904, which was acquired by the Society in 1899 as an adult, I found a vein ending on the fat-body posteriorly which ran over the liver (2. e. ventrally of it), but did not draw blood from that organ anywhere, and emptied itself into the vena cava anteriorly shortly after that vein had freed itself from the liver. It is plain that this vein is that which I call umbilical in the young Anacondas. I have no record of such a vein in Python, nor is one figured by Jacquart + nor by Gadow ¢ in Boa.
It seems quite certain that this vein is the homologue of that vein in Birds which passes between the lobes of the liver, recurring in the falciform ligament. Hochstetter has proved § that the vein in question, variously termed ‘anterior abdominal,” ‘“ epi- gastric,” and “umbilical,” is the persistent umbilical vein of the embryo. It follows therefore that it cannot be the homologue of the anterior abdominal vein of the Lacertilia, which has been shown to bea new structure, having nothing to do with the fetal umbilical vein. Thereare therefore among Sauropsida two morphologically distinct veins or systems of veins which convey blood along the ventral surface from the posterior region of the abdomen to the liver or to the hepatic vein, and which are undoubtedly super- ficially similar, so much so that embryology alone has been able to decide the question of their distinctness.
The coincidence of these two veins in Hunectes solves the problem so far as concerns that species. At present so little is known of the venous system in the Ophidia, so few types have been examined from this point of view, that so far the Anaconda is the only snake in which the two forms of abdominal vein have been met with together. It is impossible therefore to build up
* Since this paper was written I have found the vein in a second adult male E. murinus.
+ Ann. Sci. Nat. (4) iv. 1855.
+ Bronn’s Thierreich, Bd. vi. Rept. Abth. iii. Schlangen.
§ Morph. Jahrb. xiii. 1888, p. 575.
a*
20 MR. F E, BEDDARD ON THE [Jan. 16,
much in the way of an attempted explanation without further facts. In the meantime the facts suggest that Hunectes offers a transitional state of affairs between the retention of the umbilical vein as the vein of the abdomen and its replacement by the sub- sequently developed anterior abdominal. It is exactly analogous to the relations between the posterior cardinals and the vena cava. In the primitive Ceratodus we have one cardinal persisting in its entirety and at the same time an undoubted vena cava posterior *. In higher types the posterior cardinals are more or less rudi- mentary, and the vena cava alone is concerned with the circulation of the region of the body formerly served by the cardinals.
Replacements of this kind are familiar to morphologists in connection with many organs.
It is not without significance, in my opinion, that this state of affairs has been discovered in a snake, and especially in an un- doubtedly primitive snake. That the Squamata form one group is probably the opinion of every zoologist at present. It is further clear that no existing group of Lizards is much nearer to the snakes than any other. The origin of the Ophidia must have been from some earlier type. This may land us some way back in the history of a group which with Hatteria appears to me to represent the archaic reptilian structure more than any existing group of Reptiles. It is possible that in the extensive fat-body of snakes we have the cause of the origin of the double anterior abdominal veins. The only fragment of evidence, however, which points to a large fat-body as a character of the ancestral Squamata is its large size in Amphisbenat. But this evidence is not to be neglected. The growth of the abdominal veins would render the umbilical superfluous, they taking on its function of drawing blood from the body-walls.
System of Anterior Venee Cave.—The veins of the anterior region of the body consist of four main trunks, which of course unite in pairs to form the two superior cave. The left tracheal unites with the left anterior vertebral and each right-hand vessel. It is noteworthy that the two anterior vertebral veins run super- ficially at equal distances from the median anterior vertebral artery. Each vessel lies in a furrow between longitudinal bands of musculature. The vertebral vein receives a large branch from the parietes just before joining the tracheal vein; the conjoined vein then receives just before its entrance into the heart the azygos, which is only present upon this side of the body. The azygos is short, and only collects blood from four intercostal spaces. Of the four branches which constitute it, one is especially large, and comes off exactly opposite to the point of entrance of the azygos into the Ductus Cuvieri. In the second specimen the azygos was much the same.
Epigastric Vein.—This vein has already been referred to in
* See W. B. Spencer, in “The Macleay Memorial Volume”’ published by the Linnean Society of New South Wales, 1893.
+ v. Bedriaga, Arch. f. Naturgeschichte, Jahrg. 50, 1885, pl. iv. fig. 2, Fk.
1906. ] ANATOMY OF THE OPHIDIA. 21
connection with its perforation by the umbilical aperture. It is as well developed as in the Ophidia generally, and lies, as in other snakes, immediately above the middle line of the body. Its posterior connection I did not see in either of the specimens. In the region of the liver, this vein has two connections with the hepatic portal system, and two only that were visible, which were identical in both specimens and which therefore may be looked upon as distinctive of the species. One branch from the vein joined the main portal stem a little behind the point where it reaches the liver. The other branch of the epigastric connected with the hepatic portal system lies a long way anteriorly and enters the liver-substance approximately in the middle line, a very little way behind the anterior termination of the liver. I saw no other veins passing from the epigastric to the liver. But in an adult Hunectes murinus there were four or five veins entering the liver between these two. It may be that the existence of two only in the young Hunectes noteus is a mark of immaturity.
Afferent Renal Veins—The caudal vein emerging from the tail receives branches from the cloaca, of which I am not able to give a particular account, as this region got damaged in displaying the course of the main trunk of the vein. The vein runs close to the large intestine on the left side, and receives in its course to the left kidney a series of veins from the parietes to the left of the dorsal median line (text-fig. 5, p.17). I counted altogether twelve of these before the vein enters the kidney, of which it is the afferent renal. On the right side the afferent renal seems to have no direct connection with the caudal vein, but the anastomoses between veins in this region are so numerous that there is doubtless an actual connection between them. Still there is no such direct. continuity as obtains on the left side. It is also to be remarked that the right renal afferent vein is a very distinctly smaller vein than its fellow of the left side, and in correspondence with this the affluents from the parietes which join it are much less conspicuous than those of the left side.
The figure (text-fig. 5, p. 17) shows the series of veins from the parietes which enter the left renal afferent vein. And an inspection of this figure will render unnecessary a detailed de- scription of the veins in question. It may be remarked, however, that there is a tendency for them to be connected together by a longitudinal trunk which is not present throughout the whole extent of the series, but gets to be more pronounced anteriorly. These parietal veins moreover vary in importance, some being much more slender than others. The most prominent of the whole series enters the afferent renal just before the latter enters the left kidney. This vein is connected with a very conspicuous longitudinal trunk (text-fig. 6, p. 23) which runs along the whole length of the kidney, being stouter at the two ends of that organ and more slender in the middle; it is, however, nowhere deficient. There is a general correspondence between the numbers of these
22 MR. F. E, BEDDARD ON THE [Jan. 16,
parietal veins on the two sides of the body; but, as has been remarked, the most noteworthy difference is that the veins of the right side are more slender than those of the left, and this applies also to the section of the parietal system which runs along the outer border of the kidney.
In the second specimen, the longitudinal vessel running to the outside of each kidney was not marked, at least on the right side of the body. ‘The same large vein enters the afferent renal just before the latter reaches the kidney. Towards the anterior end of the kidney the veins of five or six intercostal spaces join together and form a single trunk, which does not extend beyond the kidney but appears to plunge into the substance of that organ, There is thus an accessory renal portal system formed which is exactly like that which characterises Amphisbena*. These vessels are represented in the first of the two specimens which I dissected; but in that individual they are connected with the thinner anterior section of the kidney and with a vein which runs along that region of the kidney. In the second specimen, the vein distinctly opens into the posterior thicker region of the kidney and some way behind its anterior termination, In this example, moreover, the right afferent renal is traceable back along the intestine for a much shorter distance than in the other specimen. The posterior continuation indeed assumes the form of an in- conspicuous branch of the stout parietal vein which joins the renal afferent just before its entrance into the kidney.
The afferent renal trunk is also partly fed from the intestinal walls. Slender branches enter the transversely running affluents of that vein.
it is evident that the renal afferent system shows the same asymmetry that we see in the hepatic-portal system ; that is to say, that the system of longitudinal parietal vessels connected with the afferent veins are developed only upon the left side of the body.
Afferent Suprarenal Veins.—These veins arise, as in other reptiles, from the dorsal parietes, near to the middle, and are a continuation of the series which supply the liver in front and the kidneys behind. They are, however, unlike the hepatic series, developed upon both sides of the body. These vessels are con- nected with a slender vein which runs from the anterior end of the broader region of the kidney along the ‘‘ mesonephros” to the neighbourhood of, and behind, the gall-bladder. This vessel, which exists in other reptilest, is, as [ think, a portion of the right posterior cardinal, the corresponding vessel on the left side of the body being of course the left cardinal vein. I am not positive that this vessel is continuous all the way along the mesonephros ; but it is to be found at any rate for considerable stretches. Into this longitudinal vein open the afferent vessels from the parietes. I counted five of them altogether, of
* P.Z.S. 1905, vol. ii. p. 485. t+ E.g. Amphisbena, cf. Beddard, P. Z.S. 1905, vol. ii. p. 486.
1906. ] ANATOMY OF THE OPHIDIA. 23
which two at any rate are fed each from two intercostal spaces. Inasmuch as the actual suprarenal body does not extend along
Text-fig. 7.
Text-fig. 6.
MN Sy eS iii
Text-fig. 6.—Kidneys of newly-born Anaconda (Hunectes noteus), showing their lobulation and veins (a & a), which run parallel with each, receiving affluents from body-wall. L.R.aff., R.R.aff. Left and right renal afferent veins.
Text-fig. 7—Liver and portal veins of young Anaconda (Hunectes noteus).
L. Liver; P. Portal vein running along surface of liver and receiving, p, branches from parietes, which unite on their own account to form a continuous lon-
gitudinal trunk, and branches, St., from stomach.
24 MR. F. E. BEDDARD ON THE [Jan. 16,
the whole of the mesonephros, and as the blood-vessels arising from the parietes do extend along at any rate very nearly the whole of the mesonephros, as does the longitudinal vein into which they pour their contents, it would appear that these suprarenal afferent portals are also concerned with the blood-supply of the mesonephros, vid the remains of the posterior cardinal. The suprarenal portals are thus not veins especially destined for the suprarenal circulation, but originally merely the parietal branches of the cardinal. On the left side I found only three of these veins.
Precisely the same series of modifications appear to have pro- ceeded in the case of the liver and of the parieto-hepatic portals. In the Anaconda for example, and for the matter of that in all snakes, as it appears, that have been hitherto examined ana- tomically, the portal vein extends along the liver nearly to its anterior end. This is shown plainly in the figure of the circulatory system of the Python given by Jacquart*. Into this portal, which runs along the lower surface of the liver, open all or most of the vessels bringing blood to the liver from the parietes. In the same way in certain Lizards (for instance, Amphisbena, Ophisaurus, and Hatteria) there are at least considerable traces of the same forward extension of the portal. Finally we get the stage which characterises the majority of the Lacertilia, so far as existing knowledge allows us to say, in which the portal enters the liver at its posterior extremity and is not continued forward as a continuous trunk. In these lizards the parieto-hepatic portal veins enter the liver directly, instead of indirectly through the portal vein.
Dorsal Parieto-hepatic Veins.—These veins are entirely developed upon the left side of the body in both specimens. They are, as in other snakes, very highly developed, and a great portion of the blood of the whole body must be contained in them. I describe them only in one specimen; they appeared to be much the same in the other. The first of these veins, advancing from behind forwards, joins the portal vein about on a level with the extreme end of the liver, one lobe of which reaches considerably further back than the other. It is one of the largest of the dorsal parieto- hepatic veinsand on reaching the neighbourhood of the body-wall divides into a forwardly running and a backwardly running branch. Just before this division the vein receives twigs from the stomach. The backwardly running branch supplies seven intercostal spaces. The forwardly running branch supplies nine intercostal spaces before the second trunk arises which joins the portal vein just in front of the end of the shorter liver-lobe. Asin other snakes, the portal vein runs superficially along the liver, giving off twigs right and left to the liver itself and receiving the dorsal parieto-hepatic vessels. Of these vessels (see text-fig. 7, p. 23) I counted nine in addition to the two that have already been described. At their
* Ann. Sci. Nat. (4) iv. p. 321.
1906. ] ANATOMY OF THE OPHIDIA. 25
dorsal extremities these vessels are put into communication with each other by a continuous longitudinal vessel which is an extension forwards of the first dorsal parieto-hepatic already mentioned. Some, if not all, of the dorsal parieto-hepatic veins (which vary considerably in calibre) are joined by branches from the stomach. In addition to these special branches reach the stomach from the parietal longitudinal trunk. Although the parietal longitudinal trunk which collects blood from the parietes and transmits it to the liver runs on the left side only and has no fellow on the right side, it receives intercostal twigs from the right side. _It is to be noted that the continuous longitudinal trunk lying on the parietes is characteristic of the Boide in general, though I am not able at present to assert that it differentiates them from other snakes.
Note upon certain Structural Differences between the Species of Anaconda, Kunectes murinus ard Kunectes notzeus.
Hitherto the species Hunectes noteus, the Southern Anaconda, has been distinguished from its ally the more common form by its lighter colour and by the markings generally, into which I have no occasion to enter, as they have been dealt with by its describer, the late Prof. Cope *.
I have found in dissecting examples of the two species that there are certain differences in the viscera which seem to distinguish also the two species from each other. Upon one of these differences T lay more stress than upon the other, because I have been able to verify it in two examples of Hunectes murinus and three examples of Hunectes noteus, including among the three the two young specimens which form the subject of the present communi- cation. ‘This difference concerns the pancreas and spleen. In Eunectes noicus the pancreas is a large bilobed gland which lies in close contact with the duodenum and remote from the spleen, which is on a level with the front end of the gall-bladder.
In two specimens of Hunectes murinus I find the following arrangement of these two viscera:—In both the pancreas 1s divided into two parts, of which one is situated, as is the whole pancreas in H. noteus, close to the intestine. There is also another piece of pancreas lying in front of and in contact with the spleen, which itself has much the same position that it has in £. notceus. The duct from the anterior part of the pancreas runs to and buries itself in that piece of the pancreas which lies in juxta- position to the gut. In addition to this difference, I find in both specimens of H. mewrinus one or two splenculi in the neighbourhood of the spleen and anterior portion of the pancreas. I presume these to be splenculi on account of their colour and general appearance. A dissociation therefore of the pancreas into discrete portions is accompanied by that of the spleen. Upon this anatomical difference I lay some stress, inasmuch as it is to be found in more than one example of each species.
* P, Acad. Nat. Sci. Philad. 1862, p. 70.
26 MR. F. E, BEDDARD ON THE [Jan. 16,
Another apparent difference I mention, although I have only noted it in one specimen of each of the two species under consider- ation, as it could hardly be ascertained in the case of the newly-born young. This fact concerns the gall-bladder. In Humnectes noteus the gall-bladder gives rise to two ducts lying side by side and arising independently from the bladder, These branch and anastomose in a moderate way with each other and with the hepatic duct. The latter is single, but in the neighbourhood of its junction with the cystic ducts it gives off twigs which forma network alongside of the main duct which is obviousassuch. Threeducts pierce the pancreas on theirway to open intothegut. In #. murinus, onthe otherhand, four cystic ducts arise from the bladder and form in the same way but a slight rete. There are, however, two distinct hepatic ducts running side by side from the liver. Three of the cystic ducts join almost immediately to forma single duct, so that the difference from #. noteus is not so very marked.
In showing difference in the structure of the pancreas between different species this genus is like Python, where the discrete multilobate pancreas of “ P, bivitiatus”* contrasts with that of some other species.
Résumé.
It may be convenient to briefly recapitulate the main facts in the foregoing pages.
(1) The young of Hunectes noteus are produced alive and with considerable remains of the yolk-sac.
(2) They possess a mesonephros quite continuous with the kidney, and (female) a mesonephric duct extending along the greater part of the mesonephros and reaching beyond it anteriorly.
(3) The umbilical vein of the newly-born young persists in the adult (Hunectes murinus) and is quite independent of the anterior abdominals.
(4) The anal “claws” in Hunectes noteus (and in some other Boide) offer distinct sexual characters, differing in form in the two sexes. These characters are recognisable in the newly-born young.
(5) The suprarenal portal vessels open into a continuous slender trunk running along the mesonephros and ending on the kidney posteriorly. Their “ portal” character is thus secondary.
(6) The vessels which collect blood from the parietes and join the portal systems, whether of the liver, kidneys, or suprarenal bodies, are mainly, and in the case of the liver exclusively, affluents of the left-hand system of veins and arise exclusively or almost so from the left parietes.
(7) Both the liver and the left kidney are supplied from a continuous longitudinal parietal vessel which receives the branches from the body-wall and transmits the blood by a series of branches to the portal vein or to a longitudinal vein connected with the
* See fig.in Bronn’s Thierreich, Bd. vi. Abth. iii. pl. exxxii. fig. 6 (copied from Poelman). The identity of the species seems uncertain.
1906. | ANATOMY OF THE OPHIDIA. 27
kidney. These longitudinal parietal vessels are upon the left side of the body.
(8) As in Hunectes murinus, the left anterior abdominal trunk only is connected with the afferent renal of its side*. The right-hand trunk arises from a plexus on the gut.
(9) The azygos vein is short and developed on the right side only.
(10) Attention is called to anatomical differences between the two species of Hunectes, which concern the division of the pancreas and spleen in #. murinus, these organs being in one piece in E. noteus.
(2) Some Notes upon the Venous System of Python sebe.
It might perhaps be supposed that after the apparently exhaustive survey of Jacquart t hardly anything concerning the venous system of Python remained for description. Nevertheless M. Jacquart has not dealt fully with a few points of which the importance was perhaps less apparent at the time when he wrote than at present. Since M. Jacquart’s memoir, which is abun- dantly illustrated, nothing concerning the vascular system in this genus has been published except a few notes by myself = ina paper dealing mainly with the arteries of a number of genera of Ophidia. Those notes, however, refer to Python spilotes. My present communication refers, as did the memoir of Jacquart, to Python sebe. -
L have dissected during the past year or two three individuals of this snake, of which one only was specially favourable for the study of the venous system, owing to its fresh condition and the turgescence of the veins. It was possible in this individual to follow the smaller branches of the veins with ease, and no injection could have produced so favourable a state of affairs for examining the relations of veins.
Having so recently studied in detail the anatomy of the venous system of the Anaconda §, my object has been to compare and is to set down the differences and resemblances between these two genera of Boidse—types as they are of the two subfamilies, Boine and Pythonine, into which systematists have divided the family. The validity of comparisons having a purely classificatory aim is of course to some extent affected by the undoubted fact that species of the same genus among the Boide may show rather important differences in their veins, as I have been able to demonstrate in the case of Hryzx ||.
Afferent Renal Veins and Posterior Cardinals.—In Amphisbena 4
* It is necessary to emphasise this agreement between two species of the same genus, since in Eryx there are specific differences in this respect between Hryx jaculus and E. conicus, as | have shown (P. Z. 8. 1904, vol. ii. p. 119).
+ Ann. Sci. Nat. loc. cit.
+ P. Z.S. 1904, vol. 1. p. 362.
§ See above. || P. ZS. 1904, vol. ii. p. 107.
@ P. Z.S. 1905, vol. ii. p. 485.
28 MR. F. E, BEDDARD ON THE [ Jan. 16,
and Hunectes* I have described a vein running along the sperm- duct and receiving branches fromthe parietes which are the equivalents of the supra-renal portal vessels of other Reptiles. This vein in the two genera mentioned is no doubt the equivalent of a vein described and figured by Hochstetter in Varanus T which also accompanies the sperm-duct. In both Amphisbena and Hunectes, as it appeared to me, the vein gradually died away anteriorly and arose posteriorly from the substance of the kidney, being not in any direct communication with any of the principal longitudinal veins of the body. It appeared to me nevertheless to be probably the homologue of the posterior cardinal veins of the embryo, diminished in size and functionally replaced by the renal efferent veins or ven cave posteriores or inferiores. It was the general relations of this vein on either side to the mesonephric region and to the parietes which led to this opinion. The conditions which obtain in Python sebe amply confirm this point of view, and, as I think, settle the matter as certainly as it can be settled in the absence of embryological data.
The surface of the kidney shows in this specimen the course of the various vessels which traverse it in the clearest fashion. The single renal artery comes to lie on the kidney close to the anterior end, and can be traced back beyond the kidney to the ureter. The efferent renal vein begins quite near to the posterior end of the kidney, and runs forward, increasing in volume. It is quite distinct, of course, from the afferent renal, which runs not only to the extreme anterior end of the kidney, diminishing in volume as it proceeds, but is continued beyond ; it becomes, in fact, the vein which I have already spoken of as the probable equivalent of part of the posterior cardinal. The anatomical facts which have just been detailed seem to me to prove that this inter- pretation of the vein is the just one. It is quite possible that the examination of rather better material of the two genera to which I have referred in comparison with Python might show a continuity in their case also. In any case it is quite obvious in Python sebe and beyond the possibility of error. The only instance of this forward extension of the afferent renal in another snake (Zamenis gemonensis) has been recorded by myself =, where, however, it is short and plunges at once into the body-wall, this portion being, of course, the equivalent of the parietal branches of Python and Hunectes.
Umbilical Vein.—I have pointed out that in Hwunectes the umbilical vein is not merely a vein of the fcetal circulation, but that it persists in the adult, at least in Hunectes murinus, where T noted the existence of the vein before the study of the newly- born LHunectes noteus enabled me to fix its homologies. It becomes, therefore, a matter of interest to enquire how far this vein is represented in other Snakes. I have found in Python
* See above p. 21. + Morph. Jahrb. xix. 1892, pl. xvi. fig. 17. {f P. Z.S. 1904, vol. ii. p. 117.
1906. ] ANATOMY OF THE OPHIDIA, 29
sebe what I believe to be the remains of this vein. Along the course of the vena cava posterior, about midway from the anterior termination of the liver and the entrance of the vein into the auricle, is a branch of the vena cava, which is of some thickness and runs for a very short distance ventrally. Here its calibre lessens abruptly, and it becomes continuous with slender veins which run to the ventral parietes and are a part of the epigastric system. It is the large size and the sudden alteration in the calibre of the part of this affluent of the epigastric which joins the vena cava which lead me to infer that this end portion is the persistent umbilical. It is, however, further away from the liver than the umbilical vein is in Hwnectes. I am not able, therefore, at present to do more than suggest the homology which further facts may substantiate *.
Anterior Abdominal Vein.—The condition of this vein in Python sebe presents one feature of considerable interest. In an earlier communication T I pointed out that the observations of Jacquart as well as myself tended to show that it is only among the Boine Snakes that the anterior abdominal vein is directly connected with the renal afferent veins, as they always are among the Lacertilia. In Humnectes (both H. murinus and EH. notewus) it is only one of the two roots of the anterior abdominal which is thus connected, and that of the left side. The right origin of the anterior abdominal is from a plexus of venules upon the gut. 1 quoted Jacquart to the effect that this also is the arrangement in Python. Waving examined Python sebe with great attention in regard to this important matter, I am able to state that that snake differs from Hunectes and agrees with Hryx jaculus in that both roots of the anterior abdominal are connected each with one of the two renal afferents. The veins in question can be easily observed on dissection to lie dorsally of the forwardly-directed rudiment of the pelvic arch, which thus conceals a portion of each vein when seen after the serpent is opened in the usual way from the ventral surface. Python, at least P. sebe, is therefore quite Lacertilian in the disposition of these vessels, and the opinion that the Boide generally come nearer to the common starting- point of both Ophidia and Lacertilia is strengthened by this observation.
In view of accumulating facts concerning the venous system of Snakes, it is important to notice all variations so as to arrive at the normal characteristics. I may therefore mention that, as Jacquart and I have previously stated, the anterior abdominal of Python sebw divided to form two tubes during its course. In the individual upon which I report here, the division commenced at a point about opposite to the middle of the left kidney. The tubes reunited behind the gall-bladder. The anterior abdominal, as in other individuals, unites with the portal a little way in front of the gall-bladder It is to be
* See below, p. 35. + P.Z.S. 1904, vol. ii. p. 116.
30 MR. F. E. BEDDARD ON THE [Jan. 16,
noted, therefore, that Python differs from Hunectes in that the anterior abdominal of the latter remains double for some distance forwards after its origin, while in Python the roots join at once to separate and rejoin later.
Azygos Vein.—As in Eumectes, the azygos vein of Python sebe is retained upon the right side only, but it is less extensive in the former than in the latter snake. The azygos vein just before it enters the right heart receives a stout branch running up verti- cally from the parietes and a thinner branch coming from the neck-region. Posteriorly the vein passes backwards, and soon divides into two trunks which are thin. The more dorsal of these runs close to the body-wall and receives twigs from each intercostal space. The upper branch runs in close connection, or rather contact, with the lung, for it does not appear to receive any branches from that viscus. After the termination on the parietes of the lower branch of the azygos, this upper branch bends down and supplies, or rather receives, blood from the inter- costal spaces some way down the body, though not so far as the region of the hepatic parietal vessels, which will be dealt with immediately. It takes up blood from the next intercostal space to that which furnishes the last branch to the lower of the main branches of the azygos. Thereis thus no break in the circulation of this region of the body.
Vessels of Neck.—Although the facts concerning the vessels of the neck have been made known by Jacquart, 1 may take this opportunity of pointing out that in their disposition there is an obvious difference from the corresponding vessels of Hwnectes. In the latter snake all the four typical vessels are present *, viz., two running along the trachea, and two anterior vertebrals ; whereas in Python (at least in P. seb) there are only three of these main venous trunks present, the left vertebral being absent. There is also a disproportion between the two tracheal or jugular trunks, the left being very much smaller in Python. T did not observe such a difference in Hwnectes. This series of facts shows that it is not possible to place the two genera of Boide in ascending relations to each other in respect of the venous system; for while L’wnectes is more primitive in the arrangement of the main veins of the neck, this genus is less primitive in the fact that the anterior abdominal has only one posterior connection with the veins of the posterior region of the body, whereas both are present in Python.
Hepatic Portal Veins.—These veins are constituted in Python sebe upon the plan which is to be found in Hunectes and Hrya. In all of these genera the parietal system (7. e. longitudinal veins running along the parietes and collecting the blood from the several intercostal branches before rendering it up to the liver circulation) is much developed. There are, however, differences in the way in which this system is formed in these various Boine
* Only three are figured by Gadow in Boa madagascariensis (loc. cit. pl. cxxxv.).
1906. ] ANATOMY OF THE OPHIDIA. 31
Snakes. I have already dealt with Hryx* and Hunectest. I do not find that Jacquart’s illustrations conform absolutely to my own observations; but it may be, of course, that this system shows some variations. I find that, as in other Boide, the system of longitudinal parietal vessels is chiefly developed upon the left side of the body, but not so exclusively so in Python as in Hunectes. The left parietal vessel commences posteriorly at about the end of the lung, that is to say very considerably behind the liver. Anteriorly it ends at about the level of the bifurcation of the aorta into its right and left moieties.
Between these two points the vessel is almost, if not absolutely, continuous. It receives the intercostals as they emerge from the parietes, and sends off branches to the liver and to the alimentary canal in this region. These branches are at least partly in- dependent of each other; that is, separate branches pass to each viscus or perhaps rather from one to the other. The veins which pass to the liver enter the anterior extension of the portal vein lying upon the liver, and there are at any rate nine of them. Anteriorly the longitudinal parietal ends in a bifurcation, and behind this there is another branch, which, like one half of the anterior bifurcation, lies upon the right side of the body, distri- buting its branches to the intercostal spaces of that side of the body. In Humectes, although the left longitudinal parietal vessel is the only one developed, it receives twigs from both sides of the body, but there is no development of a longitudinal trunk or trunks upon the right side. As in other Snakes, a particularly strong branch from the longitudinal parietal passes up to the portal trunk just before the latter reaches the liver.
The main points in the foregoing notes upon the veins of Python sebe to which I desire to call attention are the following :—
(1) The double origin of the anterior abdominal vein, one root from each renal afferent vein, as in Arya jaculus and in the Lacertilia without exception.
(2) The prolongation of the renal afferent vein beyond the kidney along the sperm-duct, this region being, like the renal afferent, a persistent posterior cardinal.
(3) The existence of a trace of the umbilical vein, which persists in its entirety in Hunectes.
(4) The paired main trunks of the neck distinguish Hunectes from Python, where, as has been also shown by others, there are only three trunks and the paired jugulars are not symmetrical in size.
(3) Some Notes upon Ilysia scytale.
The following notes are the result of the examination of a single specimen of Jlysia seytale which has been in my possession for some time.
# P. Z.S. 1904, vol. ii. p. 118. + Above, p. 24.
32 MR. F. E, BEDDARD ON THE [Jan. 16,
It measures 19? inches in length.
The base of the heart lies 44 inches from the tip of the snout. The liver, which is 7 inches long, commences just at the heart as in Vipers. The gall-bladder lies 14 inches behind the tip of the snout. The kidneys are elongated, and not particularly shortened as in Eryx. The right kidney, 23 mm. long, ends 21 mm. in front of cloaca. The left kidney, 21 mm. long, ends 11 mm. in front of cloaca. The right testis begins 10 mm. behind the gall-bladder; the left testis begins 9 mm. behind end of right testis.
The lung of Jlysia is single*, and the vascular part, which is of considerable thickness, extends for a considerable way down the liver, being some 2} inches in length. The windpipe, as is usual, is formed of incomplete rings, a fibrous fold connecting them along its entire length. Just before the trachea opens into the lung there is a minute orifice which represents the 2nd bronchus; but there is no approximation to an equality between the two bronchi such as exists in the Boine snakes. The end of the trachea, that is, of course, of the only functional bronchus, extends some little way into the lung before it disappears. This disappearance is not quite abrupt; the rings of the bronchus cease just before the end to be circular, transversely-arranged cartilages ; they anastomose with each other, and finally assume a honeycomb disposition, precisely like the lining-membrane of the ensuing lung. Still the bronchial region can be distinguished from the pulmonary by its bluish colour.
Alimentary Canal.—The condition of preservation of the spe- cimen which I have dissected, and the comparatively empty alimentary canal, render it possible to give an accurate account of the ruge and plications of the different regions, which is not always so easy. There are, moreover, obvious differences between Llysia and some other snakes, both in the structure and pro- portions of the several regions of the alimentary tube. The wsophagus extends to the posterior end of the liver, where it more or less suddenly passes into the stomach. Internally the demar- cation is quite abrupt. It is shown, in fact, by the different nature of the folds of mucous membrane which line the two sections of the anterior part of the alimentary canal. The stomach in its anterior part is lined by three, and three only, thick longitudinally-running folds. At the junction of stomach and cesophagus these thick folds disappear as such, and are either nearly or quite continuous, with at least six similar but much smaller folds. There is thus a perfectly obvious demarcation between cesophagus and stomach. The stomach itself is 56 mm. (or about 2 inches) long, and is plainly divisible into two regions : the first of these is much the larger and measures 48 mm.; the second is about coextensive with the gall-bladder, which is attached to it and measures 8 mm. The larger anterior region of the
* As Mr. Butler (P. Z.S. 1895, p. 704) and others have noted.
1906. | ANATOMY OF THE OPHIDIA. 33
stomach is, as already mentioned, traversed by three thick folds, which show here and there an interesting trace of a reticular arrangement. There are occasional short branches of the folds, which, although they do not reach another fold, indicate an approach towards, or a reminiscence of, a reticular arrangement. Towards the posterior end of the anterior part of the stomach the folds lose their importance and smaller folds between them appear. But the whole series of folds dies out upon a cushion- like elevation. Then follows the shorter posterior region of the stomach, which is entirely free from ruge of any kind. The small intestine is short as compared with that of some other serpents. Not counting its windings, it measures in a straight line from the end of the stomach to the beginning of the large intestine 51 mm., or 2 inches. About half of this consists of a closely-folded section, which commences immediately after the circular valve separating the stomach from the intestine. Then follows a straight section, which just before its opening into the dilated commencement of the large intestine bends once or twice abruptly upon itself, thus constituting another shorter coiled region. The intestine is lined with fine wavy folds. The large intestine is also 51 mm. in length to the anus.
Vascular System.—Though it is impossible to elucidate the arrangement of the vascular trunks in a spirit-preserved specimen in a thorough fashion, I have nevertheless been able to ascertain certain facts which are of importance from a systematic point of view. Ihave been able to follow the abdominal branches of the aorta, and find that the first hepatic branches are disposed as follows:—The first branch which arises supplies the stomach, and reaches that organ at about its middle. On a level with the gall- bladder a second equally stout branch is given off which divides into two trunks: one of these runs forward and is gastric ; the other has a posterior course and is intestinal. The next branch of the aorta is that of the right testis; it is of course very slender. T did not discover the corresponding branch to the other testis. The next artery to arise is the right renal, which enters the kidney near to its anterior end. A rectal artery comes next and is followed by the left renal, which is also the only artery of this kidney and enters it near to its anterior end. Then follow three rectal arteries.
Intercostal Arteries—In the arrangement of these arteries Tlysia plainly shows its affinities to the Boide. In the last- mentioned family * the intercostal arteries are paired structures, which is not the case with, at any rate, many Colubride and Viperidee. In Jlysia the same paired structure is plainly visible; furthermore, it is to be noted that the intercostal arteries are regular in their arrangement—that is, that they supply the inter- costal spaces continuously, a pair of arteries corresponding to each vertebra. This is, of course, also a Boine character. It is
* Jacquart, Ann. Sci. Nat. (4) iv.; and Beddard, P. Z. S. 1904, vol. i. p. 388, & P. Z.S. 1904, vol. ii. p. 108. Proc. Zoou. Soc.—1906, Vou. I. No. III. 3
34 MR, F, E. BEDDARD ON THE (Jan. 16,
important also to note that the intercostal arteries arise from the aorta as single trunks and bifurcate later, The arrangement of these arteries, in fact, is precisely 4s in Python, as figured by Jacquart.
Among the Lacertilia it is very general, if not universal, for the mesenteries to contain muscular strands, which in the case of the dorsal mesentery arise from the vertebral musculature and are often inserted on to viscera. These bands of muscle are made up in all cases where they have been examined of unstriped fibres. I find in /lysia a structure which is to be regarded as the equivalent of these bands of muscle in the Lacertilia. This consists of a single band of fibres of about the same calibre as one of the larger arteries—for which indeed, or for a vein, I at first mistook it. The nacreous glitter, however, of the strand shows its muscular nature, which appearance was confirmed by microscopic examination. The fibres of which it is composed are plain fibres. This band of muscle arises from the centra of two vertebra by strands from each near to the posterior end of the liver. It runs obliquely forwards, and ends in an attachment tothe upper surface of the liver. It avoids the alimentary canal, passing to the right of that tube. The existence of only this one straight band of fibres of unvarying calibre throughout is different in detail from what is found among the Lacertilia.
From this necessarily imperfect survey of the anatomy of Llysia, it is possible to abstract two characters in which this genus, placed in a Family of its own, resembles the Boide, and one other in which it departs from the structure of that group.
Liysia agrees with the Boide in the following points :—
(1) In the regularity and paired character of the intercostal arteries. 2) In the fact that each kidney is supplied by a single renal ; y Py y s artery.
On the other hand, /lysia differs from the Boide (excepting Ungalia*) chiefly in the fact that the lungs are not normal paired structures but that one is rudimentary.
The other features which I have been able to note in this survey of certain points in the anatomy of this genus are, in the present state of our knowledge, indecisive of the affinities of the genus.
(4) Zhe Structure of the Young Bitis nasicornis, with Notes on other Vipers.
A considerable number of young Vipers of this species were born in the Gardens in November last, some of which were alive, whilst some had been born dead. J examined several of the latter. The young snakes were enveloped in the amnion, and the other
* See Cope, On the Lungs of the Ophidia,” Proc. Amer. Phil. Soc. xxiii. 1894, E. 220
1906. ] ANATOMY OF THE OPHIDIA. 35
foetal membranes were in much the same condition as in the ease of the newly-born Hwmnectes noteus described above. The “ navel” (text-fig. 8) is, however, in this Viper much smaller than in the Anaconda, The actual patch of skin uncovered by scales only occupied the extent of two of the ventral scales, which were here divided into two. On either side a small number of the ventral scales were divided in the middle line, but there was no interval between the two halves of each scale. Moreover, only two scales anterior to the navel and four behind it were thus divided. The conditions, therefore, are distinctly different from those obtaining in Hunectes. Furthermore, the position of the navel differs. In Bitis only 10-14 scales intervened between the navel and the cloacal orifice: the actual distance was 14 mm., the whole snake measuring some 92 inches*, The embryonic veins were apparent,
Text-fig. 8.
Region of umbilicus in newly-born Bitis nasicornis. U. Umbilical sac.
as in Hunectes. The umbilical vein can easily be traced from the “navel” to the vena cava inferior, which it enters towards the anterior extremity of the liver. As in Hunectes, this vein has no relation whatsoever to the anterior abdominal vein that I could discover. It has, however, an obvious connection with the epi- gastric vein, which leads me to introduce the matter here, not as a contribution to the anatomy of the Vipers, but as explanatory of the anatomical facts which I have just referred to in the Python *. Close to the union of the umbilical vein with the vena cava a branch ascends from the epigastric to join the umbilical. This seems to me to be a fact supporting the inference which I have put forward
* In Vipera berus also the navel is close to the cloacal aperture + Above, p. 29. 3”
36 MR. F. E, BEDDARD ON THE [Jan. 16,
as to the partial persistence in the adult Python of the embryonic umbilical vein.
Trachea and Lung.—tin this Viper the tracheal lung is not in any way marked off from the ensuing thoraco-abdominal portion of the lung. The latter extends as an efficient respiratory organ for some distance down the liver. The tracheal or bronchial gutter ends at a level with about the middle of the heart. There is no trace of a second lung. 1t is necessary to emphasise these various facts, since they differ among the Viperide. The total absence of a second rudimentary lung in this species has been already noted by Butler*. In addition to the species which he mentions, | may note here that Lachesis gramineus has a rudimentary left lung. In Ancistrodon piscivorus the lung is vascular for about an inch behind the heart, and the tracheal gutter is continued for about the same distance, being thus more extensive than in itis nasi- cornis. In Lachesis gramineus the vascular part of the lung is less extensive posteriorly. Cawsus rhombeatus differs in some respects from these other Vipers. Like all other Solenoglypha, it possesses the tracheal lung; but it differs from some other Vipers in the fact that the tracheal section of the lung is the only part which is vascular, the rest being anangious. The trachea is continued down the lung from the very beginning of the tracheal lung as a gutter down to near the end of the liver.
Intercostal Arteries.—In dealing with certain facts concerning the vascular system in Lachesis gramineust, | pointed out that, contrary to what is to be found in many Snakes, the intercostal arteries perforate the body-wall accurately in the middle line and singly. This feature, I am now inclined to believe, is distinctive of the Viperide, for I have since found a similar state of affairs in Causus rhombeatus and Ancistrodon piscwwvorus.
Veins and Arteries of Lung.—Dr. Gadow, in a figure{ of the pulmonary arteries and vein of Crotalus, represents these vessels as passing forward from the heart. This direction is of course in conformity with the tracheal lung of that snake, where possibly the respiratory organ lies entirely in front of the heart. In the young itis both artery and vein bifurcate. The artery bifurcates some little way after its origin, and the anterior branch supplies the tracheal lung, while the posterior branch supplies that portion of the lung which lies behind the heart. The pulmonary vein shows the same general distribution. There were two main branches, one anterior and one posterior. The direction of emergence from or of entry into the heart, as the case may be, was rather lateral than definitely anterior or posterior. I deal ona later page $ with these facts with reference to the original form of the lung in the Squamate reptiles.
Veins of Neck (text-fig. 9, p. 38).—Contrary to what is to be found in many Snakes, there are only two main veins of the neck in Bitis
* P. Z.S. 1895, p. 705. + P.Z.S. 1904, vol. i. p. 366. f In Bronn’s Thierreich, Rept. Bd. vi. Abth. iii, Taf. exxxvi. fig. 2. § Below, p. 41.
1906. | ANATOMY OF THE OPHIDIA. 37
nasicornis. Hach of these two is closely accompanied by an artery. The larger enters the heart in common with the azygos, and runs forward on the right side of the body as far as the head. It is much more slender in its most anterior region than near the heart. This vein gives off to the right a series (I counted seven in the most fully developed specimen) of branches to the parietes, which immediately plunge into the thickness of the body-wall after a very short free course. From the left arise a smaller number of veins which have a longer course. Of these I found not more than three. They arise at right angles from the main trunk, and cross the body to the left side of the dorsal median line. Here they appear to become continuous with a longitudinal trunk which runs a considerable way towards the head and posteriorly as far as a little way down the liver. This vein runs much further from the dorsal middle line than the right jugular and the right azygos. The branches, soon after leaving the jugular, send off a twig to the cesophagus.
The longitudinal vein of the left side of the body, supplied by branches from the right jugular, represents, as I imagine, the left anterior cardinal, the right vein being the persistent right anterior cardinal. Their mutual asymmetry with reference to the median lines of the body is remarkable. It is possibly to be accounted for by the pressure exerted by the trachea and lung pushing the vein further away from the middle line. The right azygos vein appears at first sight to be of less extent than a more careful examination shows it to be. In three of the specimens which I examined this vein, which is of considerable calibre, appeared to end at about the level of the ventricular apex; but in a fourth example it was distinctly continued back by a very slender prolongation to a point quite on a level with the middle of the liver. It is note- worthy that the azygos not only gives off branches to the intercostal spaces along which it runs, but that close to its origin—or rather debouchement into the auricle—it sends out a forwardly-running branch, which ought, so to speak, have been furnished by the anterior cardinal of that side of the body.
In addition to the right jugular, which is a vessel as thick as any in the body, and the rudimentary left jugular, which is prac- tically a branch of it, though morphologically, as I imagine, a distinct vein, an internal jugular runs along the trachea, This vein, like the right jugular or anterior cardinal, is closely accompanied by an artery. The artery, however, like the vein, is thinner than are the pair which run along the body-wall on the right side. Whether there is a corresponding and less developed vein for the opposite side of the trachea, I am unable to say.
In the arrangement of these anterior veins Bitis stands at the opposite pole from the Anaconda, whose anterior veins have been considered above. In the latter serpent all four veins, viz. both anterior cardinals and two internal jugulars, exist. In the Viper the reduction is striking. Intermediate conditions are offered by Boa, in which, according to Gadow’s figures, the veins of the neck
38 MR. F, E, BEDDARD ON THE [Jan. 16, are three. This reduction is in consonance with the generally
received opinion as to the much modified character of the Viperide as compared with other Serpents.
Text-fig. 9. Text-fig. 10.
Text-fig. 9.—Veins of neck of newly-born Bitis nasicornis. Az. Azygos vein; H. Heart; R.V.V., L.V.V. Right and left vertebral veins.
Text-fig. 10.—Veins in region of kidney of newly-born Bitis nasicornis.
a. Anterior termination of testis in a fibrous band; Z.K. Left kidney. Sr.v. Suprarenal portal vein; Sy. Suprarenal body; 7. Testis; V.R.aff. Afferent renal; V.R.eff. Efferent renal.
1906. | ANATOMY OF THE OPHIDIA. 39
Veins of Kidney, Suprarenal Bodies, and adjacent region.—The kidneys measure from 29 mm, to 32 mm. in length, and are divided into about 20 lobules. The afferent renal vein extends to within 3 mm. or so of the anterior end of the kidney, giving off a branch between each lobule. In one specimen, at any rate, the renal afferent vein gives rise to a large branch (see text-fig. 10, p. 38) which leaves the surface of the kidney some way before its anterior end, so that the rest of the renal afferent anterior to this point of origin appears to be a branch of this, the main trunk. This vessel is clearly a persistent posterior cardinal, which runs forward in close contiguity to the suprarenal gland. It extends beyond this gland and ends in two branches to the parietes. These latter seem to me to be the afferent suprarenal veins. There is furthermore another afferent suprarenal vein, which also joins the cardinal, but towards the posterior end of the suprarenal body. This vein collects blood from the lateral parietes, and runs parallel with the kidney. Whether it does or does not communicate with the afferent renal behind the kidney, I do not know. In another specimen this vein was quite as well or even better developed; but it was con- tinued directly into the vein running along the suprarenal body, which I have presumed to be the posterior cardinal vein of this side of the body. There was no connection, that I could ascertain, with the afferent renal.
The suprarenal veins are very conspicuous and two in number ; they run from the anterior part of the suprarenal body over the testis, and open into the efferent renal vein.
It will be seen that thé vein which runs along the parietes beside the kidney is precisely that of Hunectes described above™ ; but in Bitis I have not been able to ascertain the presence of a posterior connection with the afferent renal. It is, furthermore, plainly to be compared with the vein occupying a similar situation in Chameleo, which has been described by Hochstetter ap cnaxel myself ¢, and which I have figured §.
Hepatic Veins.—So many of the observations upon the veins of Reptiles have been made upon a single example, that it is not always certain how far the appearances described represent the normal. Iam therefore careful here to describe the course of the veins in all of the specimens of this Viper that I have had the opportunity of studying. The result is to show that the variations are not very great (so far, of course, as the small number of examples allows of such a statement), and that therefore the arrangement of these veins at any rate is of use for systematic purposes. The portal vein (text-fig. 11, p. 40) offers no remarkable features. It reaches the liver at the junction of right and left lobes, as in other Serpents ; and from that point until the anterior end of the liver it runs superficially, receiving branches from both the dorsal and ventral parietes. Of the dorsal parieto-hepatic veins, one is absolutely constant throughout the whole series of
* p. 22. + Morph. Jahrb. xix. p, 462. t P.Z.S. 1904, vol. ii. p. 8. § Loe. cit. fig. 14, p. 8.
bo bo
40 MR. F, E. BEDDARD ON THE [Jan. 16,
individuals. This vein reaches the portal shortly after it has reached the liver, collects blood from several intercostal spaces, and runs back to nearly the level of the posterior end of the liver. Itis apparently generally present in the Ophidia and is not merely a characteristic vein of the Vipers.
Text-fig. 11.
Portal veins of newly-born Bitis nasicornis.
Ep. Epigastric ; H. Heart ; L. Liver ; p. Parieto-hepatic veins, two of which receive branches from stomach (S¢.), represented as cut oft at the end; P.V. Portal vein; U. Umbilical vein; V.c.7. Vena cava posterior.
This vein received, in the most fully-developed specimen which I examined, two veins from the stomach. There were in this
1906. ] ANATOMY OF THE OPHIDIA. Al
example three other veins of the same kind, of which two received branches from the gut; there was also an independent gastro- hepatic vessel, as is shown in the accompanying figure (text- fig. 11, p. 40), which represents the liver-veins of this example. The epigastric vein sends, at any rate, two branches to the liver, which are very anterior in position: one of these receives a branch from the stomach before entering the liver. In another example there were only three dorsal parieto-hepatics. It was in this example that a branch from the epigastric joined the umbilical vein, as referred to above. In a third specimen, I saw only two parieto- hepatic vessels, arising, as in the others, from the left side of the dorsal median line. In this and other cases the differences may not be real, but due to absence of blood in the vessels at the time of examination.
The origin of the hinder mesenteric vein in the Ophidia has been variously stated, the different modes of origin described possibly corresponding to the different species and genera examined. Hochstetter* describes and figures Yropidonotus natrie (with which he finds Coluber wesculapiz to agree) as possessing a mesenteric vein which arises from both afferent renals, the two branches combining to form the single vein. T have not been able to ascertain to my satisfaction the arrange- ment of these veins in Bitis nasicornis; but in another Viper, Ancistrodon piscivorus, 1 have found that each renal afferent vein gives off a branch, and that these join to form the mesenteric vein running along the lower surface of the large intestine. The arrangement characteristic of this Viper is therefore precisely that of Tropidonotus and Coluber.
(5) Considerations respecting the Primitive Structure of the Lungs in the Squamata.
Hatteria (or, indeed, most Lacertilians) on the one hand, and such a snake as Causus rhombeatus on the other, represent the two extremes of modification of the Squamate lung. In the former the lungs are paired and equal, and are effective breathing-organs throughout : they are separated from the glottis by a long stretch of trachea, and by two equisized bronchi into which the trachea divides some way in front of the lungs. In the Viper, on the other hand, the trachea opens into the lung but a short way behind the glottis, down which it is continued as an open gutter; at, or about, the level of the heart the lung becomes anangious and is a mere air-sac; while there is no trace of a second lung, or of a division of the tracheal gutter into two bronchial tubes. It is undoubtedly the prevalent opinion that of these two extremes, that represented by Hatteria is near to the primitive Sauropsidan lung, while the lung of Oausus represents the most modified type. Paradoxical though it will appear, there are reasons founded upon anatomical
* Morph. Jahrb. xix. 1893, p. 489, pl. xvi. fig. 19.
42 MR. F. E, BEDDARD ON THE [Jan. 16,
facts which necessitate a reconsideration of this view, and which tend to destroy its apparent obviousness.
Cope *, to whom our knowledge of the headwards extension of the lung is mainly due, though the fact of this extension in the Viper was known fifty or sixty years earlier, terms this section of the lung the ‘tracheal lung,” and after a survey of the leading groups of Ophidians found it to occur in the principal subdivisions of the order or suborder. He found this tracheal lung in Ungalia among the Boide, ‘in the Solenoglypha without exception,” and in several Colubrines, to which I myself have added the Hamadryadt. The occurrence of these tracheal lungs so widely among the Ophidia suggests a retention of a character rather than its independent development in the several groups. So far, however, one can do no more than incline to the former view. There are, however, other facts. In the first place, among Snakes generally the rings of the trachea, where there is no tracheal lung, are incomplete posteriorly, leaving a gap filled in with soft tissue. This soft tissue is continuous with the lung-tissue where the latter commences, in these cases near to the heart.
it might be held—if the matter ended here—that the non- junction of the tracheal rings posteriorly had no more significance than the failure to join posteriorly of the tracheal rings in the Cassowary ¢ or in Man§. But a few cases seem to show that this failure to join is of meaning as the last term in a series. For in some Serpents, e.g. in Lioheterodon, there is not merely a failure to unite posteriorly among the tracheal rings, but the membranous space left is of wide dimensions, much wider than the actual trachea, and fully as wide as the tracheal lung where that organ is developed. Moreover, in this snake there are traces of a develop- ment of diverticula of the cavity such as are to be met with in a much more fully developed condition in the Hamadrya These facts therefore afford some evidence that the tracheal lung was formerly more widely spread among the Ophidia than it is now; that 1t is not a new structure in those forms where it occurs, but an archaic structure so far, at any rate, as Snakes are concerned.
Jt will be observed, moreover, that there is a distinct relation between the development of the neck part of the lung and the asymmetry of the lungs. This relationship, however, does not after all amount to a great deal; for the only Serpents in which there are a pair of well-developed thoracic lungs are the Boide. It is nevertheless noteworthy that among these primitive Snakes, as they are held to be, the genus Ungalia, which possesses the tracheal lung, is, like the Colubrine Snakes, without more than a rudiment of one of the lungs. The only allied form in which
this asymmetry of the lungs is known to exist is J/ysia; but in Tlysia there is no development of the tracheal lung. My object,
* Proc. Amer. Phil. Soc. 1894, p. 217.
+ P.Z.S. 1903, vol. 11. p. 319. t Forbes, P. Z. 8. 1881, p. 783. § Treatises on aman Anatomy.
|| Beddard, P. Z. 8. 1903, vol. ti. p. 319.
1906. ] ANATOMY OF THE OPHIDIA. 43
however, being rather anatomical than physiological, this mode of compensation—as it may be considered to be—will be left aside. For it is, I think, impossible to hold, in the present state of our knowledge at any rate, that the unpaired condition of the lung is the primitive one for Snakes, and that the minute rudiment of the second lung in many Vipers and Colubrines is an incipium of a second lung. Still the question does not appear to me to be absolutely settled, for reasons which I hope to investigate more fully later.
But although it may be said that there is some evidence that among the Ophidia the existence of a tracheal lung is not an innovation but an inheritance, the case would seem at first sight to be quite different among the Lacertilia. If the assumption that the Lacertilia form one order with the Ophidia, and the theory which I seek to prove concerning the origin of the lungs in the Squamata be probable, there should be evidence of a positive kind among the Lacertilia of the existence of traces of a tracheallung. The most positive piece of evidence is that furnished by Prof. Wiedersheim, who has described in Amphisbena fuliginosa what appears to be a persistent tracheal lung *, the existence of which, however, has not been confirmed for other speciest. It is noteworthy, too, that in various Lizards the tracheal rings are far from meeting posteriorly ; in Lacerta, for instance, there is a very wide membranous interval posteriorly, at the edge of which only appear the tips of the tracheal rings. Furthermore, in many Lizards—this is particularly well seen in Varanus—the lung ex- tends forward a good way beyond the entrance of the bronchi into the lung. The arrangement in such a lizard as Varanus is quite reminiscent of the disposition of that organ to be seen in Heterodon platyrrhinos, where the tracheal lung is not traversed by a tracheal gutter, but extends forward along the intact trachea as a con- tinuation forwards of the thoracic lung.
Were it not for the numerous cases of a tracheal lung attached to the trachea throughout, this condition in Heterodon would probably have been compared merely with the slight forward extension of the lung in many Lacertilians, in which the bronchus enters at the side rather than at the base of the lung. Such a comparison would indeed be correct, but it would not be so far- reaching as I believe there are grounds for regarding it. Besides, this incomplete comparison of facts, as I regard it, would leave it an open question as to whether the lungs in the Squamata were not derivable from the type shown in Hatteria, and to which a forward extension had been afterwards added. As it is, there are further facts which enforce the position taken up by me in this communication. I have pointed out, in describing the
* ‘Lehrbuch der vergleichenden Anatomie der Wirbelthiere.’
+ Beddard on Amphisbena, P. Z.S. 1905, vol. 11. p. 489.
t This statement of course assumes the validity of Prof. Cope’s view that the headward extension of the lung in Hetevodon is the homologue of the tracheal lung in, €. g., the Viperide.
44 ON THE ANATOMY OF THE OPHIDIA. [Jan. 16,
pulmonary artery and vein of Sitis nasicornis*, that both artery and vein bifurcate soon after leaving, or just before entering, the heart. One branch goes to, or comes from, the anterior tracheal part of the lung, while the other branch has a similar relation to the thoracic part of the continuous lung. It seems to me that this anatomical fact explains two other facts which have been a little difficult to me hitherto.
The trachea in Snakes, and in certain Lizards at any rate, is closely accompanied by arteries or an artery which is one of the systemic branches. This carotid artery is concerned with the blood-supply of the windpipe and adjacent organs and regions. In some cases, however (probably much more generally than I am at present in a position to know), the trachea is accompanied by arteries which arise not from the systemic arteries but from the pulmonary. Ihave shown this to be the case in Gerrhosaurus tT and more recently in Hatteriat. In both of these Saurians the artery in question is most clearly a branch of the pulmonary, and equally clearly lies alongside of the windpipe anteriorly. A careless dissection would fail to show this, as I consider it, highly important point. It is, however, plain when the artery is ‘properly followed out in an injected specimen.
Now the pulmonary artery is, it is hardly necessary to say, a respiratory artery: it is concerned, that is to say, not with the nutritive supply of the lung-tissue ‘but with the oxygenation of the blood. The tissues of the lung receive their nutritive supply from elsewhere. Branches from the aorta supply this need which have no relation whatever to the special respiratory arteries and veins. This is, of course, universally true of the higher vertebrates. It seems therefore that the persistence of a branch of the pulmonary artery supplying the trachea, taken in conjunction with the bifurcating pulmonary artery of the Viper with its tracheal and thoracic portions of the lung, is a fact which decidedly points in the direction of a previous respiratory function of that part of the respiratory passage which it now supplies.
The assumption upon the various facts which have been briefly dealt with in the course of the preceding remarks, that the most primitive type of Squamate lung is most nearly preserved in certain Serpents, is recommended by certain general considerations.
Whatever may be the views as to the phylogeny of the Squamata, it can hardly be disallowed that Reptiles generally have emerged from an Amphibian or Dipnoan form. On this view, the com- mencement of the lung far forward in the body is intelligible, for the earliest condition known, that represented in the Dipnoi, shows a lung at first (or always, Ceratodus) unpaired communicating directly with the exterior through the glottis and mouth-cavity.
* Above p. 36.
t+ “On the Anatomy of the Yellow-throated Lizard,” P. Z.S. 1904, vol. i. p. 263, text-fig. 37.
t “On the Vascular System of Hatteria &c.,” P. Z.S. 1905, vol. ii. p. 462.
1906. ON THE TEETH OF CREODONTS. 45
3. On the Minute Structure of the Teeth of Creodonts, with especial reference to their suggested resemblance to Marsupials. By CHartes 8. Toms, M.A., F.R.S., Vice-Pres.Z.S.
[Received January 15, 1906.} (Text-figures 12-25.)
hat the Creodonts, though obviously not Marsupials, never- theless present resemblances to them has been noted by many observers. Filhol (1) has discussed the question, and Matthew (2) uses the expression ‘“‘ pseudo-marsupial characters of the Mes- onychidze,” while he also says of Pachycena that “in its dentition this species approximates the Marsupial dental formula.” liydekker (3) goes a little further, and says ‘these and other fossil forms, such as Sorhywna, seem to indicate an intimate relationship between the Polyprotodont Marsupials and the Creodont Carnivores represented by Hyenodon.” Wortman (4) also speaks explicitly upon the same point; whilst frequently alluding to marsupial resemblances in their osteology and den- titions, he says: “ By taking the more primitive members of the existing marsupials as the basis of our comparisons, I am convinced we shall be able to arrive at a very much clearer understanding of what the ancestors of the Creodonts were like ” ; and “ Present evidence points to the fact that the two groups of the Creodonts probably arose side by side from Mesozoic Marsu- pials,” and further “that they were derivatives or offshoots of any pre-existing group of Placentals is exceedingly unlikely.”
Such speculations being rife, 1t occurred to me that it would be interesting to ascertain what evidences of affinity the minute structure of their teeth might afford, and by the kindness of Dr. Matthew, of the American Museum of Natural History, i have been enabled to make sections of the teeth of a number of representative species of Creodonts, whilst Dr. Smith Woodward has kindly furnished me with a fragment of a premolar of Borhyena.
But, before detailing the results of an examination of these teeth, it seems desirable to say a few words upon the nature and value of the evidence to be derived from the histological structure of teeth, the more so because this line of investigation has been but partially pursued and its results appear to be not well known to the majority of naturalists.
It might have been expected that there would be but little variety of structure in the teeth of animals belonging to the same great groups, for it is not easy to see how this should be affected by the ordinary processes of selection. It might have been thought that so long as a tooth was strong enough, sharp enough, and well adapted in external form to its work, its structure would
46 MR, C. 8. TOMES ON THE [Jan. 16,
matter little and would remain constant. But it was shown by my father, the late Sir John Tomes (5), that by a mere examina- tion of sections of the enamel it was possible in the case of Rodents not merely to pronounce that the enamel was that of a Rodent, but in a large number of instances to refer it correctly to a particular family of Rodents, or to a group of families.
Tn the more simple forms of enamel, the enamel prisms all pass outwards from the surface of the dentine to the outer surface of the enamel, and are, with very slight exception, exactly parallel with one another.
But in the Rodents contiguous layers of enamel prisms start off from the dentine at different angles, the layers alternating in this respect, so that if the section embraces in its thickness more than one layer, as such sections almost invariably do, patterns are produced by the crossing of the prisms, and these patterns are constant and characteristic of many of the families.
Similarly, my father showed that the enamel of Marsupials (6) presented characters very unusual in Placental mammals, and therefore almost characteristic of Marsupials, whilst the Carnivora also presented well-marked enamel characteristics.
It therefore seemed to be well worth while, in view of the uncertainty of the position of the Creodonts and of their relation to recent Carnivora, and possibly to the Marsupials, to apply this test of their affinities.
As bearing upon the subject generally, I may mention that I myself examined the teeth of a number of genera of the Gadide (7), a family selected as being both fairly numerous and at the same time compact, with the result that I found that the enamel was alike in all, but that the dentine presented marked variations upon a common type of vasodentine, and that these peculiarities coincided with their zoological arrangement, and not with the functional development of the dentition. Thus some of the largest teeth presented the simplest, and almost degraded, struc- ture common to them and their immediate relations, whilst some teeth, reduced so as to be almost rudimentary, retained the com- plexity of structure characteristic of their zoological relations. These, however, are the only papers I have met with in which this line of research had been followed out to any extent.
In what may be termed the normal arrangement of the dental tissues of placental mammals, the tubes of the dentine end by branching and becoming very fine, or by entering minute globular or angular spaces within the boundaries of the dentine (see text-figs. 15, 16, & 19, pp. 50, 51, 53); but it was shown by my father that in Marsupials the greater number of the dentinal tubes, instead of so ending, became continuous with tubes which traverse the enamel. This is true of all Marsupials, recent or extinct, which were examined by him or by myself at later dates, with the solitary exception of the Wombat, in which this does not happen, though, as might be expected from what has already been said, the precise extent to which it happens
1906. TEETH OF CREODONTS. 47
and the patterns produced vary in different families of the Marsupials.
Were there no more than this to be said, we should be pro- vided with a criterion of marsupial affinity both certain and easy of application. But unfortunately the case cannot be fully stated quite so simply. Whilst it remains quite true that all marsupial enamels present this character of penetration by the dentinal tubes, the converse is not quite true. Thus Hyrax has an enamel so richly penetrated by dentinal tubes that it might be easily taken to be a marsupial enamel, though in this respect it stands quite alone among placental mammals. But traces of this peculiarity are to be found in much reduced degree in certain Insectivora, notably in the Shrews; this occurrence in Insectivora may possibly be interpreted as a survival from some marsupial form of ancestor. But this explanation is not available for all cases: in very reduced degree the character has been found in the Jerboa, in some Carnivora, and even in Man, though in Man the rarity of the occurrence and its irregularity when it does occur suggest that it is pathological, or at least that it is a reversion towards something which has disappeared longago. And investi- gations of my own (8) into the development of enamel, and especially of marsupial enamel, distinctly point to this penetration of the epiblastic enamel by tubes continuous with those of the mesoblastic dentine being a primitive character, to which some slight tendency to revert has not been quite lost by placental mamuals.
Hence, in the interpretation of the occurrence of this character a different value appears to attach to negative and _ positive results : if we find no tubes at all in the enamel, we shall, I think, be quite justified in saying that no near affinity with the Marsupials canexist. On the other hand, if we find rudimentary traces of this penetration, we shall not be justified in attaching great importance to it as an evidence of marsupial affinity, though if we find an abundant penetration we shall have a character which, so far as is known, is peculiar to Marsupials and to Hyrax.
Having thus cleared the ground as to the value of the evidence, it remains to describe in slightly greater detail what is met with in the Marsupials, in Carnivora, and in Creodonts.
MarsupraAt ENAMEL.
A general character of marsupial enamels is the simplicity of the course pursued by the enamel prisms; each prism pursues, as a rule, an almost straight course from the dentine to the enamel surface, and where marked curvatures do occur, all of the contiguous prisms pursue the same curve, so that no patterns are produced by neighbouring prisms crossing one another. Where, however, the tubes are very abundant, the enamel prisms can hardly be seen at all, and we have to take the tubes as indicative of their course.
48 MR. C. S. TOMES ON THE [Jan. 16,
The enamel is most richly tubular in the Diprotodont group ; though it must not be thought that the tubes are sparse in the Polyprotodonts. In a large number*of instances there is a slight dilatation at the point of junction of the dentinal tube with the enamel tube, a sort of clumsy joint in fact (text-fig. 12); but this is not an invariable character, the tube sometimes passing on with no mark at the point of junction. Where, however, the enamel thins out towards the neck of the tooth, the tubes in it, whether it be that of a Diprotodont or a Polyprotodont, become few or none, so that it is necessary to be careful not to select the enamel to be examined from this situation.
As illustrations of typical marsupial enamels I have selected that of Hypsiprymnus (text-fig. 12), of Thylacinus (text-fig. 18, p. 49), and of Dasyurus (text-fig. 14, p. 49); those interested in the subject
All the figures, though drawn from actual slides, are semi-diagrammatic.
Hypsiprymnus.—Longitudinal section of dentine (D) and enamel (KE). The tubes in the enamel reach its outer surface, or nearly reach it. Slight dilatations mark the passage from the dentine to the enamel.
will find figures from other genera in the paper of my father’s already referred to. With regard to these and the other figures illustrating this paper, I may say that for the sake of clearness they are semi-diagrammatic. Though all have been drawn from actual sections, fewer tubes have been drawn than actually exist in a given area, and all indications of structure, other than those with which we are immediately concerned, have been left out.
In the Diprotodont Hypsiprymnus (text-fig. 12) the enamel tubes are seen in their greatest development. Starting, usually with a dilatation at that point, from the dentinal tubes they traverse the entire thickness of the enamel, turning a little to one side as they approach its periphery, and some of them branching off almost at right angles. It must, however, be understood that had the section been taken from the thin enamel near the neck of the tooth, fewer tubes, and finally no tubes at all, would have been seen. Where the tubes are very abundant, as in this case,
1906.] TEETH OF CREODONTS. AQ
it is difficult to see the outline of the enamel prisms, which, however, where traceable pursue a nearly straight course.
When, however, we come to the enamel of Zhylacinus (text- fig. 13), we find that the tubes thin out and are lost before they reach the exterior of the enamel, even where this is thickest, though sections may be found in which they penetrate further
Text-fig. 13.
Thylacinus.—Longitudinal section of dentine and enamel. ‘The tubes reach halfway through the enamel.
Text-fig. 14.
Dasyurus.—Longitudinal section of dentine and enamel.
than in that figured. And where there is a tolerably abundant passage of tubes, the so-called granular layer, which marks the exterior of Placental dentines (cf. text-fig. 15, p. 50), is absent or but little conspicuous. Outside the region of the tubes the enamel prisms are fairly distinct and are seen to be straight; the straight prisms may also be traced by careful illumination right in to the
Proc. Zoou. Soc.—1906, Vou. I. No. IV. 4
50 MR. ¢. S, TOMES ON THE [ Jan. 16,
dentine surface. Towards the neck of the tooth enamel without tubes is, as in the Diprotodonts, to be found.
Dasyurus enamel resembles that of Thylacinus pretty closely, though the tubes generally reach further through the enamel (text-fig. 14, p. 49).
Tar ENAMEL OF CARNIVORA.
As an illustration I have selected that of the Hyzna (text- figs. 15 & 16), though it may be premised that the enamel patterns of Carnivora are fairly constant. As one would expect from analogy, they are not quite identical in all: thus in the Dog group they are simpler, and where the enamel is thin the prisms become quite straight. Where, however, the enamel is thicker, the patterns are easily identifiable as similar to those found in, for example, the Felidee, though the curvatures are less pronounced,
Text-fig. 15.
Hyena.—Longitudinal section near apex of cusp. A few dentinal tubes pass a little way into the enamel. Most of the dentinal tubes branch and terminate in the minute spaces of the granular layer. The enamel prisms are arranged in alternating bundles, and pass nearly at right angles to one another.
Two figures are given to show the difference in pattern when the enamel is viewed in a longitudinal and in a transverse section of the tooth; hence any obliquity in the plane of the section will alter the appearances. But, after a little experience, it is not difficult to discriminate between differences due to differences of plane and those due to real differences in arrangement.
It will be noticed that no prisms in this, the thicker portion of the enamel, pursue a straight course, and that all do not pursue the same course. They are, however, grouped into bundles or sheaves of prisms pursuing an approximately parallel course, whilst towards the exterior of the enamel all the bundles become parallel
1906.] TEETH OF CREODONTS. Bil
and straight. They are thus interwoven with one another in a way that is not found in any known marsupial.
Text-fig. 16.
Hyena.—Transverse section.
Text-fig. 17.
Ocelot.—The enamel prisms are not shown, but some dentinal tubes pass a little way into the enamel.
As regards the other character, namely, that of the enamel being penetrated by tubes running in from the dentine. none are 4*
52, MR. C. 8. TOMES ON THE [Jan. 16,
seen to do so in text-fig. 16; in text-fig. 15 one or two penetrate a very little way.
In text-fig. 17, the enamel of an Ocelot, I have drawn an example of a greater penetration of the enamel by dentine tubes: this, | may say, is the section which shows this to the greatest extent out of some sixty sections taken from different genera of Carnivora. But though, on the whole, this shght rudimentary degree of penetration is perhaps rather more frequent in Carnivora than in most mammalian orders, in none does it occur to an extent in the least comparable with that found in Marsupials.
THe ENAMEL OF CREODONTS.
The examination of fossil teeth presents greater difficulties than that of recent teeth. Structurally, the enamel is always well preserved, but it has in the process of mineralisation often become unduly transparent, so that careful illumination is even more essential in deciphering its structure. And the teeth are
Text-fig. 18.
Hyenodon.—Transverse section. The enamel prisms are arranged in bundles radiating so as to present a goblet form. In the dentine, not very well preserved, the excavations of a fungus are seen.
often exceedingly brittle and friable, so that it is difficult to get good sections; this can be partly overcome by imbedding the teeth in desiccated Canada balsam before grinding them down. The dentine, however, being richer in organic matter, is often very badly preserved, so that sometimes all structure has dis- appeared ; a fact which handicaps the observer in tracing the passage of tubes from it, and sometimes leaves him only able to look for characteristic appearances of tubes in the enamel itself. Moreover, many of the teeth being rare, only small bits or
1906.] TEETH OF CREODONTS. 533
damaged teeth were available for examination, so that it was not always possible to select the plane in which a section was most desirable: one had to take what one could get.
Still, if the enamel of a fossil Diprotodont be examined, there is no difficulty in seeing the enamel tubes and being absolutely sure of their existence and their course, and the direction of the enamel prisms can always be traced in a fossil enamel.
The first figure (text-fig. 18) represents the enamel of Hycenodon (Oligocene of 8. Dakota): in it no trace of penetration by tubes can be found, and the very distinct enamel pattern is closely similar to that found in a recent Carnivore (of. text-figs. 15 & 16), In passing I may call attention to the curious spaces, dark in the figure, found in the dentine, which are quite common in the dentine of fossil teeth. In recent teeth they are only known to oceur in teeth which have been lying about in a graveyard, or in others which have been lying at or near the surface of the ground. They are excavations caused by a boring fungus; it is generally believed to be one of the mould fungi and is perhaps Saccharomyces mycoderma. As it does not seem likely that even the hungriest of mould fungi could find much pabulum in a fossilised tooth, this boring presumably took place when the tooth was com- paratively fresh, and thus points to the persistence of this mould fungus from Oligocene and Eocene periods.
Text-fig. 19.
x
Mesonyx.—Longitudinal section. No tubes penetrate the enamel : there is a well- marked granular layer, the dentine being well preserved.
Mesonyx (Middle Kocene) (text-fig. 19).—In this specimen the structures are well preserved. Not only are there no notable enamel tubes, but the outer periphery of the dentine presents appearances inconsistent with penetration of the enamel: namely, the dentinal tubes fine down, or spread into tiny branches,
54 MR. C. 8. TOMES ON THE [Jan. 16,
and there is a well-defined granular layer. The course of the enamel prisms is similar to that seen in Hyenodon and in recent Carnivora.
Text-fig. 20.
\
Pachyena.—Dentiue perished. The enamel prisms ate atranged in bundles like Hyenodon, but the plane of the section is not quite the same.
Pachyena (Lower Kocene) (text-fig. 20).—The dentine structure is gone, but the enamel prismsare very distinct, no tubes are seen in it, and the prisms are gathered into bundles pursuing a course similar to that seen in Hyenodon or in Hyena.
Text-fig. 21.
Oxyena.—Dentine well preserved.
Oxyena (Lower Eocene) (text-fig. 21).— Here again similar con- ditions obtain, though the section not being in exactly the same
1906. ] TEETH OF CREODONTS. 55
plane, the course of the prisms appears to differ a little. But this is a difference purely due toa difference of plane, and not toa real
difference of course. Text-fig. 22.
Bp os A Se ty
(S ‘tl ‘
a
3 ;
Th / wet i) . . Ly
Sinopa.—Dentine perished, except in places.
Sinopa (Middle Hocene) (text-fig. 22)—Here again we get just 4 co) e) co) the same pattern, and the same absence of penetrating tubes.
ay Af — 5 ?
Borhyena.—Dentine perished, enamel well preserved.
Borhyena (text-figs. 23, 24).—In this genus we find the absence of penetrating tubes, and can distinctly recognise the carnivorous pattern in the course of the prisms. But apparently the prisms are a little straighter than in recent Carnivora, or at least in recent Felide. Itis not, however, possible to speak very positively as to this greater simplicity, as I had only a fragment of a tooth at my disposal, and the sections I was able to get were small and may not have included any of the thickest parts of the enamel, where, as has already been noted, these characters are to be found most marked. However, there is ample evidence to say that the enamel of Borhycena is essentially of the Carnivorous type, and
56 MR, C, S. TOMES ON THE [Jan. 16,
bears no more resemblance to that of the Marsupials than does that of other Creodonts.
Text-fig. D4.
Borhyena.—Dentine well preserved in places. The plane of this section is not quite the same as fig. 23, but only fragments having been available it is not possible to define the plane in which the section lay,
Text-fig. 25.
Cynodictis.—Longitudinal section. Enamel prisms all parallel; here and there dentine-tubes penetrate a little way into the enamel.
Didynictis (Lower Kocene).—Of this genus I have only two sections, but in them the enamel prisms are all parallel and pursue a course only slightly curved. The typicai carnivorous pattern is not to be found, nor is there any trace of it, so that of the Creodonts examined this and Cynodictis stand alone in this respect. Portions of enamel might be found resembling this in the teeth of the Dog, if taken towards somewhat low down upon the tooth, but the enamel is in the two sections I have obtained tolerably thick, and might have been expected to show more complexity of structure if any such exists anywhere upon the tooth,
Cynodictis (Oligocene) (text-fig. 25),—A specimen of this genus
1906. ] TEETH OF CREODONTS. 57
was sent to me by Dr. Matthew as an example of an early true Carnivore. In it the enamel prisms are almost straight and no decussation, or only the faintest trace of decussation, of the prisms of different planes is to be seen. It resembles chiefly the enamel of Didynictis, and differs in respect of its greater simplicity from that of the other Creodonts examined and from recent Carnivora. My section of Cynodictis embraces the whole tooth, so that there is no question as to greater complexity of pattern existing in any other parts of the tooth. In some of the Creodont enamels, and particularly in Cynodictis, slight indications of a rudimentary penetration of the enamel by dentinal tubes are seen, but in none does it exceed or even attain to the amount seen occasionally in recent Carnivora (cf. text-fig. 17, p. 51).
CONCLUSIONS.
The nature and the limitations of the evidences of affinity which can be derived from a study of the minute structure of teeth have already been alluded to, and it must not be forgotten that it is unsafe to build too much upon any one single character.
But, so far as the structure of their enamel may be taken as evidence, neither Borhyena, Pachyena, Hycenodon, Sinopa, Mesonyx, Oxyena, Didynictis, nov Cynodictis presents any greater resemblance to Marsupials than do the recent Carnivora. On the other hand, with the exception of Didynictis and Cynodictis, the enamel has reached just that stage of evolution found in the true Carnivora, and the enamel patterns are strikingly similar to those of recent Carnivora.
The uniformity of the patterns found in all of the Creodonts examined, excepting again Didynictis and Cynodictis, seems to point to the structure of their ename! having attained to a sort of finality ; that is to say, it was probably not undergoing any rapid evolutionary changes, a conclusion borne out by its close resem- blance to that of alien descendants, the recent Carnivora.
The absence of the peculiar stamp of the marsupial, the tubular enamel, would justify us in saying that they certainly do not stand very near to any marsupial, and that if there be a marsupial ancestor, or an ancestor common to the Marsupials and to the Creodonts, it must be sought considerably further back than any of those examined. ‘This is a somewhat disappointing conclusion : when I undertook the investigation I quite expected to find some distinct indication of mar supial relationship; that is to say, I expected to find that the general resemblance in macroscopic character of the dentitions to those of the polyprotodont Marsupials would have been accompanied by histological resemblances.
Ihave also been surprised to find that the enamels of Didynictis and of Cynodictis are actually simpler than those of the other Creodonts, and simpler than most recent Carnivora. As Cynodictis at all events appears to be nearer to the true Carnivora than are the Creodonts, the simplicity of its enamel as compared with
58 DR. J. ROUX ON THE TOADS [Jan. 16,
theirs may point to its not lying on quite the same line of descent.
Though I would not attach too much importance to it, I would again call attention to the fact already mentioned, that in Car- nivora, and still more so in Insectivora, rudiments of a penetration of the enamel by dentinal tubes occur with more frequency than in other mammals. This may possibly indicate some remote connection with the Marsupials, but the point which I wish to emphasise is that, as regards this character, the Creodonts carry us absolutely no further than do the recent Carnivora.
BIBLIOGRAPHY.
(1) Frru0L.—* Mammifeéres fossiles des Phosphorites.” Annal. Se. Géolog. viil., 1877.
(2) Marruzew, W. D.—“ Additional Observations on the Creo- donta.” Bulletin of American Museum Nat. History,
1901, (3) Lypexker, R.—Art. “ Mammalia,” Encyclop. Britannica, vol. xxx., 1902. Also Proc. Zool. Soc., 1899.
(4) Worrman.—‘Studies on Hocene Mammalia.” American Journal of Science, vols. xii., xill., & xiv., 1901-3.
(5) Tomes, J.—‘‘ On the Dental Tissues of Rodentia.” Philos. Trans., 1849.
(6) Tomzs, J.—‘‘ On the Dental Tissues of Marsupialia.” Philos. Trans,, 1850.
(7) Tomes, C. 8.—‘‘ On the Teeth of the Gadide.” Quart. Journ. Mier. Se., 1899:
(8) Tomes, C. S—‘‘ On the Development of Marsupial and other Enamels.” Philos. Trans., B. 1898.
(9) Tomes, C. 8.—‘“ On the Structure of the Teeth of Votoryctes.” Proc. Zool. Soc., 1897.
4. Synopsis of the Toads of the Genus Nectophryne B. & P., with special Remarks on some known Species and Description of a new Species from German Hast Africa. By Dr. Jean Roux, Curator in the Basle Museum of Natural History.
[Received December 11, 1905. } (Plate IL. *)
On visiting, last spring, the beautiful collections of the Museums of Paris and London, I “had occasion to examine, especially in the British Museum, most of the typical specimens of the known species of the genus Vectophryne. Whilst verifying the diagnoses, I was able to make some observations modifying or completing
* Por explanation of the Plate, see p. 68,
PZ SD LOO. wo, Pl:
J.Green del. et bth Bale & Damelsson, LY mp 1, NRCTORRIENANI, TOSI, ZAIN Win aPwil, 3 INS IMUNCIROMN IES) AWN © EIN PRR
1906. ] OF THE GENUS NECTOPHRYNE. 59
somewhat the descriptions of the authors. I was able to identify as one species two that had previously been considered different.
I add to these observations the description of a new species from German East Africa, the type of which is preserved in the Basle Museum, and conclude this paper with a key for the deter- mination of all the known species of the genus.
I am happy on this occasion to express my best thanks te Dr. Mocquard, of Paris, who was so kind as to allow me to examine the types of the species described by him. I am also much obliged to Prof. Boettger, of Frankfort a/M., and to Prof. Tornier, of Berlin. The former has been kind enough to send to me the type of his WV. exigwa which is preserved in the Senckenberge Museum; to the latter I am indebted for the loan of the types of two species in the Berlin Museum. I am particularly indebted to Mr. G. A. Boulenger, who has obligingly placed at my disposal the valuable collection in the British Museum (Natural History), and has been so kind as to verify the work done in his laboratory.
1. NecropHryne AFra B. & P. Buchh. & Peters, Mon. Berl. Akad. 1875, p. 202, pl. i. f. 5. Boulenger, Cat. of Batrach. Sal. p. 279. Hxamined : The type specimen (Berlin Museum): Cameroon. 6 specimens (Brit. Mus.): Efulen, 5. Cameroon, and Rio Benito. This species occupies quite a special place as regards its webbed fingers, the subarticular tubercles of which imitate smal! lamellee.
2. NECTOPHRYNE MISERA Mocq.
Mocquard, Le Naturaliste, 1890, no. 82, p. 182; Nouv. Arch. Muséum Paris, sér. 3, tom. 11. p. 161, pl. xi. f. 7.
Examined in Paris Museum: Type specimen, N. Borneo.
In Brit. Mus.: 10 specimens from Paka-Paka, 10,000 feet, Kina-Balu, N. Borneo.
This species has also strongly webbed fingers, but no sub- articular tubercles. Sometimes the tibio-tarsal articulation does not quite reach the tympanum.
3. NECTOPHRYNE HosiI Blgr. (Plate IT. fig. 1.) Boulenger, Proc. Zool. Soc. Lond. 1892, p. 508, pl. xxx. fig. 2. Examined in Brit. Mus. :—
Type specimen(d). Mt. Dulit, N. Borneo. 2 specimens(¢ Q). Kuala Lumpur, Selangor.
2 Lawas, Brunei.
eee: Head-waters of Sarawak R. Ios Oy Sarawak.
I, Tandjong, 8.E. Borneo. eo Akar River, Sarawak.
60 DR. J. ROUX ON THE TOADS [Jan. 16,
This is the largest species of the genus. The diagnosis given by Boulenger in 1891 was drawn up from a male specimen from Mt. Dulit, Borneo. Since then the collection of the British Museum has been increased by several specimens, especially females, which I have had the privilege to study. As is often the case with Lufo, in this species the female individuals are notably larger than the males. The distinctive characters indicated by Boulenger are generally very well marked in the female. The head is broader in comparison with the length. The canthus rostralis is well marked. The loreal region is nearly vertical and shows a slight depression in the upper part. The interorbital space, twice as broad as the upper eyelid, is very distinctly concave, as well as the part of the head situated near the parotoids. The tympanum is very distinct; it is suboval, vertically elongated, and half as long as the eye. The parotoids are well marked, pyrifor m,and begin immediately behind the eyes.
As to the limbs, we have noticed individual variations in the length, especially in the hind limbs. The fore limb is relatively long: the fingers, webbed only at the base, are bordered by the membrane, and the distal part is subtriangularly enlarged ; this peculiarity i is more appreciable in the fingers than in the toes. The hind limbs of most of the individuals observed are longer than in the type specimen. The hind limb being carried forward along the body, the tibio-tarsal articulation reaches sometimes the tympanum, sometimes the eye. The toes are generally short, entirely webbed, except the three distal phalanges of the fourth toe, which are free. The subarticular tubercles are very well developed, as well as the two metatarsal tubercles. The outer tubercle is twice as large as the inner. I have noticed the presence of a very distinct tarsal fold.
The coloration of the individuals is worthy of detailed descrip- tion owing to the marked differences between males and females.
The type specimen, a male, figured by Boulenger, is uniformly brown with some indistinct spots on the limbs; the throat is black. Two other male specimens show the same coloration, but two male specimens from Lawas, Brunei, are somewhat different. The body shows, besides the dark brown, some light brown markings, which form indistinct coarse vermiculations. The limbs are yellowish brown, and present also lighter and darker parts more or less distributed in transverse bands.
The females labelled ‘‘Sarawak,” one of which is figured on Pl. U1. fig. 1, are distinctly bicolor (yellow and black). The ground is black with small vermiculations or round yellow spots (in the latter case especially on the sides of the body).. The head, the back, and the limbs show also these vermiculations. The spots are a little broader on the anterior part and on the sides of the head. The lower part of the body and of the limbs is generally dirty grey or uniform yellowish. The border of the lower lip often shows yellow spots. The inferior part of the feet is brown. The females have generally smaller and less numerous
1906. ] OF THE GENUS NECTOPHRYNE. 61
dorsal tubercles than the males. The females from Tandjong (S.E. Borneo) and from Akar River (Sarawak) show the typical coloration above described.
A female specimen from Mt. Kuala Lumpur (Selangor) shows an interesting variation. The general colour is a dark gyvey, approaching brown. The upper part of the head, of the back, and of the tibia shows no yellow spots, but the sides of the body and of the limbs, as well as the upper part of the thighs, have round or oval spots pretty distant from one another. ‘These spots are of a fine yellow colour, with brown border. Similar but longer spots may be found on the throat and on the anterior part of the chest.
A female from Akar River (Sarawak) shows irregular and in- distinctly distributed spots. The yellow colour is prevalent on the back; the sides are marbled yellow and black; the belly is of a dirty yellowish colour.
The following are the dimensions of two individuals from Sarawak found pairing, the female in the act of spawning :—
3. : Length from snout to vent... 5°65 em, 9°8 cm. ie otpomnad! lama eeey eee (051) 6 ISU Si nlaysy Ores lnm yee saul: ACES Se)
The eggs of Nectrophyne hosii are oval, 1 millimetre in length ; they are laid in chains as in Bufo.
4. NECTOPHRYNE PARVIPALMATA Wern. Werner, Verhandl. zool.-bot. Gesells. Wien, vol. xlviii. 1898, p. 201, pl. u. ff. 7 & 7a.
Examined: the type specimen in the Berlin Museum. Habitat : Cameroon 2
5, NECTOPHRYNE EVERETTI Blgr. (Plate II. fig. 2.) Boulenger, Ann. Mag. Nat. Hist. (6) xvii. 1896, p. 450. Examined in the Brit. Mus. :— The type specimen (2): Mt. Kina Balu, N. Borneo. 1 S specimen: Mt. Penrissen, Borneo. In this second individual the tympanum is quite visible, oval. The hind limb being carried forward along the body, the tibio- tarsal articulation reaches between the tympanum and the eye.
6. NECTOPHRYNE TUBERCULOSA (Gthr.).
Giinther (Pedostibes tuberculosus), Proc. Zool. Soc. Lond. 1875, p. 576, pl. lxiv. fig. C.
Boulenger, Cat. Batr. Sal. p. 280.
Examined in the Brit. Mus. :—
2 $ type specimens: Malabar.
The hind limb being carried forward along the body, the tibio- tarsal articulation reaches the tympanum. The upper part of the limbs is also covered with tubercles.
62 DR. J. ROUX ON THE TOADS [ Jan. 16,
NECTOPHRYNE GUENTHERI Bler.
Boulenger, Cat. Batr. Sal. p. 280, pl. xviii. fig. 3. Boettger (Nectophryne exigua), Abhandl. Senck. Gesells. 1901, vol. xxv. p. 394. Examined in Brit. Mus. :— The type specimen from Matang, Borneo.
2 young specimens F y eo SP from Singapore.
2 adult He 2 adult i from Sirhassen (Natuna. Isl.).
Also the type specimen of ect. exigua in the Senckenberg Museum: Baram Riv., N. Borneo.
In this species, as in others, I have also noticed individual variations in the length of the limbs. The hind limb being carried forward along the body, the tibio-tarsal articulation reaches sometimes the eye, sometimes between the latter and the end of the snout. Jn the typical specimen it does not only reach the eye but notably behind it.
Establishing a comparison between the type of Wectophryne exigua Boettger and young individuals of Nectophryne quenthert, I have been convinced that these two species are identical.
The differences are indicated by Boettger as follows :
“ Habitus etwas weniger schlank; Trommelfell kleiner als bei NV. guentheri Bigr.”
On examining several specimens of WV. guentheri I have noticed differences not only in the respective length of the body and of the limbs, but also in the respective Himenciens of the tympanum and of the eye according to age. The young specimens have proportionally a smaller tympanum than the adults, Besides individual variations have been observed. Some measurements follow :—
Type spec. Spec. from Sirhassen.
Eye a 1:9 2 mm.
ee ee
Tympanum...... 2 i Lei. pes
Specimens from Singapore.
Adult. Adult. Juv. Juveniss. 2 eyes 1:9 4mm. als ‘ 1 ] i co) 1 33 ray 9
A young specimen of JV. guenthert in particular, which is just as large as the type of .V. eaigua, shows in the general form of the body as well as in the colour a striking likeness to the latter.
Similar black spots are distributed on the belly, and the coloration, yellow and black, on both sides of the head is identical. The limbs show also the same extent of web.
As, on the other hand, it is impossible not to recognise the existing relations between the young specimens of JV. guentheri with adults of this species, I believe that NW. ewigua may be considered a young specimen of V. guenthert. Very appreciable
1906. ] OF THE GENUS NECTOPHRYNE. 63
variations having been noticed between several specimens of the latter, the distinction drawn by Boettger between his species and that of Boulenger cannot be accepted.
8. NECTOPHRYNE MACROTIS Bler. (Plate IT. fig. 3.) Boulenger, Ann, Mag. Nat. Hist. (6) xvi. 1895, p. 171.
Examined in the Brit. Mus. :— The type specimen (2 ) from the Akar River, Borneo.
9. NECTOPHRYNE SIGNATA Bler. Boulenger, Proc. Zool. Soc. Lond. 1894, p. 645, pl. xl. fig. 1.
Examined in the Brit. Mus. : The type specimen from Rebenew Mt., Kapuas Distr., Dutch Borneo.
10. NecropHRYNE MACULATA Mocq.
Mocquard, Le Naturaliste, 1890, no. 82, p. 182; Nouv. Arch. Muséum Paris, 3° sér. t. 11. p. 162, pl. xi. fig. 8. Examined in the Paris Museum :— 3 type specimens from Kina Balu, N. Borneo.
11. NecroPHRYNE TORNIERI, sp. n. (Plate IT. fig. 4.)
Habit slender. Head moderate, as longas broad. Snout short, scarcely prominent, obliquely truncate, quite as long as the eye; canthus rostralis strong. Loreal region vertical, slightly concave in the upper part. Interorbital space broader than the upper eyelid. Tympanum exposed, vertically oval, about one-third the diameter of the eye. ‘The distance between the anterior border of the tympanum and the postericr corner of the eye equal to half the distance between the anterior corner of the latter and the nostril. Fore limb slender, equal in length to the distance between vent and tympanum. Fingers moderate, much depressed, webbed at the base, dilated and truncate at the end, first a little shorter than second. The hind limb being carried forward along the body, the tibio-tarsal articulation reaches the posterior border of the eye. ‘Toes hnlf-webbed, but the three distal phalanges of the fourth toe free. The tips of the toes less strongly dilated than those of the fingers. Subarticular tubercles well marked. Two well-developed metatarsal tubercles, the inner the larger. Skin of the upper part of body and limbs covered with numerous small round warts, irregularly distributed; the largest situated behind the tympanum and on the middle of the back ; beneath feebly granulate. The granulations are visible on the posterior part of the belly and on the under part of the thighs, and disappear on the throat.
Brown above, with darker markings, especially two pairs on the back: one between the fore limbs, the other on the sacral region. A large lateral dark band from the eye, surrounding the tympanum, caiman is hghter in colour, and extending on avon side
64 ON THE TOADS OF THE GENUS NECTOPHRYNE. [Jan. 16,
of the body. A dark streak from the end of the snout passing below the canthus rostralis, through the eye, and above the tym- panum to the commissure of the “mouth. Loreal region brown ; a light spot below the eye between yellowish-brown parts of the upper lip. Limbs brownish in colour, with darker markings arranged in indistinct large cross bars. "Sides of the body below the dark lateral band lighter than the back, more or less speckled with dark brown. Sometimes a yellowish-brown vertebral stripe extending along the middle of the back, from snout to vent. Beneath ‘entirely white or with a few small dark spots on the throat and belly.
Hab. Ukami, German East Africa. 2 specimens.
Dimensions.—From snout to vent, 27 mm.; hind limb, 38; fore limb, 20; length of head, 9; breadth of head, 9:5.
Named after my colleague, Dr. Tornier of Berlin, who has added much to our knowledge of the herpetological fauna of German Kast Africa.
The figured specimen of this new species is preserved in the herpetological collection of the Basle Museum, the other has been presented to the British Museum.
If we now consider the geographical distribution of the genus, we notice that most of the species described are from Southern Asia. Borneo is particularly rich. Not less than six species have been found on this island, and one of them has been found also in the Natuna Archipelago (Sirhassen) and Singapore. New discoveries will most likely further extend the geographical dis- tribution of the other species. But we cannot omit to state the fact that up to this date no Wectophryne has been discovered, so far as we know, in the other great islands of the Sunda Archipelago,
The genus Nectophryne has representatives also in West Africa, The fannal similarity of that district with the south-east of Asia has often been noticed (see Wallace). West Africa possesses two species, and the new species described above shows that the genus is also represented in the eastern part of the African continent.
I conclude with a synoptic table for the determination of the known species of WVectophryne, not taking into consideration doubtful species, as e. g. Wectophryne sundana (Ptrs.) (Boulenger, Cat. Batr. Sal. p. 281). I have not been able to examine the only existing specimen of this species, which is preserved in the Berlin Museum and comes from Borneo.
Key for the determination of the Species.
I. Fingers strongly webbed, very slightly dilated at the end, the inner quite rudimentary. a. Subarticular tubercles er similar to small lamellie... WV. afra. b. Subarticular tubercles absent.. SCRE ar eae acaas Anaenee ONS OCHS
II. Fingers partially webbed, more or ieee dilated at the end, the inner well developed. A. Vibio-tarsal articulation not reaching the end of the snout. a. Moesionly halt-webbedh.)./5..22teesh eee oecles cence stats -ese one e COMIENE.
1906. ] ON BONES OF THE LYNX FROM DEKBYSHIRE. 65
6. Toes more than half-webbed.
Meet ar sallat oO) Uinta nineties tect. ML nei laa chun a auiannnitn cae OSU: 2. No tarsal fold. a. Tympanum hidden .......... N. parvipalmata.
8. Tympanum visible, its diameter less than that of the eye; two metatarsal tubercles. a Fingers very slightly webbed at the base, the first
equal to 2 of the second ...... NV. everetti. ** Wingers very distinctly webbed at the base, the first equal to about 3 of the second ...... .. NN. tuberculosa.
*** Hingers 2 webbed, the web extending as a
margin to their tips; the first equal to 1 of the
second .. : NV. guentheri. y. Tympanum visible, equal to the diameter of the eye; 9 only one metatarsal tubercle .......... NN. macrotis. 3. Tibio-tarsal articulation reaching at least the end of the snout. a. Tympanum visible, equal to 2 the diameter of the eye N. signata. 6. Tympanum hidden ; tibio-tarsal articulation reaching beyond the end of thetsnout nse steal cc. be wie. I. maculata.
EXPLANATION OF PLATE II. Fig.1. Nectophryne hosii Blgr., p. 59, female. 2 nat. size. 1a. Side view of head. 2. Nectophryne everetti Blgr., p. 61, type. Nat. size. 2a. Side view of
head, x 13. 3. Nectophryne macrotis Blgr., p. 63, type. Nat. size. 38a. Side view of
head, X 2. 4. Nectophryne tornieri Roux, p. 63, type. Nat. size. 4a. Side view of
head, X 13.
5. On some Bones of the Lynx from Cales Dale, Derbyshire. By W. Storrs Fox, M.A., F.Z.S.
[Received October 25, 1905. ] (Text-figure 26.)
Remains of the Lynx have so rarely been found in the British Isles, that the recent discovery of some in a Derbyshire cave will, I hope, be considered to be worth recording. The history of the two former finds may be briefly stated. About the year 1866, the hinder portion of a skull and the right ramus of the lower jaw of this species were discovered in Pleasley Vale, on the borders of Derbyshire and Nottinghamshire, and are now in the Nottingham University Museum. Some fourteen years later a humerus anda metatarsal of the same species were found in Teesdale by the late Mr. James Backhouse, and are still in his son’s museum at York.
Thus, until the Cales Dale cave was worked, only four bones of Lynx had been found in the British Islands. I have been unable to obtain any information about the excavation in Cales Dale previous to 1897, but my own find there consists of 36 bones and teeth of Lynx, about half of this number being metapodials and phalanges.
The cave lies on the west side of Cales Dale, a small dale branching from the south side of Lathkil Dale, at a point about
Proc. Zoou. Soc.—1906, Vou. I. No. V. 4)
66 MR. W. STORRS FOX ON BONES [Jan. 16,
half a mile below the source of the river Lathkil. It is 800 ft. above sea-level, and takes the form of a narrow passage, running almost due east and west, in the Carboniferous Limestone. It possesses two entrances. The lower one is almost square in section, measuring 2 feet 8 inches across and 2 feet 9 inches high. By cra wling throu igh this, and along a passage of similar dimensions, for a distance of 62 eee a dome- -shaped chamber is reached 9 feet in height and 4 or 5 feet in diameter. It is into the side of this chamber that the second entrance opens, at about 5 feet above the floor. This second or upper entrance, almost a perfect oval in shape, is 2 feet 10 inches high and | foot 8 inches wide.
For the next 18 feet the cave consists of a passage averaging 3 feet high and 3 feet wide; it then widens out into a chamber 6 feet long and nearly 6 feet wide. It was in this chamber that the bones were found. Beyond this chamber the passage rapidly narrows to an impassable fissure,
In March 1894 I was informed that Lynx-bones had been found in the cave; but it was not until the spring of 1897 that I asked and obtained leave to excavate. In the chamber, or den, a thin layer of stalagmite was found. First, all the earth—mixed with bones and stones—lying above the stalagmite was removed ; then the layer itself was blasted, and all that had been sealed up by it was cleared away. But, unfortunately, no notes were taken as the work proceeded, so that it is impossible to say now whether any bones were found beneath the stalagmite. As the contents of the cave were dug out, they were carried to a neighbouring spring and were there carefully washed in a one-eighth-inch sieve ; and in this way even very small bones were secured.
Both Professor Boyd Dawkins, in his account of the Pleasley * Lynx, and Mr. William Davies, when describing the bones from Teesdale ft, used for comparison the skeleton of the Northern Lynx in the British Museum (1230). Accordingly, the Cales Dale bones have been compared with the same skeleton.
Of jaw-bones and teeth Cales Dale has produced :—a right ramus of the lower jaw (text-fig. 26 B); the right upper carnassial tooth, imbedded in a fragment of the maxilla (text: fig. 26 A); the right premaxilla containing its three incisors; and three canines. The ramus is incomplete, most of the bone behind the molar tooth being absent ; and the upper part of the socket for the canine is
* ‘British Pleistocene Mammalia,’ part iii. pp. 172-176 (Paleontographical Society, volume for 1868). + ‘Geological Magazine,’ volume for 1880, pp. 346-348.
Explanation of Text-fig. 26 (opposite).
Remains of Felis lynx from Cales Dale, Derbyshire.
A, A’. Right upper carnassial tooth, outer and lower veiws, p. 68.
3. Right mandibular ramus, inner view, p. 68.
C, C’. Axis vertebra, left lateral and lower veiws, p. 69.
I. Left os innominatum, outer view, p. 70.
FE, E’. Proximal end of left femur, posterior and anterior views, p. 70.
A-C, nat. size; D, E, two-thirds nat. size.
1906. ] OF THE LYNX FROM DERBYSHIRE. 67
Text-fig. 26.
Wii y
i
68 MR. W. STORRS FOX ON BONES [Jan. 16,
very much worn down. When this bone was found the canine was not in situ, but a tooth has been placed in the socket, into which it exactly fitted. 'The third premolar is missing altogether ; ; and, as the bone has entirely closed over the socket, this tooth must either have been lost some time before the death of the animal, or it could never have existed at all. The fourth premolar, when found, was separate from the jaw, but the molar was in position in the bone.
Taking into consideration the shrinkage caused by the absence of the third premolar, this ramus closely corresponds in general outlines with the Pleasley one described by Professor Dawkins. The following table of measurements, the last two columns of which are taken from ‘ British Pleistocene Mammalia,’ shows this correspondence. The measurements throughout are given in inches and tenths.
The total length of the canine now fixed in the jaw could not be measured ; and as the tip is broken off, the original height of the crown must remain uncertain, But the two odd Galles: Dale ‘anines are respectively in length: 1°65 ins. and 1:9 ins., whereas the Pleasely specimen is 1°85 ins.; and the height of the crowns of these two Cales Dale teeth is exactly the same as that of the right lower ramus of Felis lynx (borealis) in the British Museum, namely, “8 inch, as compared with °75 inch in the Pleasley animal,
Measurements of right ramus of Lower Jaw.
| | | FF. lyna Filynz. | F. lynx. (borealis). Cales Dale. | Pleasley. Brit. Mus. i | 1230 A. | { | Circumference behind M.1 ......... .... 2°37 | 22 | 2°0 | x before Pm.3........ #2714 24 | 20 Antero-posterior extent of M.1_ ...... 68 | “65 “63 | | Antero-transverse —., 3 f "25 | 26 25 | | Postero-transverse __,, Ted al Yorkie : 26 "2A, | 23 Height of crown of M.1_..........0..006.. “46 | 35 35 Antero-posterior extent of Pm. 4 ..... 5 | “49 | “46 | Antero-transverse i 3 : 18 | 19 | 19 | | | Postero-transverse — ,, 35 aa | 27 | 27 23 | Height of crown of Pm. 4 ........... “B34 | 35 | 3d
The measurements of the upper carnassial (text-fig. 26 A) and of the incisors show that there is very little difference between the Cales Dale teeth and those of F. lyna (borealis).
* As Pm.3 is lacking in the Cales Dale specimen, the measurement is taken behind
the socket of the canine. This absence of Pm. 3 accounts for the relatively small circumference here. :
Wo)
1906. | OF THE LYNX FROM DERBYSHIRE. 6
Measurements of Right Upper Carnassial Tooth = Pm. 4.
F, lyne. | FB. Lyne (borealis).
Cales Dale. | Brit. Mus. 1230 a. Antero-posterior extent .................56 73 | 72 Antero-transverse extent ........0...0...- “35 | 30 Postero-transverse extent ............... | 22 | 23 IRIGTEANY OE GROWAD coocacossvosachoancoscnsouson0 °39 “Ad
Measurements of Upper Incisors (imbedded in right premaxilla).
F. lyne. F. lynx (borealis). Cales Dale. | Brit. Mus. 1230 a.
|Lemvedany GI REINES oogosqcsnadosaasaaeseana bea bat 34 33
Maximum across crown of 1.3 oly eta 18 18 5 back to front of 1.3 ......... | “Oy 20 Height Ere Cisty Cate aia Cena cI eH | 27
There were two portions of bones of the fore limb in this cave, namely, the shaft and distal-end of a humerus and the proximal end of an ulna. As the bones of the British Museum skeleton are wired together, it is practically impossible to measure the anconeal fossa of the ulna; but, so far as could be seen, the Cales Dale fragment agrees in form with the corresponding part of the skeleton in question, though it is somewhat larger.
An axis vertebra (text-fig. 26 C) from Cales Dale agrees gene- vally with that of the Northern Lynx (B.M. 1230), and only differs to an extent which might be expected in the bone of a rather larger and more powerful animal.
Measurements of Axis Vertebra.
Cales Dale. | Brit. Mus. 1230 4. |
F. lyne. | F. lynx (borealis).
Base of odontoid process to inferior posterior
TITATESINN pacing see CLE See ear eae ee 134 Minimum transversely ..................0:002-- “BA. | “78 | Extreme length of neural spime..................] ie 7/AL | 1:45
* The inferiority of length here is due to the absence of the epiphysis.
The Cales Dale humerus appears to have belonged to a more
70 MR. W. STORRS FOX ON BONES [Jan. 16, powerful animal than /. lynx (borealis) in the British Museum and the dweller in the Teesdale cave; for in it the deltoid ridge is strongly developed, but is not mar kedly so in the other two specimens. The supinator ridge is prominent in all three. If the Cales Dale bone had been complete it would have exceeded the Teesdale one in length.
In the following table the last two columns are taken from
Mr. Davies’s article.
Measurements of Humerus.
F. lynx. | F. lynx. | F. lynx (borealis).
Cales Dale. Teesdale. | Brit. Mus. 1230 4. Transverse diameter of distal end... 1°56 | 1°40 1:50 BS trochlea...... 1:08 87 10) Smallest circumference of shaft...... 2°05 | ie 1:75
One nearly perfect left os innominatum (text-fig. 26 D) and a fragment of a right-side one do not appear to have belonged to the same animal, for the fragment seems to belong to a less recent date than the other, and to a larger animal. The bone from the left side lacks only the epiphyses on the extremities of the ilium and ischium. It has the roughened ridge above the acetabulum more pronounced than the corresponding bone of the British Museum skeleton, and is generally rather heavier in build, but in all other respects the two are exactly similar in every detail.
Measurements of Os innominatum.
| F.lynx | #F'. lynx ui ly eed | : 2 (borealis) \cales Dale | Cater Date Brit. Mins ‘ales Dale.|Cales Dale. ae 4 | 12380 a. Minimum circumference of ilium.. en 2°46 2°71 2°24, SMagictae across ilium (from pubie to osha || SSUIEACE) Mek cackh ates eons en .| “96 112 90 | Minimum circumference of jealifucn (bet weer cen | acetabulum and ischial ee. 1:96 2°05 eral | Across acetabulum <...7........... te ‘79 81 81 | Maximum length—between extremities 3 ee | ilium and ischinm | *5°90 618
* The apparent shortness is due to the loss of the epiphyses.
The hind limb is represented by two fragments of femur, namely, the proximal end and part of the shaft of one from the left side (text-fig. 26 E); and the head of another, also from the left side. The larger fragment is perfect except that the lesser
1906. | OF THE LYNX FROM DERBYSHIRE. (1
trochanter is broken off. It is distinguished from the corre- sponding bone of the Northern Lynx (B.M. 1230 a) by the greater development of the ridge which travels down the outer side of the shaft from the great trochanter.
Measurements of Left Femur.
| | FY. lyna F. lynx. | F. lynx. | (borealis).| Cales Dale.’ Cales Dale. Brit. Mus. i 1230 A. Maximum width at proximal end ............... |} 189 a | 1:62 | 45 op OR INGRGCL Sascscase qn sogdnosodone 82 83 | Sin Circumference of shaft, taken 3 ins. from| DRORTIOA NCL” 54 sa0 ow onson asc eee bes 00000 ob0aeC 2°12 nae 20
* In the B.M. specimen this was the minimum circumference.
There are five tarsal bones, including an astragalus and a ealeaneum; and also one carpal—which I take to be a pisiforim, though it differs somewhat from the pisiform of the Northern Lynx.
Of metapodial bones there are two complete metacarpals, namely, the second of the right and the third of the left manus; one complete metatarsal (mt. 5—left), and another lacking the distal end (mt. 5—right). Besides these there are two fragments, which are specially interesting owing to their size and stoutness of build. A comparison with Mr. Davies's measurements of the Teesdale and British Museum metatarsals is misleading, but 1s ~ given for what it is worth.
Measurements of Metapodials.
| i} | F. lynx, BF. Lyne, | F. Lyne, | F. lyna, | BF. Lyne. FB. lyne,| F. lyne me. 2. | me.3. | mt. 5. | mt. d. | mt. 3. | (borealis). Cales | Cales Cales | Cales Cales Ipecedale Brit. Mus, Dale. Dale. Dale. | Dale. Dale. | Sees lm o3 Ose 2 ra | LOAN .ccoongaraceosna| de || Bee BYO | one ms 3°80 4:07 Transverse diameter | (proximal end) ... BS || le “AT | 52 | 45 55 Transverse diameter | (distal end)......... “51 “46 WL