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PROCEEDINGS 


OF THE 
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS 


OF THE 


AVOLOGICAL SOCIETY | 


OF LONDON. 


1903, vol. II. 


(MA Y—DECEMBER.) 


Apr -5. ORS [ 


GQ 5s 
PRINTED FOR THE society, ‘89777 


AND SOLD AT THEIR HOUSE IN HANOVER-SQUARE. 
LONDON: 


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


COUNCIL 


1s iced b 
OF THE 


AND OFFICERS 


OF THE 


ZOOLOGICAL 


SOCIETY OF LONDON. 


1903. 


COUNCIL. 
(Elected April 29th, 1903.) 


His Grace THe Duke oF Beprorpd, K.G., President. 


Witiram T. BiLANForD, Esq., 
LL.D., F.R.S., Vice-President. 

GrorcE A. BouLEeNGER, EsqQ., | 
E.R.S., Vice-President. 

Witiram KE. pE Winton, Esq., 
Acting-Superintendent of the 
Gardens. 

Herserr Druce, Esq., F.LS. 

CHARLES DrumMonbD, Esq., 
Treasurer. 

CHARLES H. Garry, Esq., LL.D. 

FREDERICK GILLert, Esq. 

F. DuCange Gopman, Ksq., 
D.C.L.,F.R.S., Vice-President. 

Apert Gintuer, Esq., M.D., 
Pa.D., F.R.S., Vice-President. 


Carr. THE Marguis or HAmit- 
TON, M.P. 

Pror. GEorRGE B. Howns, D.Sc., 
LL.D., F.R.S., Vice-President. 

Lir.-Cou. L. Howarp Irpy. 

Sir Epmunp G. Loprr, Br. 


_ K. G. B. Meapr-Watpo, Esa. 


P. CuatMers Mircney, Esq., 
M.A., D.Sc., Secretary. 

Sir THomas Pare. 

K. Lorr Puituips, Esq. 

Davin Swarr, Esq:. 
F.R.S. 

OLDFIELD THomas, Esq., F.R.S. 

Henry Woopwarp, Esq., LL.D., 
F.R.S., Vice-President. 


M.D., 


PRINCIPAL OFFICERS. 


P. CuauMeErS Mircue, Esq., M.A., D.Sc., Secretary. 
W. E. pe Winton, EsqQ., Acting-Superintendent of the 


Gardens. 


Frank E. Bepparp, Esq., M.A., F.R.S., Prosector. 

Mr. F. H. Warernouse, Librarian. 

Mr. Joun Barrow, Accountant. 

Mr. W. H. Core, Chief Clerk. 

Mr. Georce Artaur DouBLepay, Clerk of Publications. 
Mr. Artruur THomson, Assistant Superintendent of the 


Gardens. 


LIST OF CONTENTS. 


May 12, 1903. 


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


Myr. 


My. 


Menagerie ins AjprillS Ooiiase srete setae. ioe teeter oar coy 


Frank Finn, F.Z.8. Exhibition of drawings of, and 
remarks upon, an abnormal pair of horns of the Barking- 
Weer Cer ianlas etiiet) CC! be kiss. ytd as. oe cose cre eerie 


Frank Finn, F.Z.8. Exhibition of, and remarks upon, 
specimens of the Goldfinch (Carduelis carduelis) and the 
Ruff (Pavoncella pugnax) showing variation in coloration 
On | oe ADDON ZOE) A jue Ari iam nye eet ARE eas 00. Sl Are oA EE Yet 


- F. KE. Beddard, F.R.S. Exhibition of some preserved 


and injected Diane @ te) Main inal cay eee Ne OE) ao Volpe 


A Contribution to the Study of Double Monstrosities in 


Fishes. By James F. Gemurty, M.A., M.D., Lecturer 
in Embryology, Glasgow University. (Plates I.-IV.)... 


The Metamorphoses of the Decapod Crustaceans Zgeon 


(Crangon) fasciatus Risso and Lgeon (Crangon) tri- 
sptnosus (Hailstone). By Roserr Gurney, B.A., F.Z.8. 
(Pints Vain ee ee eo ny 


Descriptions of new Species of South American Coleoptera 


of the Family Chrysomelide. By Marry JAcosy, F.E.S. 


On a Collection of Fishes made by Dr. Goeldi at Rio 


Janeiro. By C. Tare Recax, B.A. (Plates VIL. & 
RVING) Ris errectti sy ear. aiansboat Siegen Saute aie seem rats wheats 


24 


30 


Iv 


May 26, 1903. 


1. List of the Batrachians and Reptiles collected by M. A. 


bo 


(su) 


Robert at Chapada, Matto Grosso, and presented by 
Mrs. Percy Sladen to the British Museum. (Percy 
Sladen Expedition to Central Brazil.) By G. A. 
IBOULENGER os.) V le Z0 2 sie Ge eseese er cree ine eee 


. Note on some Mollusks of the Family Bulimulide from 


Matto Grosso. (Percy Sladen Expedition to Central 
Brazil.) By Enear A. Surry, .8.0., F.Z.8. ............ 


. The Present State of Knowledge of Colour-heredity in 


Mice and Rats. By W. Batveson, M.A., F.R.S., F.Z8., 
Fellow of St. John’s College, Cambridge .................. 


4. On the Marine Fauna of Zanzibar and British Hast 


The 


TD ye, 


Africa, from Collections made by Cyril Crossland im 
the Years 1901 and 1902.—Turbellaria Polyeladida. 
Part I. The Acotylea. By F. F. Larpiaw, B.A.Cantab., 
Assistant Demonstrator and Lecturer in Biology in the 


Owens College: ”\(Blate TX) hoe iss. 3. Ieee eee 


June 16, 1903. 


Secretary. Report on the Additions to the Society’s 
Menagerie in May 1903 


. Frank Finn, F.Z.8. Exhibition of, and remarks upon, 


a living hen-feathered Bantam Cock 


-- G. A. Boulenger, F.R.S. Exhibition of, and remarks 


upon, a specimen of the Frog Ceratohyla bubalus carrying 
eges on its back 


>, F. E. Beddard, F.R.S. Exhibition of a bust of the late 
pee hl (G) 


President of the Society, Sir W. H. Flower, K.C.B. 


. F. E. Beddard, F.R.S. Exhibition of, and remarks 


upon, sections of the ovary of the Thylacine 


Dionne, cocececc 


 R. E. Holding. Exhibition of some skulls and horns 


of, and remarks upon the horn-growth in, the St. Kilda 
four-horned Sheep 


A. 5. Woodward, F.R.S. Exhibition of photographs 


illustrating the exhumation of a recently-discovered 
Mammoth in Siberia 


DO OO GCOS OO ODO OOOO NOOO AO SOO OOGOD AOR 


Page 


69 


114 


114 


116 


116 


Vv 


Mr. Oldfield Thomas, F.R.S. Exhibition of, and remarks 
upon, the horns of a new form of Grant's Gazelle, 
proposed to be named G'azella granti robertsi............... 


1. Onan Extinct Species of Genet (Genetia plesictoides, sp. n.) 
from the Pleistocene of Cype us. By Dororny M. A. 
BATES a (abe ik )\at tear eeseen ene bce Peon kon dha fast 


bt 


. Description of a new Fish of the Gobiid Genus Phiacichthys 
from British New Guinea. By G. A. Bounencer, F.R.S8., 
WEES ey aCe late XGIe) 4.5 ences ciate Ce ich sate, atts ee 


. Descriptions of new Reptiles from British New Guinea. 
By G. A. Bounencer, F.R.S., V.P.Z.8. (Plates XII. 
Ci DG] TIE EN gia epi heat amines ue oman aa IC oie sd 


(su) 


4, On the Marine Fauna of Zanzibar and British East 
Africa, from Collections made by Cyril Crossland in the 
Years 1901 and 1902.—Polycheta. Part IT. By Cyrin 
CRosstAnD, B.A., B.Sc. (Plates XIV. & XV.) ......... 


5. On the Ento - Parasites collected by the ‘“Skeat Ex- 
pedition” to Lower Siam and the Malay Peninsula in 
the Years 1899-1900. By Arruur E. Suipuey, M.A., 
F.Z.S., Fellow and Tutor of Christ’s College, Cambridge, 
and University Lecturer in the Advanced Morphology 
of the Invertebrata. (Plate XOV 1.) 25.205... 


6. On the Modifications of Structure in the Syrinx of the 
Accipitres, with Remarks upon other Points in the 
Anatomy of that Group. By FmMank EH. Bepparp, M.A., 
ERS brosectormto une; Societys re es ee ee 


7. On Medusze from the Coast of British Columbia and 
Alaska. By Louis Murpacu and CressweLi SHEARER, 
eZee See (Eula best ROVE aRONIME VP ee ss ieee tans Oeaetuon 


November 3, 1903. 


The Secretary. Report on the Additions to the Society’s 
Menagerie in June, July, August, and September, 1903. 


Dr. P. L. Sclater, F.R.S. Exhibition of, and remarks upon, 
the horn of a Rhinoceros from the White Nile ......... 


The Secretary. Exhibition of, on behalf of Mr. H. Slade, a 
series of photographs of the Indian Elephant in the act 
OM COMELESS AE Mae saa sone cess Gt eae ou Renee NL LD, Bie 


164 


193 


Ma 


Mr. 


My. 


Myr. 


bo 


(SU) 


1 


~I 


Vi 


-. Henry Scherren, F.Z.S. Exhibition of, and remarks 
upon, abnormal! claws of the Crab and Lobster...........- 


R. 1. Pocock, F.Z.8. Exhibition of, and remarks upon, 
C = 7 - a 
photographs of a mounted specimen of Burchell’s Zebra. 


R. I. Pocock, F.Z.S. Exhibition of a large Geophilo- 
morphous Centipede ‘aon, WGMEVAUIOIB,, .c500c00cd00ccosconcasee 


Oldfield Thomas, F.R.S. Exhibition of specimens and 
descriptions of new species of Mammals from New 
Crm, (Peis) OUI) ase bancznosserndasee sopeeo Dosen e002 


. On some new Species of Aquatic Oligocheta from New 


Zealand. By W. B. Brnuam, D.Sc., M.A., F.Z.S., 
Hon. M.R.S. Tasm.; Professor of Biology in the 


University of Otago. (Plates XXIV—XXVI.) ......... y 


. On the Mammals collected by Ma. A. Robert at Chapada, 


Matto Grosso. (Percy Sladen Expedition to Central 


Brazil.) By Ouprirtp THomas, F.R.S. (Plate XX VIL.) 2 


. List of the Coleoptera collected by My. A. Robert at 


Chapada, Matto Grosso. (Percy Sladen Expedition to 
Central Brazil.) By C. J. Ganan, M.A., and G. J. 
IAG ROW. woe late DXOXG VHT A c- Rares bare ee eee et eee 


. On the Lepidoptera collected at Chapada, Matto Grosso, 


by Mr. A. Robert. (Perey Sladen Expedition to Central 
Brazil.) By F. A. Heron and Sir Grorce F. Hampson, 
BAGS EZ at ih ak Aa sey dsr nes) | Reser 


. Note on the Cypriote Spiny Mouse. By P. Cuatmurs 


Mrrene yt, M.A., D.Sc., Secretary to the Society 


eee eters 


. On the Muscles of the Ungulata. By Brrrram C. A. 


Winniz, D.Sc., M.D., M.A., F.R.S., Professor of 
Anatomy in the University of Birmingham, and F. G. 
Parsons, F.R.C.S., F.Z.8., F.L.8., Lecturer on Human 
and Comparative Anatomy at St. Thomas’s Hospital, 
late Hunterian Professor in the Royal College of Sur- 
geons, England.—Part II. Muscles of the Hind-limb 
and Trunk 


. On some Points in the Anatomy, chiefly of the Heart and 


Vascular System, of the Japanese Salamander, J/egalo- 
batrachus japonicus. By Franx EK. Bepparp, M.A., 
F.RB.S., F.R.S.E., Prosector to the Society 


Page 


195 


196 


244 


258 


260 


261 


vu 
November 17, 1903. 


The Secretary. Report on the Additions to the Society’s 
Menagerie in October 1903.......-.:-..:-seeeereeteetttnstes 


Mr. Henry Scherren, F.Z.5. Exhibition of, and remarks 
upon, a horn of Rhinoceros SHUDLIS chess soo ancabsosouaasacooe 


Mv. R. I. Pocock, F.Z.8. Exhibition of a piece of basalt 
containing a web of the Marine Spider Desis kenyone... 


My. RB. I. Pocock, F.Z.8. A new suggestion as to the use of 
the white rump-patches in the Ungulata ................-- 


My. BE. E. Austen. Exhibition of, and remarks upon, spe- 
cimens of the Tsetse-fly which acts as transmitter of 
“ sleeping-sickness ” in Uganda .....-..1+--s.s-seeerteeeeees 


My. Oldfield Thomas, F.R.S. Exhibition of, on behalf of 
Mr. W. E. de Winton, F.Z.S8., a drawing of a female 


Gazelle bearmg a hair-whorl on the withers .............+. 5 


Mr. C. Tate Regan. N otice of a Memoir entitled ‘* Revision 


of the Fishes of the Family Loricaritde” .........-++.-.6+ é 


1. Ow Barly Sanskrit References to the Tiger. Biya Vaivis 
RAMANAN, M.A., B.Z.S. oo... cceceeec eee e sec ee sete ec ec eeec eens 


2. On the Trachea, Lungs, and other Points in the Anatomy 
of the Hamadryad Snake (Ophiophagus bungarus). By 
Fravx E. Bepparp, M.A., F.R.S., Prosector to the 


SOGGY cngooddogdeoononc eucsdceesqusdene Coanepab do n=edeeRsbuadHac: : 


3. Report on the Fishes collected by Mr. Oscar Neumann 
and Baron Carlo von Erlanger in Gallaland and Southern 


Ethiopia. By G. A. Bovtencer, F.R.S., V.P.Z.S. 


(plievias YORIDN = OTC) Ligsadsnonlbuedae Menteur ten 


December 1, 1908. 


Mr. F. Martin Duncan. Demonstration of the Zoological 
BiOSCOPC...-..-..02-eeceeceec sete cee ee ese teaaneecteecsec ecco sene 


My. F. E. Beddard, F.R.S. Exhibition of, and remarks upon, 
a portion of the large intestine and the czecum of a Boa. 


Mr. F. BE. Beddard, F.R.S. Exhibition of, on behalf of 
Mr. G. A. Doubleday, a hairless specimen of the 
(Srotitein’ LAB ie onaseennonsopontdcodgesonqgso0eDpEaeaqcoUecdpuodcour 


Page 


315 


316 


316 


316 


a4 


Dr 


My. 


Vill 


. Walter Kidd, F.Z.S. Exhibition of a drawing of a 
Beisa Antelope showing a reversed area of hair on 
cH Rey: ava 0) tei Coie ae cles he Oe eA ma sunt cs acauaquenadgooNS 


>, Oldfield Thomas, F.R.S. Exhibition of, and remarks 


upon, a specimen of the Rodent Fornarina (gen. nov.) 
phillips and description of Heterocephalus ansorget 
(Gs WOW), cdocandessosson copessocmonanaapbao see soneHooqunccosocced 


G. A. Boulenger, F.R.S. Exhibition of a hybrid Newt 
between Molge marmorata $ and M. cristata 2 ......-.. 


Prof. E. Ray Lankester, F.R.S. Exhibition of, and remarks 


upon, two drawings representing the arrangement of 
the hair on the faces of two specimens of Okapi ......... 


Prof. E. Ray Lankester, F.R.S. Exhibition of specimens of 


ey 


Meduse from the Victoria Nyanza.......................000 


. The Mammals of Cyprus. By Dororuy M. A. Bate...... 


. On the Cause of Death of a Polar Bear recently living in 


the Society’s Gardens. By Repciirre N. SALaman, 
M.A., M.B. Cantab., F.Z.S., Acting Director of the 
Pathological Institute, London Hospital .................. 


. On the Development of the Adult Colouring in the Yellow- 


billed Cardinal (Paroaria capitata) from 8. America. 
By Arruur G. Burier, Ph.D., F.LS., F.Z.8., &e. ...... 


. On the Occasional Transformation of Meckel’s Diver- 


ticulum in Birds into a Gland. By P. CHALMERS 
MircnEy, M.A., D.Se., Secretary to the Society 


. On some Nudibranchs from East Africa and Zanzibar. 


Part IJ. By Sir C. Extot, K.C.M.G., H.M. Com- 
missioner for the East Africa Protectorate, F.Z.S8.— 
Dorididee Cryptobranchiate, I. (Plates XXXII. — 
XXXIV.) 


DOICIOOOIICCICTIUOICICOIOICIOIOIO I IOICIOOn IOICIO ICICI ICICICICICICICICICICICICICICIAC Ir Irir 


Vultures, with Remarks on the Interrelations of the 
Genera Sarcorhamphus, Gypagus, and Cathartes. By 
FRANK E. Bepparp, M.A., F.R.S., Prosector to the 
Society 


emer cece eee eseceres cr cr es esc cer eccccoage 
Fee cece ewe nese sees 


Page 


336 


336 


337 


337 


340 


341 


348 


390 


304 


. A Note upon the Tongue and Windpipe of the American 


ALPHABETICAL LIST 


OF THER 


CONTRIBUTORS, 


With References io the several Articles contributed by each. 


Arrow, G. J., and C. J. Ganan, M.A., of the British 
Museum (Natural History). 


List of the Coleoptera collected by Mr. A. Robert at 
Chapada, Matto Grosso. (Percy Sladen Expedition to 
Central, Brazils)ha Ge laitey Oxia V alin S83 ese tA: ee cel 


Austen, KE. E., of the British Museum (Natural History). 


Exhibition of, and remarks upon, specimens of the 
Tsetse-fly which acts as transmitter of ‘“sleeping- 


SIC Mess amin U fran Ceo M enn ne tee cee UM etre eames NoaSer een 


Bares, Miss Dororuy M. A. 


On an Extinct Species of Genet (Genetta plesictoides, 
sp. nu.) from the Pleistocene of Cyprus. (Plate X.) ...... 


ihegMiannatal stom Corus seas. ate sae ce ener a fais: 


Page 


244 


5316 


aN 


Bateson, Wittiam, M.A., F.R.S., F.Z.8., Fellow of St. 


John’s College, Cambridge. 


The Present State of Knowledge of Colour-heredity in 
Wie aHAl RENTS | bsassecceh-daascesgan soa0ddec cb obey onecd saan aboo5 23 


Bepparp, Franx E., M.A., F.R.S., F_R.S.E., Prosector to 
the Society. 


Exhibition of some preserved and injected brains of 
Wiermnmnalls - iiss dendeosayesecokuovesosetococscdaocusnopaaaseocnabboes 


Exhibition of a bust of the late President of the Society, 
Sir W. H. Flower, K.C.B. 


Exhibition of, and remarks upon, sections of the ovary 
of the wlhnyacime ass: 2a -)s- eee - eer etme a eee rrr er ae 


On the Modifications of Structure in the Syrinx of the 
Accipitres, with Remarks upon other Points in the 
Anatomy of that Group 


On some Points in the Anatomy, chiefly of the Heart 
and Vascular System, of the Japanese Salamander, 
Megalobatrach us aponves oot eee elatp eles lee ee eee 


On the Trachea, Lungs, and other Points in the Ana- 
tomy of the Hamadryad Snake (Ophiophagus bungarus). 


Exhibition of, and remarks upon, a portion of the 
large intestine and the cecum of a Boa 


Exhibition of, on behalf of Mr. G. A. Doubleday, a 
hairless specimen of the Common Rat ..,.................008: 


A Note on the Tongue and Windpipe of the American 
Vultures, with Remarks on the Inter-relations of the 
Genera Sarcorhamphus, Gypagus, and Cathartes ......... 
Brenuam, W. Buaxuann, D.Sc., M.A., F.Z.8., Hon. M.R.S, 

Tasm.; Professor of Biology in the University of 
Otago, 
On some new Species of Aquatic Oligocheta from New 

Zealand. (Plates XXIV.XXVL.) 


Peet eee sees ar ese n ese rere sces 


Page 


386 


Xi 


Page 
BovLencer, GeorcE ALBERT, F.R.S., V.P.Z8. 
List of the Batrachians and Reptiles collected by M.A. 
Robert at Chapada, Matto Grosso, and presented by 
Mrs. Percy Sladen to the British Museum. (Percy 
Sladen Expedition to Central Brazil.) ........................ 69 
Exhibition of, and remarks upon, a specimen of the 
Frog Ceratohyla bubalus carrying eggs on its back ...... 115 
Description of a new Fish of the Gobiid Genus 2hiac- 
ichthys from British New Guinea. (Plate XI.) ......... 124 
Descriptions of new Reptiles from British New Guinea. 
(LE RSS > GILL iha oS HS) Aine Reich Ue ai MaMa dule a 125 


Report on the Fishes collected by Mr. Oscar Neumann 
and Baron Carlo von Erlanger in Gallaland and Southern 
IDWano pie, UeeWeS! SOLO Orn) Bose shun danadunasarocooe 328 
Exhibition of a hybrid. Newt between JJolge mar- 


TOOIREMMED) (Ge EMIENG! DUE ROSEN. QO Y aclsoemoee sead Aeaeasdaceok odes sc 337 


Burter, ArraHurR GARDINER, Ph.D., F.L.S., F.Z.8. 
On the Development of the Adult Colouring in the 
Yellow-billed Cardinal (Paroaria capitata) from 8. 
BASTIEN Cal icr eel e ra2t etal aie BML ae eh AM IN WOE Cah Lh Mlk 3d0 


CrossLAnbD, Cyrin, B.A., B.Sc. 

On the Marine Fauna of Zanzibar and British East 
Africa, from Collections made by Cyril Crossland in the 
Years 1901 and 1902.—Polycheta. Part II. (Plates 
PROTA eNO VW nate Mace tua hice eee hn GR ena FRA UNG 129 


Duncan, F. Martin. 


Demonstration of the Zoological Bioscope B34 


Euror, Sir Cuarues, K.C.M.G., H.M. Commissioner for 
. the East Africa Protectorate, F.Z.S. 
On some Nudibranchs from East Africa and Zanzibar.— 
Part Il. Doridide Cryptobranchiate, I. (Plates 
BNO NO XU, NO NONGTN IAW rien A oreo nts ORR yore Ue Med a). 304 


xi 
Finn, Frayne, B.A., F.Z.8. 

Exhibition of drawings of, and remarks upon, an 
abnormal pair of horns of the Barking-Deer, Cervulus 
TMUTUITED copdrsoqopsseooesocnsacdopeesobasysccnco essccononeocDGonec 

Exhibition of, and remarks upon, specimens of the 
Goldfinch (Carduelis carduelis) and the Ruff (Pavoncella 
pugnasx) showing variation in coloration of plumage ...... 


Exhibition of, and remarks upon, a living hen-feathered 
IBeininayiny Cloyeliet oh saceceds no ghenddckauesebadssdnovonesdses 2n90-cocb00; 


Ganay, C. J., M.A., and G. J. Arrow, of the British 
Museum (Natural History). 

List of the Coleoptera collected by My, A. Robert at 

Chapada, Matto Grosso. (Percy Sladen Expedition to 

(Cloatrenil 1Bieavaill,)) (UP eise SO WIND.) ogcoboossesocarguacnoneses 


GremMitL, JAMEs F., M.A., M.D., Lecturer in Embryology, 


Glasgow University. 
A Contribution to the Study of Double Monstrosities 
in Fishes. (Plates I-IV.) 


GURNEY, Ropert, B.A., F.Z.8. 


The Metamorphoses of the Decapod Crustaceans “geon 
(Crangon) fasciatus Risso and .geon (Crangon) tri- 
spimosus (Hailstone). (Plates V. & Vi.) .........72.......-.. 


Hampson, Sir Grorce F., Bt., F.Z.S., and F. A. Heron, of 
the British Museum (Natural History). 

On the Lepidoptera collected at Chapada, Matto Grosso, 

by Mr. A. Robert. (Perey Sladen Expedition to Central 

1 ETSI) I iia tr es. LO nO eX eat a EIR 

Heron, F. A., of the British Museum (Natural History), 
and Sir Groree F. Hameson, Bt., F.Z.S. 

On the Lepidoptera collected at Chapada, Matto Grosso, 

by Mr. A. Robert. (Percy Sladen Expedition to Central 

Brazil.) 


Page 


bo 


114 


244 


258 


258 


xili 
Houpine, Ricwarp H. 
Exhibition of some skulls and horns of, and remarks 
upon the horn-growth in, the St. Kilda four-horned 
Sheep 


Jacosy, Martrn, F.E.S. 


Descriptions of new Species of South American Coleo- 


ptera of the Family Chrysomelide: 


Kipp, Watrer, M.D., F.Z.8. 


Exhibition of a drawing of a Beisa Antelope showing 

a reversed area of hair on the back ....................+-..00- 

Laipiaw, F. F., B.A.Cantab., Assistant Demonstrator and 
Lecturer in the Owens College. 


On the Marine Fauna of Zanzibar and British East 
Africa, from Collections made by Cyril Crossland in the 
Years 1901 and 1902.—Turbellaria Polycladida. Part I. 
Mhewdcotylem@s ¢ (Ue lateyEXe), Coasts 4:Bhe te woh ote 
LANKESTER, Prof. EK. Ray, M.A., LL.D., F.R.S., F.Z.S., 

Director of the British Museum (Natural History). 
Exhibition of, and remarks upon, two drawings repre- 
senting the arrangement of the hair on the faces of two 


specimens of Okapi 


Exhibition of specimens of Meduse from the Victoria 


ING EZONR ee fee. nN eee ant Scie eect We My Rl OeMw NAYS a 
Mircuent, P. Cuatmers, M.A., D.Sc., Secretary to the 
Society. 
Report on the Additions to the Society’s Menagerie in 
April 1903 


Report on the Additions to the Society’s Menagerie in 
May 1903 


Report on the Additions to the Society’s Menagerie in 
June, July, August, and September, 1903 


Page 


116 


99 


— 


114 


X1V 


Mircueny, P. Cuatmers, M.A. &e. (Continued.) Me 
Exhibition of, on behalf of Mr. H. Slade, a series 
of photographs of the Indian Elephant in the act of 
GOINGS npecnoedocoscosesseennnconaeacsoerseaeopae2epanoaasbDAeRoG*n 195 
Note on the Cypriote Spiny Mouse ........................ 260 


Report on the Additions to the Society’s Menagerie im 
(Oye nalareresl OVO Sakae en ee WU NN a oMlon goon uebdasosnootacc 315 
On the Occasional Transformation of Meckel’s Diver- 


Sromquuan shay JETS IIUFO) Al Gre “Sodacooncassoscacaenoasasoccascns 352 


Morsacu, Louis, and Cresswett Surarer, M.D., F.Z.8. 


On Medusex from the Coast of British Columbia and 


Alaska. (Plates XVIN--XXIL) 2.2.0.0... 164 


Parsons, F.G., F.R.C.S., F.Z.8., F.L.S8., Lecturer on Human 
and Comparative Anatomy at St. Thomas’s Hospital, 
late Hunterian Professor in the Royal College of 
Surgeons, England, and Brerrram C. A. WINDILE, 
D.8c., M.D., M.A., F.R.S., Professor of Anatomy im 
the University of Birmingham. 

On the Muscles of the Ungulata.—Part IT. Muscles 


Olt tine IBGimglellitrnl) armel Weta, 55sccondecccs0cevgaececcssocnsoaucs 261 


Rocockpk: i kEZ:s: 
Exhibition of, and remarks upon, photographs of a 
mounted specimen of Burchell’s Zebra .........0...........05. 196 
Exhibition of a large Geophilomorphous Centipede 
LOIN FINACTNOZAC EN aan utgaatts seen etueen ye a arvmgnce erteN ee Marrs Mh 196 


Exhibition of a piece of basalt containing a web of the 


Moarmae Spider Desiswicen vores mes-eeeen aes eee nee 316 


A new suggestion as to the use of the white rump- 


joenuCl mes) wai WN) UPDGMMENMi a caadsacnboseobiososousasauoonanssnsco sy. 316 
RAMANAN, V. V., M.A., F.Z.8., Deputy Tahsildar and sub- 
Magistrate. 


On Early Sanskrit References to the Tiger 318 


XV 


Reean, C. Tare, B.A., of the British Museum (Natural 
History). 


On a Collection of Fishes made by Dr. Goeldi at Rio 
Janeiro. (Plates VII. & VEIL.) ................ cee 


Notice of 2 Memoir entitled ‘“ Revision of the Fishes 


of the Family JOU DTRURTIROLILIEI |. Seid Soc aS Mages Eno Rb eosao on oF 


SaLAMAN, REpDcLirFE N., M.A., M.B. Cantab., F.Z.8., Acting 
Director of the Pathological Institute, London 
Hospital. 


On the Cause of Death of a Polar Bear recently living 
in the Society’s Gardens 


ScuERREN, Henry, F.Z.8. 


Exhibition of, and remarks upon, abnormal claws of 
the Crab and Lobster 


Exhibition of, and remarks upon, a horn of Rhinoceros 
STIS ass ae, beet UN REO Rt A A RC Sn ea 
Scrarer, Pump Lurtsy, M.A.. D.Se, Ph.D, E-R-S. 

E.Z8. 


Exhibition of, and remarks upon, the horn of a Rhino- 
CErOsyErombhenshnibes Naletsen ose se ae ascene nest ose sec te 


Suparer, Cresswett, M.D., F.Z.8., and Lours Mursaca. 


On Meduse from the Coast of British Columbia and 
iNinaker,  (Tellenses 2/1010) secccnseaetacseedoeepauobeneoc 
Surpiey, Arraur H., M.A., F.Z.S., Fellow and Tutor of 

Christ’s College, Cambridge, and University Lecturer 
in the Advanced Morphology of the Invertebrata. 
On the Ento-Parasites collected by the “ Skeat Expe- 


dition” to Lower Siam and the Malay Peninsula in the 
Nears US99 NGO Os (Plate NOV Is)ocesuonececs onthe oman ne sous 


Page 


318 


348 


195 


516 


194 


145 


Xvl 
Smiru, Epear A., 1.8.0., F.Z8. 


Note on some Mollusks of the Family Bulimulide from 
Matto Grosso. (Perey Sladen Expedition to Central 
[Byg2valle)) Ube doondeosdeousdeorccpgsacobr seses6-goseds or aubbeadauer ous 


Tuomas, OLpFIELD, F.R.S., F.Z.S. 


Exhibition of, and remarks upon, the horns of a new 
form of Grant’s Gazelle, proposed to be named Gazella 


grants FEO PORES RTE, TT CUB ea tran ELS ea Baa VS 


Exhibition of specimens and descriptions of new species 
of Mammals from New Guinea. (Plate XXIII.) ......... 


On the Mammals collected by Mx. A. Robert at Chapada, 
Matto Grosso. (Percy Sladen Expedition to Central 
Tsieavall,)\) (Pewee) SOLOW IIS) Soa scscabeoncedacocodsensseasc vente: 


Exhibition of, on behalf of Mr. W. E. de Winton, F.Z.S., 
a drawing of a female Gazelle bearing a hair-whorl on 


ETN @peavvalits ENGST are cs meade NaI oe hvac Ca SU AL Rc ae 


Exhibition of, and remarks upon, a specimen of the 
Rodent Fornarina (gen. nov.) phillipsi and description of 


Heterocephalus ansorget (sp. MOV.) ...-.-..220-022- +2. see seen 


Winpue, Bertram C. A., D.Sc., M.D., M.A., F.R.S., Pro- 
fessor of Anatomy in the University of Birmingham, 
and F.G. Parsons, F.R.C.S., F.Z.8., F.L.S., Lecturer 
on Human and Comparative Anatomy at St. Thomas’s 
Hospital, late Hunterian Professor in the Royal 


College of Surgeons, England. 


Part Il. Muscles of 


ae 1ebomalellinmmlo aya! Wrawoalk .. cosoaccusasoccaoscous connenoessonsen 


On the Muscles of the Ungulata. 


Woopwarb, Arruur Surra, LL.D., F.R.S., F.Z.S. 


Exhibition of photographs illustrating the exhumation 


Jee eee rene nsvvce 


of a recently-discovered Mammoth in Siberia 


Page 


70 


119 


196 


232 


ale 


336 


261 


LIST OF PLATES. 


1903.—Vot. II. 


Plate Page 
I 
JU s 
lL. Anatomy of Monstrous Fish Embryos .............. 4 
IV. 
an @xanyontlanyecya ay ayaa w in evaeeeb eovekey vavhert ici cise Gettin 24 
WADA OUAGRTTITO ICHIO Fs be Ee CAE HE OU EOD CABO S 45.00 5 l 59 
VII. 1. Peristedion altipinnis. 2. Genypterus brasiliensis... . § 
iXGe Planarians tromy Zanzibar ss see enc tinea. eee tats 99 
X. 1. Genetta genetta (L.); 2-6. G. plesictoides, sp. nov. ; 
Co, SHIPS CRO MAO, 6 ye noncoeonsoo ph DOOD ONO 121 
XI. Rhiacichthys nove-quinee ...... AiG Gab Crono ioe Oca 124 
XI. 1. Lygosomamilnense. 2. L. granulatum. 38. L. een 
XII. 1. Lygosoma pratti. 2. Toxicocalamus longissimus. 125 
Sem SUEATUCY CLUS ON acta, TN a aleNe re TUR ey Arad ai a) aya 8 ch on ws 
XIV. Fig.l. Diopatraneapolitana. 2. Onuphis holobranchiata. 
DON MARDI SO MMOCIRLOSRUM Spiele elas eerie toons 129 
XV. Figs. 7-10. Marphysa mossambica. 11,12. M. simplex, Pi 
DVS Py lo ae eyiercellat anny Spa saa. eis ie 
XVI. Ento-Parasites from Lower Siam .................. 145 
OWAGL. 
XVIII. 
XIX. \ Med Pe g i 
xx, 7 Medusze from British Columbia and Alaska .......... 164 
XXII, 
XXII. J 
XXIII. 1. Anisomys imitator. 2. Hyomys meeki ........0.5. 198 
XXIV. 
XXV.' Aquatic Oligochztes from New Zealand.............. 202 
XXXVI. f 


Proc, Zoou. Soc.—1903, Vout. II. b 


Plate 


XXVII. 
XXVITI. 
XXIX, 
XXX. 
XXXT. 


XXXII. 


XXXII. 
XXXIV. 


Page 
(CHYNS HUTA Sceocdonodebaseoodoo ood md soda DOs 232 
Coleoptera from Central Brazil ............+...-- 244 
IERVAD WEIRD Soagdoboanesosasboocoboognssoga0 
EB ATU USHCTLONG Clit mri Heliie inertial kei -t aerate aa 398 


1. Discognathus makiensis. 2. Discognathus blanfordiv. 
3. Chiloglants modjensts.....+.ceerseevvearreces 

1,2. Thordisa villosa. 3. T. crosslandi. 4. Trippa 
monsont. 5. Halgerda willeyt .......+.seserases 

1-8. Thordisa villosa. 4-8. T. crosslandi .......... L864 

1,2. Halgerda wasinensis. 8. MSclerodoris coriacea. ¢ 
4, Parasitic Copepod. 5,6. Asteronotus hemprichi. 
7. Kentrodoris rubescens.......csserecessescoees J} 


Or mh CO LS 


b> bo bo bs bo 
ore cot 


LIST OF TEXT-FIGURES. 


1903.—Vot. II. 


Page 
PEN bnornaleAmblers iol Cenmvtislius muntiac mean sane ote eens 2 
. Upper and side views of head of Norops sladem@ ............ 69 
. Sketch of anatomy of Paraplanocera aurora ..........++.... 1038 
» LOOSEN ULIOD CHIU ori hos OCH RAO CE OO UL Cin CD Somme binun doles 104 
. Anterior end of Stylochus zanzibaricus, showing the arrangement 
OMENS CERSIOCHS uo bblomniob bhlo oA tm Utieeoelne Lrplaod noloeh ool 106 
STDC OMLATCONCLLOGW inte, ice) seveeny -1i cs mii stant, aeialas sR ts fats e PHU Soe 109 
. Anterior end of Ommatoplana tuberculata, to show the arrange- 
AETIGy Olt Mere Vas DOCS aan «cheapie ster c7a wrt a wer cite ot sha eh ates ar 
. Ceratohyla bubalus, 2, carrying eggs on back................ 115 
. Variations in the horns of St. Kilda Sheep .................. LA 
. Skull and horns of Gazella granti robertst. (Front view.) .... 119 
. Skull and horns of Gazella granti robertst. (Lateral view.).... 120 
. Marphysa macintoshi. Mandibular jaw-plates .............. 138 
5 aE: Se Means! “MINOT Babe abe bes Gos Adembouaougy e 141 
. Marphysa simplec. Ends of lower or mandibular jaw-plates 
ALG Met he nventrallssideye cree res eve aera ators wan we eee a 142 
. Marphysa furcellata. Dorsal or inner aspect of ventral jaw- 
LEWES FLERE ME Oh A Amaia A Sete eS eee Rel mes eo 142 
. Syrinx of Milvago chimango, lateral view............--...50, 158 
5 \AMIB? Olt DTS URGES eHow o od ood bobo Odor boa coop one 159 
> PS Ubade Ot IONS O RATS SALT TU, noo buawadeauacoboosoanoue 161 
. Tongue of Milvago chimango, dorsal aspect ...............4.- 162 
. Tongue of Gypohierax angolensis, dorsal aspect .............. 162 
. (a) Lateral and (6) front views of horn of Rhinoceros from the 
Wi li CenINillemeeevsrersiats eemsicv ion sceuse te rotates han aL crates c hoes 194 
. Mounted Burchell’s Zebra in the City Museum, Bristol........ 197 
. Plan of genital organs in Haplotaais heterogyne.............. 224 
A Miusclestotoutemsiderofubhiohvot Elyraxne sees aes. | 260 


Hip-region of Sheep, showing double Quadratus femoris muscle. 269 
= Muscles ofannerisidevot Niehof Eyraxg.5 ses 2020s eso 274 


XX 
Page 


27. Diaphragm of Harnessed Antelope. (Abdominal surface.) .... 287 
28, Dissection of the base of the liver and lung of Megalobatrachus 
on the right side of the body, to show arrangement of 
Gboaenons Gir Oval g56cccasgeduo sco 600000. sc0mgsu00000 300 
99, Ventral view of Heart of Megalobatrachus ........+--.+++++: 303 
30. Auricular septum of Megalobatrachus, viewed from in front. ... 305 
31. Transverse section through right branch of Synangium of Megalo- 


NRTA ee LAS SAE OO ee Oe La eeONG Gyo Hilo 308 


2, Junction of vascular arches of Megalobatrachus to form dorsal 


GVEA RC erate Ra (A one eae pein Tre neatly ays Od feava ag eG 309 


2 Heart and Arterial Trunks of Megalobatrachus from the side .. 311 


34, Dissection illustrating relation of arteriz comites aorte of 
Megalobatrachus to “diaphragm” .....-..--. 6. eee ee eee 314 

. Diagrammatic sketch of skin of Gazelle to show hair-whorl on 
WulNGe been aaeMeo boon MuO BABI ke HOE yeaa ou oA Nn odo 6 317 
. Gall-bladder of Ophiophagus bungarus ..... 0.00 cece renee nees 320 
SS Dissegtronomneck ot Ophvophagese ae wierd tee alltel 323 


. A more detailed view of two air-sacs and dividing septum in 


Qa AMIGNBs 9004 50a0000t006004 Pay Bes Co n'a pro Oia O24 


. A portion of trachea from dorsal surface of Ophiophagus, showing, 


A, perforation leading into air-sac, and, B, circular thinning 
Ci MANCUEAM WNT NGI), «Gococaceonsoeocss Sobngohodacd Rand: CHD 


. Ending of trachea in lung of Ophiophagus ..... Been eo cmlad 6 326 
. Diagram of the hair-whorls and hair-streams on the fronto- 


parietal region of the head of a female specimen of Okapi in 


Sa IRICEN A CURIS ee NGG hn nin OMe SOD Eee dodo eooals 338 

2. Diagram of hair-whorls and hair-streams on the fronto-parietal 

region of the head of a small adult female specimen of Okapi 
Tim, Whe, IROWNEeMlClis MIMEEW, soon0000bon0vn0andanoncne . 339 
JEFPULTOGIS COPULIB o's Sobb0b0 Cao RED Ob SO Ob OadobdoDODS ARR PRS 343 
Meckel’s diverticulum in the Woodcock .............. ans 353 

. Dorsal view of tongue of Vultur auricularis. Dorsal yew of 
HOLA) Ohi, CIPRO HN oo ooae odsonsooderbodesooac neces: 387 

. Sarcorhamphus equatorialis; insertion of maxillo-jugal: Mandible, 
viewed from! behind hades cts sere ethene eee . 390 

. Cathartes atratus ; insertion of maxillo-jugal : Mandible, viewed 
PROM DEMING cy Weces wields cietersa tas W ecsheraihe es Beet Biotest ae 390 

. Gypagus papa; insertion of maxillo-jugal: Mandible, viewed 
pope |KSVVUACL 645466000 PRET ORIOL SRO SRST Eee Un ag 391 


LIST OF NEW GENERIC TERMS 


PROPOSED IN THE PRESENT VOLUME (1903, vot. IT). 


Page 
Anisomys (Mamm.) ............... 199 
Disparoplana (Vermes) ............ 103 
Fornarina (Mamm.) ............... 336 
Haploplana (Vermes) ...........- 109 
Eivoniys((Meamiin,)) sos csssass ses 198 


Page 
Mylacrodon (Pisces) ...........0+0 2 
Ommatoplana (Vermes) ... 111,113 
Phylloplana (Vermes) ...... 100, 107 


Sclerodoris (Moll.). 355, 360, 361, 380 


| Taupodrilus (Vermes) ............ 209 


Proc. Zoot. Soc.—1903, Vou. II. c 


ERRATA. 


ie 


Page 46, line 4 from bottom, for NEOFASCIATA ead CHALYBHOFASCIATA 


Plate XXXTYV. for Hatemrpa wassinensis read HALGERDA WASINEN 18. 


PROCEEDINGS 


OF THE 


GENERAL MEETINGS FOR SCIENTIFIC BUSINESS 


OF THE 


ZOOLOGICAL SOCIETY OF LONDON. 


1908, Vol. II. (May to December). 


May 12, 1903. 


Dr. Henry Woopwarp, F.R.S., Vice-President, 
in the Chair. 


The Secretary read the following report on the additions that 
had been made to the Society’s Menagerie in April 1903 :— 

The registered additions to the Society's Menagerie during the 
month of April were 200 in number. Of these 86 were acquired 
by presentation and 2 by purchase, 14 were born in the Gardens, 
97 were received on deposit and 1 in exchange. The total 
number of departures during the same period, by death and 
removals, was 94, 

Among the additions special attention may be called to the 
following :— 

A Bactrian Camel (Camelus bactrianus), born in the Menagerie 
April Ist. 

A Blesbok (Damaliscus albifrons), received in exchange April 8th, 

Two Green Geckos (Phelswma madagascariense), deposited by 
the Hon. Walter Rothschild, M.P., F.Z.8., April 11th. 

A large collection of South African animals presented by Col. 
A. T. Sloggett, C.M.G., April 25th. 


Proc. Zoon. Soc.—1903, Vou. Il. No. I. 1 


2 ON ABNORMAL ANTLERS OF THE MUNTJAC, [May 12, 


Mr. W. B. Tegetmeier, F.Z.S., exhibited a skin and some 
illustrations of, and made remarks on, a species of Pheasant from 
Mongolia, which had recently been described under the name of 
Phasianus hagenbecki. He suggested that it would make a 
handsome addition to our coverts. A full account of Hagenbeck’s 
Pheasant will be found in ‘The Field,’ vol. ci. p. 775 (1903). 


Myr. Frank Finn, F.Z.8., exhibited drawings, half life-size, of 
the frontlet of a specimen of the Kakur or Barking- Deer (Cervulus 


Text-fig. 1. 


Abnormal antlers of Cervalus muntjac. 


muntjac) showing a curious abnormality of 

y of the antlers (see text- 
fig. 1). The specimen had been lent to him by an a in ie 
Indian Medical Service, who had shot the deer in the valley of the 


1903.] ON COLOUR-VARIATION IN THE GOLDFINCH AND RUFF, 3 


Rydak River, north-east of the Buxa Cantonment, at an elevation 
of 5000 feet. He made the following remarks upon the exhibit 
and on other variations in Deer : 

“The specimen is remarkable as exhibiting a pair of antlers, 
springing from the bony pedicels on their outer sides, some- 
what below the ordinary antlers, which are normal in form. 
Although not alike on the two sides, these little supernumerary 
antlers have not, except for this, an unnatural appearance ; and as 
the local shikaries had informed my friend that in a part of 
Bhutan such a variation was not uncommon, it may be that we 
may ultimately see a four-horned race of the Barking- Deer. 

“While on the subject of variation in Deer, I may perhaps be 
permitted to mention two curious colour-abnormalities in these 
animals, which have come under my notice in India. In the 
above-mentioned Kakur I have seen two melanic specimens, both 
living females, one of which, to my knowledge, came from the 
Himalayas, and, I believe, the other also. Both were of a peculiar 
iron-black colour, 7. e. black of a greyish shade, like that of cast- 
iron ; this grey shade pertaining to the individual hairs, for there 
was no admixture of white ones. Also, in the Sambhar (Cervus 
unicolor) I have seen two remarkable varieties in quite young 
specimens. One of these was a very rufous specimen, as red as a 
Barking-Deer; while the other, which I saw the day after its 
birth, was spotted with white along the sides of the back, like the 
adult Barasingha (Cervus duweauceli) in its summer coat. Young 
Sambhar are commonly said to be more rufous than adults, 
though I have not noticed this in those I have seen; and Mr. W. 
Rutledge, in whose possession I saw the above rufous specimen, 
in spite of his long experience as a dealer, was so surprised at 
this one that he was doubtful of the species. As to the spotted 
fawn, there is no doubt of its rarity as a variation; I have never 
seen or heard of a similar specimen, the usual uniformity of 
the young pelage being a remarkable specific character of the 
Sambhar.” 


Mr. Frank Finn also exhibited a living specimen of the Gold- 
finch (Carduelis carduelis) showing a rare variation, and a skin of 
the Ruff (Pavoncella pugnax) showing albinism, and made the 
following remarks upon them :— 

“The living Goldfinch now exhibited is a male of the large 
Eastern race sold as ‘Siberian Goldfinches’ by bird-dealers, and is 
yemarkable in that it shows an extremely rare variation in colour. 
Behind the black band on the head there is on each side a small 
patch of glossy red feathers, similar to those of the face. I have 
never before seen such a variation either in this species or in the 
Himalayan Goldfinch, and the dealer from whom I procured it, 
Mr. A. Zache, of Great Portland Street, told me he had only seen 
one other, alsoa male. Mr. H. Blake-Knox, however, in a valuable 
paper on ‘Abnormal Plumages in the Goldfinch’ (Zoologist, 


4 DR. J. F. GEMMILL ON [May 12, 


Qnd series, 1870, pp. 2049-2052), has given several instances of 
the abnormal extension of red in this species, some forms of which 
appear to be fairly common. 

“Some attention has been directed of late to a variety of the 
Ruff which has the head and neck white or nearly so, even in 
winter plumage when the ornamental plumes are absent. The 
specimen exhibited died recently in the Zoological Society's 
Gardens, having made, at any vate this year, little progress in 
putting out a ruff. Specimens of the variety kept in the Calcutta 
Zoological Garden, however, have produced the nuptial plumage, 
which was as white as the same parts had been in winter. There 
were two of these birds, which differed in their back-feathering 
as much as other Ruffs, one being rufous-mottled, while the 
other was grizzled. The variation is evidently albinistic, as it 
may, though very rarely, affect the Reeve as well as the Ruff. 
There is such a specimen in the Indian Museum, procured half-a- 
century ago by Blyth. I myself have obtained several specimens 
of the form in the Calcutta Bazaar, which, without exception, 
were adults, as shown by the colour of their feet; so that the 
variation in question would seem to supervene late in the life of 
the individual possessing it. The white-headed birds kept alive 
did not seem at all deficient in vigour, and no doubt under 
especially favourable circumstances this very beautiful variety 
might become quite common. It has, however, the obvious 
disadvantage of being very conspicuous, which may, perhaps, 
account for its restriction to adult individuals.” 


Mr. F. KE. Beddard, F.R.S., exhibited and made remarks upon 
some preserved and injected brains of Mammalia which had been 
prepared from specimens formerly living in the Society’s Gardens. 


The following papers were read :— 


1. A Contribution to the Study of Double Monstrosities in 
Fishes. By Jamus F. Gummint, M.A., M.D., Lecturer 
in Embryology, Glasgow University *. 

[Received May 12, 1903.] 
(Plates I-IV. + ) 


This paper contains an account of the anatomy of a set of 
double monster Trout embryos, studied from serial sections and 
z ae A 5 ; ARIE 
illustrated by reconstruction drawings. For purposes of com- 
parison, a similar but shorter account of the anatomy of normal 


*% Communicated by F. G. Parsons, F.Z.S. 
+ For explanation of the Plates, see p. 21. 


12) AS. IOS) Wl, MG 


6. Bale &Danielsson, London . 


ANATOMY “OF MONSTROUS FISH EMBRYOS . 


1S VA, Sa, WNOKS, voll dl PE, 


AChor. 


Bale & Danielsson? London . 


ANATOMY OF MONSTROUS FISH EMBRYOS. 


Le 
a 


at 
= a 


ey ZS WSIONS), veil IPAL, Til 


— a} 
Bale & Danielsson, [t* London . 


ANATOMY OF MONSTROUS FISH EMBRYOS. 


TZ Se UQO)S), s7eil WL IPN INE 


We Sacc. 


Bale & Danielsson,L*¢ London . 


AN ATOMS Or MONSTROUS KISH EMBRYOS: 


rs : rie fad cm 


* é - 
le ' = ‘ 
mas Ui 
sa = 
t z 
aa as 
f 45 
, F F 
ah 
i rs 
. 
“se 


1903. ] MONSTROSITIES IN FISHES. 5 


Trout embryos at a corresponding age—three weeks after hatch- 
ing—is also given. 

The frequency of occurrence, the causation, the general appear- 
ances, and the classification of double monstrosities in fishes have 
been the subject of many papers and notices, lists of which are 
given by Prof. B. C. A. Windle in the ‘ Proceedings’ of the 
Zoological Society of London, 1895, p. 423, and by Dr. Franz 
Schmitt in the ‘Archiv fiir Entwicklungsmechanik der Organ- 
ismen,’ Bd. xiii. p. 34. The classification adopted here is on the 
same lines as that of Windle (/. c.), and has special reference to 
the material at my disposal. All my specimens were examples 
either (a) of union by the yolk-sac, or (6) of anterior duplicity. 
The former require only to be mentioned here, as each of the twin 
bodies contains a complete and separate complement of organs, 
while the latter may be conveniently divided into— 


Class I. Union in head-region : 
(a) the twin brains united at the optic lobes; 
(6) the twin brains united at the medulla oblongata. 


Class II. Union in pectoral region : 
(a) the adjacent pectoral fins not represented ; 
(6) the adjacent pectoral fins present, but united and 
reduced in size. 


Class III. Union at posterior part of body : 
(a) the two alimentary canals united a considerable 
distance in front of the vent ; 
(6) the two alimentary canals united close to the 
vent. 


ANAtoMY oF NorMAL Trout EMBRYOS. 


In normal Trout embryos of the same age as the monstrosities, 
ossification has not yet begun. ‘The parachordal cartilages are 
uniting round the anterior end of the notochord and have already 
joined with the trabeculee cranii, which, coalescing in front of the 
pituitary space, run forwards as a median flattened bar to meet 
the nasal cartilages. The pituitary space gives passage to the 
choroidal and internal carotid arteries and to the back part of the 
musculus rectus oculi externus (PI. I. fig. 1). On either side, the 
parachordals have grown upwards in the form of laminar plates, 
which meet in the mid-dorsal line over the upper part of the 
medulla, but leave a narrow V-shaped fontanelle over the lower 
part (Pl. I. fig. 2, £3). The auditory capsules are firm bosses of 
cartilage, moulded on the labyrinth, closed externally, but widely 
open towards the brain. Dorsally, they are connected with each 
other bya thin vault of cartilage roofing the cerebellum ; anteriorly, 
they are continuous with the supraorbital bars to be afterwards 
described; ventrally, they join the trabecular and parachordal 
cartilages; and externally they articulate with the hyomandi- 


6 DR. J. F. GEMMILL ON [May 12, 


bulars (Pl. I. fig. 1). The fifth nerves emerge through deep 
grooves between the trabecule and the auditory capsules, while 
the vagus and glosso-pharyngeal nerves pass out together through 
a foramen in the cartilage connecting the auditory capsules with 
the parachordals. A single opening in the floor of the capsules 
on either side gives passage to the internal jugular vein and the 
facial nerve. The nasal cartilage is connected with three pairs of 
bars: (1) the trabecule cranii, (2) the palatoquadrates, and (3) 
the supraorbitals. These last pass backwards along the dorso- 
lateral angles of the brain to join the anterior part of the 
auditory capsules on either side. Over the pineal body and the 
third ventricle the supraorbitals are connected together by a 
bridge of cartilage, but no such dzegmen exists over the cerebral or 
optic lobes, the spaces left uncovered being the anterior and the 
middle fontanelles (Pl. I. fig. 2, f1, £2). The hyomandibulars 
are connected with (1) the outer aspect of the auditory capsules, 
(2) the posterior ends of the palatoquadrates, and (3) the inter- 
hyals. The Meckelian bars are slender, and meet below the 
mouth in a symphysis. Posteriorly they articulate with the 
palatoquadrates, but not with the hyomandibular or interhyal 
cartilages. In the hyoid arches, glossohyals, hypohyals, cerato- 
hyals, and interhyals can be distinguished, The branchial 
cartilages are five in number, and have the usual forms and 
relations. The pectoral girdle is represented by a comparatively 
short bar of cartilage on either side—the coraco-scapular—and is 
far from being a complete arch ventrally. The limb-cartilage is 
an unsegmented plate continuous with the coraco-scapular bar. 
The notochord consists of pith-like tissue surrounded by a very 
firm capsule, and its anterior end is embedded in the fused para- 
chordal cartilages. In the position of each future vertebra there 
are four cartilaginous nodules, placed respectively at the dorso- 
lateral and ventro-lateral corners of the notochord, and prolonged 
into processes for the neural and hemal arches (PI. I. fig. 7). 

The anatomy of the central nervous system and of the organs 
of special sense, and of the heart and blood-vessels, is, with certain 
differences in the relative size of parts, practically the same as in 
the adult condition. As regards the aortic roots, it may be 
stated that the first root, 2. e. the first branchial vein, gives off 
the hyoid and carotid arteries and then passes backwards to join 
the second root. The resulting trunk bends inwards to the middle 
line, and, meeting with its fellow from the opposite side, forms 
the upper part of the aorta. This part is next joined on either 
side by a trunk formed by union of the third and fourth roots. 
The carotids pass forward beneath the parachordals and, traversing 
the pituitary space from below, reach the base of the brain. The 
hyoid artery arises from the first aortic root at its ventral end, 
perforates the hypohyal, runs up along the hyoid bar, and, after 
passing through a foramen in the hyomandibular, is continued 
mainly into the pseudobranch. The efferent vessel of the pseudo- 
branch passes forwards and inwards, traverses the pituitary space 


1903. ] MONSTROSITIES IN FISHES. 7 


from below, and, after running alongside the optic nerve, ends 
in the choroidal gland of the eye. Of the two cardinal veins, the 
left is usually the larger. 

The head-kidney, or pronephros, contains a single median 
glomerulus of considerable size, inside the cavity of which is a 
large vascular tuft supplied by a branch directly from the aorta. 
The Wolffian ducts, on either side, begin by a funnel-shaped 
opening in the glomerulus cavity. Then, bending forwards, they 
become convoluted, and are embedded in highly vascular lymphoid 
tissue. They then arch backwards, remaining convoluted for a 
short distance, and end in the urinary bladder (Pl. III. fig. 21 
& Pl. IV. fig. 28). The mesonephros is just beginning to develop 
in connection with their middle and posterior parts. The urinary 
bladder opens by a mesial pore situated just behind the vent. 

The intestinal canal is completely shut off from the yolk-mass, 
and there is an open diverticulum for the air-bladder. 


ANATOMY OF DouBLE MonstTRosITIES. 


I. Union in Head-region. 


The anatomy of specimens belonging to this group is very 
complex, as the region of transition from the double to the single 
condition involves the brain, the cranial nerves, and the organs of 
Special sense, as well as the cranial, mandibular, and branchial 
cartilages. The twin heads are placed symmetrically, side by 
side, and lie in the same horizontal plane. As regards the brain, 
union had occurred, in my specimens, either («) in the optic-lobe 
region, or (d) at the medulla oblongata. These two groups require 
to be described separately, as the details of their anatomy differ 
in many important respects. I shall begin with a typical example 
of (a). 

es are two notochords in front, and therefore, potentially at 
least, two pairs of parachordal cartilages, but the four cartilages 
have united to form a single basilar plate (PI. I. fig. 3, c). In 
front, the two nasal cartilages are placed widely apart; each 
contains a right and left olfactory pit, and is continuous behind 
with a pair of (now united) trabecule cranii. The two pairs of 
trabecule converge as they pass backwards; their inner or 
adjacent elements unite to form a median flattened bar, which 
joins the basilar plate formed by the parachordals. At the same 
time, the outer elements of each pair of trabecule have diverged 
from the inner elements to enclose a pair of pituitary spaces, the 
latter lymg one on each side of the median bar formed by union 
of the inner trabecular elements. There are only two auditory 
organs, and their cartilages are continuous with the outer 
trabecular and parachordal elements in the floor of the skull. 
Dorsally, the auditory capsules are connected over the cerebellum 
by a vault of cartilage, which is narrower antero-posteriorly than 
in the normal condition. Over the medulla the laminz of the 
parachordals nowhere form a complete vault. Accordingly the 


8 DR, J. F. GEMMILL ON [May 12, 


osterior fontanelle is much larger than in a normal case (Pl. I. 
fig. 4, 3). Each nasal cartilage is connected with a pair of 
palato-quadrate bars, the inner or adjacent elements of which 
converge, fuse, and end abruptly without being attached toa 
suspensorium ; the outer bars are continued backwards on either 
side, and articulate with the hyomandibulars attached to the 
auditory capsules. Articulating with the united part of the 
inner palato-quadrate bars is a small twisted piece of cartilage, 
which passes downwards in the septum between the two mouth- 
openings, and represents an inner or adjacent pair of Meckelian 
cartilages. Dorsally, each nasal capsule is continuous with a pair 
of supraorbital bars, of which the outer elements pass backwards 
on either side to join the auditory capsules, while the inner or 
adjacent elements are connected with each other and with the 
outer bars over the pineal body. Behind this the adjacent bars 
disappear, the result bemg that over the region of the cerebral 
lobes there are two small anterior fontanelles, while over the optic 
jobes there is a single large middle fontanelle (Pl. I. fig. 4). The 
hyoid bars and the branchial cartilages are normal, except that 
they diverge rather more widely than is usual. Hach arch, how- 
ever, may be looked upon as being composite, 7. e. as consisting of 
the outer elements of a double set of arches. The suspensoria on 
either side of the two periotic capsules are normal. 

As mentioned above, the notochord is double in front, two 
notochords being present as far back as the fourteenth body- 
segment. These converge together at an acute angle and finally 
unite. The condition of the neural and hemal arch cartilages is 
illustrated in Pl. I. fig. 8. Where the two notochords are some 
little distance apart, the inner or adjacent neural arches are dis- 
placed so as to form a floor for the transversely-expanded spinal 
cord, while the inner hemal arches remain as small nodules of 
cartilage. As the notochords come closer together the inner 
neural and hemal arches alike disappear, while the outer arches 
assume a normal form and position. 

The brain and its cavities show a degree of duplicity which is 
indicated diagrammatically in Pl. IT. fig. 15. There are two pairs 
of cerebral lobes and two thalamencephala, diverging forwards 
from a single composite optic lobe region. The cerebral lobes and 
thalamencephala, besides diverging, are rotated slightly in such a 
way that they lie closer together dorsally than ventrally. There 
are two pairs of olfactory nerves, two pairs of optic nerves, and 
two sets of pineal diverticula. There are also two infundibula, 
which converge as they pass downwards and backwards. Hach 
ends in a hypophysis sac after giving off the usual diverticula for 
the hypoaria (Pl. Il. fig. 17). The inner hypoarium on either 
side, owing to want of space, is smaller than normal, and lies 
above and in front of the outer hypoarium. The optic lobes show 
a remarkable transition between the double and the single con- 
dition. Their cavity and roof-parts are single, while the basal 
structures are doubled. There are thus two pairs of 3rd nerves. 


1903.] MONSTROSITIES IN FISHES, 9 


There is one pair of trigeminal nerves representing the right and 
left 5th nerves of the right and left twin heads. The succeeding 
cranial nerves are also normal, i. e. there is only a single pair of 
each. A rudiment of inner or adjacent pairs of trigeminal 
ganglia may be recognised in the form of a thin elongated 
band of tissue containing small nerve-cells and lying in the middle 
line underneath the region of the pons. This band of tissue has 
no central or peripheral nerve-fibres. The pons and cerebellum 
are single, but their internal structure shows traces of duplicity, 
especially in the case of the pons. The medulla oblongata 
is shghtly expanded transversely, but otherwise is normal. 

In the anterior part of the spinal cord there is a curious and 
interesting reappearance of duplicity, coextensive with the 
duplicity of the notochords, and with the presence, ventral to 
them, of a median composite muscular mass representing united 
adjacent lateral muscles. In this region, the spinal cord is greatly 
expanded in a transverse direction, its cavity is spindle-shaped, 
and, in addition to the usual nerve-roots, it gives off, on the 
ventral aspect in each segment, a pair of small additional motor 
roots which are distributed to the median muscular mass just 
mentioned. 

There are two pairs of olfactory organs and nerves. The outer 
eyes (right eye of right twin head and left of left head) are 
normal, but lie further back than usual, so that their optic nerves 
pass backwards as well as outwards from optic commissure to eye- 
ball. The inner or adjacent eyes may or may not be fused with 
one another. In the former case, there is usually a single lens, 
which is sometimes larger and sometimes smaller than in the 
normal case ; the sclerotic and choroid coats are single ; the retinz 
never unite, each showing its own choroidal fissure, optic nerve, 
and choroidal gland. In all cases the external rectus muscles are 
absent ; the superior obliques are absent or rudimentary, but the 
remaining ocular muscles are present in two sets. 

The heart and the ventral aorta are normal, but the dorsal 
aorta and its roots, and the choroidaland carotid arteries, require 
description. The union of the main collecting-trunks on either 
side to form the dorsal aorta is carried backwards for a very con- 
siderable distance, and takes place only at the level of union of 
the notochords. The pseudobranch on either side receives a 
branch from the hyoid artery, and its efferent vessel passes to the 
choroidal gland of the corresponding (7. ¢. outer) eye. The 
choroidal glands of the inner or adjacent eyes are supplied by 
blood which has not passed through the pseudobranch. In the 
specimen from which PI. I. figs. 3, 4, & 8 were taken, the arrange- 
ment of vessels is quite symmetrical and is indicated in PI. IT. 
fig. 12. A transverse arch vessel connects the upper aortic roots, 
and gives off a common choroidal artery which soon bifurcates. 
The two resulting vessels pass through the separate pituitary 
spaces, and are distributed to the choroidal glands of the adjacent 
eyes. 


10 DR. J. F. GEMMILL ON [ May 12, 


As is well known through the work mainly of Johannes Miiller 
(‘ Vergleichende Anatomie der Myxinoiden’), the pseudobranch 
and the choroidal gland are so related to one another in osseous 
fishes that a pseudobranch is never present in species which have 
no choroidal gland, while in rare instances only is a choroidal gland 
present where there is no pseudobranch. It is therefore of con- 
siderable interest to note that the choroidal glands of the inner 
or adjacent eyes, in the type of monstrosity under consideration, 
derive their blood-supply directly from the first aortic root. 
There is a certain amount of variation in the exact mode of origin 
of the choroidal and carotid arteries. For instance, cases occurred 
in which these vessels all arose from the first aortic root on one 
side only, instead of both roots participating equally as in the 
specimen figured (PI. IT. fig. 12). 

As regards the alimentary canal, there are two mouths, but the 
pharynx and the rest of the canal are single, the only evidence of 
duplicity being the presence of two air-bladder diverticula. 

The mesonephros, ureters, bladder, and urinary pore are 
normal, but the pronephric glomerulus is composite, or may be 
double. An example of the composite condition is figured in ~ 
PI. IV. fig. 29. The glomerulus is larger than in the normal 
case, and contains two vascular tufts between which is a median 
compartment that obviously corresponds to the fused adjacent 
halves of a pair of glomeruli, but has no Wolffian ducts in con- 
nection with it. In the specimen from which figure 30 (Pl. IV.) 
was taken the two glomeruli are separate, but one of them has 
only a rudimentary Wolffian duct arising from its inner side, 
while the other has none at all. Fig. 29 (Pl. IV.) should be 
compared with figs. 31 and 32 on the same Plate, which show a still 
greater degree of duplicity in the pronephros. 

Of composite muscles, the most important are contained in the 
median mass mentioned on page 9 as underlying the twin noto- 
chords, and innervated by the small extra motor roots of the 
composite spinal cord. This muscular mass is segmented serially 
by septa which correspond exactly with the septa of the outer 
(normal) lateral muscles. In the head-region some small and 
intricately arranged muscles are found connected with the 
cartilages which represent the reduced adjacent Meckelian and 
palato-quadrate bars. These muscles are obviously rudiments 
of adjacent mandibular and temporal muscles. It has been noted 
previously that there are no external rectus muscles in connection 
with the adjacent eyes, and that the superior obliques are either 
rudimentary or entirely absent. 


A typical specimen of the second subgroup of Class I. will next 
be described, 2. ¢. a specimen which shows the twin brains uniting 
at the medulla oblongata. 

The structure of the cranial skeleton in this type agrees with 
that of the monstrosity last described, except that the place of 
union of the skeletal elements is carried further back. This gives 


1903.] MONSTROSITIES IN FISHES. ll 


room for greater development on the part of the inner or adjacent 
elements in the twin heads (Pl. I. fig. 5). The two converging 
pairs of trabecule are separate along their whole length, and each 
unites posteriorly with corresponding parachordals. The two 
pairs of parachordals are separate in front, but posteriorly the 
adjacent elements in each pair unite, so that a single composite 
basilar plate of cartilage containing two notochords underlies the 
lower half of the medulla oblongata. The inner or adjacent 
palato-quadrates converge posteriorly and coalesce. The united 
part articulates (1) below, with a small bifid cartilage representing 
fused adjacent Meckelian bars; and (2) higher up, with a small 
cartilage representing fused adjacent hyomandibulars. The inner 
or adjacent supraorbital bars converge posteriorly, unite with one 
another, and end by becoming continuous with the roof of a small 
box of cartilage which is wedged into the apex of the angle 
between the twin heads and represents fused adjacent periotic 
capsules. This structure will be described later on, but 1t may 
be mentioned here that its roof is continuous posteriorly with a 
vault of cartilage which connects the two outer (normal) periotic 
capsules over the cerebella. In this way double sets of anterior 
and middle fontanelles are left over the cerebral lobes and mid- 
brains respectively of the twin heads (PI. I. fig. 6). 

The inner or adjacent hyomandibulars are extremely rudi- 
mentary, and are fused together to form asmall bifid piece which 
articulates (1) anteriorly, with the fused adjacent palato-quadrates, 
(2) posteriorly, with the fused adjacent periotic capsules, and (3) 
inferiorly, with a rudiment of the fused adjacent hyoid bars. The 
small artery for the supply of the inner or adjacent pseudobranchs 
passes up through the notch at the anterior end of this cartilage. 
The inner or adjacent auditory capsules are extremely rudi- 
mentary, being completely united, compressed from side to side, 
and wedged into the position above indicated. They contain a 
single distorted labyrinth, and are entered by small auditory 
nerves on either side, which are distributed symmetrically over 
the labyrinth. This composite auditory capsule is connected 
anteriorly with the fused adjacent supraorbital bars, and ventrally 
with the adjacent trabecular and parachordal cartilages on the 
inner sides of the two pituitary spaces. Behind it a small tri- 
angular opening is left, bounded on either side by the converging 
parachordals. These unite posteriorly, but leave a narrow 
foramen between them for the exit of a small nerve, which 
represents a reduced adjacent pair of vagus and glosso-pharyngeal 
nerves (PI. I. fig. 5). It will be seen from what precedes that 
there are five fontanelles, one over each pair of cerebral lobes, one 
over each mid-brain, and one over the composite medulla 
oblongata (PI. I. fig. 6). 

There are two mouth-openings placed side by side, one under 
each head, and separated from one another by a thick dorso- 
ventral septum. This septum contains (1) remains of the adjacent 
mandibular and hyoid cartilages, (2) much confused muscular 


12 DR. J. F, GEMMILL ON [May 12, 


tissue, and (3) two arteries which will be afterwards described, 
one being a continuation of the ventral aorta, and the other a 
small artery for the supply of the inner or adjacent pseudobranchs. 
The two mouth-openings lead into separate buccal cavities, but 
the cesophagus is single, the septum above mentioned ending 
opposite the second branchial cartilage. 

The mandibular apparatus may be described as consisting of a 
composite arcade underlying the two mouth-openings. The outer 
portions of this arcade are formed by normal (outer) Meckelian 
bars (i.e. right bar of right twin head and left bar of left head), 
while a small mesial portion of the arch is formed by rudimentary 
adjacent Meckelian bars (Pl. II. fig. 10, Meck’). These latter 
bars are united at their proximal ends, and there articulate with 
the fused adjacent palato-quadrates. Distally, each of the inner 
bars unites in asymphysis with its corresponding outer Meckelian 
bar. As the inner bars are exceedingly reduced in size, the two 
symphyses lie close together in the tissue of the septum sepa- 
rating the two mouth-openings. The hyoid apparatus may also 
be described as forming a composite arcade, the main part of 
which consists of the outer arches of the twin heads, while in the 
middle are interposed the fused remains of the inner arches 
(fig. 10). These remains consist of (1) a single twisted piece of 
cartilage articulating, without the intervention of an interhyal, 
with the fused adjacent hyomandibulars, and representing cerato- 
hyals ; and (2) two incompletely separated hypohyals, articulating 
below with two glossohyals which are also incompletely separated. 
Connected with the outer sides of these glossohyals are the hypo- 
hyal pieces of the outer arches (PI. II. fig. 10, BH). No adjacent 
elements are interposed in the series of branchial cartilages. 
The only evidence indicating duplicity is to be found in the second 
copular piece, z.¢. that succeeding the glossohyals. This piece is 
double anteriorly, but it becomes single opposite the articulation 
with it of the second branchial cartilage. The succeeding copular 
pieces are single, but they are a little broader than normal, 
especially in front. 

The notochords remain separate as far back as the twentieth 
somite. For the arrangement of the neural and hemal arch 
cartilages in the transition region, see above, page 8. 

There are two sets of brain cavities and masses as far back as 
the level of the fourth ventricle. The fourth ventricle is single 
posteriorly, but it bifurcates in front into two canals leading into 
the separate mid-brains. The posterior part of the medulla and 
the anterior part of the spinal cord are composite, and show the 
following characters :—(1) they are much drawn out transversely, 
and (2) they give origin to small adjacent nerve-roots. These 
roots in the _medulla are extremely ‘rudimentary, and their 
ultimate distribution could not be traced, but in the spinal cord 
the nerve-roots in question are better developed and form a 
regular series of pairs of nerves coming off from the ventral 
aspect of the cord and distributed to the somites of the median 


1903. ] MONSTROSITIES IN FISHES. 13 


muscular mass which lies ventral to the notochords (compare 
page 9). In this monstrosity, as in the one previously described, 
the anterior part of the spinal cord, though it lies nearer to the 
place of union of twin bodies, shows greater structural duplicity 
than does the medulla oblongata. 

All the outer cranial nerves belonging to the twin heads are 
normal and need no further mention. Of the inner or adjacent 
nerves, the Ist, 2nd, 3rd, and 4th are normal; the 5th and 
its ganglia are reduced in size; the 7th and 8th with their 
ganglia are very rudimentary; while only a remnant of the 
adjacent glosso-pharyngeal and vagus remains. I was unable to 
follow out the 6th pair, but the presence of well-developed 
external rectus muscles makes it probable that these nerves were 
present. As has already been mentioned, the inner or adjacent 
auditory capsules are much reduced in size, their auditory sacs 
being completely united and forming a single labyrinth, sym- 
metrical in shape, compressed from side to side, and receiving the 
two adjacent auditory nerves. The arrangement of the sensory 
epithelium inside the various parts of this labyrinth is also 
bilaterally symmetrical. A reconstruction drawing of this 
labyrinth is given in Pl. IV. fig. 27. Saccule, utricle, and 
anterior and posterior semicircular canals are all represented, but 
there is no trace of a horizontal semicircular canal. 

The heart is normal and gives origin to a single ventral aorta, 
which for a short distance upwards has a double cavity, owing to 
the presence of a median antero-posterior septum, which, how- 
ever, disappears further forwards. The gill-arteries on either 
side are normal, but in addition to them the ventral aorta gives 
rise to several small irregular branches which ramify in spongy 
tissue surrounding the ventral ends of the branchial cartilages, 
and which may be taken to represent a very rudimentary set of 
adjacent gill-arteries. But the most striking feature of the 
ventral aorta is that, instead of ending in the first gill-arteries, it 
is continued forwards and arches dorsally in the tissue of the 
septum between the two mouth-openings. Passing through the 
space between the adjacent glossohyals and the succeeding copular 
piece, it comes to lie behind the small cartilage which represents 
adjacent ceratohyals. Then, reaching the base of the skull, it 
bends backwards, and divides into two equal branches which join 
the upper aortic roots on either side (PI. IT. fig. 13). In the first 
part of its course, this continuation of the ventral aorta gives off 
(1) two inner or adjacent carotid arteries, which, after running 
forwards and outwards, pass through their corresponding pituitary 
spaces ; and (2) two arteries for the supply of the inner or adjacent 
pseudobranchs. These arteries run at first forwards and dorsal- 
wards behind the adjacent glossohyals; then curving forwards 
they unite in front of the adjacent ceratohyals ; then, separating 
again, they pass between the adjacent palato-quadrates and the 
hyomandibulars, and are distributed to their corresponding (7. e. 
inner) pseudobranchs. On either side the first aortic root gives 


14 DR. J. F, GEMMILL ON [May 12, 


off (1) a hyoid artery which sends a branch to the corresponding 
outer pseudobranch, and (2) a carotid artery. It then joins the 
second aortic root, and shortly afterwards meets the continuation 
of the ventral aorta previously mentioned. The resulting vessel 
is then joined by the third and fourth aortic roots, but it remains 
separate from its fellow so long as the notochords are separate, 
i.e. back to the twentieth somite. 

The arrangement of vessels which has just been described and 
which is illustrated in Pl. II. fig. 13 is somewhat remarkable. 
Mixing of arterial with venous blood must have taken place in no 
slight degree, and certain parts in each twin head must have been 
supplied by blood coming directly from the ventral aorta. There 
is no trace of a pair of adjacent jugular veins, and it is difficult 
to make out from sections the course of the venous blood coming 
from the adjacent sides of the twin heads. But as there is a con- 
siderable amount of spongy tissue below the base of the skull and 
in the septum between the mouth-openings, it is probable that 
the blood in question found its way into the median and the main 
jugulars. The presence of this spongy tissue no doubt indicates 
congestion. 

The pronephric glomerulus (Pl. IV. fig. 32) is composite. It is 
remarkably large, and is divided into three compartments by two 
vascular tufts each of which has an afferent and an efferent 
vessel. Normal Wolffian ducts arise from the outer compart- 
ments, while the middle compartment gives origin to a sacculated 
tubule which passes backwards a short distance to end blindly, 
and which represents fused adjacent Wolffian ducts. 

There are two buccal cavities, but the esophagus and the rest 
of the alimentary tract are single except for the presence of two 
air-bladder diverticula. 

The composite muscles described on page 9 are well developed 
in this monstrosity. 


IT. Union in Pectoral Region. 


Class IT. of the monstrosities has next to be described. In it 
the twin bodies are united in the region of the pectoral fins. All 
specimens of this Class have separate gullets, air-bladders, and 
stomachs, but a single liver, intestinal canal, and vent. The 
hearts are more or less closely united. Two subgroups of this 
Class are naturally distinguished from one another—(a@) a group 
in which union of the twin bodies takes place so far forward that 
adjacent pectoral fins are not present; and (b) a group in which 
adjacent pectoral fins are present, but are united and reduced in 
size, 

(a) Adjacent Pectoral Fins not represented.—The brain, the 
cranial and branchial skeletons are double. So also are the 
alimentary canals down to the level of the stomach. The chief 
interest in this type centres round the heart and blood-vessels. 
A reconstruction drawing of the heart and origins of the vessels of 


1903. ] MONSTROSITIES IN FISHES. 15 


a typical specimen of this group is given in Pl. IIT. fig. 19. (The 
hepatic veins are not shown because, in the removal of the yolk- 
sac, the liver had been damaged, obscuring the relations of its 
vessels.) The whole of this double heart lies inside a large com- 
posite pericardial cavity, which is prolonged a little forwards on 
either side round the origins of the ventral aorta. The ventricles 
are separate, the auricles communicate with one another, and 
there is a single large sinus venosus opening by a wide ostium 
into the auricles at their junction (PI. III. fig. 19, 7.4u). The 
sinus venosus receives blood (a) on either side from the duct of 
Cuvier formed by union of the outer cardinal and internal 
jugular veins of the twin embryos (7. e. from the right duct of 
Cuvier of the right embryo and from the left duct of Cuvier of 
the left embryo) ; (6) from two separate middle jugular veins; and 
(c) from a large trunk formed by union of the inner or adjacent 
internal jugular veins of the twin heads (PI. III. fig. 19, De’). 
This last trunk obviously corresponds to fused adjacent ducts of 
Cuvier which receive internal jugular veins only and have no 
corresponding cardinals. 

The head-kidney in this specimen is illustrated by Pl. IV. 
fig. 31, and corresponds to the description given on page 14. 

The notochords and spinal cords are still widely separate 
opposite the pectoral region, and they remain separate for a con- 
siderable number of somites behind it, but ultimately they fuse, 
so that the posterior part of the body contains a single notochord 
and a single spinal cord. 

The behaviour of the neural and hemal arches and of the 
median muscular mass corresponds to the description on page 8. 
These structures, however, may be studied to greater advantage 
in the type at present under consideration, as the whole transi- 
tional region is open for observation. The dorsal ends of the 
adjacent ‘fifth branchial cartilages are fused, but otherwise the 
twin branchial skeletons are quite separate. The ventral ends of 
the two coraco-scapular bars fail by a wide interval to meet each 
other below the pericardium. 


In a typical specimen belonging to group (0) of Class IT., in 
which adjacent pectoral fins were present but united and reduced 
in size, fusion was more complete towards the posterior (radial) 
border of the cartilage than towards the anterior border. Thus, 
near the anterior border, there was only a small bridge of cartilage 
between them; further back they approached one another, and the 
bridge was wider ; while at the posterior border they were united 
along their whole length. As regards the adjacent coraco-scapular 
bars, they were quite separate, except at their ventral ends, which 
were fused together and projected downwards into a septum 
between the two pericardial sacs. The ventral ends of the two 
outer coraco-scapular bars were very widely distant from one 
another. 

The same specimen may be used as a type for illustrating the 


16 DR. J. F. GEMMILL ON [May 12, 


heart and vessels in this group (PI. IV. fig. 20). There are two 
pericardial cavities separated by a septum of connective tissue, 
which is thin posteriorly, but in front is thick and contains the 
fused ventral ends of the adjacent coraco-scapular bars Just men- 
tioned. Auricles and ventricles are completely separate, and the 
sinus venosi communicate only by a narrow neck. Each sinus 
venosus has a pair of ducts of Cuvier, the inner or adjacent 
ducts being smaller than the outer. This difference depends 
mainly on the fact that the inner or adjacent ducts are small and 
short. They can be traced backwards inside the substance of the 
head-kidney, but are soon found to unite and to break up into 
venules in the lymphoid tissue (Pl. III. fig. 20, Vc’). 

The glomerulus of the head-kidney is shown in Pl. IV. fig. 31. 
It is greatly elongated in a transverse direction, and the tubule 
from its middle compartment, representing fused adjacent Wolffian 
ducts, passes forwards so as to lie between the two adjacent 
cardinal veins which have just been referred to. It ends blindly 
and is so much sacculated as to suggest a certain degree of 
pressure in the fluid secreted by the glomerulus. A similar 
point will be noted later (page 17), where one of the urinary 
bladders in a-double monstrosity had no urinary pore. 


III. Union at Posterior Part of Body. 


In Class I., the twin heads lie symmetrically, side by side, 
exhibiting lateral union, but in Class IT. there is a slight conver- 
gence ventrad of the sagittal planes of the twin bodies ; while in 
Class ITI. this convergence exists in a very marked degree, giving 
rise to ventro-lateral union of the twin bodies, or, even, in 
extreme cases, to what may be described as ventral union. 
Roughly speaking, the further back union of the twin bodies 
takes place, the greater is their ventral convergence. This is in 
harmony with the fact that the twin bodies are lying tangentially 
on the surface of a single small yolk-sphere. 

It will be convenient to subdivide Class III. into (a) cases in 
which union takes place well in front of the vent, and (b) cases in 
which union takes place quite close to the vent. 

In group (4) the alimentary canal is single for a considerable 
distance posteriorly, the united portion being provided with two 
dorsal mesenteries, one from each twin body; the head-kidneys 
are quite separate and are normal, but their inner or adjacent 
Wolffian ducts end blindly in the mesonephric region, while the 
outer Wolffian ducts pass backwards as the ureters of the single 
normal bladder (Pl. IIT. fig. 22). In this group, moreover, the 
ventral convergence of the twin bodies is not too great but that 
it can readjust itself at the region of transition, and allow the 
spinal cords, as in Classes I. and IJ., to unite anterior to the 
place of fusion of the notochords. 

Group (6) of Class IIT. includes cases in which union takes 


1903. ] MONSTROSITIES IN FISHES. 17 


place close to the anus. Ventral convergence of the sagittal planes 
of the twin bodies is always marked, and is often extreme enough 
to allow the notochords to unite anteriorly to the place of union 
of the spinal cords and of the dorsal edge membranes. The 
notochords are always double opposite the vent and for a consider- 
able distance behind it, but I have not met with any cases in 
which they remained separate to their extreme tips. The dorsal 
aorte unite at the same level as the notochords, and the caudal 
veins a little in advance of the aorte. On the other hand, the 
spinal cords, while they usually unite far back in the caudal 
region, in extreme cases may remain separate altogether. 

The ventral edge membranes tend to unite earlier than those 
of the dorsal edge, but in cases of pure ventral union there may 
be two composite ventral edge membranes at opposite corners of 
the quadrangular composite body, the other corners of which 
carry normal dorsal edge membranes. Apparently, during the 
concrescence of the twin bodies, the blastema for the ventral edge 
of each was kept apart in two halves, each of which, continuing 
to oceupy a lateral position, met and fused with a corresponding 
part derived from the other twin. 

Tn all of my specimens except one (PI. IIT. fig. 26) the vent is 
single, and in all of them the intestines are united close to the 
vent. Franz Schmitt, however (é.c. p. 53) refers to a case in 
which, apparently, no such union had taken place. 

Very great variation is found in the arrangement of the ureters, 
bladders, and urinary pores in this group. All my specimens have 
two bladders, which sometimes communicate with one another 
and sometimes are quite separate. In all cases, the right ureter 
of one twin and the left ureter of the other open into one of the 
bladders, while the two remaining ureters go to the second 
bladder. Thus, each bladder receives a right and a left ureter 
derived from different embryos, and, except in cases of symmetrical 
ventral union, the ureters which go to the one bladder may be 
recognised as the inner or adjacent pair, while those which go to 
the other may be recognised as the outer pair. In such cases, 
the first bladder les anterior and ventral to the second, with 
which also it frequently communicates, especially when destitute 
of an external opening. 

Attention may be drawn to figs. 23, 24, 25, 26, on Plate IIT., 
which are reconstruction diagrams illustrating the principal 
variations referred to above. Figure 23 was drawn from a 
specimen in which the bladder (4Z’) in connection with the inner 
or adjacent pair of Wolffian ducts has no urinary pore, but opens 
into the bladder (BZ) in connection with the outer pair of 
Wolffian ducts (WDa, WDb). Figure 24 illustrates a case in 
which the first bladder (BZ') had no opening and was enor- 
mously expanded, as were also the lower ends of its ureters. 
Figure 25 is from a specimen in which the bladders (BZ! and 
BL) were separate and had urinary pores which opened in the 
mid-ventral line, one behind the other. It will be seen that 


Proc. Zoou. Soc.—1903, Vou. II. No. II. 2 


18 DR. J. F, GEMMILL ON [May 12, 


in this case the two bladders lie in the same plane and have 
corresponding right and left sides. But the right side of BL’ 
+s in connection with a left Wolffian duct (Wda'), while the 
left side is in connection with a right Wolffian duct. Such a 
transposition is exceedingly rare in double monstrosities. Figure 26 
was taken from a case of symmetrical ventral union. Two 
vents and two urinary pores were present, and as they opened 
laterally in pairs towards opposite sides, they lay in a plane at 
right angles to the sagittal plane of the twin bodies. This 
arrangement has many parallels in teratology, e.g. mn cases 
of ischiopagous double monstrosity*. It preserves the natural 
correspondence between rights and lefts in the ducts and bladders, 
which, as has just been seen, is inverted in the case from which 
figure 25 of Plate III. was taken. 


GENERAL, 


With the rarest exceptions, all double monstrosities in fishes 
are examples either of anterior duplicity or of union by the yolk- 
sac. The explanation of this remarkable fact seems to me to be 
as follows :—In all these cases, two centres of gastrulation form on 
the edge of the blastoderm at a greater or less distance from one 
another. The spreading of the blastoderm over the yolk-mass 
goes on freely all round except at and near the primitive streak. 
There, changes take place which lead to increase in length of the 
embryonic axis, and which are interpreted by many as con- 
erescence. If the two centres of gastrulation happen to be near 
one another, the whole of the blastoderm edge separating them 
eventually will be used up in the process of concrescence; the 
later formed parts of the embryonic axes will be drawn closer and 
closer to one another, until in their turn the axes themselves 
coalesce. The degree of union will be in inverse proportion to 
the original distance from one another of the two centres of 
gastrulation. Should the two centres of gastrulation be so far 
apart that the middle portion of the intervening blastoderm edge 
is not involved in concrescence but is left free to extend over the 
yolk-inass, the two embryonic axes will be independent along 
their whole length, and the only structures which connect them 
will be the blastoderm and the yolk-sac. According to this view, 
double monsters showing anterior duplicity are the result of what 
may be called primary fusion, that is, conerescence of their 
growing embryonic axes. 

In birds, typical concrescence can occur only during the earliest 
stage of formation of the primitive streak, 7.e. so long as the 
groove of the sickle and knob is open. Any subsequent concres- 


cence can take place only by the incidental drawing in and 


utilisation of lateral blastema at the growing zone. This process 


* J. F. Gemmill in ‘ Journal of Anatomy and Physiology,’ vol. xxxvi. p. 263. 


1903. ] MONSTROSITIES IN FISHES. 19 


cannot exercise the same compelling influence in approximating 
the growing embryonic axes as it may do in fishes. The question 
of the origin of the different kinds of double monstrosities in 
birds and mammals is complicated by other factors and cannot be 
discussed fully here ; but the considerations suggested above may 
throw some light on the fact that practically all double monstros- 
ities among fish with united bodies show anterior duplicity, 
whist in mammals and birds there are as many or more cases of 
posterior duplicity. 

As was indicated previously (p. 16), both subgroups of Class I. 
exhibit simple lateral union. It may be inter esting, in these 
cases, to compare the behaviour, as regards union, of various 
mesial and lateral organs. Of the ‘three primitive axial structures, 
the notochords are the last to unite, and the alimentary canals the 
first, while the neural axes are intermediate (pp. 7, 8,10). It 
may be taken as a general rule, in monstrosities of this type, that 
structures and parts of structures which lie nearest the noto- 
chords retain evidence of duplicity longest. Thus, the optic lobes 
mentioned on p. 8 have single roof parts, while their basal 
structures are double; the composite spinal cord (pp. 9, 12) has 
additional nerve-roots coming off from its ventral aspect : there 
are two air-bladder diverticula in a case where the alimentary 
canal was single up to the mouth (p. 10), while there is only a 
single liver in a case where the alimentary canal was double 
down to the pylorus (p. 14); duplicity of the dorsal aorta is 
coextensive with duplicity of the notochord, while the heart and 
pericardium are single (p. 9); the cartilages of the neural and 
heemal arches are in double sets for many somites in a case 
where all the branchial cartilages are single (p. 8). 

The slowness with which the notochords unite may be due in 
part to their small size and to the nature of their tissue, but is 
probably to be referred mainly to their central position and to 
the fact that they are flanked by the bulky mesoblastic somites, 
so that primary fusion is deferred as long as possible and secondary 
fusion is prevented. The early union of the neural axes and of 
the alimentary canals, and the earlier union of their dorsal and 
ventral walls respectively, may be explained in part by primary 
frst ion, 1f one remembers that the dorsal wall of the neural axis 
is formed from the outer edges of the neural groove, and that the 
ventral wall of the alimentar: ‘y canal is for a long time incomplete. 
But such facts as the very marked increase of duplicity in the 
spinal cord as compared with the medulla oblongata in Class I. 
(a) and (4) (pp. 9, 13), indicate that, in addition “to the primary 
fusion of conerescence, secondary fusion has played some part in 
moulding organs at the transitional region. The greater 
simplicity of the medulla is explained, in part by its greater size 
as compared with the spinal cord, but chiefly by the fact that 
the notochords are closer together at theix anterior ends (where 
they are surrounded by the - parachordal cartilages) than el are 
in the cervical region where the median muscular mass serves to 


o* 


20 DR, J. F. GEMMILL ON [May 12, 


press them widely apart. The growth in bulk of this mass which 
causes this pressure can only have occurred at a relatively late 
stage. Almost perfect coalescence of the medulle has thus been 
allowed through secondary fusion, while the upper parts of the 
spinal cords have been kept markedly composite. Further, there 
is reason to believe that nearly all the composite mesenchymal 
structures occurring at the transitional region have been produced 
through secondary fusion. 

The monstrosities grouped under Class III. (p. 16) do not 
exhibit symmetrical lateral union, owing to their marked ventral 
convergence, and, accordingly, what has been said above regarding 
the behaviour of different organs at the transitional region does 
not apply to this Class. 

Tn all cases except those of pure ventral union (for an example 
of which see p. 18), the change from the double to the single 
condition is effected at the expense of the imner or adjacent 
elements, while the outer elements become the right and left sides 
of the single region of the body. This rule has long been recog- 
nised as holding good in similar double monstrosities among the 
higher vertebrates. For the sake of easy reference, it may be 
useful to tabulate here the principal composite structure, produced 
by fusion of adjacent elements, and mentioned in the descriptive 
part of this paper :— 


Trabeculee cranii, page/, Plate L., fig. 3, d. 
Parachordal cartilages, ,, 7,11, ,, IE BI Dy G. 
Vertebral do., TeEes ss fig. 8, Va’, Ha’. 
Palato-quadrate do., He tory delle ra EES, 3), Dai, 
Supraorbital do., Aatcowel Wien gee figs. 4, 6, s’. 
Periotie do., ie lle Me mer, 19, 
Hyomandibular do., panel ly . Mee, 8), 
Meckelian do., Au teal yt one ee fig. 3, g', 9,10, Meck’. 
Hyoid do., plas) Plates 1s toe Oeics rs ele 
Pectoral fin do., Pealisy i ier II, (GUL 
Jugular veins, by Los welate Lille iinc 9 Se) Ole 


Cardinal veins, oy Lb, fe: fig. 20, VC’. 

Pseudobranch arteries, ,, 9, Plate IL., fig. 12, A.Chor’, 
fig. 13, Aq’. 

Pronephric glomerulus, ,, 10, 16, Plate IV., figs. 29, 31, 32. 


Wolffian ducts, 45 LG, =. figs. 31, 32. 
Muscles, nos 
Hyes, 


Internal ear, i 
Optic lobes, ey || labia INI es, 1b, TZ 
Medulla oblongata, eal Hs fig. 16. 
Spinal cord,  @ 
1 
Pe] 


Cranial nerves, 


b) 


€ 


, Plate lV., fig. 27. 


As a rule, where adjacent muscles persist, there are also 
corresponding motor nerves. But the small muscles described as 


1903.] MONSTROSITIES IN FISHES. 21 


surrounding the fused adjacent Meckelian and hyoid bars in 
Class I. (p. 10) are destitute of nerves. 

Development of parts may take place without the presence of 
nerves of general sensation. Thus in Class I. (a) no trace exists of 
adjacent trigeminal nerve-fibres, yet the inner sides of the snout 
and head are well developed wherever they have sufficient 
space. 

On the whole, the anatomy of the series of monstrosities which 
has been described in this paper is remarkably symmetrical, both 
twins contributing equally to the sum of structures in the 
transitional region, in which, moreover, the law that union takes 
place between homologous structures alway holds good. 


EXPLANATION OF THE PLATES. 


(Reconstruction Diagrams.) 


Puate I. 


Fig. 1. Cranial and hyomandibular skeleton of a normal Trout embryo, seen from 
above, after removal of the roof-cartilages by a section supposed to pass 
horizontally through the nasal and periotic cartilages (pp. 5, 6). 

a, olfactory pit and cartilage. 

trabeculz cranii. 

ec, parachordal cartilages. 

d, placed just in front of pituitary space. 

é, 33 behind pituitary space. 

f, palato-quadrate bars. 

g, Meckel’s cartilage. 

h, hyoid bar. 

i, hyomandibular. 

k, placed in floor ot periotic capsule. 

m, pituitary space with external recti muscles passing through it. 

n, notochord embedded in the parachordals. 

Fig. 2. Roof-cartilages of skull of a normal Trout embryo, seen from above (p. 6). 
(For lettering see under fig. 6.) 

Fig. 3. Cranial and hyomandibular skeleton of a double-monster embryo, seen from 
above as in fig. 1. The duplicity affects the anterior cranial structures only 
(pp. 7, 8). Lettering as above; see also under fig. 5. 

Fig. 4. Roof-cartilages of skull of the double monster illustrated in fig. 3. For 

lettering see under fig. 6. 
5. Cranial and hyomandibular skeleton of a double monster exhibiting a 
slightly greater degree of duplicity than that of fig. 3 (pp. 11, 12). 
In figs. 3 & 5, a, b, c, &c. represent the same structures as in fig. 1; 
',g', 7, k’ ave the inner or adjacent (reduced) structures corresponding to 
the fully-developed outer ff g, i, & (i.e. palato-quadrate, Meckelian, hyo- 
mandibular, and periotic cartilages). 
Fig. 6. Roof-cartilages of skull of the double monster illustrated in fig. 5. 
Fl, f2, £3. Anterior, middle and posterior fontanelles. 
7, cartilage of roof of nasal pits. 
s, supraorbital bars (s’, adjacent reduced pair). 
, cartilage of roof of periotic capsule. 
w, parachordal cartilage. 
x, y, tegminal cartilages above third ventricle and cerebellum respectively. 

Vertebral cartilages m normal Trout embryo (p. 6). 

5 5 any double monstrosity near the transitional region. 

Spe. Spinal cord. 

Neh. Notochord. 

Na. Neural arch cartilage (Na’, adjacent reduced neural arch 
cartilage). 

Ha. Hemal arch cartilage (Ha’, adjacent reduced hema) arch 
cartilage). 


ie?) 
mis 
gg 
o 


iS 


WIS 


LD) by) 


2 DR. J. F. GEMMILL ON [May 12, 


Prate II. 


Fig. 9. Ventral ends of the Meckelian, hyoid, and branchial cartilages of the 
monstrosity illustrated in figs. 3 and 4 (see pp. 7, 8). 
Fig. 10. Ventral ends of the Meckelian, hyoid, and branchial cartilages of the 
monstrosity illustrated in figs. 5 and 6 (pp. 11, 12). 
Meck. Outer Meckelian bars. 
Meck Inner or adjacent Meckelian bars united and greatly reduced. 
GH. Glossohyal. 
BH. Hypohyal. 
DH. Ceratohyal. 
BH’. Inner or adjacent hypohyals partly united. 
LH’. 5 53 ceratohyals united. 

TIL, 1V, V, VI, VII. The branchial cartilages. 

Cop. Copular pieces, the anterior one being bifid. ahs ; 
Fig. 11. Transverse section of a composite pectoral fin, from a monstrosity in which 
union took place just behind the pectoral tins (see p. 19). 

Scap., Cor. Scapular and coracoid parts respectively of the coraco- 
scapular bars. The ventral ends of the coracoid parts 
are united. 

CL. The limb-cartilages united for the greater part of their 
length. 
R. The rays (not cartilaginous). 
Fig. 12. Diagram of the aorta and its roots in the monstrosity referred to on p. 9. 
Ao’. Aorta, the two limbs uniting further back. 
I, U, UL, 1V. Aortic roots, 7. e. branchial veins. 
Car. Carotids. 
Aa. Artery to pseudobranch, a branch of the 1st branchial. 
Psbr. Pseudobranch. 

A.Chor. Choroidal artery. 

A.Chor.’ The arteries for the choroidal glands of the inner or adjacent 
pair of eyes. They arise by a single stem from the 
middle of a vessel connecting the aortic roots on either 
side, and go to the choroidal glands without passing 
through a pseudobranch. 

Fig. 18. Diagram of ventral aorta, aorta and aortic roots in the monstrosity 
illustrated in figs. 5, 6, 16 (see p. 18). 
Lettering as above, with in addition : 
TA. Ventral aorta and its branches 1, 2, 3, 4. 
Car.’ Inner or adjacent carotids, giving off Aq’ arteries to the inner 
or adjacent pseudobranchs Psbr.’ 

It will be seen there are two sets of carotid and afferent pseudobranch 
arteries, the inner sets being derived directly from the ventral aorta. The 
ventral aorta arches dorsally in the septum between the two mouths of 
the monstrosity, reaches the base of the skull, and then divides into two 
limbs which are continued backwards to join the aortic-collecting roots on 
either side. 

Fig. 14. Beline ff cee cavity of brain of normal Trout embryo. Lettering 

as in fig. 16. 

Fig. 15. Outline of central cavity of brain of the monstrosity illustrated in figs. 3, 4, 
: & 9 (p. 8). Lettering as in fig. 16. a 
Fig. 16. Outline of central cavity of brain of the monstrosity illustrated in 

figs. 5, 6, 10 (p. 12). 

C.F.B. Cavity of the hemispheres. 


C3rd V. ne 8rd ventricle. 
COL. 3 optic lobes. 
CIt. 3 iter a tertio ad quartum ventriculum. 
C4th V. i 4th ventricle. 
Spe SS spinal cord. 


Fig.17. Outline of transverse section of central cavity in anterior part of optic lobe 
ee of the ee ale ig. 15. There are two infundibula and two 
sets of hypoarial cavities, while the main optic lobe cavity is sine . 8). 
COL. Cavity of optic lobes. ; eo ee 
Inf. Infundibulum. 
Hyp. Hypophysis sac. 
ya. One of the hypoarial cavities. 


1903. | MONSTROSITIES IN FISHES, 23: 


Fig. 


Fig. 


Fig. 


Prate III. 


18. Diagram of heart §c. of normal Trout embryo. Lettering as in fig. 20. 
19. Diagram of heart §c. of the monstrosity illustrated in figs. 5, 6, 10, and 16: 
(p. 15). Lettering as in fig. 20. 
20. Diagram of heart of the monstrosity referred to on p. 16. 
V. Ventricle. 
Au. Auricle. 
SV. Sinus venosus. 
DC. Duct of Cuvier. 
DO’. Inner or adjacent ducts of Cuvier. 
VJ. Jugular vein. 
VC. Cardinal vein. 
VC’. Inner or adjacent cardinal veins. 


Fig. 21. Diagram of Wolffian duct § wrinary bladder of anormal Trout embryo (p. 7). 
Lettering as in fig. 26. 
Fig, 22. es 3 5 in the monstr osity referred to on 
p. 16. Lettering as in fig. 26. 
Fig. 23. ss - 5 in the case referred to on p. 17. 
Lettering as in fig. 26. 
Fig. 24. ‘ is 53 in the case referred to on p. 17. 
Lettering as in fig. 26. 
Fig. 25. ¥ %s = _ inthe case referred to on p. 17. 
Lettering as in fig. 26. 
Fig. 26. 93 *5 3 in the case referred to on p. 18. 
BL. Bladder. 
BL’. Bladder formed by the inner or adjacent pair of Wolffian 
ducts. 
WD. Wolthan duct. 
WDb, WDb’. Right and left Wolffian ducts belonging to one of the twins 
in each of the figures. 
WDa,WDa'. Ueft and right Wolffian ducts belonging to the other twin 
in each of the figures. 
Int. Intestine. 
R. Rectum. 
V. Vent. 
P, Urinary pore. 
Prats IV. 
Fig. 27. Lateral view of the composite auditory sac present in the monstrosity 


Fig. 


illustrated in figs. 5, 6,10. (See pp. 11, 13.) 
Sace. Saccule. 
Utr. Utricle. 
a.se. An anteriorly placed semicircular canal. 
psec. A posteriorly 


. 28. Diagram of head-kidney of a normal Trout embry 0 (p. 7). 
GL. 


Glomerular tuft of vessels. 

A, Afferent vessel. 
V. Efferent vessel. 
C. Capsule of glomerulus. 

CWD. Commencement of Wolffian duct WD. 

HK. Coils of Wolttian duct im head-kidney. 
29. Onapaasie head-kidney in a double monstrosity (p. 10). Lettering as in 
g. 28. 
Two elomerular tufts are present in a cavity divided into three chambers. 
O. Mesial chamber. 


g. 30. Head-kidney in a double monstrosity of same general type as last, but here 


the glomeruli have not united together, though the adjacent halves of each 
give origin to no Wolffian duct or only to a short blind one (p. 10). 
Lettering as in fig. 28. 


Figs. 31 & 32. Head-kidney y glomeruli of other two double monstrosities showing a 


degree of duplicity greater than that illustrated in fig. 29 (pp. 14, 16). 
‘A blind sacculated tube comes off from the middle compartment of the 
composite glomerulus and represents united adjacent Wolffian ducts. 


24 MR. R. GURNEY ON THE [May 12, 


2. The Metamorphoses of the Decapod Crustaceans Ligeon 
(Crangon) fasciatus Risso and Algeon (Crangon) 
trispinosus (Hailstone). By Rosert Gurney, B.A., 
E.Z.S. 

[Received March 2, 1903. ] 


(Plates V. & VI.*) 


During a residence of nearly a year at Plymouth, I was able to 
obtain and study a very great variety of Decapod larve, and 
among them those of five species of Crangonide. Of these five 
species, which it will be convenient here to consider as all 
belonging to the genus Crangon, three, namely Crangon vulgaris, 
C. spinosus, and C. nanus, have already been fully described by 
Sars, and C. vulgaris has also attracted the attention of a number 
of other workers in this field of research. The remaining two 
species, Crangon trispinosus and (. fasciatus, have remained 
hitherto unrecognised. The identity of these larvee was placed 
beyond all doubt both by hatching them from the egg and by 
observing the last moult to the adult form. Attempts at rearing 
the larve hatched in the Laboratory were quite unsuccessful, 
though with the same methods (plunger jars) and under 
apparently similar conditions I was able to rear the larvee of 
Pandalina brevirostris right through to the postlarval form. 


Description of the Larve. 


I. CRANGON FASCIATUS (Risso). 


Length of larva on hatching, exclusive of the rostrum 7, 1°8 to 
2:05 mm. Average of 24 specimens 2°70 mm. The rostrum 
measures about ‘17 mm. at this stage. 

The general form of the body is slender, the abdominal segments 
being not distinctly narrower than the thorax. The carapace 1s 
prolonged anteriorly into a slender, pointed rostrum, reaching 
about half the length of the peduncle of the first antenna. Below, 
its margin is evenly arched and without teeth, ending in front, at 
the base of the second antenna, in a blunt process. There is no 
fold of the carapace over the eyes, so that the latter appear as 
prominent facetted areas of the carapace itself (see Williamson, 
1901, p. 113). In front of the eyes, and on either side of the 
rostrum, is a small knob-like process (see Pl. V. fig. 1). I can find 


a Hox explanation of the Plates, see p. 30. 

s here, so in the case of C. trispinosus the rostrum is left out of a 
‘ § ccount, as 
measurements from the anterior edge of the eye to the edge of the lateral lobe of ihe 
tail-plate give a truer basis of comparison for body-length of different species. 


DZ§.1903,vol.U. PV. 


Bale aDanidsson LS sculp. 


R.Gdel. 


CRANGON LARVA. 


124, S08), vac ULE, VAL 


R.Gdel. oy Presley icles sent 
; CRANGON LARVA. 


1903. ] METAMORPHOSES OF DECAPOD CRUSTACEANS. 25 


no mention of these structures in the descriptions of any Crangon 
larvee, though they are certainly present both in OC. trispinosus 
and in C. nanus. It seems to me quite possible that they repre- 
sent the frontal sensory processes which have been found on 
certain Cirripede nauplii for example, and which have been 
supposed to be the vestiges of a pair of preoral appendages. 

In the abdomen, the first and second segments are characterised 
by the lateral expansion of their epimera (PI. V. fig. 1), which 
makes them appear considerably broader than the succeeding seg- 
ments. The third segment bears dorsally two strong backwardly 
directed spines. In the fourth segment these spines are represented 
by a pair of very small knobs, while in the succeeding segment 
they are shifted somewhat laterally and have the form of a pair 
of long, downwardly curved, blunt processes. The tail-plate is of 
the usual Crangonid type, with seven strong ciliated spimes on 
either side. These spines, however, are somewhat peculiar in 
having quite blunt terminations, a feature which persists in the 
later stages also. 

The first antenne are unusually long—about two-thirds the 
length of the carapace. The inner flagellum, which is a direct 
prolongation of the stem, is ciliated and very bluntly pointed. 
This blunt termination looks at first sight like the result of an 
injury; but as all the larve were the same in this respect, this 
condition is clearly normal. The scale of the second antenna 
(Pl. V. fig. 3) is long and very narrow, with seven internal sete, 
two sete and ashort spine terminally, and two short external sete. 
The first maxilla (PI. V. fig. 4) has a two-jointed palp, and in other 
respects this appendage and the first maxilla also conform exactly 
to the characteristic crangonid type. 

The exopodite of the first maxillipede (Pl. VI. fig. 6) has three 
apical sete, and two sete on its external edge, differing in this 
respect from Crangon vulgaris, which has but one external seta. 
In the endopodite I can detect only three joints, and in the same 
way in the other maxillipedes I can only make out four. 
Williamson shows the palp of the first maxillipede in Crangon 
vulgaris as distinctly divided into four joints, but though there 
are certainly four small lobes, I cannot trace the separation of the 
second segment into two, nor of the third segment in the case 
of the other two pairs. 

The more advanced larvee of Crangon fasciatus are compara- 
tively rare in the tow-net collection, but four stages are to be 
distinguished in my material. That corresponding to stage iv. of 
C. vulgaris, i.e. with the pleopods represented by short buds, 
I have not found, though there can be but little doubt that it 
occurs. 

In the last stage (stage v.) the length of the body is about 
3°5-3'°8 mm., the rostrum extending about -2 mm. beyond the 
eyes. The form of the rostrum, which acquires its definitive 
shape in the second stage, differs from that of the first stage. 


26 MR. R. GURNEY ON THE [May 12, 


The basal part between the eyes is broad, and distally it con- 
tracts suddenly and tapers to a sharp point. In one example this 
narrow part was found to be bifid down to its base, though the 
postlarval form moulted from it was in no way abnormal. The 
general form of the carapace is unaltered. 

Tn the abdomen the laterodorsal spines of the fifth segment are 
longer, and end in asharply down-curved hook. I have drawings 
of a larva taken in July 1901 at Plymouth, in which this spine 
is of an unusual length, being about two-thirds the length of the 
long sixth segment. 

The tail-plate, now distinctly separated from the sixth segment, 
is much narrower at the end compared with its total length, 
and its sides are not straight, but conspicuously curved. There 
are still seven sete on each side of the posterior margin, but a 
short way up the side on either hand is a minute knob, repre- 
senting the eighth seta found in other species. 

The antenne (PI. VI. fig. 14) are now still more elongated, the 
flagellum of the second antenna now considerably exceeding the 
scale in length. The latter has now a large number of marginal 
setze. 

Such changes as have occurred in the maxill are unimportant. 
In the maxillipedes five joints can now be detected in the endo- 
podite of the second and third, but that of the first remains to all 
appearance three-jointed. All the thoracic appendages, as well 
as the pleopods, are now present, and have the usual Crangonid 
form. The first five pairs of thoracic appendages have exopodites. 
Asregards the gills, not having any example of stage iv., I cannot 
offer any observations as to their origin, but in the last stage 
they are well developed, and the five posterior pairs are distinctly 
pleurobranchs. In this respect C. fasciatus and also C. tri- 
spinosus differ from C. vulgaris, in which Williamson describes 
the gills as having the position of arthrobranchs at this stage. 

The foregoing account shows that the larva of C. fasciatus 1s 
readily distinguishable from all other Crangon larvee yet described. 
The larva most closely approaching it seems to be one described 
by Claus (1861, taf. 11. fig. 1). This larva possibly may be that 
ot C. sculptus, and differs from that now under consideration in 
its much more compact body and the relative length of its dorsal 
abdominal spines. The length of the antenne, slender body, and 
shape and size of the abdominal spines are distinctive of the larva 
of C. fasciatus. 

The next moult leads at once to the postlarval stage, in which 
the adult form is assumed. The young shrimp has the broad, 
depressed form of the adult, with its characteristic square rostrum 
and bright, somewhat banded, colourmg. The only important 
point of difference lies in the sculpturing of the carapace. In this 
first postlarval stage the carapace is smooth except for a short 
anterior median ridge, with two or three blunt prominences. 


1903. | METAMORPHOSES OF DECAPOD CRUSTACEANS. 27 


TI. Grancon Trisprvosus Hailstone.* 


Length of the first larva 1:8 to 20 mm.; average for twenty- 
five specimens being 1:9 mm. The body is light greenish yellow 
in colour, with a conspicuous branching chromatophore placed 
dorsally in the middle of the thorax. In general form the zovxa 
resembles that of C. nanus, the body being comparatively thick- 
set and tapering gradually backwards. 

The lateral edges of the carapace are deeply arched below and 
perfectly smooth except for a blunt spine-like prominence below 
the base of the second antenna (PI. VI. fig. 7). Anteriorly it is 
prolonged into a short pointed rostrum barely half the length of 
the peduncle of the first antenna. The carapace is not folded off 
over the eyes, so that the latter appear to be a part of it. The 
same pair of small outgrowths in front of the eyes as were 
described in C. fasciatus also occur in C. trispinosus (Bis Wae: 
fig. 8). No trace of them can be detected in later stages. 

The abdominal segments have their posterior dorsal edges 
usually evenly rounded, without spines, but the fifth segment 
sometimes has a pair of short dorso-lateral spines, and traces of 
them can often be detected. The tail-plate is of the usual form, 
the spines being sharply pointed, and not blunt as in C. fasciatus 
(Pl. VI. fig. 8). 

In the form of the appendages C. trispinosus differs so little 
from Cl. nanus, which has been fully described by Sars, that it is 
not necessary to describe them in full. The first antenne of 
C. nanus, however, differ from those of C. trispinosus in that the 
inner flagellum is armed with a few minute spines in addition to 
the long cilia which are common to both (Pl. VI. figs. 9 & 10). 
The second antenne in the two species agree in all respects, but 
both species differ from C. echinulatus, which is in other respects 
closely similar, in the much narrower form of the scale. 

In the later stages of the larva the body becomes more compact 
and thick-set, having an appearance very characteristic of this 
species and of C. manus, from which, however, it is easily distin- 
guishable by the absence of spines from the abdominal segments 
and from the lower edges of the carapace. 

The rostrum is now very broad at the base, contracting suddenly 
near its end and continuing asa short, sharp spine. As far as 
the scanty material at my disposal for the study of the later stages 
goes, the spines occasionally developed on the fifth abdominal 
segment of the first larva are lost with the first moult. . 

The gills appear in the fourth stage of development. At this 
stage five gills can be seen placed, as it appears to me, in the 
position of pleurobranchs—that is to say, they are well above 
the apparent attachment of the legs (PI. VI. fig. 12). As the legs 


* Claus has figured (1884, taf. vii.) a Crangon larva which almost certainly belongs 
to this species. The absence of abdominal spines, the form of the rostrum, and the 
presence of an exopodite on the second pereiopod identify the larva with C. tri- 
Spinosus. 


28 MR. R. GURNEY ON THE [May 12, 


are six-jointed, one might assume that a seventh is fused with the 
body-wall, in which case the gills might be interpreted as arthro- 
branchs or podobranchs at will. Still, in the common acceptance 
of the term, they certainly seem to me to be properly designated 
as pleurobranchs. 

I have been unable to detect at this stage the small arthro- 
branch found in the last larva and in the adult on the third 
maxillipedes. 

The first postlarval stage differs from the adult in that there is 
but a single median tooth on the dorsal surface of the carapace, 
instead of three teeth transversely arranged. For this reason 
the young shrimp at this period is hardly distinguishable from 
C. nanus; but the latter is somewhat smaller, and the single 
median tooth of the carapace seems to rise from a short median 
ridge. 

Sars figures a small posterior tooth in this ridge, corresponding 
to the second tooth of the adult, but I have been unable to detect 
it in my specimens. The possession of peculiarly distinct larval 
forms with sudden transition to a uniform postlarval type is one 
of the most remarkable features in crangonid metamorphosis, and 
the two forms described are no exception to the rule. 


Conclusions. 


Our knowledge of Decapod metamorphosis is for the most part 
very fragmentary, and for this reason it has been impossible to 
use the larval stages as evidence of relationship. We have, how- 
ever, an exceptionally full knowledge of the metamorphosis of the 
Crangonide ; and it will be interesting to see if a comparison 
of the larvee will throw light on the classification of the group. 
The eleven species of Crangonide which have been recorded as 
British have been divided up among five genera :—Crangon Fabr., 
Cheraphilus Kinahan, Ageon Risso, Pontophilus Leach, and 
Sabinea Owen; though Ortmann in his revision of the family 
joins Cheraphilus, dgeon, and Pontophilus into one genus Ponto- 
philus. The genus Sabinea may here be left out of account. 

Looking, then, at the larve of the species that have been 
described, we find them falling, as it seems to me, naturally into 
three groups :— 

1. C. vulgaris and C. allmanni: characterised by a one-jointed 
maxilla-palp and the absence of an exopodite on the second leg 
in the Mysis-stage. 

9 Ybge ia cle i L + 

2. C. trispinosus, C. nanus, C. echinulatus, and C. fasciatus : 
characterised by their two-jointed maxilla-palp, possession of five 
pairs of exopodites in the Mysis-stage, form of the rostrum, and 
arrangement of the abdominal spines. 

e : ria lia 

3. C. sptnosus and C. norvegicus: distinguished from the second 
group by their extremely elongated body form, shape of the 
rostrum, possession of a single long median spine on the third 
abdominal segment, and by the form of the tail-plate. 


1903. ] METAMORPHOSES OF DECAPOD CRUSTACEANS. 29 


C. fasciatus cannot be separated generically as a larva from 
the other species named. A characteristic of the group to which 
it belongs is the paired arrangement of the abdominal spines. It 
is true that they are practically absent in C’ trispmmosus, but their 
occasional appearance on one segment shows that their loss is a 
recent one. Classifying therefore on the basis of the larval form, 
we must merge the genus Cheraphilus into Ageon, leaving 
the genus Pontophilus to include P. spinosus and P, norvegicus. 

This arrangement seems to fit the adults as well as it does the 
larvee. In the first place, Ortmann seems to have made an error in 
including all the above-mentioned forms in a single genus 
Pontophilus. He gives as characters of this genus “ Gills seven ; 
six pleurobranchs (2, 4, l,m, 1, 0), and one rudimentary podo- 
branch 2”; but I have found the following gill-formula to hold 
for Cheraphilus trispinosus, C. nanus, and dgeon fasciatus :— 


a. Daa Cron 
feeeeae irene Lae NE 
We Walie esta Ep | 
[AIOE call, aS | 
| NADO coo) Lae |) ll | 
VANE ss. ae wa ea 
25 Bare actallpgesatle il 
(Pd: 2) erga sad ea anil 
[ees ED UB eat Merce early aL 
| XII | a | =6+2 or 3 Ep 


In Pontophilus spinosus there are seven gills, there being a 
podobranch upon the second maxillipede. 

Tf this statement holds good for the remaining species, then the 
two genera Cheraphilus and Mgeon should be separated from 
Pontophilus. As regards their relation to one another, the only 
important differences between them seem to lie in the shape of | 
the rostrum and the direction of the bend of the apices of the 
gills (Spence Bate). Such differences do not seem to me of 
sufficient importance to allow of generic distinction, and I have 
therefore in the heading of this paper treated the two species 
dealt with as both belonging to the genus dgeon Risso. 


Literature referred to in the foregoing paper. 


Bats, C. Spence.—Report on the Crustacea Macrura collected 
during the ‘ Challenger’ Expedition. 1888. 

Cuaus, C.—Zur Kenntniss der Kreislaufsorgane der Schizopoden 
und Decapoden. Arb. Zool. Inst. Wien, v.. pp. 271-318 (1884). 

Cuaus, C.—Zur Kenntniss der Malacostracenlarven. Wiirzb. 
Naturwiss. Zeits. 1. pp. 23-46 (1861). 


30 MR. MARTIN JACOBY ON NEW [May 12, 


Krvanay, J. R.—Synopsis of the Species of the Families Crango- 
nide and Galatheide which inhabit the Seas around the 
British Isles. Proc. Roy. Ivish Acad. viii. pp. 67-80 (1862). 

Orrmann, A. E.—A Study of the Systematic and Geographic 
Distribution of the Decapod family Crangonidee. Proc. Acad. 
Nat. Sci. Philad. 1895, pp. 173-197. 

Sars, G. O.— Bidrag til Kundskaben om Decapodernes Forvand- 
linger. III. Fam. Crangonide, Arch. f. Math. og Naturvid. 
xiv. pp. 132-195 (1890). 

Srmppine, T. R. R.—South African Crustacea. Report of the 
Government Biologist, Cape Town, 1901. 

Wiuramson, H. C.—On the Larval Stages of the Decapod 
Crustacea—The Shrimp (Crangon vulgaris Fabr.). 19th 
Ann. Rep. Fish. Board Scotland, pt. 111, pp. 92-119 (1901). 


EXPLANATION OF THE PLATES. 
PuatE V. 


Crangon fasciatus (p. 24). First larva, dorsal view. > 60. 
The same, lateral view. X 68. 

Second antenna. X 100. 

First maxilla. 185. 

Second maxilla. 185. 


3) 


OR to 


Prats VI. 
Fi 


99 


6. Crangon fasciatus (p. 24). First maxillipede. X 124. 

7. Crangon trispinosus (p. 27). First larva, lateral view. X 65. 

8. = re dorsal view. X 65. 

9. ys op end of first antenna. 140. 

10. Crangon nanus (p. 27). End of first antenna. X 150. 

11. Crangon trispinosus (p. 27). 1st pereiopod of the last larva. X 60. 

12. 3 Pereiopods 2 to 5 showing gills. Stageiv. > 100. 
13. f Telson of first postlarval stage. X 100. 

14. Crangon fasciatus (p. 24). Last larva, first antenna. X 72. 


3. Descriptions of New Species of South-American Coleoptera 
of the Family Chrysomelide. By Martiy Jacosy, 
HIBS” 

[Received April 9, 1903.} 


The following paper is based on the material which has 
gradually accumulated in my collection during several years, and 
which contains examples of species of which I have been unable 
to find any descriptions. The fine insects of the genus Doryphora, 
of which hitherto about 400 species were known, are here further 
augmented by more than 30 new species. A few others are de- 
scribed from Central America which were not known to me at the 
time of the publication of the ‘ Biologia’ of that country, and 
these belong to the genera Desmogramma, Stilodes, Deuterocampta, 
Cosmogramma, Zygogramma, Prosicela, Leptinotarsa, Calligrapha, 
and Hlythrosphera. 

* Communicated by the SECRETARY. 


1903. ] SOUTH-AMERICAN COLEOPTERA. 31 


DoRYPHORA BREVIFASCIATA, Sp. Nov. 


Flavous, the antenne black, the head with two greenish-black 
spots; thorax with a few fine spots, the base with a narrow 
transverse greenish band; elytra widened posteriorly, geminate 
punctate-striate, a sutural band, narrowed posteriorly, the lateral 
margins, and a short transverse band below the middle greenish 
eeneous. 

Length 9 millim. 

Head with a few minute punctures, flavous, the vertex with 
two large subquadrate metallic green spots; antennee extending 
beyond the base of the thorax, black, the terminal joints strongly 
widened, the basal joints flavous below; thorax of usual shape, 
the surface with a few fine punctures, flavous, the base with a 
narrow transverse, deeply concave band, not extending to the 
lateral margins, dark metallic green; scutellum metallic green ; 
elytra wider at the base than the thorax, gradually widened 
posteriorly, strongly geminate punctate-striate at the inner disc, 
the sides irregularly punctured below the middle, flavous, the 
suture with a metallic green band which is suddenly narrowed 
below the middle, each elytron with a short transverse band at 
the latter place not extending to either margin and the lateral 
margins metallic green; belowand the legs flavous, the mesosternal 
process short. 

Hab. Brazil. (A single specimen.) 

In coloration this species comes near D. jucunda Stal, but in 
the latter the head and underside are metallic zeneous, and the 
elytral transverse band extends across the entire disc and is 
placed higher up. 


DoryPHORA SPHRICA, Sp. NOV. 


Subglobular, testaceous, the antenne and tarsi black ; thorax 
strongly punctured at the sides ; elytra strongly geminate punctate- 
striate near the suture, irregularly punctured at the sides. 

Length 9 millim. 

Head very closely and finely punctured, labrum testaceous, 
apex of mandibles black ; antennz with the terminal joints strongly 
widened, black, the three basal joints flavous below; thorax 
somewhat narrowed anteriorly, the sides feebly rounded, the 
anterior angles distinct, but not pointed or produced, the disc 
convex, finely and sparingly punctured at the middle, strongly 
and closely semirugose punctate at the sides; scutellum smooth, 
shining ; elytra very convex, deeply punctured in double and 
treble rows at the inner disc, irregularly so at the sides ; underside 
and legs pale fulvous, the tarsi black, the mesosternal process 
very short. 

This is a species of almost subglobular shape, of which I un- 
fortunately do not know the “habitat,” and of which a single 
specimen is contained in my collection; there will be no difficulty 
in recognising the species on account of the uniform coloration, 
the black antenne and tarsi, in connection with the shape. 


32 MR. MARTIN JACOBY ON NEW [May 12, 


DoryPHORA NIGROVARIANS. Sp. Nov. 


Black; thorax sparingly and minutely punctured ; elytra finely 
geminate punctate-striate, flavous, a transverse curved band below 
the base, another deeply dentate one below the middle not extending 
to the suture, a small spot at the sides near the apex, and the 
suture black, the latter widened into a sharp triangular mark 
near the apex. 

Length 10 millim. 

Of somewhat flattened appearance, the head sparingly and 
finely punctured, with a small fulvous spot at the vertex ; antenne 
with the terminal joints broadly flattened, black, the lower three 
joints flavous below ; thorax rather short, the sides strongly 
rounded anteriorly, the extreme anterior angles fulvous, the 
surface obscure greenish black, very sparingly and minutely 
punctured, the lateral margins accompanied by a row of deep 
punctures ; scutellum black ; elytra bright yellow, finely punctured 
in double or treble rows, except near the apex where the 
punctuation is more irregular, the suture widened near the apex 
into an anchor-shaped pointed mark, black; below the base is a 
transverse black band which curves upwards at the shoulders, 
this band is widened near the suture, below the middle is another 
deeply dentate transverse band which touches the lateral but not 
the sutural margin, lastly a small black spot is attached to the 
lateral margin near the apex; underside and legs black, the 
mesosternal process short and broad. 

Hab. Brazil. 

I do not know the exact locality of this well-marked species, of 
which I possess a single specimen. 


DoryPHORA SEMIVIRIDIS, Sp. Nov. 


Bright metallic green, the elytra fulvous; thorax strongly 
punctured at the sides; elytra semi-regularly and very deeply 
punctured. 

Length 8 millim. 

Head metallic green, the vertex with a small fulvous spot; 
antenne with strongly widened terminal joints, black, the lower 
three joints flavous below; thorax short, more than twice as 
broad as long, the sides straight at the base, strongly rounded 
anteriorly, the angles not produced, the surface bright metallic 
green, impunctate at the middle, strongly punctured at the sides ; 
scutellum metallic green; elytra with the greatest elevation at 
the middle, pale fulvous, almost foveolate-punctate, the punctures 
arranged in two or three rows near the suture, more irregularly 
so at the sides, their epipleure fulvous ; below and the legs metallic 
green, the mesosternal process very short. 

Hab. Costa Rica. 

This must be a rare species, as I have not come across it during 
the progress of the Central- American monograph in the ‘ Biologia’; 
it will find its place near D. signiceps St&l, from which it differs 


1903. ] SOUTH-AMERICAN COLEOPTERA, 33 


in the non-convex elytral interstices and the more irregular 
punctation. 


DorRYPHORA PARAGUAYENSIS, Sp. NOV. 


Metallic green, the labrum and a spot at the vertex testaceous ; 
thorax short and strongly transverse, the sides rugosely punctured, 
the extreme lateral margins testaceous; elytra flavous, strongly 
geminate punctate-striate, the suture, the lateral margins, and a 
discoidal stripe, from the base to below the middle, metallic 
green. 

Length 10 millim. 

Head sparingly and finely punctured, metallic green, a central 
spot at the middle of the vertex and the labrum testaceous ; 
antenne extending to the base of the thorax, black, the basal 
joints fulvous below, terminal joints widened; thorax at least 
three times broader than long, the sides rounded, the anterior 
angles not produced but acute, the dise sparingly punctured, the 
sides strongly rugose-punctate, the extreme lateral margins and 
the anterior angles testaceous, the surface bright metallic green , 
scutellum of silky green appearance ; elytia not wider at the base 
than the thorax, having their greatest elevations placed at the 
middle, flavous, each elytron with about ten rows of deep 
punctures, placed mostly in double rows, those near the suture 
and the last submarginal row consisting of single punctures only, 
a sutural band gradually narrowed towards the apex, the lateral 
margins and a discoidal broad stripe also greatly narrowed 
posteriorly and abbreviated at the apex, metallic bright green ; 
underside metallic green, coxee and mesosternal process flavous, 
the latter very short. 

Hab. Paraguay. (A single specimen.) 

Of nearly similar coloration as D. vinula Stal, but with geminate 
punctate elytra, differently coloured thorax and underside. 


DoryYPHORA FLAVIMANA, Sp. Nov. 


Head and thorax flavous, each with four greenish spots, minutely 
punctured ; elytra finely geminate punctate-striate, flavous, the 
sutural and lateral margins, a spot on the shoulders, a transverse 
dentate band below the base, and a transverse large spot below 
the middle dark sneous; underside and legs flavous, antennze 
black. 

Length 10 millim. 

Head with a few very minute punctures, flavous, with four 
greenish spots placed subquadrately, another very small spot 1s 
situated at the edge of the clypeus; antenne extending beyond 
the base of the thorax, slender, black, lower two joints flavous 
below, terminal joints but slightly widened; thorax short, of 
usual strongly transverse shape, the sides straight at the base, the 
surface with a few minute punctures, flavous, narrowly margined 
with greenish eneous ; the dise with four greenish spots placed 
transversely, the two middle ones larger and of elongate shape ; 


Proc. Zoou. Soc.—1903, Vou. II. No. III. 3 


34 MR. MARTIN JACOBY ON NEW [May 12, 


scutellum fulvous; elytra evenly convex, light flavous, punctured 
in double rows, the punctures fine, rather irregularly placed here 
and there, all the margins narrowly, and a humeral spot dark 
zneous, below the middle isa broad transverse band with strongly 
dentate edges and not extending to the sides, another transverse, 
slightly curved band is placed below the middle touching the 
lateral margins but not the suture, the latter is likewise of 
eeneous colour and widened angulately opposite the posterior band, 
with which it is nearly connected at that place ; elytral epipleure, 
underside and legs flavous, the tibize streaked with black at the 
outer margin, mesosternal process short and straight. 

Hab. Vilcanota, Peru. (A single specimen.) 

In the Museum of the Stettin collection another specimen from 
Bolivia is contained, sent to me for examination, which I cannot 
separate from this species, but the elytral band at the base is not 
dentate but concave at the anterior margin and is connected with 
the humeral spot, and the posterior band is reduced to a trans- 
verse spot not extending to either margin ; in other respects there 
are no differences of importance. 


DoRYPHORA DIVERSIPES, Sp. NOV. 


Head and thorax flavous, the former with a triangular black 
mark, the latter with two narrow black bands, impunctate ; elytra 
obscure neous, closely punctured, the lateral margins flavous ; 
legs and underside testaceous, marked with black. 

Length 9 millim. 

Of very moderately convex shape ; the head impunctate, flavous, 
with a greenish-black V-shaped mark at the vertex; antenne 
short and feeble, blackish, the basal joint and the base of the 
following two or three joints flavous, terminal joints not longer 
than broad; thorax with feebly rounded lateral margins, not 
narrowed anteriorly, the anterior angles not mucronate, the disc 
impunctate, flavous, with two narrow blackish bands from the 
base to the apex, one at each side, and slightly curved, shape of 
the thorax twice and a half broader than long ; scutellum flavous ; 
elytra not wider at the base than the thorax, obscure brownish 
zeneous, finely and closely punctured, the punctures near the suture 
arranged in indistinct double rows, the lateral margins narrowly 
flavous, with a single row of deeper punctures; underside 
testaceous, the sides of the breast andthe abdomen partly black, legs 
flavous, femora and tibiee with a black streak below, tarsi entirely 
black ; mesosternal process flavous, very short. 

Hab. Peru. (A single specimen.) 


DoRYPHORA AMICTA, Sp. Nov. 


Underside bluish black, upperside metallic violaceous ; head 
and thorax obscure bluish, finely punctured; elytra closely semi- 


punctate-striate, the interstices minutely granulate and aciculate 
here and there. 


Length 12 millim. 


1903. ] SOUTH-AMERICAN COLEOPTERA. 35 


Of subquadrate, parallel shape; the head sparingly and finely 
punctured, dark blue, subopaque, labrum testaceous, stained with 
black; antennz extending below the base of the thorax, black, 
apex of terminal joints fulvous, the latter joints widened but 
longer than broad; thorax of usual shape, the sides strongly 
rounded anteriorly, the anterior angles mucronate, the disc very 
minutely and rather closely punctured, of the same colour as the 
head ; scutellum black, shining ; elytra finely punctured in closely 
approached rows, which are fairly regular even at the sides, the 
interstices minutely granulate and partly aciculate, their epipleuree 
blue without, testaceous within, the mesosternal process long and 
straight. 

Hab. Peru. (A single specimen.) 

This species cannot be referred to D. prasina Erichs. : the elytra 
are violaceous and the rest of the insect is dark blue, not green; , 
the sculpturing of the thorax and elytra is different, and the latter 
are neither widened posteriorly nor produced at the apex as Stal 
states in his description of the species. The punctation of the 
elytra also differs from that of the other blue species from the 
same locality. 


DoryPHORA SEMINIGRA, Sp. nov. 

Black, the elytra dark metallic green, subopaque ; thorax very 
closely punctured at the sides; elytra with closely approached, 
double, irregular rows of punctures. 

Length 15 millim. 

Head closely punctured round the eyes, black; antenne with 
the terminal five joints widened, slightly longer than broad, 
entirely black; thorax twice as broad as long, the lateral margins 
straight at the base, rounded near the apex, the anterior margins 
acutely pointed, the surface flattened at the sides, the-latter finely 
rugosely punctured, the middle sparingly and very finely punctate, 
black, opaque ; scutellum black; elytra dark green, moderately 
shining, strongly punctured in irregular double rows, closely 
placed ; underside and legs black, mesosternal process short and 
stout. 

Hab. Peru. (A single specimen.) 

Larger than D. prasina Erichs., of different coloration, and 
with the sculpturing of the elytra stronger and geminate, the 
latter have their greatest elevation at the middle. 


DoRYPHORA BISTRIGUTTATA, Sp. NOV. 


Greenish black, the thorax closely and strongly punctured, with 
a fulvous spot at the anterior margin; elytra very closely and 
strongly punctured at the sides, semipunctate-striate near the 
suture, greenish, each elytron with a round spot near the scutellum, 
another near the apex, and a transverse band at the middle 
fulvous. ° 

Length 14 millim. 

Head finely punctured between the eyes, opaque, greenish ; 


36 MR. MARTIN JACOBY ON NEW [May 12, 


antenne short and filiform, black, the lower two joints fulvous 
below, the apex of the last joint likewise of this colour, terminal 
joints slender, not widened ; thorax rather more than twice as 
broad as long, the sides straight at the base, rounded anteriorly, 
the anterior angles acute but not produced, the disc closely rugose- 
punctate at the sides, less closely punctured at the middle, dark 
greenish, with a small fulvous spot placed at the middle of the 
anterior margin; scutellum black; elytra with the greatest 
elevation near the base, strongly deflexed posteriorly, more 
strongly punctured than the thorax, the punctures larger at the 
sides near the base and crowded, the imner disc more regularly 
punctured in rows, dark greenish, with a rounded spot near the 
scutellum, a more elongate and larger spot near the apex, a trans- 
verse band at the middle (not extending to either margin), and 
the lateral margins from the base to the middle fulvous; the 
latter colour also extends to the epipleure in the same degree; 
mesosternal process long and slightly curved. 

Hab. Colombia. (A single specimen.) 

Much smaller than D. fulgora St., and with entirely differently 
shaped fulvous markings. 


DoRyYPHORA SEXSPILOTA, sp. nov. 


Testaceous, the underside and legs metallic greenish black; 
thorax impunctate, the disc greenish, the lateral margins flavous ; 
elytra geminate punctate-striate near the suture, irregularly 
punctured laterally, the sutural margin, a small spot on the 
shoulder and two others near the suture, before and below the 
middle, metallic greenish. 

‘Length 10 millim. 

Head with a few fine punctures, green, opaque ; antenne short, 
black, the lower joints fulvous below, terminal joints slightly 
widened ; thorax with feebly rounded sides, the anterior angles 
pointed but not produced, the middle of the dise impunctate, the 
sides finely punctured, flavous, the middle portion greenish, 
opaque, in shape of a broad transverse band, the sides of which 
are concave and acutely separated from the flavous portion ; 
scutellum metallic green ; elytra strongly and very closely punc- 
tured, the punctures from the suture to the middle of the disc 
arranged in double rows, the sides irregularly punctate, the 
interstices raised into smooth longitudinal narrow lines, the suture 
narrowly metallic green, the shoulders with a A-shaped green 
spot, two other spots are placed between the third and fourth 
interstices before and below the middle; underside and legs 
Srey black, shining, the mesosternal process very short and 
blunt. 

Hab. Sao Paulo, Brazil, (Three specimens.) 


DoRYPHORA SUBMETALLICA, Sp. nov. 
4Hneous, the head and thorax opaque, nearly impunctate ; elytra 


1903. | SOUTH*AMERICGAN COLEOPTERA, 37 


regularly punctate-striate, testaceous, the basal margin, a broad 
sutural band, connected with a transverse band before the middle 
and extending downwards, and the lateral margins narrowly 
zeneous. 

Length 12 millim. 

Head with a few very minute punctures, dull eneous ; antenne 
black, the lower two joints fulvous below, the terminal joints 
widened (the last two joints wanting); thorax twice as broad as 
long, the lateral margins straight, the anterior angles acute, 
pointed, the surface of the same colour as the head and similarly 
sculptured, opaque; elytra with their greatest elevation near the 
base, punctured in very regular rows, the punctures rather elon- 
gate in shape, piceous, the ground-colour flavous, the lateral 
margins very narrowly and the suture broadly dark zneous, the 
base with a narrow eneous stripe extending to the middle of the 
width of the elytra, followed by a broad transverse eneous band 
which at each end curves downwards at right angles to below the 
middle, this downward branch is rather suddenly constricted at 
its middle; underside and legs metallic eneous, shining; meso- 
sternal process moderately long and straight. 

Hab. Peru. (A single specimen.) 


DoryPHORA ECUADORIENSIS, Sp. NOV. 


Black, thorax closely and finely punctured; elytra flavous, 
strongly, closely, and irregularly punctured, the suture greenish 
black, a transverse spot at the shoulders of triangular shape, a 
small sutural spot, another below the middle, and a curved elongate 
stripe at the sides bluish black. 

Length 10 millim. 

Head closely and finely punctured; antennz short, black, the 
basal two joints flavous below, terminal joints gradually and but 
little widened, the last joint long; thorax black, with a slight 
greenish tint, the anterior angles acute but not tuberculate, the 
dise very closely punctured at the sides and the base, the punctures 
of different size, middle of the dise sparingly punctate, the sides 
with an obsolete depression ; scutellum triangular, greenish black ; 
elytra rather pointed at the apex, flavous, very closely and strongly 
punctured, especially so at the sides, the punctures of piceous 
colour, irregularly placed with some indications of rows near the 
suture, the latter greenish black, the shoulders with a transverse 
triangular spot, nearly connected with another small sutural spot 
below the base, a curved elongate, posteriorly narrowed stripe at 
the sides, and another small subsutural spot, both below the 
middle, blackish; underside and legs greenish black, the meso- 
sternal process rather short. 

Hab. Ecuador. (A single specimen.) 

This is not a variety of D. instabilis Stal, which the species 
resembles somewhat, as the elytral sculpturing as well as that of 
the thorax are quite different. 


38 _ MR. MARTIN JACOBY ON NEW | May 12, 


DoryYPHORA HISTRIONICA, Sp. NOV. 


Greenish black, the head and thorax sparingly punctured ; 
elytra regularly punctate-striate, the sutural and lateral margins 
greenish black and accompanied by a flavous, narrow stripe. 

Length 9 millim. 

Head with a few fine punctures; labrum testaceous; antenne 
black, the basal two joints fulvous below, terminal joints widened ; 
thorax of usual shape, the sides straight, the anterior angles 
scarcely pointed, the surface finely and sparingly punctured at 
the middle, more strongly so at the sides; scutellum blackish ; 
elytra evenly convex, finely and regularly punctate-striate, each 
elytron with ten rows of punctures, a sutural and a more narrow 
lateral stripe greenish black, the third and the ninth interstice 
flavous, joined at the apex, the rest of the surface fulvous; the 
mesosternal process stout and curved. 

Hab. Bolivia. (A single specimen.) 

A small species, well distinguished by its system of coloration. 


DoRYPHORA FRUHSTORFERI, Sp. Nov. 


Head black ; thorax impunctate, black, the sides and a spot at 
the anterior margin flavous; elytra finely geminate punctate- 
striate, flavous, the sutural and lateral margins and two rounded 
spots placed transversely below the middle black; underside 
black, legs and antennee fulvous. 

Length 10-11 millim. 

Head entirely impunctate, black, opaque; labrum flavous ; 
antenne extending to the base of the thorax, fulvous, the terminal 
joints transversely widened, the last one more elongate; thorax 
more than twice as broad as long, the sides slightly rounded, the 
surface entirely impunctate, black, a spot at the middle of the 
anterior margin and the sides (in shape of a subtriangular large 
spot) flavous; elytra very finely and regularly geminate punctate- 
striate, flavous, the sutural and lateral margins narrowly black, 
below the middle a black spot is attached to the lateral margins 
and a larger spot is placed between it and the suture; the under- 
side black, the legs fulvous, with the posterior femora more or 
less stained with piceous, mesosternal process short. 

Hab. Espirito Santo, Brazil. 

Of this very distinct species I received two specimens from 
Mr. Fruhstorfer. 


DoryPHORA VIRIDIORNATA, Sp. NOV. 


Dark fulvous; thorax minutely punctured, with two broad 
metallic green spots or bands; elytra strongly punctured in double 
rows, pale fulvous, each with a transverse broad metallic green 
band at the base and a triangular spot below the middle. 

Length 9 millim, 

Head extremely finely punctured, fulvous; the vertex with two 
small greenish spots; antenne entirely fulvous, the terminal 


1903. ] SOUTH-AMERICAN COLEOPTERA, 39 


joints distinctly widened ; thorax strongly ,transverse, the sides 
rather strongly rounded before the middle, the anterior angles 
pointed, the surface very finely and subremotely punctured, 
fulvous, the entire sides occupied by a large subquadrate metallic 
green patch ; scutellum fulvous; elytra with deep but remotely 
placed punctures arranged in double rows, the punctation much 
finer towards the apex, flavous or pale fulvous, with a broad 
transverse metallic green band at the base, not extending to either 
margin, and another triangular broad spot below the middle; 
underside fulvous, stained with greenish neous, legs entirely 
fulvous, the mesosternal process rather short. 

Hab. Peru. 

The metallic green elytral patches are separated by the narrow 
fulvous margins and at the middle by a slightly broader division 
or fulvous band; those of the thorax are divided at the middle 
by a narrow, anteriorly widened fulvous band, the sides are 
likewise more broadly fulvous. 


DoryPHORA HONDURAENSIS, Sp. NOV. 


Fulvous, the thorax with four green spots placed transversely, 
sparingly punctured ; elytra strongly punctate-striate, a strongly 
curved transverse band at the base, another straight band below 
the middle, and a spot near the apex metallic green. 

Length 12 millim. 

Head with a few minute punctures, fulvous ; antenne extending 
to the base of the elytra, fulvous, the terminal joints thickened ; 
thorax with the lateral margins strongly rounded anteriorly, the 
angles acute, the surface sparingly but strongly punctured, fulvous, 
with four black spots, placed transversely, two smaller ones at the 
sides round and two larger elongate ones at the middle; scutellum 
fulvous; elytra with rows of deep punctures which run in pairs 
at the sides but single at the inner disc, the ground-colour fulvous, 
at the base a strongly angulate band of green colour is placed ina 
slightly oblique direction and not extending to either margin, 
another band parallel to the first is situated below the middle and 
a transverse spot near the apex ; underside and legs fulvous, the 
mesosternal process very short. 

Hab. Honduras. (A single specimen.) 

This species resembles in its system of coloration and pattern 
D. viridifasciata Jac. (Biol. Centr.-Amer., Coleopt. vol. vi. pt. 1), 
but differs in the entirely fulvous head and in the shape of the 
first elytral band, which is almost semicrescentic, the other band 
is also much narrower, slightly angulate near the lateral margins, 
and both are of slightly oblique direction. 


DorRYPHORA NEOFASCIATA, sp. nov. 


Fulvous ; head and thorax finely punctured, each with four green 
spots; elytra geminate punctate-striate, flavous, a broad trans- 
verse band near the base, curved upwards and including a flavous 
spot and a transverse spot below the middle, greenish ceneous. 


40 MR. MARTIN JACOBY ON NEW [May 12, 


Length 10 millim. 

Head finely and sparingly punctured, fulvous, with four small 
green spots placed quadrately ; antenne black, lower two joints 
flavous below, the terminal joints gradually widened; thorax of 
usual shape, the anterior angles not produced, the surface finely 
and remotely punctured, fulvous, the anterior and posterior mar- 
gins and four spots placed transversely on the disc, metallic green ; 
scutellum dark fulvous; elytra geminate punctate-striate, flavous, 
the lateral margins, the suture, a curved transverse band at the 
base, greatly dilated at the suture and curved upwards at the 
shoulders, and an oblique spot below the middle greenish 
eeneous; underside and legs fulvous, mesosternal process short 
and straight. 

Hab. Colombia. 

Somewhat similarly coloured as D. gerstaeckert Stal, but with 
the head and the thorax fulvous and spotted with green. 


DorYPHORA TERMINATA, Sp. NOV. 


Black, thorax with a few fine punctures ; elytra finely punctate- 
striate, the interstices slightly convex, black, the apex flavous 
with a smail black spot. 

Length 12 millim. 

Head impunctate; antenne black, the terminal joints rather 
strongly widened ; sides of the thorax slightly constricted at the 
base, feebly rounded, the anterior angles produced outwards into 
a small point; elytra with distantly placed irregular rows of fine 
punctures, black, the apex in shape of a broad triangular patch, 
flavous, including a small black spot on each elytron; legs and 
underside black, mesosternal process long and curved. 3 

Hab. Costa Rica. (A single specimen.) 

Of entirely black coloration with the exception of the flavous 
apex of the elytra. 


DoryPHORA BAHIAENSIS, Sp. nov. 


Pale fulvous, the antenne (the basal four joints excepted) 
black ; thorax finely and sparingly punctured ; elytral punctation 
arranged in irregular double rows, fine, the pnnctures piceous, 
surrounding small palish smooth spots. 

Length 12 millim. 

Head impunctate, fulvous; antenne rather short, black, the 
basal four joints fulvous, the terminal joints very broadly flattened ; 
thorax more than twice as broad as long, the sides straight, the 
anterior angles acute but not pointed or produced, anterior 
margin deeply concave behind the eyes, the disc finely and sparingly 
punctured, not more closely but a little more strongly so at the 
sides 5 scutellum fulvous ; elytra with their greatest elevation at 
the middle, rather finely punctured, the punctures near the suture 
here and there arranged in irregular double rows which are more 
plainly marked near the apex, the interstices with numerous 


1 903. SOUTH-AMERICAN COLEOPTERA. 4] 


slightly raised smooth pale spots or patches of irregular shape ; 
below coloured like the upper surface; the mesosternal process 
long and stout. 

Hab. Bahia, Brazil. (A single specimen.) 

Distinguished by the unicolorous upper and under sides and by the 
broadly flattened terminal joints of the antenne and their colour. 


DoRYPHORA IMITANS, Sp. Nov. 


Below black, above fulvous ; thorax flavous, minutely punctured ; 
elytra finely and irregularly punctured; antenne black, the 
anterior femora flavous. 

Length 12 millim. 

Head nearly impunctate, palpi black; antenne with the 
terminal joints distinctly widened, black, the basal joint flavous ; 
thorax scarcely more than twice as broad as long, the sides 
straight at the base, rather strongly obliquely narrowed anteriorly, 
the anterior angles pointed, the disc finely and rather closely 
punctured, the punctures of unequal size ; scutellum fulvous ; elytra 
regularly convex, finely punctured in irregular double rows near 
the suture, the sides irregularly punctate, fulvous, the extreme 
basal margin piceous, the interstices smooth ; below black, the 
anterior femora entirely, the others more or less fulvous below ; 
the mesosternal process short and broad. 

Hab. Colombia. (A single specimen.) 

‘The thorax in this species is less transverse in shape and more 
suddenly pointed anteriorly than in D. bahiaensis ; the elytra ave 
differently sculptured and the mesosternal process is much shorter. 


DorYPHORA FLAVOANNULATA, Sp. NOV. 


Oblong, below black, above and the legs dark fulvous ; thorax 
remotely and strongly punctured at the sides; elytra regularly 
punctate-striate, an interrupted ring-shaped band from the base 
to the middle and an entire similar ring near the apex, flavous , 
antenne and the last abdominal segments pale flavous. 

Length 9 millim. 

Head minutely punctured, pale fulvous, labrum testaceous ; 
antennee extending beyond the base of the thorax, pale fulvous, 
terminal joints subquadrately widened ; thorax nearly three times 
broader than long, the sides straight, rounded near the anterior 
angles only, the dise finely and remotely punctured, the sides 
more strongly so, but little flattened; scutellum pale; elytra 
oblong and parallel, moderately convex, regularly and moderately 
strongly punctate-striate, fulvous, sometimes with a slight greenish 
tint, a ring-shaped band, interrupted at its inner and posterior 
margin, extending from the base to the middle and another nearly 
round and uninterrupted ring near the apex, obscure flavous; below 
black, with the exception of the fulvous legs and similarly coloured 
mesosternal process, the latter long and stout, slightly curved, 
last abdominal segment flavous. 


42, MR. MARTIN JACOBY ON NEW [May 12, 


Hab. Peru. (Three specimens.) 

Somewhat resembling D. estuans L., but the underside black 
and the elytral markings in shape of two rings; in all my speci- 
mens the anterior band is interrupted near the suture and at the 
posterior edge, but this may not be always the case. 


DoryYPHORA SANGUINIPENNIS, Sp. NOV. 


Black, the tibize metallic greenish ; head with one, thorax with 
two obscure fulvous spots at the sides, the latter closely punctured ; 
elytra reddish fulvous, finely and irregularly punctured, the 
interstices minutely aciculate. 

Length 14 millim. 

Oblong, parallel, the greatest convexity at the base of the 
elytra; the head very finely and closely punctured, greenish black, 
with a triangular fulvous spot at the vertex ; antennz extending 
to the base of the elytra, bluish black, the terminal joints gradu- 
ally and strongly widened, but slightly longer than broad ; thorax 
twice as broad as long, the sides flattened, the anterior angles 
not produced, the disc closely, finely, and somewhat rugosely 
punctured at the sides, much more sparingly so at the disc, the 
latter blackish, with a small fulvous spot at each side; elytra 
scarcely wider at the base than the thorax, closely, irregularly, and 
vather finely punctured, the interstices finely aciculate ; meso- 
sternal process rather short and stout. 

Hab. Provinz Goyaz, Brazil. (A single specimen.) 


DorYPHORA STAUDINGERI, Sp. nov. 


Greenish black, the last four joints of the antennz pale flavous ; 
thorax minutely and closely punctured; elytra closely punctured 
in double or treble rows, flavous, a broad transverse band at the 
base, a narrower one below the middle, the suture, and numerous 
small spots below the second band greenish: black. 

Length 11 millim. 

Elytra with the greatest height at the base, from there to the 
apex deflexed; head finely punctured, opaque, greenish black, 
labrum testaceous ; antennze extending nearly to the middle of the 
elytra, the basal two or three joints below and the terminal four 
pale yellow, the latter widened, longer than broad, the other 
joints black; thorax nearly three times broader than long, the 
sides rounded anteriorly, the anterior angles produced into a point, 
the surface closely and finely punctured, Jess closely so at the disc, 
the sides with a small fovea, the colour similar to that of the head ; 
scutellum shining, blackish ; elytra strongly punctured in closely 
approached treble rows, the margins of the bands irregularly 
notched, both bands of oblique direction, the first one placed 
immediately below the base, double the width of the second one 
which is situated immediately below the middle, both bands not 
quite extending to the lateral margins, the second one gradually 
dissolved into numerous small spots which fill out the “posterior 


1903. | SOUTH-AMERICAN COLEOPTERA. 43 


portion of the elytra; the suture likewise greenish black; the 
epipleure blackish ; the mesosternal process long and stout. 

Hab. Colombia. (A single specimen.) 

At once to be separated from other nearly similarly marked 
species by the colour of the antenne and other details. 


DoryPHORA STERNALIS, Sp. Nov. 


Greenish black ; thorax narrow, subquadrate, extremely closely 
and distinctly punctured; elytra geminate punctate-striate, 
flavous, with the suture and three narrow elongate stripes on 
each greenish ; mesosternal process extremely long. 

Length 11 millim. 

Head rather closely punctured, labrum fulvous, stained with 
black; antennz with the terminal joints twice as long as broad, 
black ; thorax twice as broad as long, the lateral margins perfectly 
straight from the middle downwards, almost slightly concave, the 
anterior angles pointed; the disc broadly flattened at the sides, 
extremely closely punctured, less crowded at the middle, the 
interstices slightly wrinkled; elytra distinctly wider at the base 
than the thorax, strongly geminate punctate-striate near the 
suture, closely and irregularly punctured at the sides, flavous, the 
suture and three posteriorly connected stripes, which are wider 
than the spaces separating them, metallic greenish, the outer 
interstice near the lateral margins of double the width of the 
others; the mesosternal process extremely long, extending to the 
head. 

Hab. Ecuador. (A single specimen.) 

This species has the longest mesosternum of any Doryphora I 
am acquainted with ; it is very pointed and slightly curved. 


DoryPHORA NIGROVIOLACEA, Sp. Nov. 


Metallic greenish black below, above blackish purplish, sub- 
opaque; thorax obscure greenish, nearly impunctate; elytra 
extremely finely punctured, with some obsolete more regular rows 
of punctures here and there. 

Length 11 millim. 

Head impunctate; antennz extending beyond the base of the 
elytra, black, the basal two joints fulvous below, terminal joints 
widened, longer than broad; thorax rather more than twice as 
broad as long, the anterior angles pointed, the surface entirely 
impunctate, of silky appearance; elytra very finely and sparingly 
punctured, with some punctures placed in distant rows; meso- 
sternal process moderately long. 

Hab. Venezuela? (A single specimen.) 

This is a very sombre-coloured species, of purplish black, with 
very finely punctured elytra, of which the rows can only be seen 
if examined carefully; 1 know of no other similarly sculptured 
species. The greatest height of the elytra is at the base if the 
insect is examined sideways. 


44 MR. MARTIN JACOBY ON NEW [May 12, 


DoRYPHORA VIGINTIPLAGIATA, Sp. NOV. 


Black, above flavous, head and the disc of the thorax greenish 
black, the latter extremely minutely punctured ; elytra strongly 
and semiregularly punctured, each elytron with ten obscure zeneous 
elongate spots (4, 3, 3) placed transversely below the base and 
below the middle. 

Length 12 millim. 

Oblong, the greatest convexity placed at the middle of the 
elytra; the head sparingly and finely punctured, with a small 
flavous spot at the vertex; antennz nearly extending to the base 
of the thorax, black, the lower four joints flavous below, terminal 
joints distinctly widened ; thorax more than twice as broad as 
long, the sides rounded anteriorly, the angles acutely pointed, the 
surface flattened at the sides, exceedingly finely and closely 
punctured ; the dise, in shape of a large transverse patch, greenish 
black, the anterior and lateral edges of the patch sinuate, the 
rest of the surface flavous; scutellum black; elytra closely and 
strongly punctured in irregular rows, which are slightly arranged 
in pairs near the suture, flavous, with ten elongate blackish-eeneous 
spots arranged in three transverse rows, one below the base 
consisting of four spots, of which one occupies the shoulders, the 
second row below the middle composed of three spots, the outer 
one of which is strongly transverse in shape, and the third row of 
three small spots near the apex. 

Hab. Espirito Santo, Brazil. (A single specimen.) 

This species might easily be mistaken for D. fasciato-maculata 
St., which it resembles almost entirely in coloration and pattern, 
but is much larger, the thorax is much more transverse and is 
devoid of the small spot placed at each side of the larger patch as 
in the above-named species, the elytra have the punctation less 
regularly and less distinctly geminate. In all specimens of D. 
Fasciato-maculata I have seen the thoracic central patch extends 
quite to the anterior margin; in the present insect the latter is 


flavous like the sides, in both species the suture is narrowly 
eeneous as well. 


DoRYPHORA LATERALIS, sp. Noy. 


Olivaceous green, more or less stained with testaceous below, 
antenne entirely testaceous ; thorax very minutely and sparingly 
punctured ; elytra strongly punctate-striate, the extreme basal 


margin and a sutural stripe dark fulvous, the ninth interstice 
obscure flavous. 


Length 15 millim. 

Subglobular, deflexed posteriorly, the head impunctate, the 
labrum testaceous ; antennz slender, the terminal joints scarcely 
thickened, entirely testaceous; thorax twice and a half broader 
than long, the sides straight at the base, rounded anteriorly, the 
anterior angles broad and produced but not mucronate, the disc 
convex, dull green, subopaque, with some minute punctures, the 


1903.] SOUTH-AMERICAN COLEOPTERA. 45 


extreme margins pale; scutellum greenish brown; elytra very 
regularly punctate-striate, the punctures deep and closely placed, 
the interstices shghtly convex, impunctate, olive-green, the suture 
piceous or dark fulvous, the ninth interstice obscure flavous, costate 
posteriorly ; below greenish, the tarsi testaceous more or less 
mesosternal process stout and straight. 

Hab. Espirito Santo, Brazil (/ruhstorfer). 

Much larger than D. dejeani Germ. or the allied species, and 
at once distinguished by the piceous sutural stripe. I know only 
a single specimen of this species, which is not an immaculate 
variety of D. piceo-maculata Jac., on account of the totally differ- 
ent elytral sculpture. 


DorYPHORA SEXPLAGIATA, Sp. nov. 


Dark violaceous; antennz black; thorax extremely finely and 
closely punctured, the angles fulvous; elytra scarcely more 
strongly punctured, the punctures partly arranged in rows, the 
interstices finely aciculate, each elytron with three flavous spots, 
one at the base, one at the margin at the middle, and the third 
near the apex. 

Length 12 millim. 

Below black with a bluish gloss, above violaceous; head with a 
few fine punctures; antenne black, the terminal joints sub- 
quadrately widened, one-half longer than broad; thorax rather 
flat, the sides straight, the anterior angles slightly tuberculiform, 
fulvous, the surface extremely finely and rather closely punctured ; 
scutellum black, impunctate; elytra with the greatest convexity 
at the base, deflexed at the posterior portion, very finely punc- 
tured in closely approached, irregular rows, violaceous, with a 
transverse spot near the scutellum, another of oblique shape at 
the middle of the lateral margins, and a third, subtriangular spot 
near the apex, bright flavous; mesosternal process long and straight. 

Hab. Peru. (Two specimens.) 

This species must not be confounded with D. secemaculata Jac., 
also from Peru, in which the elytra are either greenish or brown 
and are strongly punctate; the spots, although similarly placed, 
are also of different shape. 


DorYPHORA CYANEOFASCIATA, Sp. Nov. 


Dark fulvous below; head and thorax fulvo-zneous, minutely 
and closely punctured, opaque; elytra regularly and closely punc- 
tate-striate, testaceous, with a broad transverse oblique band 
below the base and a narrower band below the middle, as well as 
the suture, metallic blue, apical portion with minute bluish spots. 

Length 12 millim. 

Head minutely punctured, obscure eneous ; labrum and palpi 
fulvous; antenne blackish, the lower three or four joints fulvous, 
terminal joints distinctly widened ; thorax coloured like the head, 
twice as broad as long, the sides broadly flattened, rounded from 
the middle to the anterior angles, the latter mucronate, produced 


46 MR. MARTIN JACOBY ON NEW [May 12, 


in front of the eyes, the surface very minutely and evenly punc- 
tured; scutellum obscure fulvous; elytra strongly and closely 
punctate-striate at the anterior portion, much more finely so 
towards the apex, where the punctation 1s less regular, of pale 
flavous ground-colour with two metallic blue bands, the first 
below the base and of double the width of the second one, 
extending obliquely to the lateral margins where it is much 
narrowed, the second band immediately below the middle and 
running parallel with the first, the suture likewise narrowly 
margined with blue, and the space between the second band and 
the apex filled with numerous small bluish spots; elytral epi- 
pleure, the underside, and legs dark fulvous with zneous gloss; 
mesosternal process very long and stout. 

Hab. Peru. (A single specimen.) 

In general markings this species resembles D. eneofasciata, 
but the elytral bands extend quite to the margins and are of 
different shape and size, and the anterior portion of the elytra is 
devoid of spots; the sculpture also is quite different. The same 
remarks apply also to the species when comparing it with 
D, bifasciata Fab. 


DoryYPHORA FLAVOFASCIATA, Sp. Nov. 


Fulvous, the intermediate joints of the antenne black; thorax 
remotely and strongly punctured ; elytra strongly punctate-striate, 
a spot near the scutellum, another near the shoulders, an oblique 
subsutural and another submarginal longitudinal band, flavous. 

Length 12 millim. 

Head impunctate ; antenne extending beyond the base of the 
thorax, the lower five joints fulvous, the terminal two joints 
flavous, the others black, terminal joints slightly thickened, 
scarcely longer than broad ; thorax three times broader than long, 
the sides strongly flattened, the lateral margins straight at the 
base, rounded in front, anterior angles acute, the disc rather 
strongly and remotely punctured; elytra regularly convex (highest 
before the middle), finely but rather regularly punctured in closely 
approached rows, more strongly punctured at the base, with an 
elongate spot near the scutellum, a submarginal band to which 
another spot is joined above the shoulders, and another subsutural 
band abbreviated anteriorly and posteriorly, flavous; below entirely 
fulvous, mesosternal process very long and nearly straight. 

Hab. Colombia. (A single specimen.) 

Of nearly similar coloration as D. chevrolati St., but much larger, 
the thorax broader and flattened, and the elytral punctation 
quite different. 


DoRYPHORA ZNEOFASCIATA, Sp. nov. 


Greenish black below; thorax obscure neous, very finely 
punctured, the lateral margins flavous; elytra irregularly punc- 
tured, pale flavous, with numerous small zneous spots and two 


1903. | SOUTH-AMERICAN COLEOPTERA, AT 


transverse metallic green bands, one below the base, the other, of 
oblique shape, at the middle. 

Length 10-11 millim. 

Head minutely and closely punctured, obscure eneous, opaque ; 
antenne short, black, the basal two joints below and the apex of 
the last flavous; thorax about twice as broad as long, the sides 
straight at the base, rounded anteriorly, the anterior angles 
mucronate, extending to the end of the eyes, the extreme lateral 
margins flavous, the dise coloured like the head, very closely and 
finely punctured, the punctation even throughout ; scutellum 
black; elytra pale testaceous, closely, strongly, and irregularly 
punctured, covered with numerous, closely placed, green, small 
spots of irregular shape and size, below the base a narrow trans- 
verse metallic green band extends across the suture but not to the 
lateral margins, another similar band of strongly oblique direction 
is placed at the middle and is slightly abbreviated at each end; 
mesosternal process long and robust. 

Hab. Peru. (‘Two specimens.) 

Several similarly marked species are known, but they are either 
of much larger size or devoid of the thoracic devote margins ; in 
D. bifasciata Fab. and D. maculata Oliv. the anterior elytral 
bands are curved upwards and the suture is likewise provided with 
an eeneous stripe; the elytral epipleure in the present species are 
entirely flavous. 


DoryYPHORA FASCIATIPENNIS, Sp. Nov. 


Greenish eeneous, the thorax remotely punctured ; elytra closely 
geminate punctate-striate, a subsutural and marginal longitudinal 
narrow stripe, connected by a transverse one below the middle, 
flavous. 

Length 8 millim. 

Head finely and sparingly punctured ; antenne greenish eneous, 
the terminal joints moderately widened, twice as long as broad ; 
thorax sparingly punctured at the middle, the sides impunctate ; 
elytra evenly convex, strongly and closely geminate punctate- 
striate, the first two rows near the suture, single, the flavous 
bands narrow, the sutural one placed on the third interspace, the 
lateral one close to the margin which remains of the ground-colour, 
both bands are joined at the apex and connected below the middle 
by another short transverse band; the mesosternal process very 
short, claw-joints unarmed. 

Hab. Balzapamba, Ecuador. 

A small beetle, resembling in its markings several species of the 
genera Zygogramma and Cosmogramma, but a true Doryphora. 


STILODES FLAVOFASCIATA, Sp. Nov. 

Black, the labrum, palpi, and basal joints of the antenne flavous ; 
thorax opaque, nearly impunctate; elytra similarly sculptured, 
black, a narrow transverse and finely divided band at the base, 


48 MR. MARTIN JACOBY ON NEW [May 12, 


another below the middle, and a third near the apex, bright 
flavous. 

Length 10 millim. 

Head impunctate, black, opaque and of silky appearance ; 
labrum flavous ; antenn extending to the base of the elytra, the 
lower joints flavous, the terminal three or four joints piceous, one- 
half longer than broad; thorax strongly transverse, the sides 
straight, the anterior angles not produced, the surface sculptured 
like the head, with a few minute punctures ; scutellum black, 
shining; elytra with extremely feeble rows of punctures, which 
are entirely obsolete below the middle, black, with three narrow 
transverse bright yellow bands, the first at the base and divided 
by a very narrow perpendicular black stripe at the shoulders, the 
second narrow band below the middle, slightly constricted near 
the sides, the third short and broader band of oblique direction 
near the apex, elytral epipleur flavous ; underside and legs black, 
shining, the trochanters dark fulvous; metasternal epipleure 
nearly impunctate. 

Hab. Espirito Santo, Brazil. 

Closely allied to S. transversofasciata Jac., but of entirely 
different sculpturing and with the elytral bands of different shape. 


STILODES FLAVILABRUM, Sp. Nov. 


Blackish blue, the labrum flavous; head impunctate; thorax 
finely punctured, opaque; elytra strongly punctate-striate, flavous, 
a broad band at the base, another below the middle, not extending 
to the lateral margins, and the apex, more or less, bluish black. 

Length 10 millim. 

Of rather elongate shape, pointed at the apex, the head opaque, 
impunctate, greenish; the labrum flavous; antennz rather 
slender, blackish, the lower three joints flavous below ; 
thorax about one-half broader than long, the sides straight, the 
surface opaque, of silky appearance, finely and subremotely punc- 
tured, greenish black; scutellum black, shining; elytra with 
regular and deep rows of punctures, the punctures much finer on 
the flavous portion and not always evenly placed, the sutural and 
lateral margins, two broad transverse bands before and below the 
middle, and the apex to a small extent, bluish-black; underside 
and legs shining blue-black, the last abdominal segment with a 
small fulvous spot at each side; the metasternum impunctate at 
the sides. 

Hab. Sao Paulo, Brazil. 

Like the preceding species closely allied to S. transversofasciata 
Jac., but with both elytral bands laterally interrupted, the thorax 
very finely and the elytra much more strongly punctured, and the 
general size much larger. 


STILODES FRUHSTORFERI, Sp. Nov. 


Black, the antenne fulvous; thorax sparingly punctured, with 
two flavous spots anteriorly ; elytra punctate-striate, a spot near 


1903.] SOUTH-AMERICAN COLEOPTERA, 49 


the scutellum, another at the lateral margins before the middle, 
and a transverse spot near the apex, flavous. 

Length 9-10 millim. 

Head sparingly punctured, black, opaque; labrum and antennee 
fulvous, the terminal joints transversely widened; thorax strongly 
transversely convex, the sides rounded, the anterior angles not 
produced, the surface sparingly punctured, black, opaque ; scu- 
tellum impunctate, black; elytra finely punctate-striate, pointed 
posteriorly, black, opaque, with an oblong spot near the scutellum, 
a subquadrate spot at the lateral margin before the middle, and 
another transverse one near the apex, flavous; underside and 
legs black, shining, apex of the tibie clothed with fulvous 
pubescence. 

Hab. Espirito Santo, Brazil (fruhstorfer). 

Of this distinctly marked species I have six specimens which I 
received from Mr. Fruhstorfer. 


STILODES ECUADORIENSIS, Sp. nov. 


Testaceous, the head greenish black; thorax sparingly punc- 
tured; elytra regularly punctate-striate, the punctures placed on 
narrow black stripes, the fourth stripe abbreviated ; underside 
greenish piceous. 

Length 10 millim. 

Head metallic greenish, finely and sparingly punctured ; labrum 
flavous; antenne slender, greenish black, the basal joint, the 
outer ones below and the two or three terminal joints entirely 
flavous; thorax twice as broad as long, the sides perfectly straight, 
the posterior angles acute, the surface remotely and strongly 
punctured, testaceous ; scutellum metallic green ; elytra with rows 
of deep punctures, the latter not always regularly placed, the 
second, third, and fourth, and the sixth, seventh, and eighth rows 
metallic greenish black in shape of narrow stripes, the fifth row 
of this colour near the apex only, the others all more or less 
interrupted posteriorly ; underside stained with blackish; legs 
entirely testaceous. 

Hab. Santa Inéz, Ecuador (Haensch). 

Of this neatly marked species two specimens are contained in 
my collection. 


STILODES (?) GEOMETR#, sp. nov. 


Flavous, two spots at the head, the dise of the thorax, and the 
breast black; thorax finely punctured; elytra closely and irregu- 
larly punctured, flavous, with a narrow oblique olivaceous black 
band from the shoulders to the middle and another at the sides to 
the apex, both bands joined at the suture and at the shoulders, 

Length 10 millim. 

Head closely punctured, black, this colour divided into two 
large spots by a narrow central flavous stripe, the space in front 
of the eyes and the labrum likewise flavous,; antennz entirely 

Proc. Zoou, Soc.—1903, Vou. II. No. LV. 4 


50 MR. MARTIN JACOBY ON NEW | May 12, 


flavous, the terminal joints strongly transverse ; thorax more than 
twice as broad as long, the sides straight, the anterior angles 
produced forwards, the dise finely and closely punctured near the 
base, less closely so at the sides and anteriorly, the first-named 
portion flavous, the disc black, this colour in shape of a sub- 
quadrate patch the sides of which are concave, the anterior edge 
slightly divided at the middle ; scutellum black; elytra somewhat 
widened posteriorly, closely, irregularly, and rather finely pune- 
tured throughout, flavous, with two oblique narrow blackish 
stripes joined at the shoulders, the first extending below the 
middle and connected down the suture with the lateral stripe, 
the flavous ground-colour therefore forms a large triangular patch 
at the anterior portion and another one at each side, the suture 
is likewise black for a short distance at the base; the breast 
black, the abdomen and legs flavous ; metasternum scarcely raised, 
claw-joint bidentate. 

Hab. Marcapata, Peru. 

This interesting species scarcely fits into any of the groups as 
arranged by Stal, on account of the entirely irregularly-punctured 
elytra, at the same time the other characters agree with Stilodes ; 
from any of the species the present one is distinguished by the 
system of coloration and the rather curious pattern of the elytra. 
I received a single specimen from Herr Bang-Haas, of Dresden. 


DEUTEROCAMPTA PUNCTICOLLIS, Sp. NOV. 


Fulvous, the apical joints of the antenne, the knees, and tarsi 
black ; thorax strongly and closely punctured; elytra regularly 
punctate-striate, fulvous, the sutural margins narrowly black. 

Length 8 millim. 

Head finely punctate, fulvous; antenne black, the basal two 
joints fulvous, terminal ones transversely widened ; thorax of the 
same shape as that of D. scwtellata, but more closely punctured, 
the punctures at the sides scarcely stronger than those at the disc ; 
scutellum fulvous; elytra regularly punctate-striate, each elytron 
with 10 rows of punctures, the subsutural row abbreviated before 
the middle, the extreme sutural margins black ; the knees and the 
tarsi black, the rest of the under surface fulvous. 

Hab. Brazil ¢ 

Closely alhed to D. scutellata, of which it may possibly be a 
variety, but the thorax more closely and evenly punctured, the 
scutellum and underside fulvous, the legs also differently coloured. 


DEUTEROCAMPTA SCUTELLATA, Sp. Nov. 


Underside and legs, the terminal joints of the antenne, and 
the scutellum black, above fulvous ; thorax strongly punctured at 
the sides, elytra regularly and strongly punctate-striate. 

Length 8 millim. 

Head impunctate, fulvous; antennez short, black, terminal 
jomts transverse, broader than long, the lower two joints fulvous; 


1903.] SOUTH-AMERICAN COLEOPTERA. 51 


thorax slightly narrowed anteriorly, the sides obliquely rounded, 
the surface strongly and closely punctured at the sides, sparingly 
so at the middle; scutellum black; elytra with very regular and 
deep rows of punctures, the latter evenly placed, the subsutural 
row not extending to the middle, the colour like that of the head 
and thorax; underside and legs black, the metasternal parapleure 
strongly punctured, third tarsal joint simple. 

Hab. Brazil. 

Ditters from D. luteola St. in the black scutellum and regular 
elytral punctation, from PD. nigrimana St. in the unspotted 
thorax, colour of the underside, &e.; also in the same way from 
D. obscurimana St. D. flavida St. has the elytra geminate- 
punctate. I know only of a single specimen of this species, 
contained in my collection without detailed locality. 


° 
DESMOGRAMMA STALI, Sp. nov. 


Dark fulvous, the thorax with a transverse obscure greenish- 
eeneous band, remotely punctured ; elytra strongly punctate-striate, 
the third and the ninth interstice, as well as the basal margin, 
flavous, the other interstices alternately fulvous and greenish 
eeneous ; underside greenish, legs fulvous. 

Length 8 millim. 

Head sparingly punctured, fulvous, the space between the eyes 
greenish eneous; antenne with the terminal five joints strongly 
widened, black, the lower seven joints and the apex of the last 
one fulvous, third joint very elongate; thorax more than twice 
as broad as long, the sides straight, the surface remotely and 
rather strongly punctured, dark fulvous, with an obscure trans- 
verse greenish-zneous band placed at the anterior margin and 
not extending to the sides, to this band another spot is joined 
at the middle and extending to the base ; scutellum flavous; elytra 
with regular rows of deep punctures, dark fulvous, the second, 
fourth, sixth, eighth, and tenth interstice greenish eeneous, the 
third and ninth as well as the basal margin bright flavous, joined 
at the apex; below greenish, the margins of the abdominal 
segments and the legs fulvous, the femora stained with eneous : 
claw-joint simple, prosternum raised anteriorly, metasternum 
truncate. 

Hab. Brazil. 

Of this species, quite distinct in its system of coloration from 
any of its allies, I have asingle specimen without detailed locality ; 
the submarginal flavous stripe is slightly curved at the middle 
and has a single row of punctures posteriorly ; the elytral epipleure 
are fulvous. 


DESMOGRAMMA PERUANA, sp. nov. 
Black, head and thorax finely punctured; elytra regularly 
punctate-striate, flavous, a sutura band and the lateral margins 


more narrowly black. 
4% 


52 MR, MARTIN JACOBY ON NEW [ May 12, 


Var. Head and thorax fulvous. 

Length 7 millim. 

Head scarcely perceptibly punctured, with a slight greenish 
tint ; antennz black, the basal joint flavous below, terminal joints 
gradually widened, the last joint more elongate ; thorax with 
straight sides, the anterior angles pointed, the surface very finely 
and sparingly punctured ; scutellum blackish, elongate; elytra 
wider at the base than the thorax, finely and very regularly 
punctate-striate, flavous, the suture (to the extent of the third 
row) and the last marginal interstice black; underside and legs 
black, claw-joint simple, prosternum tuberculiform and raised 
anteriorly, metasternum also raised and truncate in front. 

Hab. Marcapata, Peru. 

The elytral punctures are extremely closely placed, and the 
sutural band is rather abruptly truncate immediately below the 
base and greatly narrowed round the scutellum, gradually so 
towards the apex; the marginal stripe does not quite touch the 
sutural one at the apex, and extends to the base but not to the 
basal margin. Three specimens are contained in my collection, 
one of which has the head and thorax fulvous. 


DESMOGRAMMA INCLUSA, Sp. Nov. 


Aineous, the labrum flavous, the antennz and tarsi dark 
fulvous ; thorax with the anterior margin and two oblique stripes 
flavous; elytra strongly punctured anteriorly, flavous, the base 
with a broad tranverse zeneous band, including a flavous spot. 

Length 12 millim. 

Head zeneous, with a few fine punctures; labrum flavous, palpi 
and antenne fulvous, the latter with strongly-dilated terminal 
joints ; thorax with the lateral margins straight, the disc very 
sparingly punctured, greenish eneous, this colour divided into 
three spots by two oblique narrow flavous stripes which join the 
similarly-coloured anterior margin, the darker portions are in the 
shape of a subquadrate spot at each side and a larger ~-shaped 
spot at the middle; elytra slightly wider at the base than the 
thorax, strongly punctate-striate at the base, more finely so and 
irregularly at the lower portion, flavous, the extreme sutural 
margins and a broad transverse band at the base, not extending 
to the lateral margins and including a flavous spot near the seu- 
tellum, greenish eneous; elytral epipleure flavous; underside 
zneous, legs fulvous, prosternum truncate and raised anteriorly. 

Hab. Rio Janeiro, Brazil 2 

_ The thorax in colour and pattern agrees with that of D. par- 
tite St., but the sculpturing and markings of the elytra are 
cae the markings somewhat resembling those of D. fenes- 
rata St. 


DESMOGRAMMA DECEMPUSTULATA, Sp. Nov. 


Black, the antenne, tibie, and tarsi fulvous; head and thorax 


1903.] SOUTH-AMERICAN COLEOPTERA. 53 


minutely punctured, elytra strongly punctate-striate, each with 
five flavous or fulvous round spots, 2, 2, 1. 

Length 10 millim. 

Head with a greenish tint, opaque, minutely punctured; labrum 
and palpi fulvous; antenne extending below the base of the 
elytra, fulvous, terminal joints gradually transversely widened ; 
thorax about twice as broad as long, the sides straight, the surface 
coloured like the head, black, with a greenish tint, finely but not 
very closely punctured ; scutellum black ; elytra wider at the base 
than the thorax, black, rather strongly and regularly punctate- 
striate, the punctures arranged in double rows at the sides, each 
elytron with five flavous spots, of these, two are placed at the base 
(on the shoulder and near the scutellum), the outer one being 
larger than the other, two at the middle, placed transversely, and 
one at the apex; underside blackish, the femora near the base, 
the tibie almost entirely, and the tarsi fulvous, the last abdominal 
segment with a fulvous spot at each side, the prosternum tubercu- 
lately truncate anteriorly. 

Hab, Espirito Santo, Brazil. 

I received a single specimen of this species from Mr, Fruh- 
storfer. 


CosMOGRAMMA UNICINCTA, Sp. nov. 


Black, with a slight bluish gloss; head and thorax finely and 
sparingly punctured; elytra regularly punctate-striate, black, 
with a violaceous tint, the basal margin and a sublateral narrow 
band bright flavous. 

Length 6 millim. 

Head finely and rather closely punctured ; antenne black, the 
terminal joints thickened but rather long, the lower two joints 
flavous below; thorax with the anterior angles produced into a 
small tooth, the sides straight, the surface finely and sparingly 
punctured, the punctures not stronger at the sides; scutellum 
black; elytra with a slight purplish or violaceous tint, finely and 
regularly punctate-striate, the tenth interstice and the basal 
margin bright flavous, the last interstice and the epipleuree of the 
ground-colour ; claw-joint unarmed, the metasternum truncately 
produced. 

Hab. Peru. 

The single flavous lateral stripe separates this species from 
most others placed in this genus, from which it further differs in 
the non-dentate claws; C. patricia Erichs. is, however, of similar 
coloration, but dull black, opaque, and with scarcely perceptible 
elytral punctation. 


CosMOGRAMMA PERUANA, Sp. Nov. 


Black, the thorax closely and strongly punctured; elytra 
opaque, finely punctate-striate, the third and the last interstice, 
as well as the basal margin, golden yellow, the margina] band 
with a row of dark punctures. 


o4 MR. MARTIN JACOBY ON NEW {May 12, 


Length 7 millim. e 

Head finely punctured, black, the last joint of the palpi larger 
than the preceding one; antenne black, the lower two joints 
fulvous below, the terminal joints wanting; thorax only about 
one-half broader than long, the sides straight, the anterior angles 
almost rectangular, not produced, the disc rather convex, closely 
and rather strongly punctured, black, shining; elytra opaque, 
with fine but distinct rows of punctures, which are less regular at 
the sides, black, with two narrow bright flavous bands placed on 
the third and last interstice and joined at the apex, the lateral 
band with a single row of punctures and gradually narrowed 
posteriorly, where the black margin assumes a wider shape than 
at the base, the bands are connected at the base by a narrow 
transverse stripe; metasternum slightly transversely raised, the 
claw-joint dentate, claws separated. 

Hab. Peru. 

The present species differs from its allies placed in Cosmo- 
gramma by the distinctly punctate-striate elytra, the rather 
narrow and closely-punctured thorax, and the colour of the elytral 
bands; CO. fulvocincta St. has fulvous antenne and legs, similarly 
coloured elytral bands, and a differently sculptured thorax. 


CALLIGRAPHA ANNULATA, sp. nov. 


Metallic dark green, the basal joints of the antenne fulvous; 
thorax strongly punctured at the sides only; elytra dark fulvous, 
finely and irregularly punctured, a sutural narrow band, two spots 
at the base, three before the middle, a ring-shaped mark below 
the latter, and two other small spots at the apex metallic green. 

Var. One or more spots at the anterior portion wanting, the 
ring-shaped mark interrupted. 

Length 6 millim. 

Of narrow and elongate shape, metallic dark green, the head 
with a few fine punctures, the antenne with strongly thickened 
terminal joint, black, the lower three joints fulvous, the basal 
joint «neous above; thorax rather short, metallic green, strongly 
punctured at the sides only, the dise nearly impunctate ; scutellum 
ceneous ; elytra finely and irregularly punctured except near the 
suture, where two rows of punctures accompany the metallic- 
green stripe; the latter is gradually and slightly narrowed 
towards the apex ; of the spots one is placed on the shoulders, 
another near the scutellum, both of elongate shape; these are 
followed by three small spots (when present) placed triangularly ; 
below the middle is an irregular metallic-green ring, sometimes 
interrupted, and at the apex two small spots are situated. 

Hab, Bolivia. 

I know of no other similarly marked species which is less 
convex and rounded than most of its congeners ; in the variety, © 


of the three spots placed in a triangle, only the outer one is 
present. : 


1903. ] SOUTH-AMERICAN COLEOPTERA. 59 


PROSICELA MACULATA, sp. noy. 


Metallic blue or green ; thorax finely and irregularly punctured ; 
elytra flavous, strongly punctured in very irregular rows, the 
suture narrowly, a spot on the shoulders and another below the 
middle, metallic blue. 

Var. The humeral spot absent. 

Length 8-10 muillim. 

Head remotely punctured, antenne nearly extending to the 
middle of the elytra, black, the last joint elongate; thorax 
irregularly and finely punctured, more closely so near the base ; 
scutellum smooth; elytra wider at the base than the thorax, 
strongly punctured, the rows irregular and often doubled, the 
suture (to the extent of the first row of punctures) metallic blue, 
gradually narrowed towards the apex, the shoulders with a small 
blue spot, the disc below the middle with another short, somewhat 
obliquely placed, elongate spot. 

Hab. Marcapata, Peru. 

Closely allied to P. flavipennis Erichs., but with more strongly 
and more irregularly punctured elytra bearing two spots each, 
which are absent in Erichson’s species. 


PROSICELA INORNATA, Sp. Nov. 


Metallic dark green, the antenne black, elongate; thorax very 
finely and subremotely punctured ; elytra pale fulvous, finely and 
rather closely geminate punctate-striate. 

Length 10 millim. 

Very closely allied to the preceding species, but the thorax 
more finely punctured ; the elytra geminate-punctate and without 
any metallic sutural stripe or spots; the sides of the thorax are 
perfectly straight. 

Hab. Prov. Huallaga, Peru. 

Sunilar in coloration to P. simplicipennis Jac., but the thorax 
much more transverse, and the elytra more strongly punctured 
and in double rows. 


ZYGOGRAMMA INTERSTITIALIS, sp. nov. 


Greenish eneous ; thorax very strongly punctured at the sides, 
less so at the middle, lateral margins flavous; elytra very strongly 
punctate-striate, interstices finely punctured, basal margin, third 
and last interstice flavous, joined at the apex. 

Var. Antenne and tibiz more or less fulvous. 

Length 6 millim. 

Head finely punctured, the labrum fulvous; antenne with the 
lower four or five joints fulvous, the rest black, strongly trans- 
versely widened; thorax with flavous lateral margins, very 
strongly but sparingly punctured at the sides, somewhat closely 
but less strongly so at the disc; elytra with deep and rather 
remotely placed punctures regularly arranged, the sutural rows 
finer, the third and tenth interstices flavous, the latter with a deep 


56 MR. MARTIN JACOBY ON NEW [May 12, 


row of punctures, joined to the third row at the apex, the basal 
margin likewise flavous, the rest of the interstices finely punctured 
and transversely wrinkled; claws jomed at the base, the third 
joint dentate at the apex. 

fab. Brazil. 

I know of no other species of Zygogramma with flavous thoracic 
margins in which the elytra are so strongly punctured or the 
interstices finely so; the subsutural flavous stripe gradually 
approaches the suture towards the apex: in the variety the 
antenne and legs are fulvous, and the femora are spotted with 
zeneous, but other differences I cannot find. The punctation of the 
thorax varies much in depth and number of punctures, and the 
elytral wrinkles are also more or less distinct in different specimens. 


ZYGOGRAMMA ARGENTINENSIS, Sp. Nov. 


Dark greenish, the antenne black ; thorax minutely punctured, 
the sides more strongly so; elytra with three narrow fulvous 
vitte, the discoidal ones joined at the apex, the marginal one 
with a row of punctures, the subsutural rows geminate. 

Length 9 millim. 

Larger than any of the species placed in this genus described 
by Stal, of an opaque greenish colour, the head minutely 
punctured, the terminal joint of the palpi larger than the pre- 
ceding one; antennze black, the basal two joints Havous below,~ 
terminal jomts widened ; thorax very finely punctured, the sides 
more strongly so; scutellum impunctate; elytra regularly 
punctate-striate, the subsutural row double (the first row con- 
sisting of a few punctures only), the third, fifth, and ninth 
interstices flavous, the discoidal bands joined at the apex, the 
fourth interstice rather wider than the others, the marginal 
flavous vitta with a row of piceous punctures, elytral epipleurz 
greenish ; claw-joint dentate, claws joined at the base. 

Hab. Salta, Argentine Republic. 

The larger size, greenish coloration, and the wider fourth inter- 
stice separates this species from its allies; the flavous bands are, 
as usual, joined together by the similarly coloured flavous basal 
margin. ‘There are three specimens before me. 


ZYGOGRAMMA CURVATO-LINEATA, Sp. nov. 


Dark fulvous, the breast greenish eneous ; thorax finely and 
sparingly punctured ; elytra flavous with a metallic lustre, each 
elytron with the following black markings: three short stripes at 
the base, the inner two joined at the apex, a semicircular band 
below the middle, another one within the outer band, and a 
V-shaped mark near the apex; legs fulvous. 

Length 8 millim. 

Head impunctate, fulvous; antennze with the terminal joints 
moderately thickened, longer than broad, entirely fulvous; thorax 
very sparingly and finely punctured at the base and sides, fulvous; 
scutellum black; elytra punctate-striate near the suture, finely 


1903.] SOUTH-AMERICAN COLEOPTERA. ° 57 


and irregularly punctured at the flavous portion, the dark 
markings also surrounded or bounded by deeper punctures, the 
suture, to the extent of the second row of punctures, black, in 
shape of a narrow, anteriorly widened stripe which does not quite 
extend to the base; of the dark markings three short longitudinal 
slightly curved stripes are placed at the base, of which the inner 
two are joined at the apex, these stripes do not extend to the 
basal margin but the inner ones nearly reach the middle, they 
are followed by a crescent-shaped narrow band at the sides, 
within which a smaller nearly ring-shaped mark is placed, near 
the apex another V-shaped spot is situated, at the middle of the 
lateral margins there is another smaller black spot, and the elytral 
epipleure are of the same colour; the abdomen and the legs are 
fulvous ; claws united at the base. 

Hab. Costa Rica. (In my collection and that of the British 
Museum.) 

The peculiar elytral markings differ from those of any other 
species of the genus; the flavous ground-colour has a golden hue, 
which is probably more brilliant in the living insect. 


ZYGOGRAMMA BRASILIENSIS, Sp. NOV. 


Reddish fulvous; thorax strongly punctured at the sides, 
minutely at the disc; elytra finely punctate-striate, black, the — 
third, fifth, and the ninth and tenth interstices, as well as the 
epipleuree, flavous, the two inner flavous stripes connected at 
the apex. 

Length 6 millim. 

Closely allied to 7. flavolorata Stal and Z. myops Stal, and to a 
few other species with flavous elytral stripes and epipleure, but 
at once distinguished from them by the fulvous head, thorax, and 
legs ; the head impunctate ; antennz short, fulvous, the apical five 
joints black, strongly transverse ; thorax strongly transverse, closely 
and strongly punctured at the sides, fulvous, the basal margin 
sometimes piceous; scutellum black; elytra finely and regularly 
punctate-striate, the basal margin and three narrow longitudinal 
stripes flavous, the discoidal ones joined near the apex, occupying 
the third and fifth interstices, the outer stripe with a row of 
punctures, the extreme lateral margins black, the epipleure 
flavous. 

Hab, Espirito Santo, Brazil. 

Three exactly similar specimens are contained in my collection. 


ZYGOGRAMMA BURMEISTERI, Sp. Nov. 


Greenish black ; thorax narrowed anteriorly, strongly punctured 
at the sides; elytra striate-punctate, the interstices slightly 
convex, the base of the first, the third and fifth entirely, and the 
tenth flavous. 

Length 7 millim. 

Head very minutely punctured ; antennz with short, transverse 
terminal joints, broader than long, black, the basal two or three 


58 ON NEW SOUTH-AMERICAN COLEOPTERA. [ May 12, 


joints fulvous ; thorax with the sides straight, obliquely narrowed 
anteriorly, greenish, the sides strongly and closely, the disc 
minutely punctured ; seutellum ovate, black; elytra with deep 
and regular strize which are closely and finely punctured, those 
near the suture impunctate, the basal margin and three longi- 
tudinal bands, as well as a short subsutural stripe, fulvous, the 
two inner stripes joimed at the apex and connected at the same 
place with the lateral stripe, which has a row of black punctures. 

Hab. Salta, Argentine Republi. 

Readily distinguishable from any other species by the anteriorly 
narrowed thorax, the convex elytral interstices, and the punc- 
tured strie (not punctured rows), also by the additional short 
subsutural flavous stripe at the first interstice near the base. 


ZYGOGRAMMA CRULEO-VITTATA, Sp. Nov. 


Dark fulvous, the breast and abdomen greenish piceous, apical 
joints of the antenne black; thorax strongly punctured at the 
sides only ; elytra with three dark blue and three flavous longi- 
tudinal bands each, the dark bands closely punctured, the flavous 
ones joined at the apex. 

Length 7 millim. 

Head finely and sparingly punctured, dark fulvous, labrum 
flavous, mandibles robust, strongly punctured and pubescent ; 
antenne black, the basal six joints fulvous, terminal joints 
strongly dilated, the last one oblong-ovate; thorax more than 
twice as broad as long, the sides straight, closely and strongly 
punctured, the disc very finely punctate; scutellum fulvous ; 
elytra with two regular rows of punctures near the suture, the 
latter and two longitudinal stripes dark blue, the discoidal stripe 
abbreviated posteriorly, the alternate interstices flavous, im- 
punctate, joined at the apex, the basal margin also flavous, the 
marginal flavous band with a single row of punctures, the elytral 


epipleurz geneous, except near the apex; legs fulvous, claws 
united at the base. 


Hab. Paraguay. 
The colour of the antenne and the blue elytral bands principally 
distinguish this species, which has its most nearly allied form 


probably in Z. 5-virgata Stal, from which it differs in the details 
given above. 


LEPTINOTARSA PARAGUAYENSIS, Sp. nov. 


Metallic green, the labrum and antenne fulvous; head and 
thorax impunctate, subopaque; elytra regularly punctate-striate, 
the interstices slightly convex and finely aciculate. 

Length 9 millim. 

Head entirely impunctate, of silky appearance, the labrum 
fulvous; the antenne with the terminal five joints widened, 
entirely fulvous; thorax with strongly rounded sides, rather 
convex, the angles not produced, the surface sculptured like the 
head, without a trace of punctation; scutellum shining;. elytra 


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1903. ] ON FISHES FROM RIO JANEIRO. . 59 


with the usual ten rows of extremely closely placed punctures, 
the sutural row short, the interstices feebly convex and finely 
transversely aciculate; the underside more metallic, the tarsi 
fulvous below, mesosternum scarcely raised, its epipleuree smooth ; 
claw-joint simple, tibiee with longitudinal sulcus. 

Hab. Paraguay. 

This species is placed in the genus Leptinotarsa on account of the 
sulcate tibize, which have the suleus extending to half their length ; 
the colour of the antenne and the slightly convex elytral inter- 
stices will easily distinguish the species. 


ELYTHROSPHZRA CUPREATA, Sp. nov. 

Ovate, pointed posteriorly, cupreous, variegated with metallic 
green; antenne black; thorax deeply and irregularly punctured ; 
elytra deeply foveolate punctate, the punctures arranged in rows, 
cupreous, the suture metallic green. 

Length 10 millim. 

Apterous; the head finely punctured, with a central fovea, the 
vertex metallic green, the lower portion reddish cupreous ; labrum 
and palpi black; antenne rather long, black, the terminal two 
joints elongate, thickened, the basal joint metallic green; thorax 
one-half broader than long, the sides straight, the disc deeply 
foveolate and partly confluently punctured, cupreous, with a 
metallic green band at each side; elytra very convex, widened at 
the middle, the apex pointed, each elytron with ten rows of fovez, 
regularly placed, the sutural and lateral margins green, the disc 
cupreous ; underside coloured like the upperside, the tibize metallic 
green, the tarsi black. 

Hab. Rocco Nova, Parana, Brazil. 

Of this very distinct species a single specimen is contained in 
my collection. 


4, On a Collection of Fishes made by Dr. Goeldi at Rio 
Janeiro. By C. Tarr Recay, B.A. 
{Received April 28, 1903.] 
(Plates VII. & VIII.*) 


The collection of Fishes made at Rio Janeiro by Dr. Goeldi 
contains examples of one hundred and twenty-five different species, 
four of which are described below as new to science, one of these 
belonging to a new genus. As most of the species represented 
have been recorded either from Rio Janeiro or from not very 
distant points on the Atlantic coast of 8S. America, it would be 
superfluous to give the full list; in a few cases, however, the 
occurrence of a species at Rio Janeiro has been thought worth 
special notice, and the opportunity has been taken to add some 
notes and to give diagnoses where it seemed useful. Dr. Goeldi 


* Kor explanation of the Plates, see p. 68. 


60 MR. C. TATE REGAN ON‘ [ May 12, 


has presented the types of the new species and other desiderata 
to the British Museum, whilst the greater part of the collection 
has been sent to the Museum at Berne. 


RatD#£. 
RAIA CYCLOPHORA, Sp. Nov. 


Snout with an obtuse triangular projection of moderate length. 
Anterior border of pectoral emarginate. Hye-diameter 33-4 
times in the distance from their anterior margin to the tip of 
snout and equal to interorbital width. Mouth strongly curved, 
36-38 rows of teeth in the upper jaw. Body smooth, except for 
a series of 10-11 spines on the dorsal surface of the tail, and, in 
the male, a double series of curved spines on each pectoral. 

Male with claspers extending to below first dorsal fin. 

Uniform brownish, with a conspicuous black circle on each 
pectoral near the middle of its base. 

Description based on two examples from Rio Janeiro—a female, 
480 mm. in total length, and a male, measuring 410 mm, 


MURANIDA. 
MuR2NA HELENA Linn. 


Five specimens from Rio Janeiro are all dark brown in colour, 
with numerous small white spots on the head, body, and fins. 
Examplesfrom the Mediterranean inthe British Museum Collection 
are similarly coloured, and it seems probable that J/. insudarum 
Jordan & Davis, from the Galapagos Is., which is said to differ 
from MW. helena in having this system of coloration, in reality may 
not be distinct. 

ATHERINIDA. 


ATHERINICHTHYS BRASILIENSIS Quoy & Gaim. 


Of two examples one has four dorsal spines, the other five. In 
the original description it is stated that the lower jaw is shorter 
than the upper, so that Messrs. Jordan and Evermann are 
incorrect in referring this species to Chirostoma, which is dis- 
tinguished by “the very long and strong mandible, which 
protrudes beyond the upper jaw.” The Atherinichthys brought 
from Mexico by Sallé and named A. brasiliensis by Dr. Giinther, 
is a distinct but allied species, which I propose to name A. sallei*. 


* ATHERINICHTHYS SALLEI, sp. nov. :—Depth of body rather less than length of 
head, 5 times in total length. Snout much shorter than eye, the diameter of which 
is 25 times in length of head and equal to interorbital width or length of post- 
orbital part of head. Lower jaw somewhat shorter than upper ; maxillary extending 
to vertical from anterior margin of eye. Sc. 43/10. D. IV,18; A. 1119. Spinous 
dorsal commencing above origin of anal; anterior rays of soft dorsal and anal 
produced, longest anal rays equal to depth of body ; pectorals falcate, as long as head ; 
origin of ventrals equidistant from posterior opercular margin and first anal ray ; 
caudal emarginate. A sharply defined silvery lateral band as broad as a scale. 
Description based on a single example, 75 mm. in total length, from Mexico. 

This species agrees in every respect with Messrs. Jordan and Evermann’s definition 
of Menidia. It resembles A. brasiliensis in the disposition of the fins and number of 
rays, but the latter species has a much longer head, longer snout, smaller eye, &c. 
and very indistinct lateral band. ie 


1903.] FISHES FROM RIO JANEIRO. 61 


SERRANIDA. 


Many authors have regarded Serranus flaviventris Cuv. & Val. 
as the female of S. auriga Cuv. & Val. Messrs. Jordan and 
Evermann give descriptions of three species, viz. : S. dispilurus 
Giinther, S. subligarius Cope, and S. auriga Cuv. & Val., adding 
S. flaviventris to the synonymy of the last ; but in a footnote 
they state that very probably these three species are identical. I 
have arrived at the conclusion that 8. dispilurus and S. subligarius 
belong to the synonymy of S. flaviventris, which is very different 
from S, auriga, as may be seen from the short diagnoses given 
below. 


SERRANUS AURIGA Cuv. & Val. 


Depth of body about equal to length of head, 23-22 times in 
total length. Snout as long as eye, the diameter of which is 33— 
33 times in the length of head and twice the interorbital width. 
Maxillary not extending to below middle of eye, the width of its 
distal extremity } the diameter of eye. Preeoperculum with vertical 
posterior and horizontal inferior limb, the angle rounded ; lower 
opercular spine stronger and further back than upper. D. X 12- 
13, commencing slightly in advance of axil of pectoral, third spine 
_ very elongate. A. III 7, second spine not, or but little, longer 

than third, its length about 24 times in that of head. Sc. 48-52 


6-7 3 : > ours ad: 52 row 
jaz not extending on to upper sui face of head ;_54 rows between 


soft dorsal and lateral line. Two dark blotches on lower half of 
body, the anterior including ventrals, the posterior extending on 
to anal, a light area between them. 

Diagnosis based on four examples from Rio Janeiro, the largest 
150 mm. in total length. 


SERRANUS FLAVIVENTRIS Cuvy. & Val. 

Dules flaviventris Cuvier & Valenciennes, ui. p. 113 (1829). 

Centropristis brasiliensis Brisout de Barneville, Rev. Zool. 
1847, p. 131. 

Centropristis dispilurus Giinther, Proc. Zool. Soc. 1867, p. 99. 

Centropristis subligarius Cope, Proc, Ac. Philad. 1870, p. 120. 

Dules auriga Steimdachner, Sitzb. Ak. Wien, xcvi. I. 1888, 
p. 57, pl. 1. fig. 2. 
 Serranus auriga (part.) Boulenger, Cat. i. p. 287 (1895). 

Depth of body about equal to length of head, 24-3 times in 
total length. Snout as long as or a little longer than eye, the 
diameter of which is 4-44 times in the length of head and 13-13 
times the interorbital width. Maxillary extending to well beyond 
middle of eye, the width of its distal extremity 3 the diameter of 
eye. Preoperculum evenly rounded in the whole extent of its 
posterior margin; lower opercular spine not stronger and not further 
back than upper. D. X 12-13, commencing somewhat behind 
axil of pectoral, third spine not elongate, fourth or fifth highest. 
A. III 7, second spine stronger and longer than third, its length 


62 MR. C. TATE REGAN ON [May 12, 
about 21 times in that of head. Sc. 43-46 ace extending on 
head to posterior margin of eye, 4 rows between soft dorsal 
and lateral line. 5-7 dark vertical bars on upper part of body, 
extending on to dorsal fin; a light area on lower part of body in 
front of anal; a pair of conspicuous black spots on base of caudal. 

Diagnosis based on four examples (including the types of 
C. dispilurus) from the W. Indies, the largest 98 mm. in total 
length. The smaller size of the specimens described makes it still 
more notable that the eye is smaller and the mouth extends farther 
back than in S. @wriga, whilst other differences are sufficiently 
numerous. 


PRISTIPOMATIDA. 


Diagramma cavifrons Cuv. & Val., from the coast of Brazil, 
has been redescribed by Boulenger as D. goeldii; this was 
doubtless due to Giinther having erroneously placed this species 
in Pristipoma, so that it appears from the British Museum 
Catalogue that the genus Diagramma is exclusively Indo-Pacific. 
The synonymy of this species is as follows :— 

Diagramma cavifrons Cuvier & Valenciennes, v. p. 290, pl. 123 

1830). 
ae ane cavifrons Giinther, Cat. 1. p. 286 (1859). 

Genyatremus cavifrons Gill, Proc. Ac. Nat. Sai. Philad. 1862, 
p- 256. 

Genyatremus luteus* Jordan & Fesler, Proc. Ac. Nat. Sci. 
Philad. 1889, p. 504; Jordan & Evermann, Fishes N. Am. ii, 
p. 1342 (1898). 

Diagramma goeldi Boulenger, Ann. Mag. Nat. Hist. (6) xx. 
1897, p. 294. 


MyYLAcropon, gen. nov. 


Body compressed. Scales moderate, cycloid; lateral line 
complete, concurrent with the dorsal profile, the tube straight, not 
extending the whole length of the scale. Mouth moderate, 
protractile ; the maxillary slipping for most of its length under 
the preorbital, exposed distally, without supplemental bone. A 
series of conical teeth in each jaw, posteriorly becoming shorter, 
rounded and molar-like ; internal to these anteriorly one or two 
series of small rounded molars; no teeth on tongue or palate. 
Head scaly ; nostrils close together, rounded, the anterior largest ; 
no pit below the chin; preoperculum serrated ; operculum not 
spmate. Gillmembranes united far forward, free from the 
isthmus; seven branchiostegals; pseudobranchie present ; gill- 
rakers rather short. Two dorsals continuous at the base, with 
XT, 113 rays; anal with III 10 rays; both with a scaly 
sheath at the base. Pectorals asymmetrical, with 18 rays, 
the upper rays longest; ventrals below pectorals, each with 
a strong spine and a scaly axillary process; caudal emarginate. 


* The identity of this species with Lutianus luteus Bloch is extremely doubtful. 


1903. | FISHES FROM RIO JANEIRO. 63 


Air-bladder large, with two long lateral horns anteriorly and 
with a series of compartments along each side: anteriorly 
attached by a strong unpaired muscle running forward above the 
cesophagus. 7 


MYLACRODON GOELDII, sp. nov. (Plate VII.) 


Depth of body 25—-2+ times in total length, length of head 34- 
32 times. Snout as long as eye, the diameter of which is 4 times 
in the length ‘of head, interorbital width 3; times. Lower jaw 
included within the upper; maxillary extending to below anterior 
quarter of eye ; preorbital ae 12-13 gill-rakers on lower part 
of anterior arch. Sc. 55-60 = igi Upper part of head, cheeks, and 
opercles scaly; snout, Jaws, and preorbital naked. D, XI, I 13, 
commencing somewhat behind the oll sus spine very small, 
second short, third longest and equal to ?-2 the length of head, 
thence decreasing ; afiterior soft rays longest, equal to seventh 
spine. <A. IIT 10, first spine short, third "38 of the length of 
second, which is equal to half the length of head, Pectorals scaly 
at the base, $$ the length of head, ventrals a little longer, not 
extending to vent ; caudal strongly emarginate. Dark grey above, 
silvery below, fins blackish. 

Description based on two examples from Rio Janeiro, 225 and 
230 mm. in total length. 


GERRIDA. 


The East Indian genera Gazza Riippell and Liognathus Lacep. 
(Zquula Cuv.), until now placed with or near the Carangide, are 
without doubt very nearly allied to Gerres, which they most closely 
resemble not only in external features but also in their internal 
anatomy. ‘The relations of the genera of this family are shown 
in the subjoined key :— 


A. Scales moderate or large; gill-membranes free from 
the isthmus. 
D. IX-X 10; A. TI-WIL 7-9 ooo... cece cce ese 1. Gerres Cuvier. 
ae IX-X 15-16; A. V 13-14 . pea ... 2, Pentaprion Bleecker. 
. Scales small; ill anaaalpaaes narrowly actin. to 
isthmus; D. VIII 15-16. A. III 14. 


Teeth in jaws minute ............0....... ee. 8. Liognathus Lacepéde. 
Teeth in jaws rather strong, compressed, pointed... 4. Gazza Riuppell. 


The distinction of the species of the genus Gerres is a matter of 
some difficulty. Dr. Goeldi’s collection includes examples of a 
Gerres belonging to the section with preoperculum and prxorbital 
serrated and “with dark longitudinal lines along the rows of scales, 
and which I identify with G. patao Poey. Nearly all the 
American species of this section resemble each other very closely 
in proportions of the head and body, size of the eye, extent of the 
mouth, and number of scales and fin-rays; and it appears that 
the most constant characters which can be used for specific dis- 
tinction are the number of gill-rakers, the length of the second 


64, MR. C. TATE REGAN ON [May 12, 


dorsal and anal spines and of the pectoral, the number of rows of 
scales between the lateral line and the sheath at the base of the 
dorsal fin, and the number of anal rays, JIL 7 or III 8. Messrs. 
Jordan and Evermann give the number of anal rays for one 
species as III 7-8, but I have been unable to find a single case 
throughout the genus where the number of anal rays is variable 
in a species, and there is good reason for supposing that in this 
instance two different species have been confounded. The same 
authors regard G. patao Poey as identical with ’G. brasilianus 
Cuv. & Val., which is said to be only doubtfully distinct from 
G. lineatus Humboldt, in the synonymy of which G. axillaris 
Giinther is included. Other species regarded by them as valid 
are G. brevimanus Giinther, G'. pluniert Cuv. & Val., G. embryx 
Jordan & Starks, and G. mexicanus Steind. 

After examining all the available material in the British 
Museum Collection and referring to the original descriptions, I 
have arrived at conclusions somewhat different from those of the 
authors above mentioned, and the relations of the American 
species of this section which I regard as valid are shown in the 
following key and short revision :— 


Preorbital and praoperculum serrated ; dark longitudinal lines along the rows of 


scales. 
I, Anal with III 8 rays. 
A. Pectoral # length of head ................. 1. G@. brevimanus Giinther. 


B. Pectoral as long as ora little longer than head. 
a. Second dorsal spine as long as or a little 
longer than head. 


Second anal spine % length of second dorsal spine ... 2. G. plumieriCuv. & Val. 
Second anal spine 3 length of second dorsal spine ... 3. G. meaicanus Steind. 
6. Second dorsal spine 2-4 length of head. 
About 15 gill-rakers on lower part of anteriorarch... 4. G@. lineatus Humboldt. 
About 12 gill-rakers on lower part of anterior arch... 5. G. axillaris Giinther. 
Oe, Jelinek yinnilay JOO ¢/ sens) ceo csnseoemocssanecooracsacecce (Gh Gh ynemiI THAR 


GERRES BREVIMANUS Giinther., 


This species is at once distinguished by its short pectoral ; the 
body is a little less elevated than in its allies; second anal spine 
4 the length of second dorsal spine, which is +the length of head ; 
11 very short gill-rakers on lower part of anterior arch. 

A single example from Chiapas, West coast of Mexico. 


GERRES PLUMIERI Cuv. & Val. 


The elongate second dorsal spine and the second anal spine 
nearly as long are diagnostic ; pectoral as long as head or a little 
longer; 13-14 very short gill-rakers on lower part of anterior 
arch. 

Atlantic coasts of Tropical America, Lake Yzabal. 


GERRES MEXICANUS Steindachner. 


This species, from the R. Teapa, apparently differs from the 
preceding only in the shorter second anal spine. 


1903. | FISHES FROM RIO JANEIRO. 65 


GERRES LINEATUS Humboldt. 


The British Museum possesses only a single example of this 
species, from Mazatlan. The pectoral is a little longer than the 
head; the second anal spine nearly equals in length the second of 
the dorsal, which is a little less than ? the length of head. 
15. very short gill-rakers on the lower part of anterior arch ; 
6 scales between first dorsal spine and lateral lime, and 4 rows of 
scales between dorsal sheath and lateral line for nearly the whole 
length of the dorsal fin. 

Apparently this species occurs also on the Atlantic coasts of 
Tropical America, whence it has been described as G. brasilanws 
Cuv. & Val.and Gerres embryx Jordan & Starks. There is nothing 
in the descriptions of these species which does not apply to 
G. lineatus, except that Messrs. Jordan and Evermann give the 
number of gill-rakers on the lower part of the anterior arch as 11 . 
for G. brasilianus, but probably this number has been taken from 
specimens with 7 soft rays in the anal, which are here regarded 
as belonging to another species, viz. G. patao Poey. 


GERRES AXILLARIS Giinther. 

Three examples of this species from Chiapas agree in having 
5 scales between the first dorsal spine and the lateral line, and only 
3 rows of scales between the dorsal sheath and the lateral line for 
the whole extent of the sheath. There are 12 gill-rakers on the 
lower part of the anterior arch, which, although short, are longer 
than in G. lineatus and nearly equal to + the diameter of eye. 
In other characters similar to the preceding species. 


GERRES PATAO Poey. 


The resemblance of this species to G. lineatus is most remark- 
able, and it is only to be separated from it by the anal with III 7 
rays, and the fewer and somewhat longer gill-rakers, of which 
there are about 11 on the lower part of the anterior arch, about 
1 the diameter of eye in length. Some examples have the pre- 
maxillary groove densely covered with small seales as far forward 
as the anterior margin of the eye, but in others these are deciduous, 
and in some even the premaxillary groove shows no trace of having 
been scaly, so that this is an unsafe character to use in specific 
diagnoses. 

Hab. Atlantic coast from Cuba to Bahia. 


TRIGLID®. 
PRIONOTUS BEANIL Goode. 


This species has hitherto been known only from one specimen, 
from off Trinidad, and the taking of another at Rio Janeiro seems 
worth recording. 


PERISTEDION ALTIPINNIS, n. sp. (Plate VIII. fig. 1.) 
Depth of body 54-52 times in total length, length of head 
Proc. Zoou. Soc.—1903, Vou. II. No. V. 5 


66 MR. C, TATE REGAN ON [May 12, 


about 22 times. Snout, including processes, 13 times as long as 
eye, the diameter of which is 4 times in the length of head, 
interorbital width 42 times. Interorbital space concave; no 
spines on snout. Anterior processes longer than broad, flat, 
rounded, with denticulated edges; from each a ridge running 
backwards, ending in a rather broad, flat, somewhat obtuse 
preopercular spine; a short ridge below eye, without spine ; 
a short feeble spine above posterior part of eye, a stronger 
one behind it near posterior margin of head, below which 
another, somewhat weaker; operculum with well-developed spine. 
Margin of lower jaw with groups of short tentacles and a pair of 
longer fringed tentacles which extend back scarcely beyond the 
cleft of mouth. Gill-rakers moderate, about 16 on lower part of 
anterior arch. 27 scutes in 4 longitudinal series, the 3 upper series 
with strong recurved spines, the spines of the lower series quite 
rudimentary. D VIII, 17, the spines slender, the third longest 
and equal to 2 the length of head, thence decreasing; soft rays 
increasing in length to about the sixth, which is as long as the 
longest spine, thence decreasing. AI 16, about 2 the height of 
dorsal; pectoral half as long as head; ventrals extending to anal ; 
caudal emarginate. Greyish (in spirit); pectorals dark. 

Description based on two examples from Rio Janeiro, the larger 
190 mm. in total length. 

This species is very distinct from others so far described, and 
it is difficult to say which should be considered its nearest ally. 


CARANGID&. 


The American species of the genus Scombroides Lacep. (Chori- 
nemus Cav. & Val.) belong to the section Oligoplites Gill, distin- 
guished by having 4—5 dorsal spines, no pterygoid teeth, cheeks 
with sclerous plates attached to the suborbitals, and linear scales. 
There has been considerable difference of opinion as to the number 
of species which ought to be recognised. 

Three examples of S. saliens Bloch from Rio Janeiro agree in 
every particular with others in the British Museum Collection 
from the Pacific Coast of Mexico and Ecuador. ‘This species is 
readily distinguished by the anterior dorsal with 4 spines, the 
deep body (depth 23-3 times in total length), and the wide mouth 
(the maxillary extending well beyond the eye). S. palometa Cuv. 
& Val., hitherto regarded as a variety or subspecies of S. saliens, 
is really quite distinct, the depth of the body being contained about 
32 times in the total length and the maxillary only extending 
to below the posterior margin of the eye, or a little beyond. This 
species has hitherto been recorded only from the fresh or brackish 
waters of Lake Maracaibo, Venezuela, but the British Museum 
Collection contains a small example from Lake Yzabal, a large 
inland lake communicating by the Rio Dulche with the Bay of 
Honduras. /S. altws Giinther and S. sawrus Bl. Schn. each has 
five spines in the anterior dorsal; in the former the body is deep, 


1903. ] FISHES FROM RIO JANEIRO, 67 


as in S. saliens, and the maxillary extends to below the posterior 
margin of the eye; the latter has a more slender body than any 
other species of this section, and the maxillary not reaching the 
posterior margin of the eye; like S. saliens, it occurs on both the 
Atlantic and Pacific coasts. <A fifth species, S. mundus Jordan & 
Starks, from the Pacific Coast of Central America, is said to 
differ from S. altws in having a larger mouth; an example received 
from Dr. Jordan under this name has only four spines in the 
anterior dorsal, and is identical with S. saliens: it may be that 
the number of spines is variable in this species. 


ScOMBRIDA. 


American ichthyologists distinguish between Scombromorus 
regalis Bloch and S. maculatus Mitchill; but the supposed 
differences are so few, so trivial, and, judging by the few specimens 
I have been able to examine, so inconstant, that there can be but 
little doubt that the two are not distinct, and that the latter 
should be added to the synonymy of the former. American 
authors also agree in considering Cybium immaculatum Cuv. & 
Val. to be a synonym of Scombromor us cavalla Cuv. & Val. Iam 
inclined to think, however, that two small examples in the 
British Museum Collection should be referred to S. immaculatus, 
and that they are specifically distinct from S. cavalla, two large 
examples of which are included in Dr. Goeldi’s collection. The 
number of fin-rays is the same and the course of the lateral line 
similar in both species, and the hrnseineinees will be apparent from 
the following short comparison :— 


ScoMBROMORUS CAVALLA Cuv. & Val. 


Depth of body 53-53 times in total length, length of head 
42 times; depth of head (at level of upper angle of gill- opening) 
iE times in its length, diameter of eye 6-63 times, Gill-rakers 
nearly equal to 3 eye- diameter, 8 on lower part of anterior arch. 
Jaws with about 15 teeth on each side. Body dark above, silvery 
below, with traces of dark oval spots on the sides. 

Two specimens, 560 and 620 mm. in total length, from Rio 
Janeiro. 


ScOMBROMORUS IMMACULATUS Cuv. & Val. 


Depth of body 34-4 times in total length, length of head 5 times ; 
depth of head 1+ times in its length, diameter of eye 45-44 times. 
Gill-rakers knob-like rudiments, 8-9 on lower part of anterior 
arch. Jaws with 8-10 teeth on each side. Body dark above, 
silvery on sides and below, without spots. 


Two specimens, 150 and 185 mm., from San Domingo. 


OPHIDIID.E. 


The genus Genypterus Philippi 1s disting uished from Lep- 
ophidiwm Gall by the stronger dentition, the outer series of teeth 
5* 


68 ON FISHES FROM RIO JANEIRO. [May 12, 


in the jaws being large and pointed, and the palatines having a 
single series of strong teeth, The scales are small, regularly 
arranged, deciduous; all the species have a strong opercular 
spine, and in none is a spine on the snout developed. 


GENYPTERUS BRASILIENSIS, sp. n. (Plate VIII. fig. 2.) 


Depth of body about 7 times in total length, length of head 
about 42 times. Snout a little longer than eye, the diameter of 
which is 5-6 times in the length of head, interorbital width 
about 7 times. Lower jaw shorter than upper; maxillary ex- 
tending beyond posterior margin of eye, the width of its distal 
extremity equal to diameter of eye. Scales on upper surface of 
head extending as far as posterior margin of eye; cheeks and 
opercles scaly ; interorbital space, snout, and jaws naked; 15-18 
rows of scales between anterior dorsal rays and lateral line. 
About 8 gill-rakers on lower part of anterior arch, the upper 
of moderate length, graduating to rudiments below. Dorsal 
beginning above posterior third of pectoral when laid back; 
pectoral 3 times in length of head, posterior ray of ventral twice. 
Greyish above, silvery below. 

Description based on five examples from Rio Janeiro, the largest 
440 mm. in total length. 

Apparently no species of this genus has hitherto been described 
from the Atlantic coast of America. In all other species the 
dorsal commences above the middle of the pectoral when laid 
back. 

PLEURONECTIDA. 


SOLEA FONSECENCIS Giinther. 


The occurrence of this species at Rio Janeiro is interesting, as 
it has been previously recorded only from the Pacific coast of 
Mexico and Central America. I have carefully compared the 
single example in Dr. Goeldi’s collection with others from the Gulf 
of Fonseca and the Rio Presidio, and am unable to detect the 
least difference between them. 


LopHip#. 
LopHtius PISCATORIUS Linn. 


A small specimen from Rio Janeiro is exactly similar to others 
of this species from both sides of the North Atlantic. 


EXPLANATION OF THE PLATES. 
Puate VII. 
Mylacrodon goeldi (<2), p. 63. 


Prats VIII. 


Fig. 1. Peristedion altipinnis (x*), p. 65. 
2. Genypterus brasiliensis (x4), p. 68. 


1903.] BATRACHIANS ELC. OF THE PERCY SLADEN EXPEDITION. 69 


May 26, 1903. 


G. A. Boutenesr, Esq., F.R.S., Vice-President, 
in the Chair. 


The following papers were read :— 


1. List of the Batrachians and Reptiles collected by M. A. 
Robert at Chapada, Matto Grosso, and presented by 
Mrs. Percy Sladen to the British Museum. (Percy Sladen 
Expedition to Central Brazil.) By G. A. BouLENGER, 
Dace, WioleAasr 

[Received April 25, 1903. ] 


BATRACHIA. 
1. Buro ryPpHontus L. 


bo 


EUPEMPHIX NATTERERI Stdr. 


3U) 


Hya nasica Cope. 


HYLA SENICULA Cope. 


HYyLaA VENULOSA Laur. 
LEPTODACTYLUS TYPHONIUS Daud. 
PALUDICOLA SIGNIFER Gir. 


CERATOPHRYS CRISTICEPS FE. Miill. 


HYLODES GOLLMERI Ptys. 


REPTILIA. 


LACERTILIA. 
1. Norops SLADENI4, sp. n. (Text-fig. 2.) 


Text-fig. 2. 


Res 
iS 


sve 
3s 


Se, 
= 
is 


Upper and side views ot head of Norops sladenie, enlarged two diameters. 


Very closely allied to WV. auratus Daud., but less slender in 


70 ON MOLLUSKS OF THE PERCY SLADEN EXPEDITION. [May 26, 


form; head broader, less than twice as long as broad ; scales on 
the snout tri- to quinquecarinate, upper labials more numerous 
(6 or 7 to below centre of eye), scales on the body smaller (84 to 
88 round the middle of the body), and the dorsals relatively smaller 
and passing more gradually into the laterals. The hind limb does 
not reach beyond the ear. Greyish or coppery brown above, with 
an ill-defined lighter vertebral line, which may be flanked on each 
side by a series of small blackish, oblique transverse spots ; belly 
pale golden, gular region bright yellow. 


millim. millim. 
Total length ......... Wye Hore) lam brn) aaa 23 
verdes. ous pve 14 End tliminy eee 37 
Width of head ...... 8 AB] oye Wena irate i ae Ory 8 11 
TBO” sosccasnoocas00a9 43 ADE RS oe ONO a raat 1 15, 


Three female specimens. 

. PoLyCHRUS ACUTIROSTRIS Spix. 
LiocEPHALUS cADUCUS Cope. 
TROPIDURUS SPINULOSUS Cope. 
. Hoptocercts spinosus Fitz. 

. AMEIVA SURINAMENSIS Laur. 


, MaButa AURATA Schn. 


ota nr wp 


. MABUIA FRENATA Cope. 


OPHIDIA. 
9. LioPHIs ALMADENSIS Wael. 


10. APOSTOLEPIS ASSIMILIS Reinh. 


2. Note on some Mollusks of the Family Bulimulide from 
Matto Grosso. (Percy Sladen Expedition to Central 
Brazil.) By Hpear A. Smrrg, I.8.0., F.Z.8. 


[Received April 25, 1903. | 


Some specimens of Lulimulide, all of which are in perfect 
condition, have recently been presented to the British Museum 
by Mrs. Percy Sladen. They were collected at Corumba, Matto 
Grosso, by Mr. A. Robert, and two of the species are useful as 
coming from a locality different from that whence the specimens 
already in the Collection were obtained, and the third is of 
interest as being new to the Museum. 


BuLmuuvs Montivacus d’Orbigny. 


Bulimulus montivagus VOrbigny ; Pilsbry, Man. Conch. ser. 2, 
vol x12) p590) pla xii ties: 208030) ipl xiv fess U4 Hor volematgs 
p: 147. 


1903. ] ON COLOUR-HEREDITY IN FANCY MICE AND RATS. is 


Only a single specimen was obtained by Mr. Robert. It agrees 
exactly with the figs. 14 and 15 above cited, the shell there 
depicted also having been collected at Corumba. On comparing 
this specimen with d’Orbigny’s types, I have come to the con- 
clusion, in agreement with Mr. Pilsbry, that the change of name 
proposed by M. Ancey for this shell was not necessary. 


BuLIMuULUS CORUMBAENSIS Pilsbry. 


Bulimulus corwmbaensis Pilsbry, Man. Conch. ser. 2, vol. xi. 
p. 68, pl. xiv. figs. 3-8. 

The colour, as described by Pilsbry, is fairly typical, but the 
ground-colour may vary from “opaque white” to a vinous tint, 
and “the dark brown or corneous longitudinal streaks ” are some- 
times replaced by pellucid stripes. The form also is variable, 
some specimens being broader and more robust than others. 
Mr. Pilsbry described this species from specimens collected at 
Corumba by Mr. Herbert H. Smith. 


DryMaus paciLus dOrbigny. 


Drymeus peecilus, Pilsbry, Man. Conch. ser. 2, vol. xi. p. 285, 
pl. xlix. figs. 49-57. 

This species varies considerably in size, form, and colour, Six 
of the seven Matto Grosso specimens are whitish or yellowish, and 
most have upon the body-whorl seven more or less interrupted 
dark zones, the four uppermost being sufficiently disconnected to 
appear like rows of spots, whilst the three lower ones are but 
very slightly interrupted. Pilsbry’s figure 50 represents this form, 
except that the spire is hardly acuminate enough. His fig. 53 
is rather like the seventh specimen, which, however, is still more 
darkly coloured. 


3. The Present State of Knowledge of Colour-heredity in 
Mice and Rats. By W. Bateson, M.A., F.RS., F.Z8., 
Fellow of St. John’s College, Cambridge. 


[Received May 26, 1903. | 


With the revival of interest in the experimental study of 
variation and heredity which has followed the discovery of 
Mendel’s work, Mice have naturally been chosen by several investi- 
gators as a subject for experiment. To the breeder mice offer 
attractions from their small size, cheapness, healthiness, and rapid 
rate of multiplication. They have further the great advantage 
that the same male can be simultaneously tested with several 
females. They are, however, short-lived, 13 years being a high 
limit of the breeding age. There are also more serious drawbacks. 
They are extremely addicted to eating their young. It is not 
easy to label a live mouse in a permanent fashion, and special 


72 MR. W. BATESON ON COLOUR-HEREDITY — [May 26, 


methods have to be adopted for tracing the identity of the indi- 
viduals, which in such work is indispensable. Another difficulty 
arises from the fact that mice present few readily estimable features 
of structure. Also, though there are now many types of colour, 
few of them as yet exist as pure strains, and hence it is not easy 
to obtain reliable material with which to begin the experiments. 
Nevertheless, in spite of these drawbacks, the subject is a good one, 
and there can be no doubt that our knowledge of heredity can be 
rapidly extended by experiments on mice. As regards Rats the 
case is similar, save in one respect, in which there is a very 
remarkable difference, namely that the colour-types of fancy rats 
are as yet extremely few. For this reason, though the scope of 
experiment is reduced in the case of rats, some serious complica- 
tions are eliminated, and certain fundamental questions, as, for 
example, the relation of pied to self-coloured varieties, could 
probably be studied more easily in rats than in mice. 


As a great deal of work on these species is now being done, it 
has seemed to me useful to codify the chief information already 
at our disposal, and to state as carefully as is yet possible some 
of the more immediate problems presented by the existing facts. 

It would greatly assist discussion of these problems if uniform 
names could be used for the colour-types. An attempt is there- 
fore made to suggest such names, and to indicate how the types 
may be recognised. The specimens examined for this purpose 
have been obtained chiefly from Mr. J. Wilson Steer, of 45 Raleigh 
Road, Hornsey, N., and from Mr. Atlee, of Royston, Cambs., and 
T am greatly indebted to both of these well-known breeders of 
mice for information and assistance. The microscopical examina- 
tion and discrimination of the types was carried out by Miss F. 
M. Durham. This work is only in a preliminary stage and, it is 
hoped, will form the subject of a separate communication. 


Microscopical examination shows the hairs of mice to contain 
numerous minute medullary spaces separated from each other by 
bridges of keratin. These spaces are arranged in longitudinal 
rows, the number of which varies from one to four (perhaps 
five), thick hairs having usually more rows than thin hairs. The 
pigment is deposited in two ways:—(1) massed in the proximal 
walls of the medullary spaces, and (2) scattered in the external 
cortex. Since air bubbles out of the spaces when reagents are 
applied, the spaces probably open to the exterior. 

The pigments in wild W. musculus or sylvaticus ave readily seen 
tobe of three kinds :—(1) Densely opaque black. (2) Less opaque 
brown. (3) Transparent yellow. The chemical nature of these 
pigments and their possible relations to each other seem to be quite 
obscure. If the hairs are cleared of air, the three kinds of pigment 
can be recognised. On treating with 40 per cent. aqueous solution 
of potash, the yellow dissolves atonce. The brown disappears much 
more slowly, but is rather more soluble than the black, which can 


1903. IN FANCY MICE AND RATS. 73 


withstand the treatment more than 24 hours, though ultimately 
it also (and the keratin) disappeavrs. 

The brown may be present in both medulla and cortex; the 
black is chiefly deposited in the medulla, but may be cortical also, 
while it is doubtful whether the yellow is ever present except in 
the medulla. 

All these pigments may coexist in the same hair; but hairs are 
found with only black and brown, others containing only black and 
yellow. Other types possibly occur. The lighter colour is mostly 
peripheral (in hairs which contain other colours), but brown often 
is present in the cortex at levels where the medulla contains black. 

In J. sylvaticus the condition is similar, but the amount of 
black is less. 

The different colour-types of fancy mice are due to the presence 
or absence of one or more of these pigments in various amounts. 
Both the yellow and the brown may exist separately, without any 
other pigment being discoverable, but, so far, no mouse has been 
seen having black only, some brown being always associated with 
black. 

Each chief type of coloration, black, brown, and yellow, exists 
in at least two forms—the one more wétense, the other more 
dilute. The dilution, which affects both medulla and cortex, 
seems to be due to greater scarcity of the pigment-granules, not to 
diminution in their size. 

The following list includes all the types examined, though some 
probably remain to be seen. Waltzing mice, so far, have not been 
examined. The fanciers’ names are generally retained, as on the 
whole distinctive and practical. Owing, however, to the ambiguity 
in the use of the term “fawn” to denote both “ yellow” and 
colorations containing other pigments, the term ‘“ yellow” is used 
for the type containing yellow pigment only. 

1. Ordinary Cinnamon (or Agouti). The colour of AZ. musculus, 
having same three pigments. Exists in at least two strains, one 
rather darker than the other. This is doubtless the “ grey” of most 
writers. Pied forms and strains common. 

2. Golden Agouti. Like (1) but yellower. Contains brown and 
yellow, without black. 

3. Sable. This rather striking type is like (1) on the back, but 
with yellow hairs interspersed at sides. Flanks almost wholly 

ellow. Pied with white this colour gives the so-called “ tricolor.” 

4. Blue-and-tan. Not examined microscopically. [Probably 
sable in which black is diluted. ] 

5. Chocolate= Plum. Contains brown alone. May be pied. 

6. Silver-fawn. A diluted form of (5). Many hairs have 
colourless tips. 

7. Yellow. Contains yellow only. Often called “ fawn,” though 
this term is also applied sometimes to colour containing brown or 
black. When dark pigment is present in association with pre- 
dominant yellow the colour is spoken of as “dingy” or “sooty 
fawn.” 


74 MR. W. BATESON ON COLOUR-HEREDITY [May 26, 


8. Cream. Diluted yellow. 

9, Black. Both black and brown present, without yellow. 
The bases of the hairs are the darkest, and the black does not 
extend to the tips of the large contour hairs, which are brown. 
Hairs behind the ears or on belly are a stilllighter brown. Com- 
plication arises from the fact that at least two kinds of black 
exist, known as “ black” and “ sable-bred black,” viz. thrown by 
sables. These two kinds probably differ in their heredity- 
properties. Pied forms common. 

10. Blue. Diluted form of (9); both black and brown pigments 
coexisting. Blues may be thrown by the “blacks” (not sable- 
bred) and then breed true. Pied forms exist. 

11. Albino. No pigment in any part. As albinos, however 
produced, breed true to the albino character generally, if not uni- 
versally, individuals of dissimilar origins are often mixed together. 
One strain at least, that of Mr. Atlee, is recognised in the fancy 
as having special features of size and shape, and has been kept 
distinct for many generations. 

12. Black-eyed white. Strains of this type have been indepen- 
dently produced twice, perhaps oftener. The degree of pigmenta- 
tion in the eye varies in at least one strain, some eyes being full 
black, others looking blackish red. Whether the type ever breeds 
quite true we cannot say. In our experience offspring with small 
black marks occur (compare phenomenon seen in albino Guinea-pig, 
p. 76). 

13. Variegated. In these, irregular small spots of black or 
chocolate occur on a white ground. Such forms are quite distinct 
from the ordinary piebald and Dutch-marked (viz. like the Dutch 
rabbit) combinations of colour with white. 

In comparing colours care must be taken that specimens are of 
similar age and in similar moult-stages. Differences of intensity 
of colour are of course characteristic of different strains, and 
probably intermediates can be found; but there is no doubt of 
the practical distinctness of each of the forms enumerated. 
‘“Brindling,” viz. lighter or even white hairs distributed as 
ticking, occurs in some of the coloured varieties, as in rabbits, but 
we have not been able to examine specimens, 

As to the age and mode of origin of the several forms little is 
known certainly. Severalconditions are plainly due to resolution of 
compound characters, such as often follows crossing in animals and 
plants. The blue, the black-eyed white, and the variegated are 


certainly productions of the last few years ; the rest (? sable) have 
existed for a long time. 


The question how far J/. sylvaticus has been used in the pro- 
duction of the varieties is a very important one. The experiment 
was suggested many years ago in ‘ Fancy Mice’ and has probably 
been often tried. Mr. Atlee has given me a most circumstantial 
account of a eross with this species made by him nine years ago 
on black-and-white does, and I feel no reasonable doubt that it 


1903. ] IN FANCY MICE AND RATS. 15 


was actually made. The first generation were “agoutis” of very 
large size. Later generations gave amongst others a strain of 
blue, and of black-eyed white. A strain of agouti has also been 
saved from it. He tells me that this formerly had the white feet, 
a character he carefully bred out. Such a cross may have affected 
the whole race of fancy mice at the present day, Our search for 
structural characters referable to sylvaticus, however, has failed 
to show any case of one pair of pectoral mamme (as in sylvaticus) 
or any case of long hind foot. All specimens examined were pure 
musculus in these features. On the other hand, a feature some- 
times seen in fancy mice, and greatly valued by exhibitors, is a 
large eye, much exceeding the size in an ordinary musculus. But 
this eye, though large, is still smaller than that of sylvaticus. 
Nevertheless the large eye is a modern feature in the fancy, and 
I think it not impossible it may have been derived from a 
sylvaticus ancestor. Further experiment alone can decide this 
question. 


In order to appreciate what follows, the reader must have some 
acquaintance with at least the outline of the Mendelian principles 
of heredity. In theirsimplest expression these principles, as they 
are exhibited for instance in the experiments of Cuénot (12), are 
easily comprehended; but when we pass from these simplest 
phenomena to the more complex facts elsewhere witnessed, we 
soon reach difficulties which our experimental evidence is as yet 
only adequate to elucidate tentatively and in part. 

Cuénot experimented by making reciprocal crosses between 
albino, pink-eyed, fancy mice, and wild grey mice (JZ. musculus). 
He was careful to use wild mice in order to be sure that his 
coloured form was pure. As a result he obtained always and 
without exception grey mice. In Mendelian terms, grey is 
therefore dominant over albinism, which is called by contrast 
recessive. The first filial generation thus produced, which we 
may conveniently call F,, when bred inter se, gave a total of 
198 greys and 72 albinos, constituting the second filial genera- 
tion, or F,. The ratio of dominants (D) to recessives (R) is here 
2°75 to 1, a fairly near approach to the ratio 3:1, which on 
the simplest form of the Mendelian hypothesis is to be expected. 
In other words, the facts are, as Cuénot stated*, in agreement 
with the supposition that in the formation of the gametes of the 
hybrid F,, there is complete segregation of the grey colour from 
albinism, and that in both male and female hybrids there are on 
an average equal numbers of gametes produced bearing each of 
these two characters. 

According to the same hypothesis, the grey mice in F, should 
consist of pure or homozygous greys (DD) and of heterozygous 
greys (DR) in the proportion of 1 : 2. 

Cuénot tested this to some extent by breeding the F, dominants 


* Cuénot’s paper seems to be the earliest application of Mendelian principles to 
animals. 


76 MR. W. BATESON ON COLOUR- HEREDITY [May 26, 


inter se, and found that some pairs gave the expected mixture, 
while others gave dominants only. Qualitatively therefore the 
result is the normal one. It is not stated that the “ extracted” 
albinos were tested, but there is little doubt that, in accordance 
with almost universal experience, they would have produced 
nothing but albinos. 

A leading fact illustrated by Cuénot’s experiments, viz. the 
recessive nature of albinism, is borne out by the whole series of 
experiments under review. The fact is true of albinos in mice, 
rats, guinea-pigs * (Cumberland, 13; Castle, 7), and rabbits 
(Castle), so far as experiments have reached. Cases of the pro- 
duction of albinos by coloured rabbits (e.g. Polish by Dutch, 
albinos by silver-greys) are frequent in the fancier’s literature. 
The contrary, the production of coloured animals by albinos, is 
not, so far as I know, illustrated by a single case, with the 
following exception. In the later editions of ‘ Fancy Mice’ 
(Upeott Gill), Dr. Carter Blake, formerly secretary of the 
Anthropological Institute, commenting on the statement that 
albino mice of whatever parentage produce nothing but albinos, 
writes (p. 16) that a pair of albinos produced some brown-and- 
white, some plum, some grey, and some albinos. If this result 
occurred under all precautions, it stands alone. 

Nevertheless we should be cautious in declaring the result 
impossible, for in Mendelian experiments the observer must be 
on the look out for the appearance of a character, elsewhere a 
definite dominant, as the consequence of crossing two dissimilar 
recessives. Not only may a dominant colour be produced by 
crossing two forms having a recessive colour,—e. g., purple flowers 
by crossing the white Datura levis with white D. ferox; purple 
flowers in Sweet-Pea by crossing white “Emily Henderson” 
round-pollened form with the long-pollened form of the very 
same white variety ; purple flowers in the Stock by crossing two 
white varieties: —butalsoa dominant structural character, hoariness, 
may be produced by crossing glabrous (recessive) stocks of dif- 
ferent colours, e.g., red and cream, or red and white+. In each of 
these cases the appearance of an atavistic character occurs as a 
consequence of the union of gametes bearing dissimilar characters ; 
but the character in which the reversion appears is of a class 
different from that in which the parental differentiation was seen. 

The same may very possibly be true of animals also. But in 
each of the cases known, the two varieties united, though alike 
bearing the same recessive character, differ obviously in some 
other respect ; and we know that the cross-bred raised by their 
union is a heterozygote, 7.e. a zygote formed by the union of 
dissimilar gametes. It is, I think, scarcely likely that Carter 
Blake’s case of the mice is really to be so regarded, and on the 
whole the hypothesis of error is more probable; but the possibility 


* Pe pare ae Gy : : ; ; 
* Small “smudges” are said to occur irregularly in albino cavies, however pure. 


+ This statement is based on results of experiments made by Mi ; 
as yet unpublished. : Aiea eee 


1903. ] IN FANCY MICE AND RATS. aa 


that colour may be influenced by structure, and structure by 
colour, must be remembered. 

Naturally we may inquire whether albinism in Man is not a 
similar recessive, Castle has given evidence pointing in this 
direction. The occurrence of albinism in the families of first 
cousins (see Day, Seligmann, &c.) is consistent with this view; 
but there are a few recorded cases of the occurrence of albinos in 
the offspring of albinos breeding with normal parents, where the 
hypothesis that the normal parent was DR is not at all easily 
admissible. No case of the union of two human albinos is known 
tome. The matter cannot here be further discussed, and the 
reader must refer to the literature, the most important paper 
being that of Cornaz*. 

There are a few cases on record where the production of albino 
offspring by animals and plants must almost certainly be regarded 
as the occurrence of a new and original variation, though the 
cause of such sporting is entirely unknown. 

We here encounter the first problem calling for experimental 
study. What zs an albino? We know that it appears to form 
no pigment; but such a body has other characteristics also. 
While the blood of pigmented animals shows intravascular 
clotting on the injection of nucleo-proteids, that of the albino is 
declared to be unaffected. The mountain hare is said in this 
respect to behave as a normal in its summer coat, but as an 
albino after the winter change. How these differences are 
related to the want of pigment we do not know. Such an 
inquiry offers a wide field for experiment. In particular, we 
ought to know how the albino or the normal behaves towards 
the nucleo-proteids of an albino, and so forth. However this 
may be, there seems to be but little doubt that the albino-bearing 
gametes can generally segregate that character entire, as they 
divide from the colour-bearing gametes of the hybrid zygote 
of any colour; and if we knew more clearly what is the real 
physiological difference between colour-secreting and albino 
organisms, we might get a clearer conception of the nature of such 
segregation. 

We may consider next the work of Crampe, which is on a 
large scale and relates entirely to Rats. His latest paper is dated 
1885 (10), and consequently is pre-Mendelian in treatment. He 
bred nearly 14,000 rats, and made elaborate records and tables of 
conclusions. Many of these observations are readily available 
so far as they provide simply qualitative as opposed to quantita- 
tive evidence ; but after many attempts I have not succeeded in 
unravelling the material enough to group the statistics in Men- 
delian form. Though only a sketch can be here given, many of 


* The student of albinism who refers to the paper of Legrain (Bull. Ac. Méd. 
Bruxelles, ix. 1866) should remember that it is the curious instance cited by Darwin 
(Lite and Letters, i, p. 106) as a deliberate invention, 

+ The figures given (10) pp. 555 & 612 are the likeliest, but even these obviously 
contain certain heterogeneous elements now not distinguishable with confidence. 


78 MR. W. BATESON ON COLOUR-HEREDITY [ May 26, 


these qualitative observations are of great value and will provide 
a basis for future work. 

Tn rats the Mendelian rules, in their simplest form, are plainly 
inadequate to express the facts, and we soon meet a number of 
deductions of specific application, each needing full investigation. 
Crampe’s account is long and difficult to follow. At first sight 
also it seems not wholly consistent in certain particulars, but the 
conclusions here summarised seem well established. 

Breeding albino rats with wild J. decumanus, Crampe found 
that F, might have one of two forms, being either a self-grey like 
the wild type, or grey with white marks. Unfortunately no precise 
description of this and of the other broken-coloured rats is given, 
and we do not know the precise extent and distribution of the 
white*. According as F, presented the first or second form, the 
subsequent offspring produced from F, bred iter se, differed. 
The whole series of colours presented by such offspring is arranged 
by Crampe in seven types, thus :— 


. Self-grey. 

. Grey with white marks. 
. White and grey. 

. White (albino). 

5. Black-and-white. 

6. Black with white marks. 
7. Black without marks. 


H> CS bo 


The self-coloured grey in F, gave in their posterity all the 
types except 3 and 5, but F, of type 2 gave all seven types. 

The nature and cause of the heterogeneity in F, is as yet 
unexplained. Such an occurrence is, however, not rare. In my 
own poultry experiments for example, the dark feathers scattered 
in the white F, raised between a brown and a white breed may 
be either chequered or plain black. According as one or the 
other form appears in F, the posterity probably differ, though 
this point is not yet established in the case of poultry. 

The existence of two classes in F, indicates in all probability 
the existence of two classes of gametes, either in the wild 
decumanus or in the albinos, but in which we cannot say. From 
the evidence, it seems that both forms of F, could be produced 
by the same pair of parents, but I cannot find the fact explicitly 
stated. Both forms occurred in F, not only when decuwmanus 
was crossed with albino, but also when it was crossed with type 3 
and with type 5. 

Only the albino could cause all seven types to appear in progeny 
(F, &e.) raised from a cross with the wild type. 

The albino was recessive to all the other six types, and albinos 
of whatever parentage gave nothing but albinos when bred 


* First crosses shown me by Miss Douglas were grey except for an irregular but 
small amount of white on the chest and belly. I take this to be Crampe’s type 2. 


1903. ] IN FANCY MICE AND RATS. 79 


iméer se. ‘This point was elaborately tested. Crampe states that 
albinos true-bred for some generations behaved differently from 
extracted albinos, the former being, as he says, merely “absorbed,” 
v.@. vecessive, on crossing with colour; while extracted albinos 
gave, as I understand him, a mixture of ancestral forms when 
they were crossed with other types. This part of his paper 
(10. pp. 573-5) is difficult to follow ; and I cannot find any example 
showing precisely the nature of the distinction he means to 
emphasise so far as albinos are concerned. We must here await 
fresh experiments. We readily see, however, that though in 
respect of its albinism we may regard the albino as always the 
same, it may obviously be retaining other characters derived from 
various progenitors. Accordingly we find, as will appear, albinos 
apparently of the same species manifesting different properties in 
crossing. I suspect, however, that Crampe is here extending to 
the albino a generalisation really based on a mistake arising from 
misconception respecting the phenomenon of dominance. [See 
note added p. 97.] 

We may now, though the evidence is imperfect, consider the 
significance of the appearance of these many new forms in F,. 
This phenomenon is a most usual result of a cross between 
distinct varieties. It is the source of the majority of our new 
garden varieties, and of many at all events of the colour-varieties 
of domestic animals. In general terms we can declare that the 
result of the cross—the “asymmetrical fertilisation,” to speak 
strictly—is the production of a diversity of gametes. Pending 
histological research, we cannot tell the origin of the characters 
borne by these gametes; but from many circumstances it seems 
inevitable that they must be regarded as created in such a case 
partly by resolution of the character brought in by the dominant— 
which we therefore call a compound character, and partly by the 
imperfect segregation of that compound or of its components from 
the recessive character (and its components if it be also resoluble). 
In most cases the process of resolution is not complete for all the 
gametes ; and some of the gametes are bearers of the wholly or 
partly unresolved character, just as all the colour-bearing gametes 
were in Cuénot’s simpler case. The Mendelian hypothesis leads 
us to believe that the actual numbers of each type of gamete 
will be on the average definite, and that the union of any two of 
them will give rise to a zygote of definite character. 

The number of types of gametes and their several properties 
can only be determined on a minute analysis of each member of 
the series of zygotes by exhaustive breeding. No such evidence 
is yet complete in any one case, but we see already in certain 
cases that some of the F, are homo- and some hetero-zygous, and 
we are beginning to suspect the ratios of the gametic forms in a 
few simple cases. 

Returning to Crampe’s evidence, though the ratios are 
quite uncertain, we find that the several types had different 
properties. 


80 MR. W. BATESON ON COLOUR-HEREDITY [May 26, 


On breeding specimens of each type iter se he found the 
following behaviour :— 


Type 1 might give Types 1 2 4 6 7 
” Za, ” ” LiMo OWL De On ial 
3 ; , Bees) 
” 5 29 ) 
29 4 y) 9? 79 4 z 
9? 2 ) 7 Fi 2 : a 
” es ” ” ae 3) 6 f 
7 4 Suhel 


In other words, each type is dominant to albino and the wild 
type is dominant to all. The grey forms are dominant to the 
black. The black-and-white of type 5 is recessive to type 6, 
(black with white markings), but the self-coloured black does not 
contain type 5. ‘These are some of the deductions from the table. 
The peculiarities of types 3 and 5 are especially noteworthy and 
call for fresh experimental study. 

It appears that types 3 and 5 could be ultimately bred true. 
As to 6 and 7 the evidence is not very clear; but as I understand 
the account, neither was completely freed from throwing the 
other. The breeding in these types was the least successful and 
extensive. Possibly they are illustrations of the d/cttel-rassen of 
de Vries. It is especially noteworthy that the grey-and-white 
type 3 and the black-and-white type 5 do not give rise to self- 
grey gametes or to self-black gametes, a fact found again in 
mice. We see therefore that there are gametes for black-and- 
white and for grey-and-white, each of which may behave as a 
single character and dominate over albino. 

Similarly when pure black-and-white was crossed with the wild 
species, all the coloured types might appear in F, but no albinos 
(10. pp. 555-6). Therefore, in this very important case, when 
black-and-white of type 5 appeared in the posterity of such a 
cross, they were all homozygotes and produced only their own type 
(p. 555). This fact may furnish a useful basis for a new experi- 
ment. In strict accordance with our expectation, Crampe found 
that black-and-whites which gave albinos when bred infer se, 
gave albinos if crossed with albinos; but when they did not 
produce them themselves, they did not when bred with albinos. 

The similar variety, grey-and-white (type 3), is always homo- 
zygous except when it appears as a dominant containing types 
4or 5, But if I rightly understand Crampe’s use of the word 
“constant,” type 5 1s not produced by crossing type 3 with the 
wild form, unless it was already brought in as recessive to type 3. 
On the other hand, type 3 can be produced (in F,) by crossing 
type 5 with the wild form. It is most desirable that the pro- 
perties of these two types (3 and 5) should be fully explored. 
They give a chance of investigating the resolving powers of a 
recessive that is not albino, and free from several complications 
attending the use of the latter. 

When we try to picture what is taking place in the resolution 


1903.] IN FANCY MICE AND RATS. 81 


effected by types 3 and 5, Crampe’s figures, though too imperfect 
and irregular to justify a positive statement, show pretty clearly 
that these particular recessives do not appear nearly so often as 
1 in 4; and consequently it is prima facie likely that some of the 
new types of gametes are formed by imperfect segregation, and 
are combinations containing elements of both the dominant and 
the recessive—a phenomenon indicated by experiments with other 
forms of animals and plants (cf. de Vries’ resolutions of néir- 
rhinwm). 

These are some of the chief deductions apparent from Crampe’s 
work. Many others will strike a careful reader and are indeed 
given by the author, but for these reference must be made to the 
original. 

From the want of details the important question of the identity 
of the several types is not easy to settle, but I think that we may 
allot Crampe’s varieties among the well-known types of rats, with 
fair confidence, as follows :— 

1. The wild decumanus. 

2. Like decumanus, but with a more or less sharply defined white 

area on the ventral surlace (together perhaps with white on the 
feet). 
2 Head and shoulders wild colour, forming the ‘“‘ hood” of the 
fanciers. This is continued in a broad stripe down the middle of 
the back to a patch on the rump. ‘The rest of the body is white. 
The coloured area may be considerably extended on to the flanks, 
and more rarely * the dorsal stripe may be broken. 

4, Albino. 

5. Like 3, but black being substituted for wild colour. 

6. Like 2, but black instead of wild colour. This type is known 
in the fancy as the “ Irish” variety. 

7. Self-coloured black. 

With repect to the kinds of pigments in rats I have as yet no 
information. The distinction between black and the wild colour 
is apparently less sharp than im mice, and both black and black- 
and-white rats have a good deal of dark brown hair, especiaily in 
the edges of the black patches of the parti-coloured, and on the 
belly of the self-coloured black. 

No doubt there is also some change with age, moulting, &c.7 

Crampe (9. p. 393) mentions the black stripe in his black-and- 
whites, and there is practically no doubt that his var. 3 and 
var. 5 are correctly referred to the hooded and striped types. He 
remarks that by selecting those with stripes so wide that the rats 
were more black than white, he got no nearer to breeding blacks. 
Similarly whites could not be bred from the whitest grey-and- 


* This is Miss Douglas’ experience. In this respect strains doubtless differ, for 
My. F. Swann tells me he formerly kept a strain in which the stripe was generally 
broken. 

+ Crampe records (9. p. 395) changes with age in piebalds from “ grey ” to black and 
vice versa, both colours appearing together in the transition. The change in the 
direction of darkening seems to be normal as the adult fur grows. In thesame place 
he mentions a rat as ‘* Gelb-grau.” 


Proc. Zoo. Soc.—1903, Vor. II. No. VI. 6 


82 MR. W. BATESON ON COLOUR-HEREDITY [May 26, 


whites. The types are in fact definite, and cannot be built up by 
cumulative selection. 

T am indebted tc Miss Douglas for much information as to the 
varieties of rats and for the loan of specimens. She tells me that 
rats coloured otherwise than the varieties named are exceedingly 
rare. Irregularly piebald or spotted rats occasionally occur, but 
she has tried recently to obtain such rats from fanciers without 
success. In her experience the type 6 breeds true or nearly so. 
Of the blacks examined by me this type had less of the brown hair 
than type 7. 

A striking feature appears from the rat-evidence, namely the 
absence of yellow, blue, chocolate, and indeed most of the varieties 
so familiar in fancy mice. On superficial examination, the 
colour of a wild rat is not very greatly different from that of a 
wild mouse. In rabbits also the yellow as well as the black forms 
are common. Yellows or yellow-and-whites are also familiar in 
guinea-pigs, fowls (buff, and pile”), and pigeons. Miss Douglas 
has heard of a pair of cream-coloured rats, but otherwise I can 
find no records of any kind of yellow in the fancy. As blacks 
are so easily produced by resolution in the rat, the absence of 
the corresponding yellow and chocolate is remarkable. One is 
tempted to inquire whether the existence of black gametes does 
not suggest that yellow or at least chocolate gametes must also 
exist. The problem of their disappearance raises many important 
questions as to selective union between gametes, and others too 
elaborate to develop now. As there are no yellows, so also there 
are no chocolates. 

Another noteworthy fact is the complete absence of blue rats. 
This particular stage in the diminution of the amount of dark 
pigment is well known in mice, rabbits, cats, and several birds, 
but it is unknown in rats. There is of course no question that 
such forms would have been preserved if they had been seen by 
fanciers. Hither yellow or blue rats would be worth several 
pounds. We may take it therefore that these particular resolu- 
tions, or perhaps mutations, cannot be produced by any of the 
means by which they have been produced in other forms. Con- 
ceivably, if some distinct species were crossed with our fancy rats, 
some of these forms might be created.’ Similarly there are no 
“ Himalayan” rats, 2. e. pink-eyed with patches of colour (blackish 
or yellow), forms well known in rabbits, guinea-pigs, and in the 
“* Japanese” waltzing mice. 

To sum up the evidence as to rats, we have clear proof of the 
segregation of certain types of gamete—the albino, the black-and- 
white, and the grey-and-white, though the ratios in which they 
are produced by heterozygotes are not yet determined. Further, 
there is proof that certain of the colour-types exhibit definite 
valency (Werthigkeit of Tschermak) and dominate over each other 
according to a regular system. Of the other colour-types one, viz., 
type 2, is almost certainly a definite heterozygote form, and is 
probably incapable of being made into a pure race. 


1903.] IN FANCY MICE AND RATS. 83 


We may next consider the further evidence regarding mice. 
By the great kindness of Mr. F. G. Parsons, I am permitted to 
include in this paper notes of 50 matings made by Mr. Parsons in 
conjunction with Dr. S. M. Copeman. These experiments were 
made with other objects in view and are still in progress, but as 
they stand they are a valuable contribution to the question of the 
inheritance of colour. The notes which Mr. Parsons has placed 
at my disposal are here given exactly as they were received. I 
have merely inserted the letters DR and DD according as the 
results show that the individual in question was a heterozygote 
containing albino, or that it was free from the albino character. 
Those not thus distinguished cannot be discriminated by the 
existing evidence. 

The following abbreviations are used: 6/., black; 6r., brown 
= chocolate; blw., pied-black; brw., pied-brown ; w., albino. 


Experiments made by Mr. F. G. Parsons. 


Mating. Offspring. 

(GD) eV cacocasoeoctooncen 2 Llores 22 (0D)183)) 1 blw 3 Ww. 

(2) Re eae sete KNW Orie eeiorte vous Ma NT Ma aac, Tw. 

(B) he ac pec ce has DEW: 3 (BXa) ) (DR) lbrw. 3w. 

Gwe Pe Be wycecccsyececcs x blw. da (DR) 2blw. lbrw. 4w. 

(BS) eile Pee eae Sawai Oh nes eins 5 w. 

(GREE ee ee es SX ORioG) 77 (BS<@) (DIR) ahbaaune 5 w. 

(ZR NESS SA Dae Se x 2 4brw. lw. 

(8) ble: OCCDD) ...... x blw. da (DR) 3 blw. 1 brw (1 eaten ?). 
(9) Sy Mb ike tact x w. dB... 2blw. 1 brw 
(10) w. D ... x blw. da (DR) Bidk, Ilion 9 as 
(a) Be WAGE Ree aee ie SILL g i A) cyte eer 5 w. (i eaten P). 
G2) ..... xX brw. dy (Bxa) (DR) 2br. 6w. 
(GIS ARR Wr resia ete velit < Qbl. Q2br. 4w. 
(GIES Ae ieticonsaentacnnaaty x blw. Be a 1 (DR) Sigs | ilo Ahwe 
(QB) Ts 2IBs . eooceds00008 66 x blw. da (DB) 2blw. 2brw. 4w. 
(1G) Gro PI — oocaeccoqzoense x blw. ga (DR) lblw. Lbrw. 2vw. (5 eaten ?). 
(Gi) aes Fees Waly Sut ye Tw. 
(18) w.2 BI (BX a) obeeas x blw. da (DR) 2 blw. 3 w. (2 eaten). 
(19) w.2 B2(BXa) ...... x oy 2 blw. 1 w. (3 eaten). 
(20) PUTA aU ene Sac xXw.de. Sw. 
(21) w.9 B3(BXa@) ...... x blw. da + (DR) 4 blw 3 w. (L eaten ?). 
(22) a eee ee x wide. Ow. 

(ZB) eR eae iets x blw.C 1(C2 Xa) (DR) Sblw. ... 2w. 
(24) blw. 2 cl (Cx a)(? DD) x blw. da (DR) 7 blw. 

25) MWA GHE s saastataees 2 bl. 4 blw. (1 eaten ?). 
(26) brw. 2 CCX a)(DR) xX blw. ga (DR) lbrw. lw. 
(27) +9 x 4 2blw. 2brw. 2w. (3 eaten). 
(28) Sp SiGe eas iblw. 4brw. lw. 
(29) » xX brw. fC?’ 2(DD) 7 brw 
(80) oe s< - 6 brw 
(31) x soo Ot NOE 
(32) bl. @ D(Dxa)(DR) x blw. 3a (DR) 5 bl. 2w 
(33) x biw. ¢ C2! (DR)(C2x a) 7 bl. 1 blw 


84 MR. W BATESON ON COLOUR-HEREDITY [May 26, 


Mating. Offspring. 
(34) bl. 2 D2(Dxa)(DR) x blw. ga (DR) 2b). 1 br. 2 blw. 
(35) 3S Siw ienmensennes 1 bl. 1 br. 3 blw. 2 w. (4 eaten). 
(36) Ps X brw.  C22(DD)(C2 a) 3 bl. 2br. 1blw. (8 eaten). 
(37) br. 2 D3(D Xa) (DR) X blw. ga (DR) Bio illo, swe 
(38) x brw. dy (DR) 3br. 2brw. 3w. 
(39) 5 Kiwi elsiseasase ces 1 bl. 1 blw. 1 brw. 2 w. 
(40) w. 9 B?! (B2x a) x blw.g a (DR is 2blw. 3w.(3 eaten). 
(41) 55 x 5 see 5 w. 
(42) blw. 2 Cl? (C!x a)(DR) X blw. da (DR) ae 7 blw. 
(43) 5 x 5 ae 4blw. 2w. 
(44) blw. 2 Cl'3 (C1xe) DR X brw.gy(BXa)(DR)... 7blw. lw. 
(45) blw. 2 D23 (D?xa) x blw. da (DR) 5 blw. 


(46) blw. 2 D33(D3xa) x brw. 6 C22(C2xa)(DD) 1bl. 1 br. 1blw. (1 died). 
(47) bl. Dt (D!xa)(DR)X w.d€ occa 1bl. 2blw 2w. 


(48) blw. 2. Cl21(Cl2 x a) blw.g a (DR)... uae 5 blw. 
(49) blw. 2 Cl22(Cl2 xa) xX blw.ga(DR) ... Pie 7 blw. 
(50) blw. 2 Cl221(Cl-22 x a) X blw. ga (DR).. A A blw. 


The mice originally introduced were 5 albino females, 2 albino 
males, 1 black-and-white male, and 1 black-and-white female, all 
of unknown extraction. t 

Several albinos were produced in the experiments. Mated with 
albinos they gave albinos only (41 im all). The original blw. g @ was 
heterozygous, containing albino, but the blw. 2 C probably did not, 
though the figures are insufficient for proof. Both of the original 
blw. on mating with albino gave rise to some brown or brown e- 
and-white offspring, and were probably giving off gametes of 
this nature. All such specimens were alike in tint except one 
which was distinctly lighter. 

The families of the albino D (expts. 10-14) are especially 
interesting ; for,as Mr. Parsons pointed out to me, all the 13 
coloured offspring by two different broken-coloured males, one brw., 
the other blw., were self-coloured, brown, or black. This result 
resembles one obtained by Castle (5. p. 542), but the suggestion 
that such a pied individual is a mosaic which throws self-colour 
gametes is not readily applicable to this case. For here the 
peculiarity evidently lies in the gametes of the individual albino, 
since with other albinos the same males gave pied offspring. As 
Castle commonly obtained such self-coloured mice from albimos 
crossed with pied, it is likely that the peculiarity may belong to 
certain strains of albinos. The detailed account of his experiments, 
which is promised, may perhaps give an indication on this point. 

Parsons’ cases give besides some indications as to the ratios of 
the gametes. It will be seen that the relation of brown to black 
is not merely that of dominant and recessive, for either may give 
either. Nor can it be supposed that the brown is a mere hetero- 
zygous form. Each colour, whether self or pied, may be 
dominant over albino, and the figures show pretty clearly that in 


* Microscopical examination of a specimen kindly sent by Mr. Parsons proved 
this colour to be “ chocolate.” 


1903. ] IN FANCY MICE AND RATS. 85 


the blw.da the albino gametes equalled the browns and the 
blacks collectively, for with various albinos he gave 27 coloured 
(q.v.), 31 albinos, a near approach to equality. As regards the 
brw. dy, the evidence is that with 3 albinos he gave 11 coloured, 
19 albinos. On the other hand, 4 heterozygous coloured females 
xw.de gave 16 coloured, 7 albinos. As the result of the 
reciprocity of these numbers, the total (adding expt. 25) of 
coloured to albinos, produced by all matings in the form DR x R, 
is exactly 59 to 59. It is, however, difficult to believe that the 
departure from equality just named is simply fortuitous, for it is 
in each case steadily maintained through a series of families. If 
the figures are grouped according to mothers (instead of fathers, 
as here) these peculiarities are partly lost, but further experiment 
may possibly indicate that different kinds of heterozygotes are 
here to be distinguished. 

Of matings in the form DR x DR there are ten cases, expts. 
26-7, 32-4, 37-8, 42-4 giving a total of 49 coloured, 14 albinos, 
the simple Mendelian expectation being 47:25 to 15°75, 

In these experiments there is also good evidence of the appear- 
ance of dominants containing no albino, for example brw. ¢ 


(C**), and probably blw.C'. 


Information given me by Mr. J. Wilson Steer and Mr, Atlee, 
and the experiments lately begun by Miss Durham and Mr. Staples 
Browne, enable me to add a few indications as to the probable 
specific behaviour of some of the colour-types in crossing, though 
these have at present only the value of hints for further 
experiment. 

The agouti (us musculus colour) is of course dominant to 
albino, but so far has not been resolved in F,, having apparently 
the same behaviour as the wild colour in Cuénot’s experiments, 
but sometimes piebalds of agouti and white appear. 

Yellow and black crossed have given sables or ‘“ dingy fawns.” * 

Chocolate x albino may give, according to Mr. Steer, the wild 
musculus colour, doubtless by reappearance of the black pigment 
in association with brown of the chocolates. Probably the strains 
used had other differences also (cf. p. 97). Miss Durham has found 
chocolate a simple dominant over albino. 

Sable x black-and-white, according to the same authority, has 
given sables, though in this case the result will probably be found 
to differ according as the black-and-white is homo- or heterozygous. 

Sables bred together have given sable, black, and dingy fawn in 
the same litter. 

Blue xalbino has given a full black, sometimes with white 
patches on tail. 

Two yellows (from an inbred strain 4 years old) gave 1 yellow 
and 2 chocolates (Steer). This last occurrence is unexpected and 
needs careful verification. 


* More recently Miss Durham has thus bred some full blacks. Reliable and 
extensive information as to the result of mating yellows with blacks is greatly needed. 


86 MR. W. BATESON ON COLOUR-HEREDITY [May 26, 


Variegated black-and-white x chocolate-and-white gave on one 
occasion (Steer) 5 black-eyed whites and | chocolate-and-white. 

As mentioned above, blue may be recessive to black and breed 
true from its first appearance, and will doubtless prove to be a 
homozygous colour. 


We may now pass to a consideration of the crosses made with 
““ Japanese” waltzing mice. The exact physiological nature of 
the waltzing habit seems to be still uncertain. Reference to the 
work especially of Cyon, Rawitz, and Zoth shows that, though 
malformation of the labyrinth is not infrequently associated with 
this condition, at least the degree of the structural malformation 
varies considerably. 

The origin of the variation is still more obscure. Mouse-fanciers 
have assured me that something like it may appear in strains inbred 
from the normal type, though I cannot find an indubitable case. 
Such an occurrence may also be nothing but the appearance of a 
vare recessive form. Certainly it is not a necessary consequence 
of in-breeding, witness von Guaita’s long series of inbred albinos. 
From analogy with other cases, we should be prepared to find 
that the existence of such a structural feature in one of the gametes 
had an effect on the colour of the heterozygote ; but the evidence, 
as we shall see, is on the whole unfavourable to this view. 

As to crossing of waltzers and albinos, the earliest evidence is 
that of Haacke, whose records are qualitative only. Crossing 
waltzers, blue-grey with white marks, and albinos, he obtained 
mice generally self-grey (?agouti), more rarely self-black. Their 
offspring occasionally had a small white mark on the ventral 
surface. 

The next large body of evidence is that of von Guaita (19), who 
used black-and-white waltzers with dark eyes (von Guaita 7 litt.) 
and an inbred strain of ordinary albinos. From this cross, F, was 
always (from 4 pairs) a self-coloured house-mouse, and was also 
like that wild type in size (being larger than the waltzer and 
smaller than the albino) and in wild disposition. F,, raised from 
F, bred inter se, consisted of albinos and 4 coloured types—black, 
grey, black-and-white, grey-and-white. The totals were 30 
coloured, 14 albinos. On the expectation of 3:1 there should 
have been 33 and 11, so that the excess of albinos is distinct, 
though the numbers are small; but when all certain cases of 
DRx DR (taking albino as R) are included, the numbers are 
117 coloured and 43 albinos, coming very near indeed to the 
expectation 120 to 40. There can therefore be no doubt that 
the heterozygotes produced on an average equal numbers of 
albino gametes, and of gametes bearing the various colour-types. 

There are only two matings certainly in the form DR R. 
These gave 23+1 coloured, 20+1 albinos, closely approaching 
the expected equality. 

In (20) Table I., from 1st and 5th pairs, we have families of 17 
coloured and 13 coloured respectively, showing pretty clearly that 


1903. ] IN FANCY MICE AND RATS. 87 


some of the coloured individuals contained no albino. As far as 
the few observations went, the extracted albinos gave only albinos. 
So far therefore the Mendelian hypothesis harmonises well with 
the phenomena. 

When, however, we begin to consider the relations of the 
several colour-types to each other, we meet some important 
problems. The original waltzers are described as black-and-white. 
Of what pigments the black was made up we do not know. 
Probably it contained both the black and the brown elements. 
However this may have been, the reversionary heterozygote 
clearly did, though whether it also contained the yellow pigment 
is not so clear. 

On examining the details as to the offspring of the several 
pairs, 1t appears that though the self-greys may, as the first cross 
proves, contain all the other 3 coloured types and the albino, the 
erey-and-white contain the albino only. Similarly the black-and- 
white can only contain the albino, so far as the evidence goes. 
But black seems to be dominant over black-and-white. 

The facts are not sufficient to make these deductions quite 
certain ; and, in particular, since the evidence in rats shows that 
grey-and-white may dominate over black-and-white, it may be 
merely from accident in the choice of individuals that no black-and- 
white was produced by any of the grey-and-white mice. 

The families from the 4th paix on Table I., and from the 3rd 
pair on Table IT., are especially interesting as giving indications as 
to the gametic ratios in a complex case, though the evidence is 
insufficient to determine these ratios. In the first case black- 
and-white x black gave 10 black, 15 black-and-white, 7 albino. 
Both parents were heterozygotes containing albino, being each 
raised from self-greys x white. From the facts it is clear that one 
parent at least was giving off gametes black, black-and-white, and 
white; and from the indication that black is dominant to black- 
and-white, it 1s probable that this parent was the black. The 
simplest supposition is, then, that the black-and-white gave off 
blw. and w., and that the black gave off bl. and blw. in equal 
numbers, and whites equal to their sum, This distribution would 
give the ratio 

1 bl, : 2 blw.: 1 w., 


and where experiment gave 


10 bl.: 15 blw.: 7 w. 
we should expect 8 bl.: 16 blw.: 8 w., 


which fits well. But in the 3rd pair on Table II. we havea blk. x 
albino giving 
7 bl. : 16 blw, : 20 w., 

where, on the hypothesis suggested, we should expect equality 
between bi. and bl.w., and the discrepancy is considerable and 
emphatic. 

Pending further experiment, the relations of bl. to blw. and to 
white cannot be stated with any confidence. Another point 


88 MR. W. BATESON ON COLOUR-HEREDITY [May 26, 


calling for elucidation is the distinction between the black-and- 
white of the original waltzers and the black-and-white of sub- 
sequent generations. F, from albino and the waltzers was the 
atavistic grey, but there is no indication that the subsequent 
heterozygotes between blw. and albino were grey; or more 
strictly there is a great deal of evidence that they were usually 
blw. The suggestion also that the atavistic colour was due to the 
union of the waltzing and non-waltzing character seems to be 
plainly excluded, because even normal albinos in later generations 
proved to be heterozygotes of waltzing and non-waltzing gametes. 
At present, therefore, we cannot declare what was the difference 
between the original pure gametes which caused the reversion 
when they were united. 

Lastly, there is a difficulty, perhaps the most serious of all, in 
the result of the union of albino x self-grey (19. p. 328, 2nd pair, 
both parents being in F,) which gave 13 grey, 3 grey-and-white. 
T see at present no suggestion as to the gametic production of the 
grey parent in this case which can be made with any probability, 
consistently with the other facts. Possibly the diversity of 
gametes lay with the albino. 


I now pass to an examination of the evidence of Darbishire, 
who crossed ordinary albino mice with waltzers. The waltzers 
used were ‘“ pale fawn ”-and-white with pink eyes, thus outwardly 
corresponding somewhat with one of the breeds of rabbits called 
“Himalayan.” The nature of the pigmentation described as 
“fawn” is not specified; but from the results, and especially 
from the distinction drawn by the author between “fawn,” 
“fawn-yellow,” and “yellow,” there can be little doubt that the 
fawn was composed of both yellow pigment and a dark pigment, 
probably black. Twenty-nine pairs were used in the production 
of F,. The offspring of 23 of these pairs, 120 individuals, had 
grey colour. All except one had more or less white or whitish, 
differing in extent. Some had more white than the waltzers, 
while in others the whitish colour only appeared on the belly. 
The pure albinos gave on the whole more fully coloured, the 
extracted albinos less fully coloured heterozygotes. The tint of 
the grey is further classified into “pale wild colour” and “ dark 
wild colour,” both being stated to be such as occur in musculus. 
Full details as to distribution are given (¢. v.). 

Of the remaining six pairs, four gave one or more mice with 
the colour-patches black (with grey brethren in three families) 
as follows :— 

Pane Patches. 
Grey. Black. 


DOT EE strrte se ce ae en Siok 1 
UEXGXON PUM eee ss) sg 1 5) 
JUDO. WALLEY Paes ea areal + 3 
BROKEN iat Stirs Srielt lar 6 2 

iil §) 


1903. | IN FANCY MICE AND RATS. 89 


Lastly two pairs gave both greys and yellows, thus :— 


Page Patches. 
he Grey. Yellow. 
D-41 LA dete emaeeae fateh en aN a 2 3 
THD OO-G Vie MetigsenenasoacoH one 5 A 

a ia 


In the original account and in the discussion of the facts by 
Professor Weldon in ‘ Nature,’ the offspring of the 29 families 
are referred to as having consisted of a mixture of greys, yellows, 
and blacks; and the fact that only certain families gave blacks 
and certain families yellows, and that no family gave both blacks 
and yellows, 1s not emphasised. We can conceive that both 
yellows and blacks might be associated with greys when “ fawns Z 
are crossed with albinos, but till the phenomenon occurs it need 
not be considered in this connection *. 

To proceed with the fundamentally important question of the 
purity of the coloured race, we are informed that the original 
waltzers were bred together for some months and gave only 
offspring like themselves. The number of individuals thus tested 
and the number of offspring raised from them are not given, but 
we may conclude that they were considerable. When, however, we 
regard this evidence of purity in the light of the facts provided 
by the six families which gave either yellows or blacks, we per- 
ceive that if “fawn” is dominant to yellow and to black, the 
occurrence of yellows and blacks in the crosses with albino is 
readily explained. We have in fact only to suppose that in 
family 27 the coloured mother, and in families 77, 78, 85 the 
coloured fathers, contained black ; and that in families 12 and 84 
the coloured fathers contained yellow ; and the results are fairly 
clear. The chance of seeing the impurity by merely crossing 
fawns together would not be very great. Most of them evidently 
were pure, and since black x yellow certainly may give a 
dingy fawn heterozygote, the impurity would probably not be 
demonstrated unless fawns containing black bred together, or 
fawns containing yellow bred together. By breeding the mother 
of family 27 with the father of 77, 78, or 85, some test of this 
suggestion might have been made. Of course we have as yet no 
direct experimental proof that fawn is dominant to black and to 
yellow; but since sables can throw blacks, and since in rats grey- 
nd-white is dominant to black-and-white, it seems very possible 
that these “fawns” may also have been thus dominant. 


% When a compound character is crossed with a recessive, it sometimes happens 
that components of the compound appear already resolved in members of F,. For 
example, I have seen the “ walnut” comb of the pure Malay fowl (which can be pro- 
duced by synthesis of rose-comb and pea-comb in a more or less stable union) crossed 
with single comb give some rose, some pea, some walnut, as well as some single. Such 
a phenomenon will probably be demonstrated to be a partial monolepsis (“false 
hybridism ”’), and zygotes exhibiting the several components will probably not 
reproduce the excluded elements in their posterity. 


90 MR. W. BATESON ON COLOUR-HEREDITY [May 26, 

The most striking fact about the F, heterozygotes (not men- 
tioned in Darbishire’s first paper) is that they all had dark eyes, 
though both parents had pink eyes. The albino showed itself a 
recessive as usual. Moreover, just as in von Guaita’s case, the 
colour of the waltzers did not behave as a simple dominant, but 
formed a specific and reversionary heterozygote. It is especially 
interesting that this heterozygote should have been so nearly the 
same *, though Darbishire’s original coloured form was ‘fawn ”- 
and-white, while von Guaita’s was black-and-white. This cer- 
tainly suggests that the completeness of the reversion may have 
been due to the meeting of some other dissimilarities than those 
indicated simply by colour and albinism (cf. Steer’s case of 
chocolates, p. 85). Other facts point in the same direction. 

Moreover, if the ‘“fawn-yellow” of Darbishire’s class 6 is 
the same colour? as the pale fawn of the original waltzers, it is 
curious to find that in F, there were black-eyed (therefore pre- 
sumably heterozygous) “‘fawn-yellows,’ when the colour grey 
would have been the natural expectation. This phenomenon 
may be compared with that seen in von Guaita’s work, where 
origimal black-and-white x albino gave greys; but in F, black- 
and-white may be a simple dominant over albino, (Compare 
also Parsons’ evidence as to chocolate (= brown) with Steer’s 
experience. ) 

The result of mating the wild-coloured F, together, as far as the 
detailed tabulation extended, was : 


Ja\MOWIXS) ah eccooeonenneandac Deaiiethene all pink-eyed. 

BYiclloweece nae aes A tind ste 3 pink-eyed. 

Fawn-yellow ......... OCR ees 3 pink-eyed. 

Pale iseyger eects 9 

Dankfomeivgeice eee eee 1 

Blacks eee ie ee D 

pevlblac perimental seers Delta Basaeeis 1 pink-eyed 
37 


Tn all, therefore, 16 were pink-eyed and 21 dark-eyed, when 
equality is expected. 

A postscript gives the number raised in F, (presumably from 
wild coloured F,) as increased to 66, and though the individuals 


are not classified according to colours, the information is given 
that there were— 


Albinos Coloured Coloured (? all) 
pk-eyed. pk-eyed. dk-eyed. 
13 ef 36 
tne expecta- | ,,. ; 9 
tion being To .2 Lore) ee 


* Darbishire’s being, however, mostly pied, while von Guaita’s were selfs. 


+ Until qualitative details of these colorations are published, their exact nature 
can only be surmised. 


{ [From a specimen exhibited by Mr. 
one of the dilutions of brown pigment. 
“ silver ” of fanciers. | 


Darbishire, I think this colour is probably 
It appeared to be a paler shade of the 


1903. | IN FANCY MICE AND RATS. 91 


Before attempting further to analyse these facts, the results of 
wild-coloured F, x albino must be given. In the body of the 
aper 88 young so produced are recorded, viz., 39 albino, 31 wild- 
coloured (18 darker, 13 lighter), 15 black, 3 yellow. The total 
increased subsequently to 205, of which 111 were albinos and 
94 had some colour in their coats, the specific colours not being 
as yet given. All the coloured individuals from this mating were 
dark-eyed, as would be expected. In accordance with the other 
results we should expect— 


105-5 albinos and 105°5 coloured with dark eyes, 
where experi- | 
paepeld : o4 : : 


ment gave 


Tt is therefore clear that if we regard the gametes of BF, as 
consisting of two kinds, colour-bearing and albino, in equal 
numbers on an average, this simple form of the Mendelian 
hypothesis fits the facts very closely, and the distribution of 
albinism and eye-colour 1s approximately what that hypothesis 
leads us to anticipate. When, however, we try to assign the ratios 
of the several colour-gametes to each other, and to determine the 
specific results of their unions, we encounter certain difficulties, 
though in all probability further experiment will enable us to 
make this analysis complete. 

So fai: we have no knowledge of the specific composition of the 
several types seen in F,, and until they are bred separately inter sé 
we can only predict the offspring with reserve. Any such predic- 
tion can only be made on the hypothesis that the regularity of the 
behaviour has been maintained, and that no original variation or 
mutation arises (as may happen for instance in peas and perhaps 
sweet-peas). We must also expect some irregularities from the 
fact already mentioned, that the several families in F, were not 
all comparable, and in subsequent generations it will be necessary 
to distinguish members of black-giving, or ot yellow-giving 
families from the rest. Subject to these provisos, we expect the 
pink-eyed coloured types to give only pink-eyed when bred inter se 
and no albinos*, but that the dark-eyed will give both pink-eyed 
and dark-eyed; and that some of the offspring of dark-eyed 
mated inter se (or with albinos) will be albinos. The pink- 
eyed coloured forms mated with pure albinos will presumably give 
all dark-eyed offspring again alve 

Similarly taking the types of F, singly, it is likely that dark- 
eyed yellows will give only yellows, perhaps creams, and albinos, 

* Unless the phenomenon seen in the albino guinea-pig occurs. 

+ In Mr. Darbishire’s third report (28) just published it is recorded that a pink- 
eyed yellow-and-white in F, bred with albinos gave 8 albinos, 11 grey-and-white, 
i yellow-and-white. The occurrence of albinos from this mating is of course a proof 
that the constitution of the pink-eyed yellow was in this case not that suggested in 
the present text. Experiments with other coloured members of F. are not yet given. 
The relation of the yellows to the rest must be regarded for the present as quite 
unknown. It may be remarked that yellow-and-white varieties both of rabbits, fowls, 


and pigeons are often peculiar in their inheritance, and rarely breed true for many 
generations.—July, 1903. | 


92 MR. W. BATESON ON COLOUR-HEREDITY [ May 26, 


while the dark-eyed blacks should give only blacks, perhaps blues, 
and albinos. The dark-eyed greys will probably give both blacks 
and yellows, though in view of von Guaita’s evidence that grey 
may exist as a new homozygote, this is perhaps doubtful. As far 
as the published evidence goes, the most probable constitution of 
the several forms in F, is as follows :— 


DE UCOOOS ar Plaats om Oa Nan ene: ONE albino x albino. 

Pink-eyed yellows............ yellow x yellow. 

Dark-eyed iycce eset s.na-men ces yellow x albinos. 

Pink-eyed fawn-yellows ... fawn-yellow x fawn-yellow. 

Dark-eyed blacks ............ black x albino (@) 

Dark-eyed greys ...........- fawn x albinos (and perhaps 
some other combinations). 

Pinl<eved: Wacsya.ss5 sence lilac x lilac. 


The absence of blacks with pink eyes is noticeable, and raises 
the question whether there is not a permanent synthesis in these 
blacks. 

Finally, we have two important problems, the nature of the 
dark-eyed fawn-yellows and of the dark-eyed “lilacs.” As they 
are dark-eyed they presumably both contain albino. But as 
regards the first, it is difficult to see what the other gamete 
can be in that case. For from F, we learn that fawn x albino give 
grey, not fawn-yellow. On the other hand, as there are black and 
yellow gametes, we ought to find their heterozeyote, which will 
presumably be fawn. But if this combination follows the rules 
of the others, the heterozygote should be pink-eyed, not dark-eyed. 
The number of dark-eyed fawn-yellows, three, is too few to make 
it likely that these are the black x yellows, which we expect to 
appear as a fairly frequent combination ; and the general indica- 
tions are quite unfavourable to the view that any considerable 
number of heterozygotes can be dark-eyed without the presence of 
the albino, though it is not impossible that such real synthesis 
may take place. 

Next the “lilacs” raise certain questions. We must suppose that 
the dark-eyed “lilacs” contain albino; but in the offspring of F, x 
albino there are no “lilacs.” As F, is giving off gametes capable 
of forming “ lilacs,” we see that the lilac x pure albino gives some 
other colour. Next, which of the groups can be supposed to 
represent the lilac-bearing gametes in their other combinations 2 
This also is a question we cannot answer. 

A similar difficulty is created by the scarcity of yellows in the 
offspring of F,xalbino. There were only 3 in 88. We might 
have expected the numbers of yellows and blacks to be equal, but 
there were 15 blacks. Moreover, all the yellows were in one 
family. So far this 1s quite mexplicable. It probably indicates 
that some of the albinos possessed powers of resolution different 
ee those of others, or conversely that some of the original 

awns were more easily resoluble than others. [ef. Cuénot’s 


new results (27), where blacks were resolved out, but apparently 
no yellows. | 


© 


1903. IN FANCY MICE AND RATS. 93 


A line of inquiry is suggested by the miscellaneous constitution 
of F,. We have seen that all members of F, are not alike, and it 
is not impossible that the greys from families which give no blacks 
or yellows may be different in constitution from greys in families 
which gave one or the other. If the individuality of the several 
parents of F, were given, this possiblity could be examined. The 
fact that an original waltzer was giving off yellow or black gametes 
might be an indication that resolution of characters had already 
begun ; and perhaps therefore the F, from different families, though 
alike grey, may be in some measure heterogeneous. In these cases 
it is most important that each individual parent and its offspring 
should be separately traceable. 

It is not impossible that some light on these questions could be 
obtained by noting the sexes in which the several forms appear. 

In view of the facts I do not understand the meaning of 
Darbishire’s statement that ‘the inheritance of eye-colour is not 
in accordance with Mendel’s results.” * So far as the experiments 
are yet recorded, the behaviour of the eye-colour is typically 
Mendelian, and follows Mendelian expectation in its simplest form. 
The occurrence of albinos is similarly Mendelian, one albino in 
four being plainly indicated as the average from F, x F, 7. 

* For further criticism of this statement see Castle and Allen (7). 

+ As regards the waltzing character von Guaita’s experiments agree with Darbi- 
shire’s in showing that it was always recessive to the normal. No individual in Fy, 
or in families produced by crossing F, with the pure normal, waltzed. In Darbishire’s 
experiments F, x F, gave § waltzers in 37 offspring, indicating 1 in 4 as the probable 
average. DRXR isnot recorded. Krom von Guaita’s matings in the form DR X DR 
the totals of families were 117 normal and 21 waltzers made up as follows :— 


Normal. Waltzers. 
36 8 
14 2 
25 a 
41 2 
i 2 
ava 21 


There is therefore a large excess of normals over the expected 3 to 1. This is 
possibly due to the delicacy of the waltzers, which are certainly much more difficult to 
rear than normals are. ‘The small numbers in von Guaita’s litters make it very likely 
that many were lost before such a character as this could be determined. On the 
other hand, we have to bear in mind that as the presence of waltzers is here the only 
proof that the matings were in the form DRxDR, it is possible that the total of 
normals should really include some families which gave normals only. 

DRX R gave 18 normal and 10 waltzers distributed in families thus :— 


Normal. Waltzers. 
3 4, 
3 2 
10 3 
2 1 
18 10 


Here the same paucity of recessives is noticeable, 
Von Guaita did not succeed in raising any offspring from extracted waltzers bred 
é. 
ON ee can be positively asserted is that the qualitative result is in full agree- 
ment with the Mendelian expectation based on the absence of waltzers in F,, and 
that it is not impossible that there may be the expected equality in number between 
D and R gametes produced by F}. 


94 MR. W. BATESON ON COLOUR-HEREDITY [May 25, 


As we have also seen, the colours taken collectively followsimple 
expectation ; F, x F, giving approximately 3 coloured to | albino, 
and F, xXalbino giving approximately equal numbers of each. 

As to the frequencies and valencies of the particular colours 
nothing can be said with much confidence as yet, beyond the 
statement that F, gives off albino gametes about equal in number 
to the various coloured gametes collectively. In a discussion of 
this subject, Professor Weldon (25) has suggested that an average of 
one albino in nine might have been expected. I can see no 
reason why this proportion should be impossible in nature, from 
F,xF,. Its occurrence would, however, be remarkable and raise 
some important problems in gameto-genesis. So far, however, it 
has not been recorded. Professor Weldon is in error in stating 
(25. p. 34) that I have already dealt (4. p. 52) with such a case 
of 1 albino in 9. The case in question was that of Antirrhinwin, 
where de Vries obtained from F, x F, four forms in the proportion 
9:3:3:1, the one being the white, which therefore occurred 
in the proportion of 1 in 16. This is the proportion Mendel 
himself conjectured might be found in a case of resolution, but I 
do not gather that he had actually observed such a case. 

No case of resolution has yet been sufficiently studied for us to 
speak with any confidence as to the ratios of the gametes or the 
nature of the process of resolution, Tschermak has had cases of 
1 recessive in 4, after resolution. In poultry I have had cases 
somewhat similar, to be described hereafter. 

In apparently all recorded cases of resolution some gametes of F, 
carry the compound character unresolved. It is not at all easy to 
suggest a scheme which shall fit both the observed facts of resolu- 
tion and those of cell-division. For example, suppose the gametes 
of F, to be 50 per cent. albino, 50 per cent. variously coloured, if 
segregation were complete. Let us consider the coloured gametes 
separately, and for simplicity assume there are only three kinds of 
them, viz. the unresolved grey, black, and yellow, the two latter 
being hypallelomorphs of grey. It is then clear that in whatever 
numbers the three types are each represented, so long as their 
sum equals the total of albino gametes, there must be more 
black character in any black gamete, and more yellow in the 
yellow gamete, than in any grey gamete; or there must some- 
where be a cell-division in which a part of the yellow and a part 
of the black have been lost. If, for instance, the hybrid bore 
gametes in the proportions 


2 grey (=black+ yellow), 1 black, 1 yellow, 4 albino, 


we recognise that unless the blacks and yellows carry double 
portions of their respective colours, part of the colour originally 
introduced into F, has been lost. Such doubling is not altogether 
inconceivable, though until histological methods are made applic- 
able to these questions of gameto-genesis the possibibity can hardly 
be tested. We note as a fact favourable to such a view, that the 
visible amount of pigment in a black or a yellow zygote is far 


1903. | IN FANCY MICE AND RATS. 95 


greater than the amount of the same pigment in the original 
compound colour. But this consideration cannot be allowed much 
weight, seeing that there may be an excess of pigment in hetero- 
zygotes produced even from two gametes apparently bearing no 
pigment elements at all (cf p.76). In the chemistry of pigment- 
ation there may perhaps be interactions and cancellings so 
complex as to make this particular problem as yet quite msoluble. 

Fuller analysis is especially needed also to determine the place 
of the pied and diluted colour-bearing gametes in the series, but 
it is fairly certain that they must be regarded as due to disinte- 
eration and imperfection of resolution of the colour from the albino 
character. 

Future experiment must decide the conditions determining 
resolution. Cuénot, as I understand his paper, got none in the 
main experiment with wild mice; but he states that he obtained 
yellows, blacks, and pieds “ accessoirement” (perhaps by intro- 
ducing some coloured fancy strain ¢). 


From this survey of evidence mostly already published, it is 
clear that Mendelian analysis provides a means of elucidating a 
large part of the phenomena. The majority of the observations 
are in accord with the Mendelian hypothesis in a simple form. 
The true solution of several subordinate problems still remains 
obscure. The value of the Mendelian analysis will be the more 
appreciated when it 1s remembered that previously the whole body 
ot facts must have been regarded as a hopeless entanglement of 
contradictions, as reference to any non-Mendelian discussion even 
of these very phenomena will show. 

As I have elsewhere pointed out, the central phenomenon in 
Mendelian heredity is segregation. The characters in simplest 
cases are treated as units in gameto-genesis. In more complex cases 
there is resolution, sometimes also disintegration and imperfect 
segregation, leading to the formation of fresh units. The gametes 
bearing these units are produced in numerical proportions which 
on an average are also definite, but as yet these proportions have 
only been determined in the simple cases. Thereis no doubt that 
further experiment will determine them in complex cases also. 

It is the object of Mendelian analysis to determine 


(1) the constitution of the several types of gamete produced by 
each type of zygote ; 

(2) the numerical proportions in which each type of gamete is 
produced ; 


(3) the specific result of the union of any two of the types of 
gamete in fertilisation. 


Though for convenience we may still speak of inheritance as 
being ‘‘ Mendelian” or “ non-Mendelian,” we are rapidly passing 
out of the initial phase of the inquiry in which such expressions 
are demanded. In our further investigations we are concerned 
not so much with the question of the applicability of the simplest 


96 MR. W. BATESON ON COLOUR-HEREDITY | May 26, 


Mendelian hypothesis to special cases, as with the formulation of 
the specific laws followed by the several characters of various 
animals and plants in gameto-genesis and in union by fertilisation. 
As in chemistry, these laws must be worked out separately case 
by case, and each as it is determmed has the value rather of fact 
than of hypothesis. 

In dealing with this class of fact, special precautions are 
necessary in order to establish the identity and purity of any 
variety chosen for experiment. From the description of the 
varieties of mice given at p. 73 it will be seen that some colours 
may be easily confounded in description, though the pigments on 
which they depend have a different chemical behaviour. This 1s 
especially the case with regard to “fawn,” “fawn-yellow,” and 
“yellow.” In such cases it is absolutely necessary that the presence 
or absence of dark pigment should be noted, and that some attempt 
should be made to distinguish the two dark pigments from each 
other. 

In all attempts to trace laws of colour-heredity, colours of the 
various parts will usually have to be reckoned with. In horses, 
for example, the general body-colour, without that of the mane, 
tail, and fetlocks is likely to be an insufficient guide to the 
heredity. In man the heredity of eye-colour cannot be sufficiently 
investigated if it be separated from the colour of the hair, and so 
forth. Fox the present, therefore, Professor Pearson’s conclusion 
that the Mendelian system does not apply to coat-colour of horses 
or to eye-colour in man should not be received without reserve. 
By neglect of the precautions named above many results may be 
described as conflicting with each other, which further analysis 
would show to be harmonious. 


[ Vote added July, 1903. 


When this communication was read I had not seen the 1m- 
portant paper of Cuénot (27) dated March 1903. He states that 
grey mice of his F, when crossed with albino gave several blacks. 
These, when bred with certain albinos produced black hetero- 
zygotes, which bred inter se gave the Mendelian 3 black to 1 albino. 
Some of these latter blacks were then homozygous, and from them 
a pure strain of blacks was raised. This strain crossed with wild 
musculus behaved as a simple recessive, giving grey F,, with 
Mendelian ratios 3 grey to 1 black, or 1 grey to 1 black in F,, 
according as F, was mated with F,, or with black. 

The fact that the original black did not appear in F, or in F 
calls for elucidation. It suggests a possible difference between the 
albinos used either in producing F, or later, some individuals 
having the power of resolving the grey, while others had not that 
power. 

Cuénot next records the new and important fact that the 
colour of the F, produced from his black strain x albino differed 
according to the class of albmo used. (1) Albinos extracted from 


1903. ] IN FANCY MICE AND RATS. a7 

A 
the cross with wild grey gave grey F,. (2) Albinos extracted 
from the cross with black gave black F,. (3) Albinos extracted 
from a cross with yellows (of complex origin) gave a mixture in 
F,, either of yellows and greys, or of yellows and blacks. ‘There 
is therefore a proof that individual albinos, though outwardly 
alike, may belong to several distinct classes, exhibiting different 
properties in their heterozygous unions (cf. Parsons’ case, p. 84). 
The resemblance of the heterozygote to the coloured type from 
which the albino was extracted is a new fact, the significance of 
which we cannot yet fully appreciate *. Cuénot tentatively makes 
the attractive suggestion that the particular colour of the hetero- 
zygote may depend on the association in the same zygote of various 
colour-constituents ; and that though the albino is white in itself, 
it may carry on such constituents from a previous coloured parent. 
Then, according as one or other of these complementary consti- 
tuents is brought in by the albino, the heterozygote will show the 
corresponding colour. The chief obstacle to this view is the fact 
that when a heterozygote shows reversion (as opposed to simple 
dominance) the reversion frequently includes various qualities, 
such as size, temperament, habit of growth, &c., as well as colour. 

A cognate problem was alluded to (p. 88) in the discussion of 
von Guaita’s facts. His original black-and-white x albino gave a 
reversionary heterozygote; yet in F, both the black-and-white and 
the grey-and-white DRs present those colours as simple dominants 
over albino, as their offspring proved. Since in this case no new 
strain was introduced, the reference to pedigree is not sufficient 
to elucidate the whole difficulty. 

The relation of the several classes of albinos to each other seems 
to be the next point for investigation, and a useful experiment 
might be made by breeding albinos extracted from one colour, with 
albinos extracted from another colour, the offspring to be then 
tested with a single pure coloured race. It is not impossible 
that the various types of albino will then themselves exhibit 
phenomena of segregation. 

The new report of Mr. Darbishire (28) and Professor Weldon’s 
comment (29) have appeared too late for adequate discussion here. 
Tt may, however, be remarked that both authors scarcely appreciate 
the Mendelian view when they state that according to it all 
albinos (or other recessives) may be treated as ‘‘in every respect 
similar.” No one, I imagine, would suppose that the similarity 
need extend to characters other than the albinism. We are 
familiar with cases in which recessives, though alike in the reces- 
sive character, are dissimilar in other respects ; and (as stated supra 
p. 76) may, when crossed together, even produce heterozygotes 
exhibiting a character known to be dominant over the particular 
recessive concerned. We should no more suppose all albino mice 
to be identical because they were albino, than all white sweet-peas 


* Possibly it is to this phenomenon that Crampe refers in the statement discussed 
on p. 79 of this paper. I cannot, however, find a case of Crampe’s exhibiting 
Cuénot’s phenomenon. 


Proc. Zoou. Soc.—1903, Vou. II. No. VII. i 


98 ON COLOUR-HEREDITY IN FANCY MICE AND RATS. [May 26, 


because they were white, or all glabrous stocks because they were 
glabrous. 

Professor Weldon’s appeal for the precise gametic formula of F, 
must at present go unanswered. Pending analysis of the various 
coloured types in F,, no one could give the statistical composition 
of the gametes of F, so far as coat-colour is concerned ; and though 
the general composition of F, agrees closely with simple Mendelian 
expectation, the particular composition of the various types is a 
question that further experiment must decide. ‘To take only one 
possibility, imperfect segregation is often seen in such cases of 
complex resolution. No criterion save the actual production of F, 
from F, can show whether any of the types of F, illustrate this 
phenomenon. When such evidence is forthcoming there is every 
likelihood that both the qualitative and quantitative composition 
of the gametes of F, will be determinable with approximate 
accuracy. | 


Bibliographical Summary. 


(Scattered through zoological and fanciers’ literature there are 
many notes on this subject not here cited.) 


ra 


Batreson, W. ‘Mendel’s Principles of Heredity.’ Cambridge, 
1902. 

— ‘Nature,’ 1905, Ixvil. pp. 462, 585; Ixviii. p. 33. 

and H.R. Saunpers. Rep. Evol. Committee Roy. Soc., 
No. 1, 1902. 

“‘ Note on the Resolution of Compound Characters by 
Cross-breeding.” Camb. Phil. Soc. Proc. 1902, xii. p. 50. 

. Caste, W. KE. “ Mendel’s Law of Heredity.” Proc. Amer. 
Ac. Sci. 1903, xxxviil. especially pp. 542-4. 


ar ar 


6. and W.C. Faraser. “ Notes on Negro Albinism.” 
Science, 1903, xvii. p. 75. 
Ue andG. M. Auuen. “ The Heredity of Albinism.” Proc. 


Amer. Ac. Sci. 1903, xxxviii. p. 603. 

8. Cornaz. Ann. Soc. Méd. Gand, 1855, xxxiii. p. 269. [A 

monograph on human albinism. | 

9. CrampE, “ Kreuzungen zwischen Wanderratten verschiedener 

Farbe.” Landwirths. Jahrb. vi. 1877, p. 384; continued 
ibid. xii. 1883, p. 389; xiii. 1884, p. 692. 

‘““Die Gesetze der Vererbung der Farbe.” Jbid. xiv. 
1885, p. 539. 

11. Curnor, L. Comptes Rendus, 1902, vol. 134, p. 779. 

“La Loi de Mendel et l’Hérédité de la Pigmentation 

chez les Souris.” Arch. Zool. exp. et gén. 1902, p. xxvii. 

13. CumBrrLAND, C. ‘The Guinea-pig.’ London (Upeott Gill), 
n. d., pp. 20-1. 

14. Darpisuire, A.D. ‘“ Note on the Results of Crossing J apanese 
Waltzing Mice with European Albino Races.” Biometrika, 
1902, ii. p. 101. 

15 —— Ibid. 1903, ii. p. 165. 


Py Anonlg OS ollie 


Figs. 3-6.A.D Darbishire delad nat. 


Huth, Lith. London 


PLANARIANS FROM ZAN ZIBAR., 


1903.] ON POLYCLADS FROM ZANZIBAR AND BRITISH E. AFRICA. 99 


16. 


Davenport, C. B. ‘‘ Review of von Guaita’s Experiments in 
breeding Mice.” Biol. Bull. 1900, 1. p. 121. [Interesting 
as a careful attempt to discuss the facts before the Mendelian 
principles were rediscovered. | 
Day, J. R. Monthly Homeop. Rev. Lond., 1897, xli. p. 148. 
[Case of albinos from parents who were first cousins. | 


. ‘Fancy Mice.’ Anonymous. London (Upcott Gill), pp. 4, 5. 


[Contains also Carter Blake’s notes, pp. 14—23.] 


. von GuaiTa. ‘ Versuche mit Kreuzungen von verschiedenen 


Rassen der Hausmaus.” Ber. naturf. Ges. Freiburg-i.-B. 


1898, x. p. 317. 
Ibid. 1900, xi. p. 131. 


. Haackxr, W. ‘ Ueber Wesen, Ursachen, und Vererbung 


von Albinismus, &e.” Biol. Centralblatt, 1895, xv. p. 45. 


. Pickrrine, J. W. ‘“ Coagulation in Albinos.” » Jour. Phys. 
) g yi 


1896, xx. p. 310. [Full references. | 


. Sexiemann, C.G. ‘A Note on Albinism,” Lancet, 1901, 


Ixxx. p. 803. 


. Weupon, W. F. R. ‘“ Mendel’s Laws of Alternative Inheri- 


tance in Peas.” Biometrika, 1902, i. p. 244. [Refers to 
Johann von Fischer’s testimony. | 
‘ Nature,’ 1903, Ixvui. pp. 512, 610; Ixviii. p. 34. 


. Zoru,O. ‘ Hin Beitrag zu der Beobachtungen und Versuchen 


an japanischen Tauzmiiusen.” Arch. ges. Physiol. 1901, 
Ixxxvi. p. 147. [Full references to Cyon, Rawitz, &c.] 


. Cutnor, L. “ L’ Herédité de la Pigmentation chez les Souris.” 
5) co} 


9 


Arch. Zool. exp. (4) 1903, vol. i. no. 3. 


. Darpisuire, A. D. “Third Report, &c.” Biometrika, 1903, 


i, pt. 111. 


. WeLpon, W. F. R. ‘Mr. Bateson’s Revisions of Mendel’s 


Theory of Heredity.” Biometrika, 1903, ii. pt. 111. 


On the Marine Fauna of Zanzibar and British Hast 
Africa, from Collections made by Cyril Crossland in the 
Years 1901 and 1902.—Turbellaria Polycladida. Part I. 
The Acotylea. By F. ‘F. Lamnaw, B.A. Cantab., 
Assistant Demonstrator and Lecturer in Biology in the 
Owens College *. 

[Received May 25, 1903. | 
(Plate IX.7 & Text-figures 3-7.) 


Mr. Crossland’s collection contains, so far as the Acotylea are 


concerned, specimens of four new genera and eight new species out 


of 


is 


a total of nine species inall. This high percentage of novelties 
not surprising when one remembers that but little is known of 


* Communicated by the SECRETARY. 
+ For explanation of the Plate, see p. 113. 


100 MR. F, F. LAIDLAW ON POLYCLADS [May 26, 


the Polyclad fauna of the Indian Ocean, and especially of the 
African coasts. 

Asa matter of fact, I believe that no shore-haunting species have 
hitherto been recorded from the East Coast of Africa, save from 
the Red Sea and from the neighbourhood of the Cape of Good Hope. 

Of the four new genera described in the present communication, 
Phylloplana is closely related to Leptoplana, whilst the other 
three are of unusual interest. 

In order to keep the paper within reasonable limits I have 
done little more than give an account sufficient, I hope, in each 
case, to render the future identification of the species a matter 
of certainty. I have not attempted to enter into any detailed 
account of the anatomy of the various species, or to deal with 
many of the interesting questions which have been suggested to 
me by their structure. I have given a list of species of one or 
two of the genera, with their distribution and some of their more 
obvious characters, as I believe such lists may have some use. 

My thanks are due to Mr. Crossland, who has permitted me to 
examine and describe this collection, and has furnished me with 
useful notes and coloured sketches of some of the species. 

T am also indebted to Mr. A. D. Darbishire for drawings repro- 
duced on Plate IX. 


Family PLANOCERIDA. 
PLANOCERA CROSSLANDI, Sp. NOV. 


‘““ White, leaf-like form. Dredged off the mainland coast in 
10 fathoms,” Slightly damaged. 


J SyaVedslalt Saga seo cups mead eared oc 0 22 mm. 
IBreaicltiatnd Ae sence pink. cea oak Oro" 
‘““Mouth”* from anterior end... 12°5 ,, 
g aperture from “‘mouth” ... 4 ,, 
Q a Benne 110) (Stree ee ann e TD) ce 
Tentacles from anterior end ... 6 


Only a single specimen collected. This species is most closely 
allied to Pl. armata mihi [5]. 

The eye-spots have an arrangement very similar to that found 
in the latter species. There is a dense cluster at the base of each 
tentacle; the paired group of brain-eyes lying in front of the 
brain is more extensive than that behind it. The epidermis 
unfortunately has entirely disappeared from the surface of the 
specimen. The muscles of the body-wal! are very similar to those 
found in Pl. armata. The brain is well defined and of moderate 
size. The gut has the character typical of the genus, viz. some 
seven pairs of large branches from the main gut, each of which 
gives off numerous smaller ramifications which do not form any 
anastomoses. The gut is without the peculiar diverticula found 
in Pl. armaia, 


* The term “ mouth” is used to mean the opening of the pharyngeal pouch to 
the exterior, 


1903. | FROM ZANZIBAR AND BRITISH EAST AFRICA. 101 


Genital Organs. 


The male apparatus consists of a short, muscular, somewhat 
barrel-shaped penis, which tapersa little towards its free posterior 
end, where it carries three large chitinous hook-like structures, 
identical in character with those found in Pl. armata. In front 
of these its entire lumen is lined with the small, very numerous 
chitinous spines so characteristic of this and allied genera. The 
penis-muscles are longitudinal and diagonal. The prostate gland 
is large, and, with the penis, is enclosed in an outer muscular 
sheath, the walls of which are made up of an inner circular and 
an outer longitudinal layer of muscle-fibres. At the distal end 
of the penis the prostatic muscles come into close contact with the 
muscles of that organ, but at the upper end of the penis a wide 
space intervenes between it and the sheath. Here the muscles 
of the penis are collected into bundles of retractor fibres, and 
traverse the surrounding sheath-cavity to join the muscles of the 
sheath. 

The prostate is closely enfolded by the inner muscle-layer 
of the sheath ; it gives off a short duct which enters the base of 
the penis after receiving the ductus ejaculatorius from the vesicula 
seminalis. Asin Pl. pellucida and Pl. armata, this duct runs for 
a short distance right inside the prostatic duct. 

The vesicula seminalis lies outside the sheath. 

The vasa deferentia are much dilated. 

The antrum masculinum, into which the free end of the penis 
projects, is lined with a ciliated non-secretory epithelium. 

Cf. von Graft’s figures of the genital apparatus of Pl. pellucida 
and Pl. simrothi |3|; also my figure of Pl. armata [5]. 

Female apparatus—The bursa copulatrix is large, and has 
thick walls composed of an outer layer of circular and an inner 
layer of diagonal radial fibres. Its walls are much folded. 
Beyond the bursa, the vagina, which is lined with ciliated epi- 
thelium, runs forwards and upwards through the large shell- 
~ glands, and then turns sharply back, receiving as it does so the 
common duct from the uteri. Beyond this it is continued back 
as the thread-like accessory vesicle, which ends blindly. The walls 
of the bursa show no trace of secretory activity. 


The following is a list of species which can be referred at present 
with tolerable certainty to the genus Planocera :— 


A. Species in which the penis is armed with large chitinous 
hooks as well as with spines. 
a. Six hooks present. 
Pl. armata Laidlaw [5]. Maldives. 
6. Three hooks present. 
Pl. crosslandi, sp. noy. Zanzibar. 


102 MR. F. F. LAIDLAW ON POLYCLADS | May 26, 


B. Species in which the penis is armed with spines only. 
a, Pelagic forms. 
+ Nervous system much decentralised. 
Pl. simrothi v. Graff [3]. 
+r Nervous system normal. 


Pl. pellucida (Martens) [3]. 
2 Pl. pelagica (Moseley) [1]. 


B. Non-pelagic. 


Pl. graffi Lang [1]. Mediterranean. 
Pl. foliawm CArsted [1]. Mediterranean, North Sea. 
Pl. reticulata Diesing [1], Sandwich Is. 


nec Stimpson. 


Planocera nebulosa Vervill [2] is probably not a member of the 
genus Planocera s. sty. 


PARAPLANOCERA AURORA, Sp. nov. (Plate IX. fig. 1.) 


Body almost circular, margin crenellate. Shore form. 


1 Des aia Fae ra Gre A RRR LONE x Abt e 15 mm 
readin yt diswue tar tee Seana coh oree TA)! 5 
““Mouth” from anterior end ......... (esas 
Tentacles 5 MaDe ike 78 3 c35.5 TO) 
¢ aperture from “mouth” ......... BOB) an 
io) 5 ai." ana al ae ene POM 
Length of receptaculum seminis...... a0) on 
Hemiaiclessaparciee eee ner ee eene Cee eeee iO), 


The coloration of this beautiful species is shown in Pl. IX. 
fig. 1. In general it is of a rich rose-pink, becoming more intense 
towards the margin, and mottled with lines and spots of yellowish 
white. The tentacles are pink, and the mid-dorsal region white. 

The gut-branches are six in number. The arrangement of 
eye-spots and appearance of the genital apparatus are shown in 
the accompanying figure (text-figure 3, p. 103). 

The genital organs have an arrangement in general precisely 
similar to that found in the other species of the genus[6]. The 
antrum masculinum is small, lined with ciliated epithelium. The 
penis is a coiled muscular tube (p.), its walls consisting of circular 
and radial muscle-fibres. Its lumen is lined with chitinous spines 
of two different kinds, those near the distal end of the penis being 
large and irregular, whilst further forward (7. e. away from the 
antrum) the spines become small and thorn-shaped. These two 
kinds of spines merge rather gradually into each other. 

Some way in front of the antrum a thin outer layer of circular 
fibres detaches itself from the wall of the penis, to form an outer 
sheath which remains attached to the ventral side of the penis 
for some distance, so that it is only at its proximal, anterior, end 
that the penis is completely free from the sheath. The space 


1903. ] FROM ZANZIBAR AND BRITISH EAST AFRICA, 103 


between them is occupied by a loose spongy reticulum of proto- 
plasm with few nuclei. At its proximal end the penis receives 
the very short duct which opens into it from the prostate (pr.), 
which with its duct is ensheathed in a continuation of the 
muscle-wall of the penis, which at this level again fuses with the 
muscles of the outer sheath. 


Text-fig. 3. 


Sketch of anatomy of Paraplanocera aurora, sp. nov. 


a.v., accessory vesicle; b7., brain; b.c., bursa copulatrix ; m., mouth; p., penis; 
pr., prostate; v.d., vas deferens; v.s., vesicula seminalis; ¢., tentacle. 


The prostate is large. There is a pair of thin-walled vesiculee 
seminales (v.s.) outside the sheath. From each of these a short 
duct runs, piercing the sheath, and uniting with its fellow to 
enter the commencement of the prostatic duct. 

The terminal parts of the female apparatus are exactly similar 
to those of P. langi, structurally and histologically. However, in 
the present species I cannot find any indications of the shell- 
glands. 


DISPAROPLANA DUBIA, gen. et sp. nov. (Plate IX. fig, 2.) 
A single specimen. 


Temes ivan veenmemmeny: Sects au NS 12°5 mm, 
Breadth ..... scitidib GORE Geter ERE B70) oy 
‘* Mouth” from anterior end ...... AsOnne, 
3 aperture from “‘mouth” ...... lk ere. 
2 3 ips STAY (By Maaco rca Be 52 Re 


The external characters of this species are shown in text-fig. 4 
(p. 104). 
There are no tentacles The eye-spots are arranged in two 


104 3 MR. F. F. LAIDLAW ON POLYCLADS [May 26, 


rows extending forward from over the brain. They are mostly of 
small size, but immediately over the brain there are on either side 
some half-a-dozen eyes of a much greater size than the rest. 


Disparoplana dubia, sp. nov. X 4 cire. 


Unfortunately the solitary specimen is not in a very satisfactory 
state of preservation. The epidermis has almost completely 
disappeared, and the body is much distended with ripe eggs, so 
that the characters of the gut are difficult to determine. The 
pharynx is folded and of the usual Acotylean type. The brain is 
protected by a sheath of unusual toughness. 


Gemtal Organs. 


Male apparatus.—The terminal parts consist of a cylindrical 
penis lined with short chitinous spines, of a small prostate gland, 
and of a muscular vesicula seminalis (Pl. IX. fig. 2). The resem- 
blance to the corresponding organs in Planocera or Paraplanocera 
is very close. The male aperture is very small and opens into a 
narrow tube which runs forwards and a little upwards. The cells 
lining it give off a granular secretion. After a course of about 
1 mm. this passage widens out to become the lumen of the penis. 
This organ is proportionately longer than in Planoceraw and a 
little coiled ; but shorter than in Paraplanocera. Its walls are 
not very stout, and the muscle-fibres which form them are con- 
tinuous with those that surround the prostate. The spines lining 
the lumen bear a close resemblance to those of Planocera; but 
the diameter of the penis is relatively less. The prostate is small, 
but similar to that of Planocera. Its duct is, joined by the ductus 
ejaculatorius running to the penis from the relatively large 
vesicula seminalis which lies in front of the prostate and receives 


1903.] FROM ZANZIBAR AND BRITISH EAST AFRICA. 105 


the two vasa deferentia at its hinder end. Its walls are thin and 
consist of circular muscle-fibres. 

The resemblance existing between these organs of Disparoplana 
and those of Planocera and Paraplanocera is so great, that we are 
compelled to assume a close relationship between these genera. 
In the shape of the penis Disparoplana approaches rather the 
latter genus, whilst in possessing a single vesicula seminalis it 
approxunates rather to the former. 

Female apparatus —The antrum femininum is large and rather 
elongated. The vagina, after running forward to receive the 
separate openings of the uteri, is continued back as the accessory 
vesicle which ends blindly after making a second turn forwards, 
ventral to the first part of its course. The shell-glands open into 
the accessory vesicle. 

The inclusion of this species in the Planoceride will necessitate 
an alteration of the definition of that family. I do not attempt 
to offer a new definition in the present paper, since it 1s necessary 
to have information concerning the anatomy of a large number of 
species the internal structure of which but little is known, before 
any useful modification of Lang’s definition can be suggested. 

However, it is permissible to point out here that the discovery 
of such a form as Disparoplana indicates, I think forcibly, not 
only that in the future our definition of the Planoceride will have 
to be altered, but also that the Leptoplanide may be a polyphyletic 
family. 

The genus Disparoplana may be defined as follows :—An 
elongated form, not provided with tentacles; the eyes arranged 
in two lines over the brain. Mouth a little in front of the middle 
of the body. Penis cylindrical, armed with chitinous spines 
resembling those of Planocera; a well-developed prostate and 
single large vesicula seminalis present. Female ducts simple, a 
small accessory vesicle present. 


STYLOCHUS ZANZIBARICUS, Sp. Nov. 


Labelled ‘“s.s. Juba.” 
A small, rather elongate form. 


Meme Glimmer entre cee tasers ect: os 10 mm. 
ISTEAOUI ee etches dicks sons Chee 
Tentacles from anterior end ......... AAS ts 
“Mouth” 8 Fie MUNN cea. mes 3 
Genital openings from hinder end... “ae 


No note as to colour, but to judge from the single spirit- 
specimen, this is mottled reddish-brown and yellow on the dorsal 
surface; the ventral surface is uniform greyish yellow. 

The eye-spots are found on and about the base of the tentacles 
and on the anterior margin, where they are of moderate size 
(see text-fig. 5, p. 106). 

I have not found any definite group of brain-eyes. 


106 MR. F. F, LAIDLAW ON POLYCLADS [May 26, 


The anatomy of the genital organs is practically identical with 
that of S. neapolitanus [1]. As in that species, the mature 
ovaries have passed to a ventral position. 


Text-fig. 5. 


Anterior end of Stylochus zanzibaricus, sp. nov., showing the arrangement 
ot the eye-spots. 


SryLocHus suESENSIS Ehrenb. ? 
Stylochus suesensis Lang. 
A single large specimen from Ras Oswemba, 10 fathoms. 


Meme tec): eaten crt cacsneae 40 mm. 
IBReadiGhie ew caer pe eereee 25 


PP) 

The specimen is much damaged and the dorsal surface badly 
broken. 

The marginal eye-spots extend completely round the body. 
This character is not stated to occur in Khrenberg’s type, but 
may well have been overlooked. In other respects, size and colour, 
so far as the latter can be determined, viz. dull yellow mottled 
with small brown spots on the dorsal side, it agrees. The ovaries 
are immature and lie among the gut diverticula. Doubtless, 
when ripe, they would shift ventralwards. The penis is squared 
at its free end. The prostate is very large and has thick 
muscular walls. 

The genus Stylochus includes at present a few rather closely 
related species, most of which have a very similar style of 
coloration, viz. a ground-colour ranging from dirty white to dull 
yellow or greyish brown; irregularly mottled with darker spots 
or streaks of brown. The tentacles lie far forward and carry 


eye-spots. Marginal eye-spots may extend completely round 
the body. 


A satisfactory grouping of the species is not at present possible, 
since one of the most remarkable features, known to occur 
in several species, viz. the ventral position of the ovaries, has 


1903. | FROM ZANZIBAR AND BRILISH EAST AFRICA, 107 


been studied only in a few cases. Consequently it cannot, 
unfortunately, be employed as one of the principal characters 
in sub-dividing the genus. 

I give below a tentative table of the species of the genus 
hitherto described, with their distribution :— 


A. Species with eyes on the anterior part of the margin only. 
a. Ovaries dorsal. 


§. pilidiwm (Gitte) (2 [ Mediterranean. Valpa- 
8. Ovaries ventral. vaiso (v. Plehn) [4]. 
Marginal eyes extremely small. 
S. neapolitanus (Delle Chiaje) ae Mediterranean. 
Marginal eyes moderate, brain-eyes absent 2 
S. sanzibaricus, sp. nov. Zanzibar. 
y. Position of ovaries doubtful. 
§. frontalis Verrill [2]. New England. 
S. limosus Diesing [1]. Japan. 
S. conglomeratus Diesing (aU Japan. 


B. Species with eyes completely surrounding the margin. 
a. Ovaries dorsal. 
Small brightly coloured species. 


8. plessisit Lang [1]. Mediterranean. 
4. Ovaries ventral. 

S. swesensis Ehrenb. ? [1]. Zanzibar. 
c. Position of ovaries doubtful. 

S. suesensis Ehrenb. [1]. Red Sea. 

S. argus Ozerniowsky [1]. Black Sea. 

S. tzebra Verrill [2]. New England. 


Since the publication of Lang’s Monograph, the following 
species have been removed from the genus :-— 
1. Stylochus littoralis (Verrill) = Planocera elliptica Girard. 


This species is now referred by Verrill to a new genus, Eustylo- 
chus, characterised by the presence of a median female anterior 
accessory vesicle, apparently similar to thatfound in Paraplanocera. 


2. Stylochus * sargassicola (Mertens) is referred by von Graff 
to the genus Séylochoplana [3]. 


Verrill’s species Stylochus crassus [2] is, I think, evidently not 
a member of this genus. 
Family LepropLANID#. 


PHYLLOPLANA LACTEA, gen. et Sp. NOv. (Plate 1X. fig. 3.) 


Shore form. “ White, with minute grey dots scattered sparsely 
over dorsal surface.” Collected 19,2.01 ; several specimens. 


108 MR. F. F. LAIDLAW ON POLYCLADS [May 26, 


The dimensions of an adult specimen are as follows :— 


IL Ssayedn ey GBguocdsoesseosodebodcdoo0e8 ax 30 mm. 
(Breadth. os0.cs. cece een erence LN oe 
“Mouth,” from anterior end... 14 ,, 
g aperture from “mouth” ... 4 ,, 
2 i spiel e senate Higa: 


This species externally bears a close resemblance to a typical 
Leptoplana, being perhaps a trifle broader and more leaf-like. 
Further, its internal anatomy shows it to be distinctly related to 
the members of that genus, but the presence of two vesicule 
seminales with thick muscular walls is sufficient to distinguish it 
from that somewhat crowded genus. The arrangement of the 
eye-spots is shown in fig. 3, Pl. IX. The pharynx is very large, 
and extends for a distance of over’8 mm. The gut-branches are 
numerous and without anastomosis. 


Genital Apparatus. 


Male organs.—The two vasa deferentia each open into the hinder 
end of the elongated and somewhat convoluted vesicule seminales 
(Pl. TX. fig. 3, v.s.). These have thick muscular walls consisting of 
circular fibres, amongst which are found a small number of oval 
nuclei. The lumen is narrow, and lined with a flattened epithe- 
hum. After running forwards for a total distance of about ‘75 mm., 
the two vesicule unite, and their lumen is continued backwards 
into a median ductus ejaculatorius, which has a length of about 
‘D>mm. For the first part of its course it has thick walls, and its 
diameter is about equal to that of either of the vesicule, whilst 
the epithelium lining its lumen is apparently of a prostatic 
character. For the last third of its course or thereabouts it 
becomes much narrower, and has much thinner walls and the 
epithelium loses its secretory character. Finally, this narrower 
part of the median ductus, which lies through nearly its whole 
length at a level dorsal to the vesicule, opens on to the base of a 
small penis, which is armed with a downwardly-curved, back- 
wardly directed stylet. The penis lies at the upper end of a 
fairly long antrum masculinum. 

Female apparatus.—The female aperture opens into a wide 
convoluted vagina, which at first has a course in general in a 
forward direction, and is provided with rather thick walls, but 
after a time its walls become thinner and the lumen wider. it 
then turns backwards, lying dorsal to the first part of its course, 
and becomes rather narrow, whilst the cells lining it, up to this 
point ciliated, lose their cilia, and have rather the appearance of 
secretory cells. When this backwardly-directed part reaches the 
level of the female aperture, it receives the two uterine ducts 
through a common opening on its ventral side, and is continued 
beyond this to end in a small, slightly muscular accessory vesicle. 
The shell-glands lie close about the aperture. 


1903. | FROM ZANZIBAR AND BRITISH EAST AFRICA. 109 


The genus may be defined very briefly as follows :—Lepto- 
planoid with flattened leaf-like body. <A pair of long muscular 
vesicule seminales, which lie parallel to the median ductus 
ejaculatorius and penis and receive the vasa deferentia of either 
side respectively at their hinder ends. 


HAPLOPLANA ELIOTI, gen. et sp. nov. (Plate IX. fig. 4.) 


Two specimens dated 25.3.01. One of them is immature; the 
other, from which I prepared sections, is in a late stage of sexual 
activity, and the centre of the body is crowded with eggs. The 
body is oblong, about 6 mm. in length, 3 in breadth, with rounded 
ends. In places it is as much as 1 mm. in thickness. 

The eye-spots are arranged in two irregular lines lying over the 
brain on either side of the middle line. They are not numerous 
(text-fig. 6). 


Text-fig. 6. 


Haploplana elioti, sp. nov., magnified and (6) natural size, 


On the dorsal side the epithelium contains a number of pseudo- 
rhabdites. On the ventral surface close to the margin there are 
numerous small rhabdites on either side, but these do not occur 
elsewhere. The basement-membrane is thin, and the muscles of 
the body-wall are very feebly developed. On the dorsal side these 
consist of a narrow longitudinal layer followed by a few circular. 
fibres. On the ventral side there is an additional inner longi- 
tudinal layer. The dorso-ventral musculature is well-developed. 
The ovaries are dorsal, the testes ventral. 

The pharynx is of a very simple type; it consists of a fold 
projecting from the middle of the wall of the oval pharyngeal 
pouch, the longer axis of the pouch coinciding with the main axis 
of the body. The opening into the gut lies rather in front of the 
opening to the exterior, which isabout at the middle of the pouch 
(see Pl. IX. fig. 4, ph.t.). 

The gut-branches are numerous and appear to undergo anas- 


110 MR, F, F. LAIDLAW ON POLYCLADS [May 26, 


tomoses, but it is difficult to determine this owing to the great 
accumulation of eggs in the lateral parts of the body. 

The “mouth” is about 1:5 mm. behind the anterior margin. 

The opening of the antrum masculinum is about 1 mm, behind 
that of the “mouth.” It is very minute. The small vasa deferentia 
are full of spermatozoa; at the level of the antrum masculinum 
they become much contorted, and finally both open into a very 
small median vesicle, which latter appears to open directly by a 
minute pore into the antrum. Thus there would appear to be no 
intromittent organ, though it is quite possible that the vesicle 
may be to some extent everted. The vesicle is provided with a 
very thin wall of circular muscle-fibres. 

The antrum femininum opens some *25 mm. behind the male 
aperture ; it is small and surrounded by a number of shell-glands. 
Dorsally it passes upwards and then backwards, receiving as it 
turns back the common opening of the two uteri. Beyond these 
it is prolonged into the very small accessory vesicle. 

The uteri extend along either side of the body not far from the 
middle line; they vary much in diameter in different localities, 
and in places contain spermatozoa. At intervals, where they 
become dilated, the eggs can be seen making their way into them 
by what appear to be simply gaps in the uterine walls. 

The whole lateral regions are crowded with large eggs, which 
lie embedded in a matrix that consists apparently of a yolk-like 
material (Pl. DX. fig. 4). 

The female aperture is of such small size that it seems impossible 
that the eggs can escape through it. Possibly the body of the 
parent ruptures after a time and allows the eggs to pass out. 
This view is suggested by the fact that in some of the sections 
eges can be seen lying in the gut itself. 

The genus may be defined as follows :— 

Body small, oval, and rather stout. No tentacles or sucker. 
Body-wall muscles feeble. The pharynx of a simple type, opening 
at the end of the first fourth of the body. Male genital apparatus 
of small size, copulatory organ much reduced. Female apparatus 
simple, with a small accessory vesicle. Eyes in two rows over 
the brain. 

The position of this curious form amongst the Leptoplanide 
depends on negative rather than on positive characters. Its 
exact affinities are doubtful, and it is probably a degenerate 
organism. 


Family CEstoPLANIpDm. 
CESTOPLANA FILIFORMIS, sp. nov. 


“15.2.01. Ribbon-shaped, about 1 in. x zim. Creamy white 
with bright yellow border, and a median stripe of the same 
colour” (ct. C. rubrocineta for colour), 

The “ mouth ” lies within 1-5 mm. of the hinder extremity. 

Evidently closely allied to the Mediterranean C. rubrocincta, 


1903.] FROM ZANZIBAR AND BRITISH EAST AFRICA. lil 


but, I think, sufficiently distinguished by its much smaller§size 
and yellow instead of red stripes. I have accordingly ventured 
to describe it as a new species, partly on account of the above- 
mentioned differences and partly on account of its different 
habitat. The genital organs, which are those of a typical 
Cestoplana, are not fully mature, but are at the same time 
sufficiently advanced to lead one to suppose that the specimen 
will not increase very largely in size. 

Unfortunately the head of the only specimen has been damaged, 
so that it is not possible to determine the arrangement of the 
eye-spots. 


OMMATOPLANA TUBERCULATA, gen. et sp. nov, (Plate IX. 
figs. 5-7.) 
One specimen, immature. ‘Prison Island, 4.6.01. White, 
about | in. long, stiff and almost harsh to the touch when alive.” 


er SIN te ser nhl os see INE Eee 26° mm. 
Bea cb ami! ee, TAs 

“ Mouth” from anterior end... 13°5_ ,, 
3 aperture from ‘ mouth” ... Oey 
2 aperture g ¢ 


The stiff texture of the specimen noted by Mr. Crossland is 
very obvious in the spirit specimen, and is due to the presence of 
large numbers of muscular wart-lke projections on the dorsal 
surface. 

The eye-spots are numerous and li scattered over the anterior 
part of the dorsal surface. Their arrangement is shown in text- 
figure 7. 


Text-fig. 7. 


Anterior end of Ommatoplana tuberculata, sp. nov., to show the arrangement 
of the eye-spots. 


The epithelium on both dorsal and ventral surfaces contains 
very numerous pseudorhabdites which stain deeply. Below the 


12 MR. F. F. LAIDLAW ON POLYCLADS [May 26, 


epithelium lies a thick basal membrane which shows no trace of 
nuclei. 

On the ventral side the muscles of the body-wall consist first of 
a very narrow outer layer of longitudinal fibres lying against 
the basal membrane. These are succeeded by an outer diagonal 
layer, which in turn is followed bya few circular fibres, and these 
by an inner diagonal layer, the fibres m this latter running at 
right angles to those of the outer diagonal layer. Lastly comes 
the thick inner longitudinal layer. 

On the dorsal surface there are first a fine outer longitudinal 
layer, next a diagonal layer, and lastly a thick circular layer. 
Hence the dorsal muscles are somewhat similar to those of 
Cestoplana, the ventval ave more highly developed (see Lang eu 
1B IG, ste, 11). 

The dorsal tubercles are more muscular and broader in pro- 
portion to their height than those of Cycloporus. They are 
covered with an epithelium, which about their base is similar to 
that of the rest of the surface of the body, but which towards 
their apices becomes flattened and loses its pseudorhabdites. 
Under the basement-membrane, which becomes somewhat attenu- 
ated on the tubercles, is a special layer of circular muscle-fibres. 
In the centre of each tubercle is a small quantity of tissue richly 
supplied with nuclei, and connected with the parenchyma of 
the body by a strand of tissue which pierces the muscles of the 
body-wall, This tissue is perhaps nervous in character (Pl. IX. 
fig. 6, x, y). 

The pharynx is large and much folded. The “ mouth” hes 
behind its posterior end, and communicates with the elongated 
pharyngeal pouch by a narrow channel. The gut-branches are 
numerous and lie at different levels (Pl. 1X. fig. 6, g’). 


Genital Organs. (Plate LX. fig. 7.) 


Owing to an unfortunate accident some of the sections in the 
region of the body where these organs occur have been lost. 
Consequently the account given below is necessarily incomplete. 

There appears to be a single genital atrium. The penis is 
provided with a true penis-sheath (cf. Cestoplana), and there is 
also a prostate gland. The vagina runs backwards and upwards 
for some little distance, then bends forwards and downwards, 
receiving on its dorsal’ side the common opening of the uteri. 
Beyond this it runs forwards and downwards towards the atrium 
(possibly opening into it 2). 

My chief reasons for placing this oval form in the neighbour- 
hood of Cestoplana ave to be found in the shape of the penis, the 
presence of a prostate gland, the backwardly-directed mouth- 
opening, and the disposition of the eye-spots. The thickness of 
the basement-membrane and the arrangement of the gut-branches 
and muscles of the body-wali are, I believe, indications which 
point in the same direction. 


1903. | FROM ZANZIBAR AND BRITISH EAST AFRICA. 113 


The genus Ommatoplana may be defined as follows :— 

Body oval, covered on the dorsal side with numerous wart-like 
tubercles. Eye-spots scattered over the anterior third of the 
body dorsally. ‘‘ Mouth” behind the pharynx. A single genital 
atrium. Penis provided with a true penis-sheath, without a 
stylet ; prostate gland present, with thick muscular walls. 
Vagina provided with an accessory vesicle. 


LITERATURE. 


1. Lane.—Naples Monogr. xi. 1884. 

2. VERRILL.—Trans. Connect. Acad. vill. pp. 459-520, pls. xl.— 
xliv. 

3. GraFF, von.—Zeitschr. wiss. Zool. lv. 1893, pp. 189-219, 
tt. vil.—x. 

4, PLEHN, von.—Jena. Zeitschr. xxx. pp. 137-176, tt. viii.—xiii. 

5. LarpLaw.—Fauna and Geogr. of the Maldive and Laccadive 
Archipelagoes, vol. 1. pt. 3, pp. 282-312, pls. xiv., xv. (1902). 

§. Larpiaw.— Manchester Memoirs, vol. xlvii. pt. 2, no. v. (1903). 


EXPLANATION OF PLATE IX. 


Hig. 1. Paraplanocera aurora, sp. nov. (p. 102). 15. 

Fig. 2. Section through the region of the male apparatus of Disparoplana dubia, 
sp. nov. (p. 103). The section is oblique, and hence it is possible to see 
the vesicula seminalis in the same section with the anterior end of the penis, 
which actually lies some distance behind it. The plane of the section is very 
nearly longitudinal. 

. 3. General anatomy of Phylloplana lactea, sp. nov. (p. 107). X 15. 

Fig, 4. Transverse section across the body of Haploplana elioti, sp. nov. (p. 109), in 

the neighbourhood of the pharynx. 


Fig. 5. Anterior end of Ommatoplana tuberculata, sp.nov. (p. 111). x 5. 
Fig. 6. Part of a transverse section across the body of Ommatoplana tuberculata 
passing through two of the tubercles. 
Fig. 7. Part of a transverse section of ditto, showing the prostate, penis, and penis- 
sheath. 
Heplanation of Lettering. 
a.m., antrum masculinum. ph.t., pharyngeal pouch. 
é., basement membrane. pr., prostate. 
ch.s., chitinous spines. ps.7., pseudorhabdites. 
d.e., ductus ejaculatorius. s., spermatozoa. 
g. gut. : sh.gl., shell-glands. 
g'., gut-branches. te., testis. 
m.¢., circular muscles. ut., uterus. 
m.d., diagonal muscles. v.S., vesicula seminalis. 
m.l.i., tamer longitudinal muscles. v.d., vas deferens. 
m.l.o., outer longitudinal muscles. a, tissue of tubercle-process of Om- 
M., Nervous tissue. matoplana, perhaps nervous in 
0.) ege. character. 
ov., Ovary. Y, gap in muscles of the dorsal body- 
Pu» PENIS. wall of Ommatoplana tuber- 
p.s., penis-sheath. culata. 
ph. pharynx. 


Proc. Zoou. Soc.—1903, Vou. IT. No. VIII. 8 


114 ON ABNORMALITIES IN DOMESTIC FOWLS. [June 16, 


June 16, 1903. 


F. Du Canz Gopmay, Esq., D.C.L., F.R.S., Vice-President, 
in the Chaiv. 


The Secretary read the following report on the additions made 
to the Society’s Menagerie in May 1903 :— 

The registered additions to the Society’s Menagerie during the 
month of May were 122 in number. Of these 14 were acquired by 
presentation, 7 by purchase, 17 were born in the Gardens, and 
84 were received on deposit. The total number of departures 
during the same period, by death and removals, was 143. 


Mr. F. Finn, F.Z.S., exhibited a living hen-feathered Bantam 
cock and the feet of a fowl showing a three-jointed hallux, and 
made the following remarks :— 

“Although there exist two breeds of fowls in which hen- 
feathering in the male is a constant character—the ‘ henny’ Game 
and the Sebright Bantam—yet the occuryence of hen-feathering 
in the male as a casual variation appears to be so rare that I 
thought the present specimen worthy of exhibition to the Society. 
It is, as will be seen, a fully adult Bantam cock of no particular 
breed, and certainly shows no traces of Sebright blood, being 
single-combed and possessing black body-plumage and a white- 
bordered neck-hackle, with no trace of the characteristic black- 
laced plumage of Sir John Sebright’s birds. It is interesting to 
recall that that gentleman made his celebrated breed hen- 
feathered by crossing into his strain a hen-tailed Bantam he 
came across casually, just as this one occurred to me. 

“Mr. W. Bateson (‘Experimental Studies in the Physio- 
logy of Heredity’: Royal Society, Reports to the Evolution 
Committee, I.) speaks of the occurrence of a chick with a long 
hallux bigeminus as an abnormality which was probably un- 
recorded. About a month before, however, in a letter published 
in ‘ Nature,’ January 30th, 1902, [ had mentioned the occurrence 
of an abnormally long hallux in a common Egyptian fowl, which 
I regarded as the homologue of the ‘ fifth’ toe of birds possessing 
a double hallux. I did not keep this specimen, but I now exhibit 
the feet of another Egyptian fowl showing the same peculiarity, 
which I obtained last year. The long halluces have each three 
phalanges, but the foot, although looking powerful, had no 
particular power of grasping, as was the case with the first speci- 
men I met with. I haveseen another case of long single halluces 
in a fowl which I believed to be Egyptian, and one also in a 
‘Silky’ fowl, this breed having usually five toes. As Egyptian 
fowls display a continuous variation from one normal hallux to 


‘ ? 5 ‘ a LAG 2ab 
the five-toed forms, more research amongst them would be 
interesting.” 


1903.] ON THE NURSING-HABITS OF A SOUTH-AMERICAN FROG. 115 


Mr. G. A. Boulenger, F.R.S., exhibited a specimen of 
Ceratohyla bubalus Espada, carrying eggs on its back (text-fig. 8), 
and made the following remarks :— = 

“A very interesting addition has been made by Mr. Ockenden, 
a zoological collector in Peru, to our knowledge of the nursing- 


Text-fig. 8. 


Ceratohyla bubalus, 2, carrying eggs on back. 


habits of Batrachians. The frog now exhibited is a female of the 
rare Ceratohyla bubalus, an inhabitant of the Andes of Ecuador, 
Bolivia, and Peru. It measures 63 millimetres from snout to 


g* 


116 MR. R E, HOLDING ON THE [June 16, 


vent, and carries on its back nine large spherical eggs, 10 milli- 
metres in diameter, each containing a little frog distinctly visible 
through the transparent membrane which at this stage constitutes 
the egg-capsule. The little one, with the abdomen, tumid with 
yolk, turned towards the back of the mother, and the limbs folded 
against the belly, is connected with the membrane by two string- 
like cords on each side, proceeding from the throat, as figured in 
Nototrema cornutum Blgr. (P. Z.8. 1898, pl. xviii.), and which 
serve to convey the blood, for the purpose of respiration, to the 
vascular, allantois-like membrane. The resemblance which the 
young bears to that of Nototrema cornutum is very striking, 
except in the breathing-organs, which im the latter, as well as in 
N. oviferum and NV. testudinewm, form funnel- or bell-shaped 
appendages. Whilst in WVototrema the eggs are protected in 
the dorsal pouch with which the mother 1s provided, the eggs 
in this Oeratohyla simply stick to the back, leaving shallow hexa- 
gonal impressions on the much-thinned dorsal skin, through which 
the neural processes of the vertebrz project to such an extent as 
to leave marks on the vitelline sacs with which they are in 
contact. 

“ A near ally of the Hemiphractide, to which family Cerato- 
hyla belongs, Amphignathodon guentheri Blgr., is provided with 
a dorsal pouch as in Vototrema. We therefore find among the 
Tailless Batrachians with teeth in both jaws the same adaptations 
for the protection of the offspring as occur among the Hylide, 
where Ayla goeldii Blgr. stands, in this respect, in the same 
relation to Nototrema as Ceratohyla bubalus to Amphignathodon. 

“The specimen exhibited was obtained at Santo Domingo, 
Carabaya District, S.E. Peru, 6000 ft. Mr. Ockenden is unable 
to give any particulars as to the conditions in which it was found, 
he not having been the actual discoverer of the remarkable 
specimen.” 


Mr. F. E. Beddard, F.R.S., exhibited on behalf of the ‘“‘ Flower 
Memorial Committee” a bust of the late President of the Society, 
Sir William Henry Flower, K.C.B., D.C.L., F.R.S., which had 
been executed by Mr. Thomas Brock, R.A., and which would 
ultimately be placed in the British Museum of Natural History. 


Mr. F. HB. Beddard also exhibited and made remarks upon 
sections of the ovary of Thylacinws which showed the immigra- 
tion of follicular cells into the ova. 


Myr. R. E. Holding exhibited some skulls of the St. Kilda four- 
horned Sheep, and made the following remarks :— 

“The small black Spanish or St. Kilda four-horned Sheep is 
interesting not only on account of the curious legend as to its 
first. appearance on the island of St. Kilda, but from its having 


1903. HORN-GROWTH IN THE ST. KILDA SHEEP. ile 


Text-fig. 9. 


Variations in the horns of St. Kilda Sheep. 


A. Ram Lamb, aged 15 months, showing retention of median or upper horns only. 
The sire and dam of this had each four horns. 

B. A well-grown St. Kilda head. 

C. Ram, showing retention of median horns and suppression of right lateral horn. 

D. Head of Ewe, showing two small horns on right side; on the left side is a knob 
apparently containing three or four points of bone covered by a single horn- 
sheath. 

E. Indian Ram (Brit. Mus.) having five horns. As many as seven or eight horns 

have heen observed. 


118 ON THE HORN-GROWTH IN THE ST. KILDA SHEEP. [June 16, 


so persistently retained its colour (dark brown and black) and 
characteristic four horns which exist in both sexes. The irregular 
development and somewhat erratic growth of these horns have 
been noted and commented upon by both veterinary surgeons and 
comparative anatomists; and some diversity of opinion exists as 
to the homology of the two pairs of horns when compared with 
the single pairs of horns as carried by some Domestic and 
Wild Sheep. The purpose of this communication is, from a 
biological point of view, to trace the cause of the duplicate pair 
of horns and to determine their homology. Text-figure 9, B, 
represents a typical well-grown head. 

“ An examination of a number of heads in the British Museum 
and the Museum of the Royal College of Surgeons and elsewhere 
shows an extraordinary variation in the form and direction, or 
‘pitch’ as it were, of the median or upper pair of horns. These 
may grow quite upright, as shown on the left side of EH, ov may be 
curved forward to a greater or less degree, but are always present. 

“The lateral or lower pair of horns, though more constant as 
regards form and more nearly approaching the typical horns of the 
Domesticated Sheep, are more subject to arrest in development 
from a variety of causes and are often absent. The apparent 
cause of the reduplication in the horns of this breed, in a consider- 
able number of instances, is the splitting or segmentation into two 
or more nodules of the centre of ossification of the frontal bone, 
as indicated by three examples exhibited and by the skull (text- 
fig. 9, E). It is upon this point or ‘ boss’ that the horns of all 
Ruminants ultimately grow, its prolongation forming the ‘ pedicle’ 
of the Cervine horns and the ‘ horn-bearer’ or core of the Hollow- 
horned Ruminants. 

‘“‘The lateral horns, on the other hand, though bearing a closer 
resemblance in form and. position to the typical horns of the 
Domestic Sheep, are variable and erratic in their development. 
Frequently only one is present (as in text-fig. 9, C), or they may be 
altogether absent, leaving only the median pair, which then 
assume a typical form (text-fig. 9, A). 

‘“‘ In-breeding, as recorded by the history of three small herds 
of these sheep, causes a reduction or arrest in development and 
often absence of the lateral horns (as in text-figs. 9, C and D), the 
median or upper pair of horns remaining constant. 

“Castration has a precisely similar effect. The totality of 
evidence gathered from the life-history of the breed would seem 
to indicate that it is the median pair of horns, notwithstanding 
their variation and abnormal appearance, which are homologous 
to those of the Domesticated or Wild Sheep.” 

Examples of three or four horns in the Domestic and Wild 
Sheep or Goat as well as in Cattle and Deer are of sufficient 
interest to the anatomist to merit record. In Proc. Zool. Soe. 
1879, p. 803, there is a good figure of a skull of the Chamois with 
four horns. Inthe Museum of the College of Surgeons there is a 
skull of a Goat with four horns. In the current volume of Proce. 


1903. ] ON A NEW FORM OF GRANT'S GAZELLE. 119 


Zool. Soe. supra, p. 2, there is a figure of a skull of a Muntjac 
showing small supernumerary horns on the pedicle, and, although 
the horns ot the Roebuek vary more than do other Cervine horns, 
examples of four horns are rare. 


Dr. A. 8. Woodward, F.R.S., exhibited photographs by 
Dr. Otto Herz, illustrating the discovery and exhumation of a 
Mammoth in the Government of Jakutsk, Siberia. He also made 
remarks on the specimen, which has now been mounted in the 
Zoological Museum at St. Petersburg under the direction of 
Dr. Salensky. 


Mr. Oldfield Thomas, F.R.S., exhibited some remarkable Gazelle 
skulls and horns from German East Africa, which had been con- 
tributed to the National Museum by Messrs. F. Russell Roberts 
and C. E. Blaine. 

The specimens belonged to the Gazella granti type, but their 
horns were each completely twisted round inwards, as shown in 
the figures (text-figs. 10, 11), so that the tips pointed backwards 


Text-fig. 10. 


Skull and horns of Gazella granti robertsi. (Front view.) 


and outwards and were very far apart. The tw isting was spread 
over the whole length of the horn, and being in the direction 
characteristic of domestic as opposed to wild Boas, gave to the 
animals a remarkably goat-like appearance. In the ‘female the 
horns were also twisted, although not so strongly as in the male. 
Writing from Mwansa, German East Africa, Mr. Roberts 


120 ON A NEW FORM OF GRANT'S GAZELLE. [June 16, 


stated among other things that all of the Gazelles, in the district 
where these were shot, were of the same kind, so that they could 
not be said to be abnormalities; that it was quite possible for the 
animal to be unknown to science, as few sportsmen had ever shot 
before in that region ; that the specimens sent, two males and a 
female, were much above the average in size, though still finer 
ones had been seen; and that the ordinary Gazella granti of the 

thi plains and Kilimanjaro was unknown there. This Gazelle 
was known as the “ Biza.” by the natives, who were Wanyumwesl. 


Text-fig. 11. 


Skull and horns of Gazella granti robertsi. (Lateral view.) 


‘The colour of this Gazelle appeared to agree in all respects 
with that of the typical form of @. granti, having the undivided 
white rump-patch, the obsolete lateral bands, and the narrow but 
distinct pygal band of that form*. The fur was somewhat more 
wavy than in ordinary specimens, 

Besides those seen by Messrs. Roberts and Blaine, two heads 
SuOSSaLn als similar characters had been obtained by Herr Oscar 
Neumann in the Loita Mountains, on the boundary of British 
and German Kast Africa, but had been not unnaturally regarded 
by him as abnormalities +, 

Tn spite of the very unusual nature of the distinguishing charac- 
ters of this Gazelle, and its identity with Gazella granti in other 
ae Mr. Thomas thought that the number of examples that 
had now been recorded, and the fact that this type alone inhabited 
certain districts and had never been found elsewhere, necessitated 


i & P. Z. = 1900, p. 806. By a lapsus calamé the name of the new subspecies 
ae vec bed (Gazella g. brightii) was erroneously written G. g. smithii in the 


t Zool. Jahrb. Syst. xiii. p. 561 (1900). 


92 AS. IOS voll, Me PX. . 


G.M. Woodwand,del . 


LOWER JAWS AND TEETH OF (1), GENETTA GENETTA, (2-6) GENETTA 
PLESICTOIDES, sp. nov. @) PoASMOMS CROIZE TT . 


1903. | ON AN EXTINCT GENET FROM CYPRUS. 12] 


its recognition as a subspecies; and he therefore proposed for it 
the name of Gazella g. robertsi, in honour of the donor of the 
specimens to the National Museum. The type would be the finer 
of the two males sent home by Messrs. Roberts and Blaine, 
B.M. No. 3.6.12.1. 

In this specimen the horns measured 233 inches along the 
(morphological) anterior curve, their tips were 233 inches apart, 
and the basal length of the skull was 9? inches. Mr. Roberts 
quoted other examples having horns 244 inches long, with a spread 
of 272 and 28 inches. 


The following papers were read :— 


1. On an Extinct Species of Genet (Genetta plesictoides, sp. 1.) 
from the Pleistecene of Cyprus. By Dororay M. A. 


BatE*. 
[Received May 12, 1903.) 


(Plate Xe) 


In October, 1901, I began my search for Pleistocene bone-caves 
in the island of Cyprus, and for the next few months confined my 
attention to carrying on some work near the Monastery of Aghios 
Chrysostomos in the north, and to excavating in several caves in 
the south-east of the island. 

In the following January I first discovered Dikomo Mandra, a 
cave containing an extensive deposit of Hippopotamus remains, 
and the largest found in the Kerynia range of limestone-hills, in 
the north of the island. However, it was not until April 1902, 
after receiving a grant from the Royal Society, that I was 
enabled to begin work here. This proved to be the only cave in 
which the remains of any carnivorous animal were found, other 
than those of the Fox stillliving in Cyprus. The remains obtained 
appear to be those of an extinct species and consist of a few limb- 
bones, a small piece of aright mandibular ramus with the posterior 
half of the carnassial, and a left mandibular ramus in which the 
incisors, the canine, and the last molar are missing. These portions 
of jaws, differing somewhat in size and wear, are evidently those 
of two individuals. 

At the back of the cave, and some feet above the floor, was a 
mass of rock and earth containing a number of Hippopotamus 
bones. Work was begun here, an attempt being made to extricate 
a skull of this animal which could be seen embedded in the matrix. 
As soon as some of this rock had been broken away, several rents 
and fissures were found filled, partly with earth which must have 
been there since the time when the cave was inhabited by these 


* Communicated by Hrnry Woopwakzp, LL.D., I'.R.S., V.P.Z.5. 
+ For explanation of the Plate, see p. 124. 


122 MISS DOROTHY M. A. BATE ON [June 16, 


extinct mammals, and partly with earth and rubbish that had 
filtered down through the cracks in the rocks above. This 
accounted for the fact that the fossils in question, which are pro- 
bably of the same age as Hippopotamus minutus, were found 
together with many teeth and bones of the Goat, several small 
rodents, and other recent species. 

On comparing the left mandibular ramus (PI. X. fig. 2), 1t appears 
to be that of a carnivore nearly allied to Genetta genetta, which is 
still found living on the opposite shores of Palestine. On the 
other hand, it presents many similarities to Plesictis croizeti of 
the Oligocene deposits of France, a mandible of which was 
originally figured by M. Pomel from a specimen found in the 
Department of Allier (Bull. Soc. Géol. France, t. iv., 1846-47, p. iv). 
A more perfect lower jawbone was procured by Dr. Forsyth Major 
from the same deposit, and both are now in the collection of the 
British Museum. The genus Plesictis is variously placed by 
different authors—by some among the Viverridw and by others 
among the Mustelide. In a work published in 1855 *, M. Pomel 
places Plesictis with the Viverride, while Mr. Lydekker, in the 
‘Catalogue of Fossil Mammalia’ (Brit. Mus. i. p. 185), includes 
the genus among the JI/ustelide, at the same time saying that ‘In 
the above mentioned general characters the genus indicates 
viverrine tendencies, and the transition to the extinct Stenoplesictis 
is so gradual that the viverrine and musteline families are 
practically united by the two genera.” 

The Cyprus fossil agrees with, and at the same time differs from, 
both G. genetta and P. croizeti, and that so impartially, that it is 
a matter of extreme difficulty to decide with which group it ought 
most properly to be associated. The scanty material adds to this 
uncertainty, which would probably be removed were the skull and 
upper dentition of this species known. However, in consideration 
of its much more recent age compared with that of the Oligocene 
fossil, it is proposed, at all events for the present, to include it 
among the Genets under the name of 


GENETTA PLESICTOIDES, sp. n. (Plate X. figs. 2-6.) 


The mandibular ramus is intermediate in size between those of 
(. genetia (Pl. X. fig. 1) and P. eroizeti (Pl. X. fig. 7), being more 
robust than that of the former and slightly less so than that of the 
latter. The anterior margin of the inner aspect of the coronoid 
process 1s somewhat deeply excavated, a feature which is found in 
neither of the last-mentioned species. The three posterlor pre- 
molars and first molar are considerably worn; the small anterior 
premolar is rather damaged and its shape consequently uncertain ; 
while the second molai is absent, though the clearly defined 
alveolus shows it to have been furnished with a single root. 

_ The lower carnassial is distinguished from that of P. croizeti, 
G. genetta, and other Viverride in the area between the three 


* Cat. Vert. Foss... . de la Loire, etc. (Paris, 1853). 


1903. ] AN EXTINCT GENET FROM CYPRUS. 123 


anterior cusps being less deeply excavated, and in the edge of the 
anterior lobe being less broadly expanded, though this last may be 
partly due to wear. This tooth also differs in the size of the imner 
cusp, which is lower than in the older form and higher than in 
G!. genetta, in which it is much reduced. The talon is small and 
has a raised posterior edge. 

The second, third, and fourth premolars are of considerable length 
antero-posteriorly, and do not narrow towards their apices to so 
great an extent as do those of P. croizeti and generally those of 
Genetta. The outer edge of the carnassial is strongly convex in 
shape, and as a result of this there is a deep angle between the 
anterior portion of the tooth and the small posterior talon. This 
is not so marked in P. croizeti, in which the position of the angle 
is filled by aridge. Although the carnassial of G. genetta does not 
show this convexity, there is nevertheless a very slightly developed 
cusp between the front of the tooth and the talon. 

Besides those characters in which @. genetta and P. croizeti agree 
together and differ from the Cyprus fossil, this last is further 
distinguished from the older form by having a less well-marked 
cingulum round the carnassial and the two posterior premolars, 
and in the crowns of these teeth being somewhat lower. Also the 
faintly marked notches on the anterior and posterior edges of the 
second and third premolars of G. plesictoides are absent in 
P. croizett. 

Below are some measurements, given in millimetres, taken from 
the left mandibular ramus of the Cyprus fossil and of the recent 
Genet, and from the two right mandibular rami of P. crovzete in 
the collection of the British Museum, both of which are from the 
Oligocene of France. 


G. genetta. |G. plesictoides.| P. croizeti | P. eroizeti. 
| (type). 
Antero-posterior length of lower! | 
CALMASSI Ali Mesy cere tate eal 75 | 8 9 8 
Greatest width of lower carnas- | 
| Ant.-post. length of pm. 4...... | 6 | 7 7 65 
Ps eo 1981, Blooadoe 6 6d 6 55 
3 ee TID Arosove 5 | 55 4°5 
From ant. margin of pm. 2 to | 
| post. margin of carnassial ... 25 | 27 27 approx. 26 
| From lower margin of ramus 
| to top of coronoid process... 24 | 245 ak 28 
| Thickness of ramus below car- | 
MASSA secs he eee eee | A, | 5 5d 5 
_ Approximate height of carnassial| 5 | 6 65 65 


The limb-bones procured belong, presumably, to G. plesictoides, 
but probably all are not those of the same individual. I have 
compared them with the corresponding bones in a skeleton of 
G. senegalensis in the collection of the British Museum. They 


124 ON A NEW FISH FROM BRITISH NEW GUINEA. [June 16, 


include the proximal halves of two humeri, which are of about 
the same size as those of the African species, while a calcaneum is, 
on the other hand, distinctly smaller, being 19 mm. in length as 
compared with 21 mm. in the recent species. This is also the 
case with the radius, of which two specimens were found, it being 
more slender and 59 mm. in total length as opposed to 61°5 mm. 
Besides these, three pieces of ulne and several portions of ribs 
were obtained. 

T have been unable to find record of any fossil Genetta, and 
among the rest of the Vivermide the only species of Pleistocene 
age appear to be V’. kariauliensis from India (Lydekk. Mem. Geol. 
Surv. Ind. 1886), and a Viverra identified by a single canine from 
the cave of Lunel Viel, France. 

Besides these, some portions of skulls found in the Quaternary 
deposits of Algeria are referred somewhat doubtfully to Herpesies 
by M. Pomel (Carte Géol. de Algérie, Les Carnassiers, 1897). 


EXPLANATION OF PLATE X. 


. Inner view of left mandibular ramus of Genetta genetta from Mt. Carmel, 
p. 122. 

. Inner view of left mandibular ramus of G. plesictoides, p. 122. 

Inner view of a right carnassial of G. plesictoides, p. 123. 

. Outer view of the specimen shown in fig. 2, p. 128. 

. Crown view of the specimen shown in figs. 2 & 4, p. 123. 

Crown view of tooth shown in fig. 3. 

. Inner view of right mandibular ramus of Plesictis croizeti Fiihol, p. 122. 
All the figures are of natural size. 


= 
we 
faa 


WED OLE 09 09 


2. Description of a new Fish of the Gobiid Genus Rhiac- 
ichthys from British New Guinea. By G. A. BoULENGER, 
vases ieee Zs. 

[Received May 13, 1903. | 
(Plate X1.*) 

RHIACICHTHYS NOVE-GUINEH. (Plate XI.) 

Depth of body nearly equal to length of head, 5 to 6 times in 
total length. Diameter of eye 6 or 7 times in length of head, 
interorbital width 3 times; snout but very slightly longer than 
postocular part of head. Dorsals VII, I 8-9; longest spine 3, 
longest soft ray 4 length of head. Anal I 8-9, longest ray as 
long as head. Pectoral about 13 length of head, ventral as long 
as head or a little longer. Caudal feebly emarginate. Caudal 
peduncle 23 as long as deep. Scales strongly ciliated, 37 to 39 
in a longitudinal series on each side, 14 or 16 round caudal 
peduncle. Dark olive above, whitish beneath. 

Total length 225 millim. 

_ Three specimens were obtained by Mr. A. E. Pratt at Dinawa, 

Owen Stanley Range, at an altitude of 4000 feet. 


* For explanation of the Plate, see p. 125. 


“WAHNIND-WAON SAHLHOIDVIHSa 


‘duit soig wrap “WATT 3® “[ep wessyg (~ 


IDO Wel IE’ SOG: Z al 


EZ, S 03 voll eae 


MinternBros ump. 


2) 15. GEAIN We sega 


J.Green delet hth. 


1.LYGOSOMA MILNENSE. 


3.L.PULCHRUM. 


P45). WIS. <7ell IMP, CuI. 


2b. wo. 3b. 
J.Green del.et lth. : Mintern Bros .imp. 
LLYGOSOMA PRATTI. 


2.TOXICOCALAMUS LONGISSIMUS. 
3.T. STANLEYANUS. 


1903. ] ON NEW REPTILES FROM BRITISH NEW GUINEA. 125 


The discovery in New Guinea of a fish of the genus Rhiac- 
ichthys Blgr. (Platyptera C. & V.), a type of Gobiide so 
admirably adapted to life in mountain torrents, is a very in- 
teresting addition to our knowledge. The type of the genus, 
hk. aspro, C. & V., which differs from L. nove-guinee in the 
larger eye situated much nearer to the gill-opening than to the 
oat of the snout, inhabits Bantam, Gelehes, and Luzon*, whilst 
a doubtful species, 2. sinensis Blkr., is founded on a Chinese 
drawing described as “ dubic exactitudinis.” 


EXPLANATION OF PLATE XI. 


Rhiacichthys nove-guinee, with upper and lower views of head and anterior 
part of body, reduced # nat. size. 


3. Descriptions of new Reptiles from British New Guinea. 


By G. A. Boutunezr, F.R.S., V.P.Z.S 
[Received May 13, 1903. } 
(Plates XID. & XTII.7) 


LiyGosoMA MILNENSE. (Plate XII. fig. 1.) 


Section Ainulia. Head large, especially in the male, body 
short; the distance between the end of the snout and the fore 
limb is contained once to once and one third in the distance 
between axilla and groin. Snout short, obtuse. Lower eyelid 
sealy. Nostril pierced in a single nasal; no supranasal ; a single 
anterior loreal; rostral forming a very long, curved suture with 
the frontonasal, which is nearly twice as broad as long; pre- 
frontals forming a median suture; frontal much teEnoreed pos- 
teriorly, longer than frontoparietals and interparietals together, 
in contact with the three first supraoculars; five supraoculars, 
first longest; 10 or 11 supraciliaries, first largest; frontoparietals 
and interparietal distinct, nearly equal in length; parietals 
forming a suture behind the interparietal; nuchals absent or 
reduced to one pair; third, fourth, and fifth labials below the 
eye. Har-opening oval, nearly as large as the eye-opening ; no 
auricular lobules. Scales smooth, dorsals lar gest, 30 or 32 round 
the middle of the body. Preanals slightly enlarged. The hind 
limb reaches the shoulder, or between the shoulder and the ear, 
Digits rather elongate, slightly compressed ; subdigital lamellee 
smooth, 35 to 37 under the fourth toe. Tail once and a half to 
once and two-thirds the length of head and body. Coloration 
very variable. Upper parts uniform brown, or with a light, 
dark-edged dorso-lateral streak joining its fellow on the base of 


* T have not been able to find on any map the locality “ Wanderer Bay” given 
by Giinther, Cat. Fish. iii. p. 138. 
+ For explanation of the Plates, see p. 129. 


126 MR. G. A. BOULENGER ON [June 16, 


the tail, or with dark brown spots forming bars across the body 
or restricted to the sides; three or four large black spots, with 
white dots between them, may be present on each side of the 
neck; a black canthal and temporal streak sometimes present ; 
lips uniform whitish, or blotched with brown or black ; lewer 
parts whitish, throat sometimes blotched with dark brown. 


3. OF 

Motalglenethi-ce-eeepacecrer 173 165 millim. 
Taleryel) ggdanndcosseqou0..q00090 20 as 
Wadthvot shea) tea epensces 14 OF ater tone 
IGM onanbouocougdosoc00dcn. 48 44, 
Words) JUAN aasanooceobsoosco: 27 2 
Jelsnavel IbboDlo) Se6cecsoccecooss0" 40 Oe 
Pall i sgus chine rep idecetres Ser: 105 104s anes 


Several specimens from Milne Bay, British New Guinea, col- 
lected by Mr. A. S. Meek. 

The nearest allies of LZ. milnense are L. concinnatum Bler., 
from the Solomon Islands, and Z. simwm Sauv., from New Guinea. 


LyGosoMA GRANULATUM. (Plate XII. fig. 2.) 


Connecting the Sections Hinulia and Otosaurus. Habit lacerti- 
form; the distance between the end of the snout and the fore 
limb is contained once and one third in the distance between 
axilla and groin. Snout very short, obtuse. Lower eyelid scaly. 
Nostril pierced in a single nasal; a very small supranasal, between 
the frontonasal, the nasal, and the first loreal, which is single ; 
rostral forming a straight suture with the frontonasal, which is 
much broader than long and touches the anterior angle of the 
frontal; latter much narrowed posteriorly, a little longer than 
frontoparietals and interparietal together, in contact with the 
three first supraoculars; seven supraoculars, first more than 
twice as long as second, seventh very small; eleven supraciliaries, 
first largest; frontoparietals and interparietal distinct, the former 
a little longer than the latter ; parietals forming a suture behind 
the interparietal; no nuchals, each parietal in contact with four 
scales; fourth, fifth, and sixth upper labials below the eye. Ear- 
opening large, oval, as large as the eye-opening; tympanum 
scarcely sunk ; no auricular lobules. 36 scales round the middle 
of the body, laterals smallest, dorsals slightly larger than ventrals, 
dorsals and laterals rough with minute granular asperities. A pair 
of enlarged preeanals. The hind limb reaches the axilla. Digits 
elongate, compressed, obtusely keeled below; 20 lamelle under 
the fourth toe. Tail about once and a half the length of head 
and body. Pale reddish brown above, variegated with dark 
brown ; sides with large dark brown spots, some forming vertical 
bars; a white dot above the tympanum, another above the 
shoulder; sides below the large spots reticulated with dark brown; 


1903. | NEW REPTILES FROM BRITISH NEW GUINEA. NDZ 


lips spotted with dark brown; lower parts whitish, throat almost 
entirely dark brown. 


Total length ...... Jhilyy tenulibwen’, |) Lk@res Ikan) Sosenneos 14 millim. 
Heads (eat: saSuies aioe DN aes Jahuadl linens) soseenee De 
Wadthwote head c) /ie ae 1 Uzi lah sat eae Cir Abare ae 
BOL; eeteseaece sas Byala Aan 


A single specimen from the Albert Edward Range, 6000 feet, 
collected by Mr. H. 8. Rohn. 
Nearest ally: Z. annectens Blgr., from New Guinea. 


LyGosoMA PULCHRUM. (Plate XIT. fig. 3.) 


Section Liolepisma. Habit lacertiform, slender; the distance 
between the end of the snout and the fore limb is contained once 
and one fourth in the distance between axilla and groin. Snout 
moderate, pointed. Lower eyelid with an undivided transparent 
disk. Nostril pierced in the middle of a rather large nasal; no 
supranasal; anterior loreal as deep as the nasal; frontonasal 
broader than long, forming a short straight suture with the 
rostral ; preetrontals meeting with their inner angles, or narrowly 
separated ; frontal small, acutely pointed behind, in contact with 
the first and second supraoculars; frontoparietal single, as long 
as the frontal; interparietal distinct, about half as long as the 
frontoparietal ; four supraoculars ; eight supraciliaries; parietals 
in contact behind the interparietal; three to five pairs of nuchals; 
four upper labials anterior to the subocular. Har-opening roundish, 
smaller than the eye-opening; no auricular lobules. 24 scales 
round the middle of the body, perfectly smooth; dersals, especially 
the two vertebral series, largest, laterals smallest. Two strongly 
enlarged preanals. The hind limb reaches the elbow of the 
adpressed fore limb. Digits slender, a little flattened at the 
base, fourth toe much longer than third; subdigital lamelle 
smooth, 22 under the fourth toe. Black above, with five white 
longitudinal lines, commencing on the snout, the vertebral nar- 
rowly interrupted behind the head and ending a little before 
reaching the sacral region; an interrupted additional line on 
each side of the body between the vertebral line and the upper 
lateral; limbs reddish, spotted with black, with regular black 
bars across the digits; tail coral-red, with a regular series of 
black spots or vertical bars on each side; lower surface of head 
and body greenish white. 


leads) 22 ee Sholay | Horewlimib 4.522. 4-2 11 millim. 
Nvidth of heady sy Ons Jabuar) Ibo), caoyonacs Le es 
TROGRY Wadeddcqdedonoc 7A i 


Two specimens, both with the tail injured, from the Albert 
Edward Range, 6000 feet, collected by Mr. H. 8S. Rohn. 
Nearest ally: ZL. pulchellwm Gray, from the Philippines. 


128 ON NEW REPTILES FROM BRITISH NEW GUINEA. [June 16, 


Lyqosoma pratt. (Plate XIII. fig. 1.) 

Section Lygosoma. Body elongate, limbs short; the distance 
between the end of the snout and the fore limb 1s contained once 
and three fourths in the distance between axilla and grom. Head 
small, much depressed ; snout moderate, obtusely pomted. Lower 
eyelid scaly. Nostril pierced in a single nasal; no supranasal ; 
vostral narrowed and produced posteriorly between the nasals, 
forming a narrow suture with the frontonasal, which is a little 
broader than long and narrowly in contact with the frontal ; 
latter as large as frontoparietals and interparietal together, much 
broader than the supraocular region, in contact with the first 
supraciliary, the first supraocular, and a very small part of the 
second ; four supraoculars; seven supraciliaries; frontoparietals 
and interparietal distinct, subequal; parietals forming a suture 
behind the interparietal; no nuchals; fourth and fifth labials 
below the eye. Har-opening oval, nearly as large as the eye- 
opening; noauricular lobules. 36 smooth scales round the middle 
of the body; dorsals, especially the two vertebral series, largest. 
A pair of slightly enlarged preanals. Limbs widely separated 
when pressed against the body. Digits short, compressed, keeled 
below ; subdigital lamelle mostly divided, 13 or 14 under the 
fourth toe. Tail very thick. Pale brown above, mottled or 
vermiculate with dark brown; head and nape blackish; two 
oblique white streaks from below the eye to the throat; belly 
white. 


Total length ...... 162 millim. | Fore limb ......... 14 millim. 
HRelsevols, ety eer te We Way iia delimibee eee 21 
Wadth of heads :.. (10)- 5 Tail (reproduced) . 75 
Bod yoni oe beta 72 


PP] 
99 


This new species, of which a single specimen was obtained at 
Dinawa, Owen Stanley Range, 4000 feet, by Mr. A. E. Pratt, is 
most nearly related to the Papuan JZ. muelleri Schleg., and 
clearly belongs to the same section of the genus Lygosoma. But 
it is also closely allied to LZ. lorie Bler., from New Guinea, which 
has been referred to the section Hinulia and evidently constitutes 
a connecting-link between the two sections. 

Having, through the kindness of Mr. Thomas Steel, of Sydney, 
had the loan of the type of Homolepida englishi De Vis, I am 
able to confirm its identity with L. mwelleri, as already pointed 
out by me in the ‘ Zoological Record’ for 1890. 


TOXICOCALAMUS STANLEYANUS. (Plate XIII. fig. 3.) 


Rostral much broader than deep, just visible from above; inter- 
nasals nearly as long as the preefrontals, which are in contact with 
the second upper labial and with the eye; frontal small, slightly 
broader than the supraocular, once and three fourths as long as 
broad, as long as its distance from the end of the snout, a little 
shorter than the parietals; one postocular; temporals 1 + 2: 


E ZS 1908) val ee ay 


Huth, Lith. London. 


C. Crossland del. 


OUN UIP ROS: LONE OS TRUAINS GSU TE vA. 


Wied. DQ UAT RUA IN IEJAIe © JL TAIN AL 


S) 
a. 


SSS, MUNIRUP NCS As IMUAIG TUNE OSs. sa. sya. 


P. Z.S 1903, vol. Il: Pl. XV. “ae 


G.Crossland del. Huth, Lith. London. 


ene? 7/0), IMU ARUP In YE SA. IMO) S S/NIMUIE) LOA. 
WZ), IML STUMP WI OS, a 5, 
13,14. M FURCELLATA nsp. 


1903. ] ON THE MARINE FAUNA OF ZANZIBAR, 129 


five upper labials, second and third entering the eye; three lower 
labials in contact with the anterior chin-shields, which are larger 
than the posterior. Scales in 15 rows. Ventrals 261; anal 
entire; subcaudals 25 pairs; tail ending in a compressed, obtusely 
pointed scute, which is obtusely keeled above. Blackish brown 
above; traces of a yellowish nuchal collar; upper lip white; two 
outer rows of scales white, each scale with a blackish central spot; 
ventrals and subcaudals white, with a black spot on each side, 
some of the ventrals with an interrupted blackish border. 

Total length 610 millim. ; tail 40. 

A single female specimen from Dinawa, Owen Stanley Range, 
4000 feet, collected by My. A. E. Pratt. 

The genus Zoxicocalamus was established by me in 1896 (Ann. 
& Mag. N. H. [6] xviii. p. 152), for a Snake from Woodlark 
Island, British New Guinea, 7. longissimus, here figured 
(BE XII. fig. 2), which differs from the one now described a 
the numbers of scales and shields (Ser 173 Vip 299=305)7 Ay 2 
C. 30-31), in the smaller eye, in the shorter internasals ind 
frontal, in the proportions of the upper labials, of which the 
third and fourth enter the eye, and in the coloration. 


EXPLANATION OF THE PLATES. 
PrarTe XII. 


Fig. 1. Lygosoma milnense, p. 125, male, natural size. 
2. nis ae p. 126, natural size. 
2a. Upper view of head, X 2. 
3. Lyg gosoma pulchr win, p- 127, natural size. 
3a Upper view of head, x 25. 


99 32 


PuatE XIII. 


Fig. 1. Lygosoma pratti, p. 128. 
Figs. 2,2a,2b. Towxicocalamus longissimus, p.128. Upper, side, and lower views 
of head and anterior part of body. 
3,3a,36. Toxicocalamus stanleyanus, p.129. Upper, side, and lower views 
of head and anterior part of body. 
All natural size. 


4. On the Marine Fauna of Zanzibar and British Hast Africa, 
from Collections made by Cyril Crossland in the Years 
1901. and 1902.—Polycheta. Part II. By Cyrin 
Crosstanp, B.A., B.Sc.* 

| Received May 25, 1903. } 


(Plates XIV. & XV.7, and Text-figures 12-15.) 


Part IJ.—THe EUNICID4. 


Before beginning my examination of the typical Nereidiform 
(i.e. Evrant) Polychetes, I investigated to some extent the 
variability of those characters usually employed for specific 

* Communicated by Prof. W. C. McIntosu, F.R.S., C.M.Z.S 


+ For explanation of the Plates, see p. 144. 
Proc. Zoou. Soc.—1903, Vor. IT. No. TX. 9 


130 . MR, CYRIL CROSSLAND ON THE [June 16,. 


distinctions, the proportions of which are described numerically 
and with great minuteness by some of the best workers. How- 
ever one may admire the care which has been employed, and 
however desirable the use of such numerical statements may be, 
my results showed clearly that they are, from the nature of the 
case, unpractical. As their employment can only render descrip- 
tions more cumbrous, and the limits of the species so defined 
more hazy even than in nature, I forego all minuteness in state- 
ments of proportion and numbers. Accurate figures, drawn to 
scale, will provide as full and unmistakable accounts of species as 
is possible by the medium of ink and paper, the extent of variation 
rarely being so great as to render a drawing thus prepared from 
one specimen not recognisable at once as a likeness of any other 
member of the same species. 

The deductions from my tables of measurements of specimens 
of Marphysa mossambica ave shortly as follows :— 


(1) The body shape is generally constant, but the numerical 
position of the widest segment may be very different in 
certain individuals. 

(2) The proportions of pro- and peristomium 7 se and iter se 
are roughly constant. 

(3) The length of the unpaired tentacle is variable within 
limits, but the number of the segment to which it reaches 
when laid along the back is not worth giving in a descrip- 
tion of the species. 

(4) The gills, as often noted, begin on different segments in 
different specimens, increase in size and complexity quite 
irregularly, and vary in the maximum number of the 
filaments they bear. 

In another examination, all the smaller gills were left 
out of account, the distance from the anterior end to the 
point where gills of approximately the full size begin to 
occur being measured. This varies from 5 to 6 cms.; the 
number of segments it contains varies from 64 to 80. 

(5) The number of teeth borne by the dental plates frequently 
varies by one on either side of the mean, 7. e. for the larger 
plates from 4 to 6. 


In Marphysa belli, a species which is characterised by the large 
size and the concentration of the gills toa few segments, these still 
vary in the same ways. ‘Two specimens, collected and preserved 
together *, differed as follows :— 

Though both are 3°5 mm. in breadth the segments in no. 1 are 
much the shorter. In no. 1 the gills do not meet over the back, 
though no. 2 agrees with the specific definition in this respect. 
No. 1 bears 17 pairs of gills on feet 13 to 30 tT, and no. 2 21 pairs 

* Lowe my opportunity of examining this species to the kindness of Prof. McIntosh. 


The specimens were dredged by the ‘ Porcupine’ in 1870 trom 81 fathoms, off Cape 
Finisterre. 


+ In all cases the first setigerows segment is counted as the first, a method of 
reckoning which is free from any possible ambiguity. 


1903. ] MARINE FAUNA OF ZANZIBAR 131 


on feet 14 to 35. The prostomium is of nearly the same size in 
both specimens, yet the unpaired tentacle, when laid along the 
back, extends beyond the anterior border of the fourth segment, 
or only up to that of the second segment. The colour of the two 
specimens differs markedly, being much duller in no. 2. 

Similar results were obtained in the case of Diopatra neupoli- 
tana, in which species such variations are much more conspicuous 
in accordance with the great development of the organs concerned. 
The resulting differences of facies have caused some synonymy, 
and have made some authors content to give quite insufficient 
descriptions. 


The OnupPHIDINA, 


The genera Khamphobrachiwm (Ehlers), Onwphis (Aud. et Ed.), 
Hyalinecia (Malm), and Diopatra (Aud. et Ed.) form a very 
well-marked group characterised by modifications for a perma- 
nently tubicolous mode of life. 

The last three genera are differently defined, however, by the 
principal authorities. All authors but Ehlers agree in separating 
the genus Diopatra because of its very characteristically formed 
and distributed gills. In other cases Ehlers would be theoretically 
right in objecting that the gills of Annelids are too variable to 
be made a basis for generic distinctions, and in urging that the 
gill-less species are derivable from either Onuphis or Diopatra, if 
the distinction between these genera is maintained. Practically, 
however, we find no form with gills intermediate in structure 
between those of Diopatra and any other Kunicid whatever; and 
this distinction is emphasised by the fact that no other member 
of this group has its largest gills confined to the anterior part of 
the body. There is no real necessity, therefore, for the confusion 
introduced by assigning the two gill-less species, fragosa* and 
glutinatri«*, and the forms with pectinate gills posteriorly, 
pourtalesu*, magnayt, and dorsalist, to the genus Diopatra 
instead of to the genera Paronuphis and Onuphis, as would have 
been done by any other writer. 

The confusion in the definitions of the genera Hyalinecia and 
Onuphis has its origin simply in the name of the former. The 
addition of a coating of mud or larger pieces of foreign material 
to the foundation of the tube secreted by the body of the worm 
cannot be regarded as a sufficient cause for generic distinction, 
however striking the difference in appearance of the tube. The 
name Hyalinwcia may be retained when not literally applicable, 
the transparency of their abode being characteristic of the great 
majority of the species. 

The character of the gills forms the basis of a distinction 
between nearly all the species of the two genera, but, as else- 
where, all stages are found between the typical comb-like gill of 


* Ehlers, “ Annelids of the U.S. Survey ship ‘ Blake,’ ”’ Cambridge, Mass., 1887. 
+ Andrews, Pr. U.S. Nat. Mus. xiv. p. 277. 


+ Ehlers, ‘ Hamburger Magal. Sammelreise,’ p. 71 (1897). 
Q* 


132 MR. CYRIL CROSSLAND ON THE [June 16, 


Onuphis and the cirvitorm gills of Hyalinecia, or even the complete 
absence of gills (Paronuphis), so that In some cases the only 
distinction available is that provided by the presence or absence 
of tentacular cirri. 

Genus DIOPATRA. 


DIOPATRA NEAPOLITANA Clap. (Plate XIV. fig. 1.) 


Since Grube’s enumeration in 1877* of eighteen species but 
few fresh descriptions have been published (Langerhans, D. madew- 
ensis +; Ehlers, D. chilensist, redescribed). This fact, and the 
very local distribution of the species given, probably indicate the 
existence of a very considerable amount of synonymy, though 
Kinberg’s seven species, assuming the correctness of his figures, 
are distinct. It is thus the more remarkable that the species 
should have so wide a distribution as from the Mediterranean to 
the East coasts of Africa and North America §. 

- The species is so abundant at low spring-tides on any sandy or 
muddy shore in East Africa, that it is strange that it has not been 
before recorded from this locality. The material of the tubes 
varies with the habitat—on a sandy beach being built of shell- 
fragments or small stones, the edges always projecting horizontally, 
on muddy shores the grass-like leaves of Zostera or any suitable 
vegetable fragments being employed. Only the projecting and 
the upper two or three inches of the buried portions are thus 
strengthened, the remainder being soft and collapsable. 

Dark green and umber-brown coloured varieties of the animal 
occur, both colours being destroyed by a brief immersion in alcohol. 
The only specimen I obtained from below the level of lowest tides 
(from 10 fathoms in Wasin Harbour, the mainland coast) shows a 
third distinct variety ||. 

The living worm was red- (not umber-) brown anteriorly, the 
colour remaining distinct after immersion in strong spirit for 
more than a year. ‘The gills were red, the colour of the blood 
not being hidden by green and brown pigments as in the 
preceding varieties. Structurally, I find this specimen to agree 
with the shore forms. The presence of similar pigmentation in 
preserved specimens obtained from Naples suggests that this 
colour variety occurs there also, and is probably the variety 
of colour which is described by Claparéde as “ ferrugineuse.” It 
is possible, of course, that green, umber, and red-brown pigments 
may occur occasionally in the same individual, though I have not 
seen cases of this in East Africa. 

The lack of perfeet completeness in Claparéde’s account, and the 
obviously diagrammatic nature of at least two of his figures, made 
it impossible to decide whether I was examining a nearly related 
species or a variety only. A comparison of specimens obtained 
from Naples with my own from Zanzibar has enabled me to 
* *Mitth. tiber die Familie der Euniceen,’ Naturw. Schles. Ges. 1877. 

+ Z. Wiss. Zool. xxxiii. pp. 513-593. 
‘Polych. des Magal. u. Chil. Strandes,’ 1901. § Andrews, loc. cit. 
|| Very numerous tubes were dredged from 3 fathoms in one spot in Chuaka Bay. 


a 
+ 
its 


1903.] MARINE FAUNA OF ZANZIBAR. 133 


supplement the earlier description and in a few details to correct 
it. As, however, a complete description, by Professor McIntosh, 
of specimens obtained by the ‘ Porcupine’ * has recently appeared, 
there is no need for me to do more than summarise the most 
definite points. 

Tf the proportionate sizes of gills and tentacles were ever 
approximately constant, surely they would be so in this species, In 
which their large size is so characteristic. I find, however, that 
the first gill, which is usually on the fourth foot, is often on the 
fifth. (Claparéde and Grube give the fifth as the first gill-bearing 
foot.) The ringing of the gill-bases, upon which stress is laid in 
Grube’s tabulation of the species, may be quite obvious or only to 
be made out by very careful examination. The anterior feet are 
ringed very faintly. The last gill-bearing segment may be any 
one between the fiftieth and sixtieth, and in one case it was the 
fiftieth on the right side, the fifty-fifth on the left. The gills, 
when laid forward, may extend either to the middle of the first 
setigerous segment or beyond the front of the prostomium. The 
buccal and first three setigerous segments are of about the same 
length, but those succeeding rapidly shorten, so that numbers four 
to eight or twelve are the shortest in the body. 

The palps are very large, together forming an area ereater than 
that of the prostomium itself. Between them is a deep and 
narrow groove bounded anteriorly by a tubercle, which leads back 
a little dorsally to the cesophagus, separation of which from the 
jaw-apparatus 1s effected by a pair of lips bearing large tubercles 
(Pl. XTV. fig. 1). 

T do not find in either set of specimens any setee corresponding 
to Claparéde’s fig. 4, pl. vi. of the ‘ Annélides du G. de Naples.’ 
The compound setz (‘‘ soies incomplétement composées ”’) of the 
first three feet have invariably two hooks, which are quite distinct. 
Tn the Naples specimens the comb-setie have few teeth, and these 
are remarkably broad and flat ; the capillary sete, like the aciculee, 
are gently bent near their ends, a fact which led Claparede to 
describe them as being bordered or ending in a lance-head. This 
is not the case, though their sides are toothed, the teeth usually 
being fine but sometimes large and coarse. 

The Zanzibar specimens agree with those mentioned above except 
that the teeth of the combs are very fine and numerous, and that 
the distal filaments of the gills are almost as longas the proximal. 
In the Naples specimens the shape of the gill agrees with 
Claparéde’s figure in the shortness of the distal filaments. These 
differences, though constant, do not warrant the creation of a 
specially named variety. 

At Prof. McIntosh’s suggestion I append an account of further 
observations on the variation of the special anterior feet, to which 
importance has been attached by systematists. Prof. McIntosh 7 


* “Notes from the Gatty Marine Laboratory,” Aun. & Mag. Nat. Hist. (ser. 7) 
vol. xii. p. 128 (1903). 

+ My thanks are due to Prof. McIntosh for showing me his preparations and for 
an opportunity of discussing them with him. 


134 MR, CYRIL CROSSLAND ON THE [June 16, 


finds in his specimen of this species from Naples that the sete 
described by Claparéde as “‘incomplétement composées ” end in a 
simple hook covered by a guard *, whereas in the ‘ Porcupine’ 
specimen a second hook is present a short distance behind the 
terminal one. Also in the former the gill is borne on the fourth 
foot, in the latter on the fifth, and in both cases, simultaneously 
with its appearance, the ventral cirrus is thickened and shortened, 
presenting an intermediate stage between the normal pointed 
organ of the first feet and the secretory pad which represents 1t 
throughout the remainder of the body. This consensus in the 
variation of three characters seems a sufficient ground for the 
separation of the specimens as two distinct varieties. 

The question, however, now arises as to whether these characters 
always vary together and in the same direction, or whether their 
variations may not occur independently and sporadically. 

As regards the difference between the sete, Claparéde men- 
tions the occasional appearance of a second hook. Both in 
Prof. McIntosh’s specimens and in my own we find that the 
young setze which do not yet project from the foot are always 
provided with the second tooth, which in my Naples specimens 
may be present or absent in those setze which are exposed. This 
indicates that, in spite of the absence of a scar in some cases and 
the unbroken condition of the guard in nearly all, the lack of the 
proximal tooth is always accidental, and is owing to its brittleness 
and to the flexibility of the delicate guard. In the Zanzibar 
specimens a distinct scar is always to be seen in those rare cases 
where the somewhat stouter hook has been lost. 

Tn three specimens from Naples the first gill was borne on the 
fourth, in three others on the fifth foot, the total number of gills 
varying between 37 and 50. Out of nineexamples from Zanzibar 
only two bear their first gill on the fifth foot, and the total 
number is between 46 and 59. The first gill is usually about half 
the length of the largest, but in two cases those borne by the 
fourth feet were only a quarter of that length. In the three 
Naples specimens, the fourth feet of which are devoid of gills, the 
same appendages bear hooked sete and ventral cirri of the normal 
form; but in the three other examples, the fourth feet of which 
possess gills, they bear simple setze and possess ventral cirri which 
are knob-like in form. Of nine Zanzibar specimens, in five cases 
the change in the ventral cirrus takes place on the first, being 
deferred to the second branchiferous foot only in the remaining 
two, and in one of these cases the foot on which this change 
occurs is the sixth. Though I have not found a ease of the 
extension of the hooked sete to the first gill-bearing foot in any 
of the Naples specimens, I find such sete to be present on these 


feet in all those from Zanzibar, even in the two cases where this 
is the fifth foot. 


* As in the lower right-hand seta of those figured by Claparéde. His other figures, 
as noted above, are optical delusions. See Prof. McIntosh’s note on the ‘ Porcupine’ 
specimen in the ‘ Annals and Magazine of Nat. History,’ loc. cit. 


1903. ] MARINE FAUNA OF ZANZIBAR. 135 


Thus, though in the majority of cases the disappearance of 
hooked setze and the change in the character of the ventral 
cirrus are coincident with the appearance of the first gill, excep- 
tions are proportionately numerous, so that variations of these 
characters cannot well be made the ground for systematic 
distinctions. 


Genus ONUPHIS. 
ONUPHIS HOLOBRANCHIATA Marenzeller. (Plate XIV. fig. 2.) 


Five specimens were collected, and in all the hind end was 
missing. The largest fragment is very nearly of the same size as 
that described by Marenzeller from Japan, viz. 4 cm. x 0°3 em., 
and consists of nearly the same number of segments, 85. This, 
with three others about half the size, was dredged from 10 fathoms 
in Wasin Harbour; the fifth, still smaller, being from the shore 
in the same locality. This last specimen is abnormal in having 
no gills on the first two pairs of feet. 

The coloration of the living animal is characteristic, the 
pattern on its dorsal surface serving to distinguish it at a glance 
from any of the numerous small species of Eunicide living 
in the same locality. The ground-colours are of a light flesh-tint 
ventrally and light yellow- -brown. dors sally, but the central part 
of the pr ostomium and a small round area in the middle of 
each segment are white. The upper surface, however, as far as 
the thir ty-fifth segment, is largely covered by markings of a dark, 
rather purple - brown (Marenzeller’s “ Dunkelbraun - violett P 
suggests an almost blue colour, which is not that present in my 
specimens). These are most numerous and closely placed at the 
bases of the feet, with the exception of the first three. On either 
side of the white central marks are slender transverse lines, three 
pairs, one of long, two of short marks to each segment. To the naked 
eye the back appears marked by two pairs of longitudinal bands, 
the outermost darker and of definite zigzag shape, the inner 
which bound the median moniliform white stripe, lighter and less 
definite in outline. The former is omitted from the first four 
segments, and the latter also is irregular there. There are dark 
marks on the prostomium just behind the bases of the tentacles, 
the ringed portions of which are themselves lightly marked. The 
fig. 2, Pl. XIV., shows this pigmentation, the peculiar proportionate 
lengths of the tentacles, &e. 

In feet, setee, gills, and other characters my specimens agree 
minutely with Marenzeller’s. The former, like their gills, are 
white and somewhat dorsally directed. The mandibular plates 
differ slightly in shape from Marenzeller’s figure, having, in the 
cutting-edge, one deep, instead of two shallow notches. 


EUNICINA, 
Genus MARPHYSA. 


The following table, which includes all the species of which 


136 MR. CYRIL CROSSLAND ON THE [June 16, 


intelligible descriptions have been published, may be useful to 
future workers :— 


A, Prostomium undivided. 
1. Gills large, confined to a few anterior segments *. 
M. stragulum, M. belli, M. adenensis. 
2. Gills of moderate size, extending over most of the 


odyis Jeno tlik bcos sehieme ea cerecneec Bammer ete M. macintoshi, sp.nov. 
B. Prostomium bifid. 
a. Capillary sete only present ..... M. mossambica. 


(3. Tentacles three times length of the prostomium. 
1, Compound! setze hooked... <2. 2i::020---0--- eee 
2. Compound sete with knife-like articulated 
LCCESN eee eee eee ete NA nee ee aera Maly LEROY CUGILLCLIEOUIEs 
y. Tentacles not twice length of prostomium. 
1, Gills of one or two filaments only : 


M. goodsiri. 


Compound setz of both kinds.................. MW. fallax. 
ie 5, with hooked appendage ... I. saxicola. 
ee ae »  knite-like appendage 

OWING Sogosseboseseoscan LL, RUMP 


2. Gills of moderate size : 


a. With hooked compound sete............... AMS aN 0c 


M, rvegalis. 
om. 5k Se M. simplex, sp. Nov. 
B. ,,  knife-like compound sete ......... Te Plcallaen pene 


Grube (‘ Mittheilungen tiber die Familie der Euniceen ” Natur- 
wiss. Schles. Ges. 1877) gives numerous species of this genus 
from Schmarda, who, however, described them under the name 
Eunice, in the definition of which genus he said that tentacular 
cirri may be present or absent. In only one case, that of Huinice 
teretiuscula, is it definitely stated that they are absent, though 
they are not mentioned in the three other species which Grube 
enumerates. But four species of J/arphysa and five of Humnice 
in one collection is so disproportionate that I am inclined to 
think that J/. teretiuscula is the only true Marphysa here. In 
any case Schmarda’s descriptions are so short and his figures so 
rough that in most instances it is not possible to compare them 
with others with any certainty. It is quite possible indeed that 
my species WM. simplex is the above-mentioned LZ. teretiwscula, 
both being from the Indian Ocean, but as certainty is not possible 
I have redescribed my species de novo. 

The following references give the best available descriptions 
(mere notes are not inserted here unless important) :— 


M. stragulum V. Grube. ‘ Annulata Semperiana,’ p. 163. 

M. belli Ehlers. ‘ American Annelids’ (U.S. Fishery Survey by 
the ‘ Blake’). 1887. 

M. adenensis Gravier. “ Polychétes de la Mer Rouge,” Arch. 
Nouy. du Muséum de Paris, 1900. 

M. mossambica (M. nove-hollandie) Grube. ‘ Annulata Sempe- 
riana.’ Gyravier, loc. cit. 


M. goodsiri 
We ae } McIntosh. ‘Challenger’ Reports, vol. xii. p. 299. 


_ * Macduffia bonhardi (McIntosh, ‘Challenger’ Reports, vol. xii.) should be 
inserted here. Its main characteristic is the possession of four pairs only of well- 
developed gills. : 


1903.] MARINE FAUNA OF ZANZIBAR. 137 


WM. acicularium Webster. ‘ Annelids of Bermuda,” Bull. U.S. 
Nat. Museum, 25, 1884, p. 319. 

M, fallax Marion et Bobretsky. Ann. Sci. Nat. (6), 1875. 

M. saxicola Langerhans, “ Einige canarischen Anneliden,” Nova 
Acta Acad. Ces. Carol.-Leop., 1881. 

M., striata Kinberg. ‘ Eugenies Resa,’ &e. (These figures without 
text are of far more value than many verbose descriptions 
without figures.) 

M. corallina Ehlers. ‘Die Anneliden des Magellanischen und 
Chilenischen Strandes. Berlin, 1901, p. 131. (Kinberg 
gives figs. of head.) 

M. regalis Verrill. “ Additions to Fauna of Bermuda,” Trans. 
Connecticut Acad. v. pt. 2, 1900. 

M. janwarti Grube. Sitz. der naturf. Gesell. Freunde zu Berlin, 
1881, p. 111. 

M. parishi Baird. Journ. Linn. Soc., Zool. x. 1870. (See note 
also by Ehlers in U.S. Fishery Survey by s.s. ‘ Blake.’) 


MARPHYSA MACINTOSHI, sp. nov. (Plate XIV. figs. 3-6.) 


Three specimens were obtained by digging in sand between 
tide-marks, on both east and west coasts of Zanzibar. The hind 
end is missing from all three, though the largest fragment has a 
length of 20 cms. Their breadth is very uniform (4 mm.), only 
the first half-dozen segments being rounder and narrower. There 
is no regular pigmentation. 

The prostomium is large and undivided, resembling in shape 
the outline of a horse’s hoof. It is fat above, but deeply grooved 
below (Pl. XIV. fig. 3). The tentacles are slender, smooth, and 
pointed, without differentiated basal portions. The median 
tentacle does not quite reach the anterior border of the pro- 
stomium, and the other two pairs, which are inserted close 
together at some distance on either side of it, are considerably 
shorter. Between the bases of the two latter can be made out 
with care a pair of smal! ill-defined eye-spots. 

The mandibles are curved outwards at their tips, their anterior 
ends marked by the lines shown in text-fig. 12, p. 138. The 
maxillary forceps are slender and strongly curved at their points. 
Great dental plates with 4 teeth on the left, 5 on the right, upper- 
most on both sides, are well developed. Curved laterals narrow, 
with 5 and 6 teeth on the left, 7 on the right. 

Feet normally developed, with rounded lip of seta-sac and long 
bunches of sete. The cirri are well developed for a member of 
this genus, their tips extending as far as the level of the seta-sac 
lip. The ventral cirrus has a thick base which almost hides the 
rest of the foot ina ventral view, to.which is joined a smaller 
cylindrical tip, the whole roughly resembling a nereid palp (see 
fig. 4, Pl. XIV.). There are three black acicule anteriorly, 
at a short distance behind the beginning of the gill-region only 
two, with an acicular ‘hook’ seta. This is bent and projects, and 
is simply bluntly pointed (Pl. XIV. fig. 6). Over the greater part 


138 MR. CYRIL CROSSLAND ON THE [June 16, 


of the body the feet are provided with this and one aciculum only, 
The ordinary setz project considerably, especially in the anterior 
feet. Here they are arranged in two distinct bundles of simple 
capillaries dorsally and compound ventrally, with a set ot shorter 
and stouter compound sete between the two. Posteriorly this 
arrangement is less definite, the bundles running into one another. 
The long capillaries are as usual slightly bent, smooth, and end in 
a very delicate point. Long and slender comb-setz occur among 
the more ventral capillaries (Pl. XIV. fig. 5) of the posterior 
segments, the teeth of which, except the two outermost, are hardly 
to be made out even by the f-in. objective. The articulated pieces 
of the compound sete are of the knife-shape. Their proportions 
are shown in Pl. XIV. fig. 5. 


Text-fig. 12. 


Marphysa macintoshi. Mandibular jaw-plates, < 45. 


The gills begin from seg. 32 to 54, attaining their full size 
more gradually in the former case. Their greatest number of 
branches may be four, five or, six, of moderate length, 2. e. they 
can be nearly made to meet over the back. Their arrangement is 
bushy rather than comb-like (Pl. XIV. fig. 6). As the gills are 
present up to the ends of all three fragments, it is most probable 
that they extend to near the anus. 

All the other species of Marphysa in which an undivided pro- 
stomium is met with are characterised by the possession of few, 
and those very large gills. The present species therefore occupies 
a conspicuous position in the genus, as indicated in the above 
table. The shortness, slenderness, and insertion of the tentacles, 
the rudimentary condition of the eyes, and the articulation of the 
ventral cirri are also prominent specific characters. 


1903. | MARINE FAUNA OF ZANZIBAR. 139 


MARPHYSA MOSSAMBICA Peters. (Plate XV. figs. 7-10.) 


Marphysa nove-hollandie Grube, Annulata Semperiana, p. 165. 

M. mossambiea Gravier, Nouv. Arch. Mus. de Paris, 1900; 
also Kinberg & Peters. 

The most complete description of this species is that recently 
given by Gravier, following on, and adding figures to, that of 
Grube. 

The species occurs abundantly in Chuaka Bay, Zanzibar, living 
in deep burrows where the sand is of some consistence, 7.¢. well 
above low-water mark. In obtaining specimens for bait, the 
natives dig pits two or three feet deep, scooping away the sand 
from the circumference until a sufficient number of the worms has 
been met with. My own specimens were obtained in this way, 
through the natives, except two younger ones, which were 
found under the bark of a tree which lay half buried in the 
sand, 

The worm is of fair size, a foot or two long, by nearly half an 
inch wide when alive. Its colour is a dark blood-red, with a green 
iridescence anteriorly, the red being obscured, however, only 
by the black gut posteriorly. No pigmentation occurs except 
certain small marks on the prostomium described below. 

The shape of the anterior end of the body (PI. XV. fig. 7) is very 
characteristic of this as of several other species. The head and 
first few body-segments are proportionately long and round in 
section, forming a cylinder, but at about the fifth they become much 
broader and especially shorter and flatter. The broadest segment 
is usually about the twelfth, after which a slight decrease in 
breadth occurs, the rest of the worm having parallel sides and being 
composed of very short and flat segments. From about half an inch 
before the anus the segments rapidly narrow, so that the hind end 
is triangular (see Pl. XV. figs. 7 & 9). The upwardly directed 
thick-lipped anus and its cirri are as figured by Gravier, but his 
figures of both head and anus suggest that the worm increases and 
decreases in breadth but slightly, and quite gradually. 

T find no reduced eyes on the prostomium. The pigment-marks 
at the bases of the tentacles and under the anterior border of the 
peristomial segment are shown in fig. 8, Pl. XV., which shows 
also the rings round the lower ends of the tentacles mentioned by 
Gravier. The tentacles are always distinctly more slender, and 
appear to arisenearer together than those shown by Gravier’s figure. 

The dental apparatus is soft and brown in the two young speci- 
mens, very hard and black in the adults, a very little white 
matter occurring on the lower or mandibular plates. The upper- 
most tooth of the right great dental plate is always smaller and 
duller than the corresponding one on the left, and those of the 
right curved lateral plate are all longer than those of the left. 

Except near the extremities of the body, the feet present the 
peculiarity of being provided with no more than the stumps of 
their setee. This, which would appear a mere regrettable accident 


140 MR. CYRIL CROSSLAND ON THE [June 16, 


if only one specimen occurred in the collection, becomes an 
interesting character when it is found that it occurs in all cases, 
even in young specimens. Besides the characteristic long capil- 
laries, short, remarkably broad comb-setee occur in some feet at 
either end of the body (Pl. XV. fig. 10). 

There are four acicule in the anterior feet, two or three in those 
of the middle region, and one posteriorly, which may or may not 
be accompanied by another which is almost colourless. 


MARPHYSA SIMPLEX, sp. nov. (Plate XV. figs. 11, 12.) 


Two specimens from between tide-marks, near Zanzibar Town : 
the larger incomplete, measuring 10 cms. by 0:45 cm., composed 
of 155 segments ; the smaller complete, 15 ems. long by 0°3 em. 
broad. 

The circular section of the body is very nearly uniform through- 
out, and the length of the segments is greater than in the case 
of the preceding species, and those immediately following the 
head are neither longer nor narrower than the rest. The body 
narrows quite gradually to the anus, the opening of which is large, 
upwardly directed, and provided with a pair of moderately long cirri 
ventrally. 

The prostomium is bilobed, the median notch being continued 
dorsally as a narrow groove to its base. The tentacles are nearly 
twice as long as the prostomium and are delicately rmged. Their 
insertion, basal parts, and the position of the eye-spots are shown 
in fig. 11, Pl. XV. No considerable portion of the prostomium 
is hidden by the next segment. The peristomium has a total length 
about equal to that of the head, the segments immediately suc- 
ceeding being about half as Jong and of the same size, or nearly 
so, as those composing the rest of the body. 

The larger pieces of the dental apparatus all show a white 
edging, especially prominent on the upper ends of the mandibles, 
which bear large plates of this calcareous matter. The great 
dental plates bear three teeth each in the larger specimen, four 
in the smaller. The teeth of the laterals are blunt, and number 
four and three on the left, six on the right. The whole apparatus 
presents no peculiarity save the structure of the mandibles, for 
which see text-fig. 14, p. 142. 

The feet are small, and contain numerous long sete, projecting 
from a seta-sac, the posterior lip of which is very broad and 
straight-edged (text-fig. 13, p. 141). All the sete are smooth, 
without toothing or borders, the capillaries being bent gently and 
pointed in the usual way, the articulated pieces of the compound 
setee being sword-like and slenderly pointed. The comb-sete are 
long and slender, very finely toothed, the outer teeth on either 
side being considerably the longer. Pl. XV. fig. 12 represents a 
group of these sete, showing their variation in the same foot, 
the distance they project from the seta-sac, &e. They are much 
larger than the somewhat similar ones of IZ. macintoshi. 

There are four or five acicule in the anterior feet; three, with 


1903.] MARINE FAUNA OF ZANZIBAR. 141 


a lighter-coloured, bent, simply bluntly pointed acicular seta, 
in the posterior. 

The sete are thus exactly like those of I/. macintoshi except in 
the greater size of the combs and the larger number of the 
acicule. 

The dorsal and ventral cirri are very short, and the latter 
are also thick. 


Text-fig. 13. 


Marphysa simplex, sp. nov. 15th foot, x 45. 


The gill-region extends over practically the whole body, the 
first thirty segments alone being without them. The gills increase 
rapidly in size, the largest having four or five filaments arranged 
comb fashion and long enough to be made nearly to meet over 
the back. In the smaller complete specimen gills of four filaments 
are found only near the beginning of the gill-region ; the remaining 
segments bearing gills of equal length but of three filaments 
or, more posteriorly, only two. 


MARPHYSA FURCELLATA, sp.nov. (Plate XV. figs. 13, 14.) 


The head and first 120 segments of one specimen were collected 
between tide-marks near the town of Zanzibar, and the whole of 
a second was dredged from 15 fathoms in Zanzibar Channel, the 
bottom being mud. The former measures 4 cms. in length and 
4 mm. in breadth, the latter 12 ems. by 4 mm. in the anterior 
third, after which the breadth decreases to 3 mm. Only the first 
few segments are round, the rest of the body being very flat 


142 MR. CYRIL CROSSLAND ON THE [June 16, 


dorsally and only slightly convex ventrally. Posteriorly it nar- 
rows gradually to the anus, which has thick lips, is dorsally 
directed, and bears ventrally four cirri, two long and two very 
short. There is no pigmentation. 

The prostomium, the insertion of the tentacles, and the position 
of the eyes resemble their arrangement in I. simplex, but the 
tentacles are much shorter (being but little longer than the pro- 
stomium) and the eyes much smaller (Pl. XV. fig. 13). 

The dental apparatus is quite of the ordinary type (see text-fig. 
15). The numbers of teeth are given by the formula 5—5 :5+4+4—7. 
All are long and sharp, a character not shared by the laterals of 
any of the preceding species. The slender maxille are straight 
proximally, bent towards each other distally. 


Text-fig. 14. Text-fig. 15. 


Text-fig. 14.—Marphysa simplex, sp. nov. Ends of lower or mandibular jaw-plates 
from the ventral side, x 22. 


Text-fig. 15.—Marphysa furcellata,sp. noy. Dorsal or inner aspect of ventral jaw- 
plates, x 22. 


The feet are normally developed, and carry bundles of set of 
moderate length. Dorsal and ventral cirri as in the preceding 
species and most others of the genus. Lip of seta-sac rounded in 
anterior feet, pointed in posterior. The sete are asin J. simplex 
except the combs, which are of two kinds, both projecting very 
slightly. The one kind has few strong curved teeth (whence the 
name of the species), the other having the usual numerous minute 
teeth, and intermediates between these extremes are frequent 
(fig. 14, Pl. XV.). There are three or four acicule anteriorly and 
one behind. The acicular seta has a simple point, is very slightly 
bent, and projects. 

The gill-region comprises the greater part of the body, continuing 
from the twentieth segment to near the anus. In both specimens 


1903. | MARINE FAUNA OF ZANZIBAR, 143 


there comes first a region of about forty segments bearing small 
gills of three or four filaments; these gills at about the sixtieth 
segment rapidly enlarge and consist of four or five filaments, 
though they never become long enough to meet over the back. 

So far as Grube’s and Baird’s descriptions go, the last two species 
correspond with MW. januarti and MW. parishi. Indeed the comb- 
sete of the latter form a distinct point of similarity between it 
and I. furcellata. But the descriptions published of these two 
forms are so exceedingly meagre that certainty is impossible, so 
that in preference to the risk of giving two worms a distribution 
from East Tropicai America to Hast Africa, I have described my 
forms de novo. 


Genus LystpIceE. 


LystpicE cotuAris Ehr., Grube. 


This species, though never abundant, occurred at most of my 
collecting-places, viz., two specimens from Wasin Harbour (one 
from between tide-marks, the other from a depth of 10 fathoms), 
two smaller ones from 3 fathoms in Chuaka Bay, and three, 
smaller still, from the shore near Zanzibar Town. 

Discrepancies of some importance occur between the descriptions 
of the species already published by Grube (Red Sea and Philippine 
collections), Marenzeller (Japan), and Gravier (Red Sea). My 
own collection shows that variations of features, usually considered 
diagnostic, occur in specimens from the same locality. 

The name ‘collaris’ obviously refers to the white ring found 
near the anterior end of the living animal. As the colour dis- 
appears from specimens which have been a few years in spirit 
(it is already becoming faint in my own after the lapse of one 
year), it has not yet been described. The ground-colour is a bright. 
yellow-brown, best developed anteriorly and gradually dying out 
at about the tenth setigerous segment. Posteriorly the body is 
nearly colourless, unless sexual products, which are pink, give it 
that colour. In one of my specimens (and presumably in that 
collected by Ehrenberg) this pigmentation is interrupted by 
numerous small white dots, and is omitted altogether from 
segments three and four, forming the above-mentioned white 
collar. In the remaining specimens, one of which is of equal size 
to this, the coloration is perfectly uniform. 

The form of the body is in life, as after preservation, flat below 
and strongly arched above throughout its length. 

The insertion of the tentacles is not noticed by former authors 
except Gravier, and in this respect, as in others, none of my 
specimens agree with his description. The tentacles, though a 
little narrowed at their bases, have no distinct basal joint, and 
though they in some cases arise from nearly the same level, yet 
the origin of the middle one is always in front of the origin of the 
other two, thus reversing the usual arrangement. The prostomium 
itself is rather longer than in Gravier’s figure. 


144 ON THE MARINE FAUNA OF ZANZIBAR. [June 16, 


All authors agree as to the characteristic form of the eyes. As 
noticed by Grube and seen in the present collection, they vary m 
breadth from narrow crescents to kidney- or bean-shape, yet never 
becoming merely oval. 

The lower or mandibular jaw-plates are most characteristic in 
constitution and form, being usually calcareous with special plates 
of brown or black horny material. Marenzeller’s figure is the 
only adequate representation of a very beautiful structure. The 
number of teeth on the left great dental plate varies from three 
to five, though usually it bears, as does always the right, four 
teeth. The curved laterals are exceptionally variable, as shown 
by the formule 3+5—5; 5+1 5; 34+1—5; 343-3; 
44+5—3, 

All authors agree in making the dorsal and ventral cirri much 
longer than those figured by Gravier. Im all the specimens I 
have examined, their tips extend as far as do the lips of the seta- 
sac, or in anterior feet a little further. 

The articulated pieces of the compound sete are not always 
the shape of an equilateral triangle, being usually somewhat longer. 
Their shafts are bent and broadened distally, and the striation and 
the toothing at the point opposite the articulation (mentioned by 
Gravier, but best seen in Marenzeller’s figure), like the bordering 
of the capillary sete, are not always distinct. 

The acicular sete always bear two distinct hooks, when not 
damaged by wear, as sometimes happens, but only in a few cases 
does their covering or winging remain. 


ZXPLANATION OF THE PLATHS. 


Pratt XIV. 


Fig. 1. Diopatra neapolitana (p. 132). Ventral view of the head after removal of 
the jaw-apparatus. 
ces., esophagus ; b.w., body-wall; a, lip of jaw-sac; 6, lip of cesophagus 
with its tubercles. 
2. Onuphis holobranchiata (p. 185). Head and anterior end, showing the 
characteristic pigmentation. 
3. Marphysa macintoshi, sp. nov. (p. 137). Head and anterior end. 
4, 25th foot of the same. 
5. Compound sete from 25th foot. Comb-sete from the 100th foot. The whole 
free length of these is represented. 
6. 160th foot of the same. 


PratE XV. 


Hig. 7. Anterior end of Marphysa mossambica (p. 139), showing the proportions of 
the body in spirit-specimens. 
8. Ditto. The front edge of the peristomium is cut away to show the pigment- 
marks on the tentacles and prostomium. 
9. Hind end, anus and its cirri. 
10. Group of comb-setz from one of the posterior feet, showing the extent of 
their projection from the seta-sac. ; 
11. Marphysa simplex, sp. nov. (p. 140). The rmeging of the tentacles is rather 
too strongly emphasised in this figure. 
12. Group of comb-sete from the 60th foot. 
13. Marphysa furcellata, sp. nov. (p. 141). Head and anterior end. 
14. Tip of seta-sac of 120th foot, shewing the short strong combs characteristic 
of the species. 


H. Wilson, Cambridge. 


ENTO-PARASITES FROM LOWER SIAM. 


1903.] ON THE ENTO-PARASITES OF THE ‘‘SKEAT EXPEDITION.” 145 


5. On the Hnto-Parasites collected by the “Skeat Expedition” 
to Lower Siam and the Malay Peninsula in the Years 
1899-1900. By Artuur EH. Suiptey, M.A., F.ZS., 
Fellow and Tutor of Christ’s College, Cambridge, and 

- University Lecturer in the Advanced Morphology of the 
Invertebrata. 


[Received May 28, 1903. | 


(Plate XVI.*) 


C'est parmi les parasites et non chez Vhomme qwil faut chercher le dernier mot 
dz la création. M Evie METCHNIKOFE, 
‘Etudes sur la Nature Humaine.’ 


The collection made by the members of the Skeat Expedition 
of Entozoa was rich and varied, and inasmuch as it was gathered 
in a land hitherto unsearched for this purpose, it is not surprising 
that it contained a large proportion of new forms. Amongst the 
more interesting of the results brought to light by the investiga- 
tion of the material are:—(i.) A new species of Tetrarhynchus 
found ina Holothurian. This is, I believe, the first record of a 
Tetrarhynchus being found in any Hchinoderm, and indeed the 
presence of Tetrarhynchide in invertebrates at all is only recorded 
for one or two Molluses and doubtfully for a species of Aphrodite. 
(ii.) An undeterminable species of Tetrarhynchus from a sea-snake. 
Here, again, we have an entirely new host, no Tetrarhynchus 
having been hitherto found in any vertebrate outside the class 
Pisces with the exception of the Chelonian Testudo mydas. 
(iii.) A considerable collection of new forms of Acanthocephala. 
The Nematodes, which have been described elsewhere by Dr. von 
Linstow, include some fourteen new species. 

As will be seen, the collection covers a wide field and includes 
representatives of the Sporozoa and all the chief groups of metazoan 
Entozoa, except the Trematoda. The Cestoda, Acanthocephala, 
Nematomorpha, Nematoda, and Linguatulida severally contribute 
one or more species to the total. 

With regard to the localities, the parasites were collected mainly 
in three or four places: Biserat, in Jalor, a province of Lower Siam, 
in Kwala Aring in the same province, and at Tremangan. 


CESTODA. 


Fam. BotHRIOCEPHALIDA. 


BOTHRIOCEPHALID LARVA, 
The snakes of Lower Siam seem frequently to harbour the large 
larval form of certain of the Bothriocephalide, of which it is 


* For explanation of the Plate, see p. 156. 
Proc. Zoou. Soc.—1903, Vou. Il. No. X. 10 


146 MR. A. E, SHIPLEY ON THE [June 16, 


impossible to identify the species or even the genus. I have 
recently dealt somewhat fully with a similar larval form from 
under the skin of a Serval from the Soudan*. These Siamese 
specimens differ from the African one in the thickness of the 
anterior end, which was cushiony, and perhaps almost as thick as 
one quarter of the transverse diameter ; and in the regularity and 
extension forward of the annuli, which extend right up to and even 
into the orifice of the single apical sucker. Specimens of these 
larve were taken in three different snakes, almost certainly 
specimens of Dipsadomorphus dendrophilus Boie, at Biserat, and 
beneath the skin of a snake from Patalung. 


Fam. TETRARHYNCHIDA. 


TETRARHYNCHUS HOLOTHURI&, n. sp. (Plate XVI. figs. 5, 
6, & 7.) 


A small collection belonging to the genus TVetrarhynchus was 
contained in a bottle labelled “ Parasites from the commonest 
Holothurian found in the sea off the Pataniriver.” The specimens 
measure some 7 mm. in length, by a maximum body breadth of 
1mm. The suckers, however, add to this last measurement in 
the region of the head. The body tapers smoothly to the posterior 
end, where there is a slight indentation at the extreme point, into 
which sections show that the two longitudinal water-vascular 
canals open, one on each side. 

The bothria are somewhat ear-shaped (Pl. XVI. figs. 5 & 7) and 
each is divided into two longitudinal halves by a median ridge, so 
that in transverse section there is the appearance of four suckers. 
The hooked arms which project from the head end in a conical 
tip, covered with very numerous spines all pointing forward 
(Pl. XVI. fig. 6). In the specimen figured, one of these spines is 
much larger than the others, but this is probably a slight abnor- 
mality. Following on this spiny end is a smooth portion, and then 
a second spiny region where the very numerous spines, all pointing 
backward, form a very firm organ of attachment. 

The tapering conical body shows no trace of strobilisation and 
is externally smooth, the only differentiation visible being the 
line of the sac into which the toothed processes are withdrawn, 
which in some cases is seen through the surrounding tissue. 

Sections reveal no trace of reproductive organs. The cuticle 
surrounds a mass of parenchyma which is looser and more vacuo- 
lated just under the cuticle, and this looseness is even more 
pronounced around the stout muscular sacs from which the four 
toothed introverts spring. The muscles of these sacs are unusually 
stout and circularly or perhaps spirally arranged. 'Two—there 
are said to be four in most adults—laterally placed water-vascular 
vessels run down the animal, and open into the terminal depres- 


* Aych. Parasit. vi. 1902, p. 604, 


1903. | ENTO-PARASITES OF THE ‘‘SKEAT EXPEDITION.” 147 


sion. The number of calcareous bodies is small, and other deeply 
staining structures, e.g. the nuclei and muscle-fibres, are but sparsely 
scattered through the tissues. There is a four-cornered nerve- 
mass well lying about the level of the juncture of the anterior one- 
third with the posterior two-thirds of the bothria, and this gives 
off four strands which pass peripherally and probably supply the 
bothria and introverts. I did not see any longitudinal nerves, 
though probably they exist. 

In his exhaustive monograph* “ Recherches sur les Tetra- 
rhynques,” Vaullegeard records his opinion that the numerous 
species of Tetrarhynchous Cestodes belong to but one genus, Tetra- 
rhynchus. Raillet has pointed out that the generic name 
Rhyncobothrius Rudolphi, 1819, has precedence of Teérarhynchus 
1809, the latter name having been given to alarvalform. In the 
present case, although it may be inconsistent with the laws of 
nomenclature, I have followed Vaullegeard, partly because every- 
one knows what Tetrarhynchus is, and partly because, in my 
opinion, needless confusion is introduced into the study of tape- 
worms by using a double nomenclature for larve and adults. In 
civilized society it is not considered necessary for a human being 
to change his name when he leaves the home of his childhood and 
sets up in a new house, and there seems as little reason for a 
young Cestode to change its name when it changes its host. 

Vaullegeard arranges the various species into two sections, one 
of which, on the type of 7. ingualis Cuv., has in its larval form 
no vesicle projecting over and protecting the head. To this 
section the cestode in question belongs. The larve which were 
collected by the Skeat Expedition were some of them wholly and 
some of them partly enclosed in cysts, but none of them showed 
any trace of the projecting vesicle. 


Tetrarhynchus holothurie, n. sp. (Plate XVI. figs. 5, 6, & 7.) 


The larval form (but not the encysted form) is 6 mm. long by 
1 mm. broad. Tapering posteriorly where there is a depression 
receiving the excretory pore. The two large bothria are sub- 
divided by a ridge. The four introverts are provided with a cap 
of hooks pointing forward at the tip and a circular band of hooks 
pointing backward, between these two toothed regions is a con- 
siderable portion devoid of hooks. 

Halitat. A common Holothurian, probably a Molpadia, taken 
off the mouth of the Patani River. 


TETRARHYNCHUS sp. (Plate XVI. figs. 8 & 11.) 


Some eight or nine small cysts about the size of very poor wheat- 
seeds were taken from the body of a sea-snake, Hnhydrina vala- 
kadien Boie. Unfortunately the position of the cysts in the body is 


* Mém. Soc. Normandie, xix. 1897-1899, p. 185. 
LO% 


148 MR. A. E, SHIPLEY ON THE [June 16, 


unrecorded, but I should judge that they lay just under the perito- 
neal lining of the body-cavity. In one or two cases the worm 
itself was emerging, or had emerged, from the cyst, but I attribute 
that to the handling the cyst received as the worms were removed 
from the body of the host. The cestodes usually were bent but 
once within the cyst, as is shown in the figure (LoTR QTE tas, 1111) 

Sections through these cysts (Pl. XVI. fig. 8) show that the 
form in question belongs to Vaullegeard’s second group founded 
on the type of Tetrarhynchus erinacews van Ben.,in which the 
larvee have a vesicle surrounding and protecting the head. This 
vesicle is clearly shown in the figure; within it the head of the 
larva and the neck, as far back as the muscular sacs into which 
the introverts are retracted, are coiled. These coils, being hidden 
by the vesicle, cannot be seen through the walls of the cyst ; 
they are, however, sufficiently numerous to permit four or five 
sections of the head at different levels to be displayed in one 
section. The head passes into the body, which has two longi- 
tudinal excretory canals and shows no sign of reproductive organs ; 
in fact, the only differentiation from the loose parenchymatous 
tissue is a layer of muscle-cells situated about halfway between 
the periphery and the centre. 

The vesicle is folded over the head lke an amnion; it is, 
however, not closed, but remains open by a pore guarded by 
thickened lips. Iam inclined to think that these lips contain 
muscle-fibres, and that the aperture can be tightly closed if 
occasion arises. According to Vaullegeard the vesicle detaches 
itself when the larva becomes sexually mature. 


The genus Tetrarhynchus is often regarded as exclusively a fish 
parasite: it has, however, been described in certain Molluses, 
e.g. Sepia officinalis and the Pearl-Oyster, and perhaps in Aphro- 
dite aculeaia, though nobody seems to have found it in that animal 
since the distinguished courtier, philosopher, parasitologist, and 
poet, Francis Redi of Arezzo, recorded it in 1664. I have found no 
record of the genus occurring in Echinoderms, so that the discovery 
by the Skeat Expedition of the larval forms ina Holothurian is a 
matter of considerable interest. This form, though not mature, is 
not enveloped in a vesicle, and presents certain features which 
allow me to suggest a specific diagnosis. 

The second form brought back from the coast of Lower Siam 
is equally new as regards its host. There has hitherto been 
recorded, so far as I can find, but one vertebrate host of the genus 
Tetrarhynchus outside of the class Pisces. This is Testudo mydas, 
in which, in 1840, Meyer described vesiculate larvae. We can now 
add a second Reptilian host in the case of Hnhydrina valakadien 
Boie, a sea-snake, belonging to the family Colubride, which is not 
unfrequently taken along the coast of India and Burmah, and 
which ranges from the Persian Gulf to the Malay Archipelago 
and Papuasia. These very poisonous ophidians are fish-eaters. 


1903. ] ENTO-PARASITES OF THE ‘‘SKEAT EXPEDITION.” 149 


ACANTHOCEPHALA. 
Fam. EcHINORHYNCHIDA. 


HCHINORHYNCHUS PATANI, nu. sp. (Plate XVI. figs. 9 & 10.) 


The encapsuled larve of Hchinorhynchide are by no means 
uncommon in snakes, and von Linstow* suggests that their corre- 
sponding adult forms are to be looked for in the bodies of raptorial 
birds. In the paper referred to, he enumerates species of these 
parasites taken from snakes the names of which he gives. 
Unfortunately many of the names are nomina nuda, and the hosts 
cannot now be identified, and the same is the case with the new 
species here described, as the name of the snake was not preserved. 

The cysts are at most 10 mm. long by 5mm. broad. The worm 
is bent twice, and the three limbs may lie in one plane or in two. 
One of the specimens had escaped from its cyst, and had already 
inserted its spiny head into some piece of tissue from which it was 
well nigh impossible to free it. Another had freed its head and 
straightened out its body, which, however, was still surrounded by 
a thin film-like sheath of tissue. The length of these specimens 
was 25 mm., the breadth of the trunk was 2 mm., of the head 
1mm. ‘The hooks were arranged in 12—14 longitudinal rows, the 
hooks of one row being at the level of the space between two 
neighbouring hooks of the row right and left (Pl. XVI. fig. 10). 
In this way the hooks in the horizontal rings also alternated with 
one another. There were six of such rings, and the second and 
third row consist of markedly large hooks, twice as large at least 
as any of the others. Altogether there were six rings visible ; 
there may have been one or more hidden by invagination, but I 
do not think so. 

It is most unfortunate that the name of the snake which 
harboured this parasite is unknown, but the parasite seems to be 
a hitherto undescribed species. The names of those already known 
to be encapsuled in snakes are recorded by von Linstow as 
follows :— 


(1.) Heh. oligacanthoides Rud., with 4—5 rows of hooks. 

(u1.) Heh. cinctus Rud., with 140 rows of hooks of similar size. 

(ii.) “ch. obligacanthus Rud., with 13 rows of hooks. 

(iv.) Heh. megacephalus Westrumb, with very numerous rows 
of hooks and the proboscis swollen in the middle. 

(v.) Heh. dipsadis von Lins., with some 12-14 rows of large 
hooks, followed by 20 rows of smaller hooks. 

(vi.) Heh. heterorhynchus Par. Proboscis anteriorly slender 
with 11 rows of hooks, posteriorly enlarged with 16 rows of hooks. 


Wedl has mentioned other larval forms found in snakes, but 
they are not in any case specifically identified. 


* Arch. Naturg. 54 Jahreang, i. 1888. 


150 MR. A, E, SHIPLEY ON THE [June 16, 


Echinorhynchus patani, n. sp. (Plate XVI. figs. 9 & 10.) 


Length 25 mm., breadth 2 mm., breadth of head 1mm. Hooks 
in 6 rings, perhaps 7 or 8, and in 12-14 longitudinal rows. The 
hooks in one ring are opposite the interspaces in the next. The 
2nd and 3rd rings composed of unusually large hooks. The worms 
were folded twice with cysts some 10mm. long by 5 mm. broad. 

Habitat. The body of an unknown snake, possibly Dipsado- 
morphus dendrophilus Boie, taken at Patani, Lower Siam. 


ECHINORHYNCHUS BUFONIS, nu. Sp. (Plate XVI. figs. 1, 2, & 4.) 


Several specimens of what I regard as the same species of 
Echinorhynchus were taken from the alimentary canal of two 
species of Bufo obtained at Patani, which have been kindly identified 
for me as Bufo melanostictus Schneider, and in all probability 
Bufo penangensis Wilson & Gray, by Dr. Gadow. 

Bufo is known to harbour the comparatively widespread 
Ech. heruca, but I know of no other species of Hchinorhynchus 
parasitic in this amphibian, and as the parasites from Patani differ 
in many respects from any descriptions which are available, I 
have established a new species. 

The worms fall into two groups. The larger, probably the females, 
measure some 15 mm.; the smaller, probably males, some 5 mm., 
but there are many intermediate in length. The greatest 
breadth of the larger specimens is 15mm. Asa rule their out- 
line and surface is smooth, but some were wrinkled either wholly 
orin part. The body is usually curved, and in the larger specimens 
markedly so. The most conspicuous feature in which this species 
differs from the majority of its congeners is that the proboscis or 
introvert is not median and terminal, but projects from the trunk 
a little way, sometimes more, sometimes less, from the anterior 
end; it usually slopes forward, but it may stand out at right 
angles to the axis of the body like the handle of a walking-stick 
(Pl. XVI. figs. 2.& 4). It is always protruded on the inner 
surface of the curve. This feature and its divergence from the 
more usual type are represented in the Gyphyrea, where Aspido- 
siphon bears the same relation to most other Sipunculids that this 
species does to other Hchinorhynchi. 

The number of hooks is comparatively small, there being some 
6-8 rings, alternately arranged with 14-16 longitudinal rows. The 
rings being alternating, the number of hooks in each ring is half 
the number of longitudinal rows. 


Hehinorhynchus bufonis, n. sp. (Plate XVI. figs. 1, 2, & 4.) 


Curved, with proboscis opening on the concave surface just 
behind the anterior end of the body, which extends beyond the 
point of emergence of the proboscis. Length 15 mm. or less; a 
number, probably males, only 5 mm. long. Few hooks, 6-8; rings 
with 7-8 hooks alternating with those of the next row. 


1903. | ENTO-PARASITES OF THE “‘ SKEAT EXPEDITION,” 151 


Habitat. The alimentary canal of Bufo melanostictus Schneider 
and of Bufo ? penangensis Wilson & Gray, taken at Patani. 


ECHINORHYNCHUS XENOPELTIDIS, nu. 8p. (Plate XVI. fig. 3.) 


Three curious specimens of Echinorhynchide were taken free 
in the body of Xenopeltis unicolor, the sole species and genus of 
the family Xenopeltide, which ranges over South-eastern Asia, 
from India to the Malay Archipelago. Unfortunately, the part 
of the host’s body infested by the parasite is unnamed, but, presum- 
ably, it was the intestine. 

The parasites are three in number, and measure respectively 
about 25 mm., 17 mm.,and 15 mm. I say about, because they 
were all coiled in a curious, angular sort of way, so that it was not 
possible to straighten them. They are plump, fleshy-looking 
creatures with an average breadth of 2°5 mm., though in one 
specimen a breadth of fully 3 mm. was attained. They hardly 
taper at all at the ends, which are truncated. 

Their most peculiar external features are two. The first is the 
colour. This, in specimens kept for some years in spirit, is a 
delicate, salmony pink, somewhat resembling a freshly peeled pink 
banana. I have never seen an Echinorhynchid with anything 
approaching this colour. The second feature is the wrinkling. 
This is very marked, and produces a very definite deepening of 
the colour. The areas into which the wrinkles divide up the skin 
become in the anterior end almost regularly quadrilateral, and a 
quite peculiar marking is the result. This is well seen in 
Pl. XVI. fig. 3. 

The only specimen f sacrificed to the razor showed that these 
wrinkles are the expression of deep narrow grooves which pene- 
trate the subcuticle almost as far as the basement membrane. 
Unfortunately the sections obtained did not clearly show the 
number of rings of hooks. They were not very numerous, perhaps 
some 8 to 12. In each of the three specimens the proboscis was 
retracted. 


Echinorhynchus xenopeltidis, n. sp. (Plate XVI. fig. 3.)3 


Length varying from 15 to 25 mm. Average breadth 2°5 mm. 
Plump forms with anteriorly many wrinkles, which showed a 
tendency to break up the surface into squarish areas. Colour, a 
delicate salmon-pink. Hooks in ? 8-12 rows. 

Habitat. Xenopeliis unicolor Reinw. Taken at Kwala Aring. 


E{CHINORHYNCHUS TIGRIN 4, 1. Sp. 


Two complete specimens and a fragment were taken from the 
intestine of a Rana tigrina Daud. The former had a length of 
10 mm. and an average breadth of 2mm. They were of a slaty-grey 
colour, and marked by transverse grooves at irregular intervals. 

The proboscis is very short and very small. It does not emerge 


152 MR. A. E, SHIPLEY ON THE [June 16, 


terminally, but rather from the side, where it is overtopped by 
the anterior end, and looks like a little head sunk in one of the 
enormous collars in vogue at the Regency period. The number 
of hooks is very small, only 4-5 rings, and but few hooks ina ring. 
At first I thought that the smallness of the number of the 
hooks indicated that we had to do with a young, or, at any rate, a 
not fully grown, individual, but the lumen of the trunk was 
crowded with ova in well-developed chitinous egg-shells. Hach 
ovum is along cell, rounded at the ends with a conspicuous nucleus 
in the centre. The egg-shell is rather more pointed at the ends, 
so that the egg with its shell forms a spindle-shaped object some 
"08 mm. in length and -02 mm. at the greatest diameter. 


Lchinorhynchus tagrine, n. sp. 


Length 10 mm., average breadth 2mm. Greyish, transversely 
wrinkled. Proboscis short, small, arising from behind the anterior 
end. Four or five rings of very few hooks. Ova spindle-shaped, 
0-8 mm. x ‘02 mm. 

Habitat. Intestine of Rana tigrina Daud. Taken at Biserat, 
Jalor. 


Some small fragments of another Hchinorhynchus, too small to 
admit of identification, were taken from the intestine of the toad 
Callula pulchra Gray. 


NEMATOMORPHA. 


Professor Camerano has kindly described the specimens of this 
very difficult group and has published accounts, with full details 
as to the structure of the cuticle on which the classification of these 
creatures so largely rests, of four species, two of which, Chordodes 
siamensis and Gordius paronw, are new. I extract the following 
from Camerano’s descriptions :— 


1. CHorDODES monTonr Camer. 


1895. L. Camerano, Bull. Soc. Zool. France, xx. p. 99. 
1897. LL. Camerano, Mem. Acc. Torino, ser. 2, xlvii. p. 387. 
1899, L. Camerano, Atti Acc. Torino, xxxiv, figs. 3, 3 a. 
1901, L. Camerano, Boll. Mus. Torino, xvi. no. 408. 


A single specimen, a male, was taken by Mr. Laidlaw whilst 
making its exit from a large Mantis captured at the foot of Gunong 
Tnas, Perak. The length could not be determined, as:the worm 
was in pieces. The colour of the spirit-specimen is white at each 
end, black and velvety in the middle. 

The cuticle showed the characteristic markings described by the 
author in the first-named specimen, which came from China, other 
specimens have been described from Perak. 


1903. ] ENTO-PARASITES OF THE “‘ SKEAT EXPEDITION,” 153 


2. CHORDODES PUNCTICUTATUS Camer. 


1895. L. Camerano, Notes Leyden Mus. xvii. 

1897. L. Camerano, Mem. Acc. Torino, ser. 2, xlvii. p. 384. 

1899. L. Camerano, Atti Acc. Torino, xxxiv. 

1901. L. Camerano, Boll. Mus. Torino, xvi. no. 408. 

A single male specimen taken on an island off Kedah measured 
23°2 mm. Its colour was a darkish brown lightening towards each 
tip. The cuticle resembled other examples described from Deli 
in Kastern Sumatra and from Perak. 


3. CHORDODES SIAMENSIS Camer. 

1903. L. Camerano, Boll. Mus. Torino, xviii, no. 437. 

A single female specimen, brownish-black in colour, but lighter 
towards the ends, 20 cm. long. This specimen was taken at 
Biserat. 

4, GORDIUS PARON# Camer. 


1905. L. Camerano, Boll. Mus. Torino xviii. no. 437. 


Three male specimens taken at Kota Bharu, varying in length 
from 17°5 cm. to 22°5 em. 


NEMATODA. 


The Nematoda have been described by Dr. von Linstow in the 
‘Archiv fiir mikroskopische Anatomie und Entwicklungsgeschichte, 
Band lx. 1903, p. 108. I here give only a list of the species 
and of their respective hosts, and must refer the reader to the 
memoir mentioned for further details. 


1. ASCARIS INFUNDIBULICOLA Vv. Lins. 


From the alimentary canal of Python reticulatus Gray, killed in 
Tremangan, 


2. ASCARIS SOLITARIA v. Lins, 

A single undeveloped female was found in the stomach of 
Dipsadomorphus dendrophilus Boie, taken at Kwala Aring. 

3. ASCARIS DIPSADOMORPHI v. Lins. 

A number of larvee found in the cysts scattered through the 
mesentery of D. dendrophilus Boie, Kwala Aring. 

4, H&TERAKIS RIMULA Vv. Lins, 

Out of Centropus sinensis Steph. 


5. HETERAKIS CIRCULARIS v. Lins. 
From the same host as No, 4, 


154 MR. A. E, SHIPLEY ON THE [June 16, 


6. CHEILOSPIRURA OPHTHALMICA V. Lins. 
From the eye of Turnix taigor Sykes. 


7. CHEILOSPIRURA SIAMENSIS V. Lins. 
From Centropus sinensis Steph., probably from the eye. 


8. OXYURUS SIAMENSIS v. Lins. 
From the stomach of Liolepis bellii Gray. 


9, OXYURUS CORONATA Vv. Lins. 


From the large intestine of Galeopithecus volans Linn., taken 
at Patalung. 


10. OxysoMA TUBERCULATUM V. Lins. 
From the alimentary canal of Megalophrys montana Wag). 


11. FILARIA LONGICIRRATA V. Lins. 
From the subcutaneous tissue of Galeopithecus volans Linn. 


12. aria scivnri (2) v. Lins. 


A single immature female found under the skin of Sciwrus 
caniceps Gray. Von Linstow regards the naming of this specimen 
as provisional. 


13. HILARIA CORYNODES v. Lins. 


From under the skin of Semmopithecus albocinereus, Kwala 
Aring ; see also von Linstow, MT. Mus. Berlin, i. 1899, p. 23. 


14, AnGIOSTOMUM BRACHYLAIMUS V. Lins. 
From Bufo melanosticius, probably from the lungs. 


15. LISSONEMA ROTUNDATA v. Lins. 


From Centropus sinensis Steph., probably from the alimentary 
canal. 


LINGUATULIDA. 


POROCEPHALUS MONILIFORMIS (Diesing). 


Synonym. Pentastoma moniliforme Diesing Denk. Ak. Wien, 
xi. 1856. 

Three specimens of this common parasite were taken from the 
lungs of a Python reticulatus Schneid., at Biserat; and six or 


seven specimens, varying a good deal in size, from another Python 
of the same species at Tremangan. 


1903. ] ENTO-PARASITES OF THE ‘“‘SKEAT EXPEDITION,’ 155 


SPOROZOA. 


Order Sarcosporip1A Balbiani. 


Several specimens of this order were found amongst the muscles 
and the tissues at the base of the tongue of a Bos bubalis killed 
at Kwala Aring. Dr. von Linstow has described these specimens 
as a new species—balbiana (Sarcocystis) siamensis—in the article 
mentioned under the Nematoda. I have figured these life-size, 
ony PIP PNGV ela tical 2s 


Lists oF Hosts CONTAINING THE PARASITES COLLECTED ON THE 
SKEAT HXPEDITION. 


ECHINODERMA. 


A Holothurian, probably a Molpadia, infested by Tetrarhynchus 
holothuric, n. sp. 


INSECTA. 


A Mantis was infested by the Gordian Worm, Chordodes mon- 
toni Camer. 


AMPHIBIA. 


Bufo melanostictus Schneider and ? Bufo penangensis Wilson & 
Gray had Hehinorhynchus bufonis, n. sp., in their intestines, and 
Angiostomum brachylaimus v. Lins. probably in its lungs. 

Callula pulchra Gray, infested by indeterminable species of 
Echinorhynchus. 

Megalophrys montana Wagl. had in its stomach examples of 
Oxysoma tuberculatum v. Lins. 

Rana tigrina Daud., infested by Hchinorhynchus tagrine, n. sp. 


OPHIDIA. 


Dipsadomorphus dendrophilus Boie, infested by Ascaris soli- 
taria v. Lins. in the stomach, a single specimen, and by Ascaris 
dipsadomorphi v. Lins. in numerous cysts on the mesentery ; 
and almost certainly by certain Bothriocephalid larve and by 
Lchinorhynchus patani, n. sp. 

Enhydrina valakadien Boie, infested.by a cystic form of Tetra- 
rhynchus whose species is not determinable. 

Python reticulatus Schneid. had its lungs infested by Poro- 
cephalus moniliformis, and another specimen contained many 
examples of Ascaris infundibulicola in its alimentary canal. 

Xenopeltis wnricolor Reinw., infested by Hehinorhynchus xeno- 
peltidis, n. sp. 


156 THE ENTO-PARASITES OF THE “ SKEAT EXPEDITION.” [June 16, 


LACERTILIA. 


Liolepis bellii Gray, contained in its stomach examples of 
Oxyurus siamensis v. Lins. 


AYES. 


Centropus sinensis Steph. contained specimens of Heterakis 
rimula v. Lins., and of 1. circularis v. Lins., and of Cheilospirura 
siamensis v. Lins., the last-named probably from the eye. A new 
genus and species, named by von Linstow Lissonema rotundatum, 
was found in this species, probably in the alimentary canal. 

Turnix taigoor contained in its eye examples of Cheilospirura 
ophthalmica v. Lins. 


MAMMALIA. 


Bos bubalis was infested at the root of its tongue with speci- 
mens of a Sareosporidian, regarded by von Linstow as new and 
named by him Laliiana siamensis. 

Sciurus caniceps Gray, had under its skin an example of a 
Filaria, provisionally named by von Linstow /ilaria sciuri. 

Galeopithecus volans Linn. was infested in its large intestine 
with Oxyurus coronata v. Lins., and in its subcutaneous con- 
nective tissue with /ilaria longicirrata v. Lins. 

Semnopithecus albocinereus was subcutaneously infested by 
filaria corynodes v. Lins. 


EXPLANATION OF PLATE XVI. 


Fig. 1. Side view of Hchinorhynchus bufonis, X 8, p. 150. 

2. Anterior end of the same, X 16, p. 150. 

3. Echinorhynchus xenopeltidis, X 8, p. 151. 

4, Section through the anterior end of H. bufonis to show the angle at which 
the proboscis leaves the body, p. 150. 

5. Tetrarhynchus holothurie, X 18, and the indented posterior end, p. 146. 

6. A single hooked arm of the same, X 100, showing the arrangement of the 
hooks, p. 146. 

7. An enlarged view of the head of the same, X 100, showing the arms and 
the bothria, p. 146. 

8. A longitudinal section through the cyst and the contained larva of a Tetra- 


rhynchus from the sea-snake Enhydrina valakadien, showing the cyst and 
the vesicle (ves.) surrounding the head, p. 147. 
9. Hehinorhynchus patani, X 3, p. 149. 

10. Head of the same, X 24, p. 149, 

11. View of one of the cysts, x 3, containing the Tetrarhynchus found in the 
sea-snake H. valakadien, p. 147. 

12. Pao ssnectmens life-size, of the Balbiana siamensis found in Bos bubalis, 
p. 155. 


1903. ] ON THE SYRINX OF THE ACCIPITRES. 157 


6. On the Modifications of Structure in the Syrinx of the 
Accipitres, with Remarks upon other Points in the 
Anatomy of that Group. By Frank H. Bepparp, 
M.A., F.R.S., Prosector to the Society. 


[Received June 16, 1903. } 
(Text-figures 16-20.) 


Although a considerable number of species of Accipitres have 
provided material for the study of the avian syrinx*, I am not 
aware of the existence of any general survey of the structure of 
the windpipe in this group of birds which is so extensive as that 
which I am able to lay before the Society. My material has been 
gradually amassed by my predecessors in the office of Prosector 
to the Society and by myself, and represents a large series of 
Accipitrine genera t, viz. the following :— 

Thrasaétus, Falco, Herpetotheres, Buteo, Milvus, Spizactus, 
Gypattus, Urubitinga, Haliaétus, Pandion, Tinnunculus, Vultur, 
Milwago, Spilornis, Hrythropus, Morphnus, Helotarsus, Leuco- 
pternis, Circus, Aquila, Circaétus, Dryotriorchis, Gyps, Hieracidea, 
Archibuteo, Geranoattus, Nisaéius, Asturina, Polyborus, Poly- 
boroides, and Gypohieraw: in all 31 genera, of many of which I 
have examined several species. 

I shall not, however, describe all of these types in detail, 
inasmuch as there are very close resemblances between many of 
them. My purpose is rather to establish the importance of the 
syrinx as confirmatory of other opinions upon the Classification 
of this Group. 

Genus Fauco. 


Of this genus I have examined syringes of the following seven 
species, viz.:—F. sacer, Ff. wsalon, F. candicanst, FP. feldeggt, 
F. peregrinus, F. biarmicus, PF. lanarius. 

Wunderlich’s description applies perfectly well to all of the 
species enumerated above. The species studied by him were 
F. peregrinus and F. subbuteo. The salient feature of the syrinx 
is the large extent of the membrana tympaniformis externa, due 
to the concave lower border of the first bronchial ring and the 
concave upper border of the second bronchial semiring. This 
characteristic feature of the syrinx of Falco is more marked in 
F. sacer than in any other species. The intrinsic muscles are 
attached to a specially thickened and apparently cartilaginous 
bar-like tract of the membrana tympaniformis externa, which 
varies somewhat in stoutness and exact position. The complete- 


* See Wunderlich, “Beitrage zur vergleichenden Anatomie und Entwicklungs- 
geschichte des unteren Kehlkopfes der Vogel.” Nova Acta Acad. C. Leop.-Carol. 
Deutsch. Ak. Naturf. Halle, xlviii. No. 1 (1884). 

+ I do not, of course, include the American Vultures, Sarcorhamphus, Gypagus, 
&c., among the Accipitres. ] 

{ Or Hierofaleo candicans. 


158 MR. F. E, BEDDARD ON THE [June 16, 


ness or incompleteness of the bronchidesmus is a feature which 
apparently differs from species to species. In /. feldeggt I find 
that the membrane is incomplete, that is to say it does not extend 
right up to the bifurcation of the bronchi. In /. sacer and 
F. biarmicus, on the other hand, the bronchidesmus is quite 
complete. Upon other species I am unable to report. 


Genus HiERACIDEA. 


This genws, represented for me by the species //. berigora alone, 
has a syrinx belonging quite to the same type as that of Falco. 
In fact a general description would apply equally well to both, an 
obvious justification of the close alliance usually asserted to exist 
between them *. 

Genus Minvaco. 


This genus, as represented by Milvago chimango (text-fig. 16) 
and J. chimachima, is essentially Falconine in the characters of 
its syrinx, and does not appear to me to come nearer to Polyborus 
than to Falco. 


Text-fig. 16. 


Syrinx of Milvago chimango, lateral view. X 2. 


The first bronchial ring is very thick, especially posteriorly, where 
it curves down and bends forward; it is to this piece that the 
intrinsic muscles are mainly attached, though they are also 
inserted into the membrana tympaniformis externa. It is, I take 
it, the homologue of the thickening of the membrana tympani- 
formis which is so characteristic of Falco. 


Genus HERPETOTHERES. 


Herpetotheres cachinnans has a syrinx which is constructed upon 
the Falconine plan. 


* T may remark, however, that the backwardly directed papilla upon the tongue 
are shorter than in any Falco. 


1903. ] SYRINX OF THE ACCIPITRES. 159 


Genus PoLyBorus. 


The syrinx of P. brasiliensis is constituted upon the plan of 
Falco, but there are a number of differences in detail. 

There is the same extensive membrana tympaniformis externa 
into which is inserted the intrinsic muscles. But the last six 
tracheal rings are more thoroughly fused into a box, this fusion 
taking place in front. At the same time the last three or four of 
these rings increase rapidly in diameter so that the lower part of 
the trachea is very wide, much more so than in Falco. The 
intrinsic muscles, moreover, are distinctly smaller in proportion in 
Polyborus. 


Genera TINNUNCULUS and ERYTHROPUS. 


The syrinx of the Common Kestrel (7. alaudarius) appears to 
me to need no special description. It is quite like that of Falco, 
as is also that of Hrythrops vespertinus, which genus perhaps hardly 
needs to be separated from Falco. 


Genus NISAETUS. 


The syrinx of Visaétus fasciatus (text-fig. 17) will serve as a 
type of the Aquiline form of this organ as contrasted with the 
Falconine. 


Text-fig. 17. 


Syrinx of Nisaétus fasciatus (from a drawing by the late Prof. Garrod): the 
left-hand bronchus seen from in front, the right from behind. 2. 


The most salient difference is the absence of a specially enlarged 
area of membrane between two of the early bronchial semirings ; 
that is to say, the membrana tympaniformis externa is not a con- 
spicuous feature of the syrinx of this bird, as it is of that of Falco. 
The ring which gives rise to the pessulus in front is separated by 
four semirings from that which receives the insertion of the 
intrinsic muscles; but some of these rings, three at any rate, are 


160 MR. F. E. BEDDARD ON THE [June 16, 


united to the pessulus. Posteriorly the pessulus is fused with 
the ring in front of that from which it originates anteriorly. 


Genus SPIZAETUS. 


The syrinx of S. ceylonensis presents no differences from that 
of S. orientalis, and the genus itself should, I think, be placed 
near to Visaétus by virtue of its syringeal characters. 

There is, however, a complete union between the rings which 
are connected with the pessulus, although the intrinsic muscles 
are inserted on to a semiring which appears to correspond exactly 
to that which bears the same muscles in Visaétus. 


Genus PoLYBOROIDES. 


T found, much to my surprise, that the syrinx of P. typicus 1s 
not like that of Polyborus, that it does not conform to the 
Falconine, but to the Aquiline, or Buteonine, type. 

The bronchial rings are close together, and there is no spacious 
membrana typaniformis externa as in Polyborus and Falco. 1 
find also that the intrinsic muscles are attached to the second 
bronchial semiring. The name of this genus is clearly unfortunate, 
as is its association with Milvago and Polyborus in the ‘ List of 
Vertebrated Animals’ *. 


Genus VULTUR. 


The two species, V. auricularis and V. calvus, which I have 
examined, show rather more differences than might perhaps have 
been expected. The differences concern the musculature of the 
organt. In V. auricularis the muscles are inserted upon the 
second bronchial semiring; in V. calvus the bulk of the fibres are 
inserted upon the same ring, but a good many bundles of fibres 
stray down a few rings beyond. 

Except for the fact that the connections of the pessulus are 
“normal,” the syrinx agrees in structure with that of Gyps. It 
is moreover cartilaginous, the pessulus alone being bony. 


Genus GyYPs, 


The syrinx of Gyps rueppelli shows one characteristic feature 
which I have not observed in any other Accipitrine bird. The 
pessulus, or threeway-piece, is not coossified posteriorly with any 
of the tracheal rings. It is only connected by membrane, and 
forms therefore a movable triangular plate, which (in the spirit- 
preserved specimen at any rate) is depressed below the level of the 
tracheal rmgs which abut upon it—thus giving a peculiar and 
unusual appearance. The pessulus itself is ossified, the rings of 
the trachea and bronchi are cartilaginous. The intrinsic muscles 


* List of the Vertebrated Animals now or lately living in the Gardens of the 
Zoological Society of London, 9th ed. 1896, p. 403. 
+ The horny papilla upon the tongue differ in the two species. 


1903.] SYRINX OF THE ACCIPITRES, 161 


are attached to the middle of the second bronchial semiring. ‘The 
first three bronchial rings are closely laid by each other and the 
tracheal rings; larger interspaces separate the ensuing bronchial 
semirings ; but there is no wide membrana tympaniformis externa. 


Genus Minvus. 


In this genus, at least in the species J. ictinuws, the bronchi- 
desmus is incomplete. There is no conspicuous membrana 
tympaniformis externa. The intrinsic muscles are attached to 
the second of the entirely free semirings, 7. e. those which are not 
fused with the pessulus. The syrinx therefore is constituted on 
the plan of that of Visaétus. 


Genus DRYOTRIORCHIS. 


D. spectabilis is a Hawk with an altogether unusual form of wind- 
pipe (text-fig. 18): that is to say, unusual as to details, for it dis- 
tinctly belongs to the Aquiline as opposed to the Falconine division 


Text-fig. 18. 


Syrinx of Dryotriorchis spectabilis. X 3. 


of the group. In this genus the bronchial character of the syrinx, 
suggested among the Aquiline forms, is more strongly emphasised 
than in any other Accipitrine genus which I have had the 
opportunity of examining. In fact this Hawk may be fairly 
described as possessing a bronchial syrinx. This syrinx is, how- 
ever, an exaggeration of the conditions occurring in such a form 
as WVisaétus. In that genus and its allies four or five rings, which 


Proc. Zoou. Soc.—1903, Vou. II. No. XI. 11 


162 MR. F. E, BEDDARD ON THE [June 16, 


are more or less incomplete, intervene between that bearing 
the pessulus and that upon which the intrinsic muscles of the 
syrinx are inserted. The same is the case with the syrinx of 
Dryotriorchis; but the tracheal rings immediately preceding that 
to which the intrinsic muscles are attached le well below the 
pessulus, are joined internally by the membrana tympaniformis 
interna, and clearly constitute a portion of the bronchi. It will 
be recollected that in other groups of birds*, some of the members 
of which possess the bronchial form of syrinx, the first semirings 
or rings of the bronchi are tracheal in character, and differ from 
the ensuing semirings which belong to the bronchi proper. In 
fact it appears as if the bronchi in the bronchial syrinx were partly 
formed by a split trachea, and partly by semirings belonging to 
the bronchi proper. 


The tongue in the Accipitres shows characters which allow of 
the division of the group into a Falconine and an Aquiline series. 


Text-fig. 19. Text-fig. 20. 


SSS SSS 
SPs 


Text-fig. 19.—Tongue of Milvago chimango, dorsal aspect. X 2. 
Text-fig. 20.—Tongue of Gypohierax angolensis, dorsal aspect. X 13. 


In several species of Falco, in Tinnunculus alaudarius, Hiera- 
cidea berigora, and Milvago chimango (text-fig. 19), the back part of 


* Owls, Cuckoos, and Goatsuckers. 


1903.] SYRINX OF THE ACCIPITRES, 163 


the tongue is covered by several rows of backwardly directed spines, 
in addition to the row of larger spines along the posterior edge of 
that organ. On the other hand, in one or more species of the 
genera Milvus, Circus, Leucopternis, Spizaétus, Archibuteo, Dryo- 
triorchis, Morphnus, Asturina, Gypohierax (text-fig. 20, p. 162), 
and Urubitinga, the tongue has only the posterior row of spines. 
Vultur is peculiar in possessing, in addition to the posterior row of 
spines, a lateral row on each side along the thick edge of the 
tongue. There are traces of them in Aguila, Nisaétus, and 
Gypaétus, and perhaps in some other forms. 


Conclusions. 


The principal conclusion of the foregoing enumeration of facts 
concerns the classification of the Accipitres. It is perfectly plain, 
as I think, that the structure of the syrinx permits of, or indeed 
necessitates, the division of the Accipitres into two families or 
subfamilies, which may be termed Falconidee and Buteonidee (or 
Falconine and Buteonine). It is important to notice that in 
insisting upon such a division, | am in complete agreement with 
Dr. Suschkin* and Mr. Pycraftt, both of whom have lately 
attempted a classification of the group on Osteological grounds. 
This satisfactory result shows the value of the form of the syrinx 
in this group, as has been shown with the syrinx of other groups, 
such as the Cuckoos. 

As to further subdivisions of these two main divisions, the facts 
at my disposal do not allow of any expression of opinion. Nor 
do I venture to lay too much stress upon the apparent agreement 
in the structure of the tongue with the facts derived from a study 
of the syrinx and bones, since the material at my disposal was not 
very abundant. 

The next most important point which I have been able to 
ascertain is the development of a bronchial syrinx in Dryotriorchis. 
This makes it more plain than ever that it is dangerous to compare 
birds belonging to different groups which happen to possess the 
bronchial form of the syrinx; for while it is possible on other 
grounds that the Goatsuckers and Owls, both of which contain 
genera which have bronchial syringes, are allied, it is not probable 
that they, the Cuckoos, and the Hawks are all nearly akin. It 
seems to me to be a form of syrinx which has been acquired 
several times and independently. 


* Zool. Anz. vols. xxii. & xxiii. 1899 & 1900. 
+ P. Z. S. 1902, vol. i. p. 315. 


11* 


164 MESSRS, L. MURBACH AND C, SHEARER ON [June 16, 


7. On Meduse from the Coast of British Columbia and 
Alaska. By Louis MurpacH and CRESSWELL SHEARER, 


¥.Z.8. 
[Received April 8, 1903. } 


(Plates X VIL-XXIL*) 


INTRODUCTION. 


The Meduse forming the basis of the present paper were 
collected by one of the authors, during a short trip to the coast of 


British Columbia, in the summer of 1900, in company with - 


Prof. MacBride, of Montreal. ‘They were obtained mostly in the 
immediate vicinity of Victoria Harbour, Vancouver Island, and 
the adjoining waters of Puget Sound. The rocky nature of 
the coast, shelving off rapidly into deep water, combined with the 
strong tidal currents of the main portion of Puget Sound which 
sweep in close to the shore, make this a very favourable point for 
the collection of pelagic life. Victoria Harbour itself, almost 
completely land-locked and several miles in length, affords shelter 
for many Medus not found in the more exposed and open waters 
of the Sound. 

We are indebted to Prof. Kincaid for kindly placing at our 
disposal a small but well-preserved collection of Medusee from 
Alaska and the region of Puget Sound about Port Townsend. 
The range of our original collection has been considerably 
extended, and we have obtained valuable material for comparison 
fyom different localities. While the number of species obtained 
is small, we have been fortunate in securing numerous examples 
of almost all of these. Where only a few individuals of a species 
have been obtained, doubt has been expressed as to identity. 
Measurements apply to material preserved in 3 per cent. formalin 
in sea-water. 

Our best thanks are due to Dr. Newcomb, President of the 
Natural History Society of Victoria, B.C., for the facilities he 
kindly afforded us in collecting and for much general inform- 
ation regarding the fauna, for which we feel deeply grateful, 
and to Mr. Adam Sedgwick, of the Morphological Laboratory, 
Cambridge, for placing the resources of the Laboratory at our 
disposal. Up 

The list of species represented in the collection is as follows :— 


Codonium apiculum, sp. nov.t 
Syndiciyon angulatum. 
Dipurena dolichogasier. 


* For explanation of the Plates, see p. 192. 


+ This list, with a brief notice of the new species, has been ublished in the 
‘ Annals and Magazine of Natural History,’ ser. 7, vol. ix. (1902). 


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MEDUSA FROM BRITISH COLUMBIA AND ALASKA. 


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Bs, 1903; vol MP Xe: 


E.Wilson, Cambridge. 


MEDUS& FROM BRITISH COLUMBIA 
AND ALASKA. 


1903. ] MEDUS FROM BRITISH COLUMBIA AND ALASKA. 165 


Turris breviconis, sp. nov. 
Hippocrene mertensi. 
Thaumantias cellularia. 
Polyorchis minuta, sp. nov. 
Proboscidactyla brevicirrata. 
DPhialidium languidum. 

9 gregartum. 
Mesonema victoria, sp. nov. 
Gonionemus vertens. 

» Agassiz, Sp. NOV. 
Muggica kochii. 


A. ANTHOMEDUS A. 
I. Coponip@ Haeckel 


Coponium Haeckel (18, p. 13). 


1. Coponium ApicuLuM, sp. nov. (Plate XVII. fig. 1 and 
Plate XXII. figs. 4 & 5.) 


Specific description.—The bell is nearly one-half taller than it 
is broad (1°5 cm. by 1 cm.). In some individuals the difference 
between height and breadth is not so marked. The diameter at 
the velum is somewhat less than it is nearer the apex. The apical 
process on the external surface is small and not abruptly set off. 
The apical canal (Stielcanal) is likewise short but always present. 
The velum is well developed. 

The four tentacles are rather stout, and in the contracted con- 
dition are two-thirds to one and a half times the diameter of the 
bell in length. They are attached to the bell-margin by large 
prominent tentacle-bulbs, brownish in colour, having each a 
distinct black ocellus. On either side of each tentacle-bulb is a 
large nematocyst-pad (¢f. Gronberg, 17). 

The stomach passes without distinction into the proboscis, 
which is cylindrical and contracted into circular ridges. The thin 
membranous end bears the small circular mouth, which is not 
lobed. The stomach is very distensible, being filled in most of 
our specimens with a mass of small Copepods. Often it is only 
the upper end, near the attachment to the bell, that is so 
distended ; when this is the case the lower end is usually con- 
tracted and collapsed, as shown partially in Pl. XVII. fig.1. The 
proboscis seems seldom to be extended beyond the velum, although, 
if required, it can be protruded a distance of over a centimetre. 

The radial canals are very slender, and terminate in circular 
openings in the upper end of the narrow stomach. Above the 
point of their entrance into the stomach is the apical canal. In 
the circular folds of the stomach are masses that look like gonads ; 
the preservation, however, was not sufficiently good enough to 


166 MESSRS. L. MURBACH AND C, SHEARER ON [June 16, 


make certain of this in sections. No medusa-buds were observed, 
although they were sought for in a large number of individuals. 
In sections, the lumen of the stomach-cavity is seen to be quad- 
rangular (Pl. XXII. fig. 5). In some examples the approximation 
of the stomach-walls forces out the corners of this quadrangle, so 
that they are thrown into folds something like the condition 
deseribed by Linko (25) in Sarsia brachygaster. 

Colowr.—Bell-margin and radial canals pale blue, proboscis 
reddish violet or light purple. Ocelli black. 

Habitat.—Victoria Harbour, Puget Sound, collected by Shearer. 

Discussion.—This Medusa is perhaps the commonest species in 
the waters of Puget Sound during the month of July. It was 
seen every day in great numbers, frequently the shoals or banks in 
places so dense as to obscure the colour of the water. The 
Medusze allow themselves to drift passively in the tidal currents, 
once in a while making a few vigorous contractions of the bell, 
then remaining quiet as before. While being carried along in 
this manner, their tentacles are extended a considerable length 
behind them, one individual having been noticed with its 
tentacles extended a distance of over 9 em., although its bell 
measured less than one centimetre in height. When suddenly 
taken up from the surface of the sea with their tentacles in the 
extended condition, they contract them quite slowly and with an 
invegular jerky motion, the animals taking two or three minutes 
to contract them to the normal length. 

The size and length of the proboscis and stomach in the 
Codonidee seem to be subject to great variation; this is markedly 
the case in this species, in which it varies from a short stump 
several millimetres long to a condition in which the stomach 
protrudes almost beyond the velum. One individual of C. api- 
culum, kept for several days in a small jar of sea-water, had a 
habit of resting about half a centimetre from the bottom of the 
jar, allowing its proboscis to drop down every now and again. 
‘As soon as this touched the bottom it was rapidly withdrawn into 
the bell, and then slowly allowed to drop down again. Many 
examples of this Medusa which have been preserved in formalin 
show interradial contractions of the bell-surface, giving it a 
cubical appearance; these contractions are not present in the 
living condition, and are caused by preservation. It may be 
contractions similar to these which Hincks (22) mentions as 
interradial on the bell of Sarsia (Codoniwm) pulchella, and 
which Haeckel calls “interradial furrows” in the same species. 
The examination of a large number of individuals shows 
considerable variation both in the size of the bell, the shape 
of the apical canal, the general shape of the proboscis, and 
in the length of tentacles. Apart from these variations, its 
specific distinctive characters would seem to be: the truncate 
shape of the bell; the small apical process on the exumbrella ; 
the short pointed apical canal, which is never knob-shaped ; the 


1903. ] MEDUS& FROM BRITISH COLUMBIA AND ALASKA. 167 


pear-shaped tentacle-bulbs, having each two nematocyst pads ; 
the relatively short proboscis and tentacles. 

Fewkes (14) has described a Medusa from the Southern 
coast of California under the name of Syncoryne rosario, the 
hydroid form of which he thinks is the Coryne rosario of Agassiz. 
Although he figures it with no apical process it is undoubtedly a 
true Codonid. It seems to be very closely related to our species, 
so far as can be determined. Possibly it is a southern variety, 
or possibly the two are identical. From Fewkes’s description it 
is difficult to definitely identify his species. While it is possible 
to determine little from his figures and description (the latter 
answering for almost any known Codonid), it seems his 
species possess a rather sharp conical bell, no apical process, and 
a proboscis a third shorter than in our species. On the other 
hand, if he identifies it with Syncoryne or Coryne rosario, then 
from what we know of Agassiz’s (2) description of this 
medusa, the bell would not be so pointed, and the proboscis 
would be longer than in our species. In a memoir published the 
same year as the paper mentioned above, but in a different 
journal (15), Fewkes names it Syncoryne occidentalis, sp. nov. ; 
no reference is made to his former paper or classification, the 
same drawing and description being used for both papers. It is 
unfortunate, after finding this Medusa in such abundance at 
different places on the Californian coast, he has not given us a 
more definite description. 

Brown (8) has remarked, in speaking of C. pulchellum, that 
while he has always found the apical process (the ‘“ Scheitel- 
Aufsatz” of Haeckel) present, the “Stieleanal” is frequently very 
difficult to distinguish and is sometimes absent. In C. apiculum 
absence of the “ Stieleanal” was never observed, although 
numbers were examined with this point in view. On the 
other hand, the apical process, or ‘‘Scheitel-Aufsatz,” was very 
poorly marked in some individuals, especially in small and young 
specimens, some of which had the sharp conical bell shown in 
Fewkes’s figure (14, pl. xxv. fig. 7). 

Agassiz (2) in his figure of Coryne rosario, p. 177, repre- 
sents it as possessing a “ Stielcanal” but no apical process, the 
top of the bell beimg rounded. Haeckel (18) divided the 
original genus Sarsia of Forbes into the two genera Codoniwm 
and Sarsia.  Codoniwm is characterised by the possession of 
an apical process and “Stielcanal,” Sarsia by their absence. 
While Haeckel places the Sarsia pulchella of Forbes under 
Codoniwm (this species, according to Brown, sometimes having 
the “Stielcanal” absent), he has retained the Coryne rosario of 
Agassiz under Sarsia, although Agassiz plainly represented it 
with a “Stielcanal.” Both these forms possess certain characters 
of each genus. 

Codonium apiculum differs from C. princeps in the shape of 
the bell, the tentacles, and the tentacle-bulbs. It also differs 


( 
168 MESSRS. L. MURBACH AND C. SHEARER ON [June 16, 


from (. princeps, figured by Grénberg (17), in the shape of the 
bell, the proboscis, and the tentacle-bulbs. It differs from 
C. codonophorwm in the shape of the bell and of the peduncle, 
this species producing medusa-buds. It differs from C. pul- 
chellum in the shape and size of the bell and the tentacles: this 
species again produces medusa-buds. Brown (8, p. 473) states 
that he has found medusa-buds in the young of Amphicodon 
fritillaria, Margellium octopunctatum, and Lnzzia blondina. He 
thinks that it is probably characteristic of the immature condition 
to produce medusa-buds, while in the adult condition there are 
gonads on the stomach. For this reason he considers it probable 
that O. gemmiferum and Sarsia prolifera, which both produce 
medusa-buds, and are regarded as distinct species, will ultimately 
prove to be only the early stages of other known species. The 
peculiar shape of the bell in Haeckel’s (18) C. conicwm (Nachtrag, 
p- 634) can hardly be compared with C. apiculum. It is also 
different from C. gemmiferwm in the size and shape of the bell, 
the tentacles, and the tentacle-bulbs. The last-named species 
also produces medusa-buds, 

The various species at present included under the family 
‘odonidee certainly need careful revision; until more is known of 
their individual life-histories it is difficult to estimate their syste- 
matic value. Undoubtedly, as Brown has suggested, many will 
prove to be immature stages of other species. Whether this 
will be the case with C. apiculum can only be said when its 
life-history is known. Without any knowledge of this it is 
difficult to find a place for it under any of the species already 


known 
Synpicryon A. Agassiz (Haeckel, 18, p. 20). 


1. Synprcryon ANGULATUM (Mayer). 

Specific description.—In outline the shape of the bell is that of 
a truncated oval, 1:5 mm. high by 1 mm. broad at the velum. 
In the preserved condition it has the peculiar quadrangular form 
mentioned by Mayer (27, p. 5) in his recently described species 
S. angulatum. The wall of the bell is thin and is usually 
covered with nematocysts, but these are absent in some examples. 
Tn all but one of the specimens there is a depression in the top 
of the bell, and this is met by a canal from the subumbrellar 
surface forming a complete tube through the apex of the bell. 
This is the remains of the canal originally connecting the animal 
with its hydroid form. 

The velum is well developed. The four tentacles are much 
contracted, apparently swelling at thei distal ends, and covered 
with rows of nematocysts arranged in a spiral manner. The 
tentacles are attached to the bell-margin by means of large 
tentacle-bulbs, which bear on their outer surface a slight elevation 
containing the ocellus (cf. Mayer, loc. cit.). 


1908. | MEDUS FROM BRITISH COLUMBIA AND ALASKA. 169 


The base of the stomach is small, gradually enlarging into a 
cylindrical proboscis, which extends from nearly two thirds to 
the whole length of the bell. In the contracted condition it 1s 
more spindle-shaped. ‘The mouth is small and circular, without 
lobes or folds. 

The radial canals are four simple tubes running from the 
highest point of the stomach to the circular canal. The latter runs 
near the inner wall of the bell and makes the quadrangle that is 
seen from the under surface in Agassiz’s figure (2, p. 178) of 
S. reticulatum. No gonads are to be distinguished. 

Colowr.—Bell light brown, proboscis yellow, ocelli black. 

Habitat.—Victoria Harbour, collected by Shearer. 

Discussion.—The presence of urticating-organs on the exum- 
brellar surface of the bell (unless they are arranged in a definite 
manner retained in the adult stage) used as a generic character 
by Haeckel (18) is invalidated by the fact that these structures 
occur in many young Meduse, e.g. Awrelia flavidula, Dinematella 
cavosa (Fewkes), Proboscidactyla flavicirrata, Gonionemoides geo- 
phila, Polyorchis penicillata, and the young of Sarsia. This is 
emphasised by Agassiz’s statement that this character is lost 
in the adult Syndictyon. The bell in one of our examples was 
entirely free from nematocysts. Agassiz (2) indicates a similar 
apical canal in his 5. reticwlatwm just freed from the polyp-nurse 
(fig. 291, p. 178). 

The absence of gonads, the presence of this apical canal, and 
the presence of urticating-organs on the exumbrellar surface of 
the bell indicate the immature condition of our examples. 
Possibly they may be the young of some other species; it has 
been placed under the species it resembles most. Syndictyon 
angulatwm was found by Mayer (27, p. 5) off Turks Islands in the 
Bahamas. , 


DipurENA MacCrady. 


1. DrpurENA DotrcHoGAsTER (Haeckel, 18, p. 25). 


Specific description.—The bell is a low truncated cone, 2 mm. 
high by 1:75 mm. broad in the region a little above the tentacles. 
From this, its widest diameter, the bell draws rapidly inwards, 
ending in a rounded cone-shaped top. The velum is narrow. 
The four tentacles are small and rather spindle-shaped, and in the 
contracted condition are surrounded by six or seven rings or 
welts of urticating-organs. The end-rings cause the appearance 
of bulbs at the base of the tentacles. 

The tentacle-bulbs are very large in proportion (15 by 1°25 mi.) 
to the other organs, somewhat crescent-shaped, seated in a 
curved depression on the bell-margin. On the outer end of 
each bulb there is an eye-spot. 

The digestive cavity is a large cylindrical tube hanging down a 


170 MESSRS. L. MURBACH AND C. SHEARER ON [June 16, 


distance of about two-thirds the length of the bell-cavity. The 
proboscis is long, and the mouth a plain circular opening. The 
radial canals run from the tubular end of the stomach slightly 
upwards and then downwards to the circular canal of the bell- 
margin. Their course is marked by refractive granules which 
look like nematocysts, but which are not regularly arranged. No 
gonads were seen; it is probable the single example obtained was 
immature. 

Colour.—Organs pale blue. 

Habitat.—Victoria Harbour, collected by Shearer. 

Discussion.—Some doubts as to the accuracy of the above 
identification are raised by several of the characters, the most 
striking being the tentacle-bulbs. Only one individual was 
obtained and that had been badly preserved. 


Il. Trarip# Haeckel. 


Turris Lesson. 


1. TURRIS BREVICONIS, sp. nov. (Plate XVIII. figs. 1 & 2.) 

Specific description.—The bell is 45 cm. high by 3°5 em. broad 
at the level of the velum. It is somewhat cubical and quite 
massive in appearance. ‘The general shape of the bell corresponds 
with Haeckel’s (18) figure of Tiara pileata (pl. 3. fig. 7). The 
velum is well developed and strong. On some parts of the bell- 
margin there appears to be but one row of tentacles; for the 
greater part, however, there are two rows, arranged in a zigzag 
manner. The tentacles are numerous, over a hundred and forty 
in all. They seem quite uniform in size when mature; the smaller 
ones are more irregular and evidently less developed. They are 
coiled and show the structure peculiar to coiled tentacles, as do those 
of Physalia, the ectoderm being very much enlarged on one side, 
while the contractile fibres of the inner side are covered with but 
a thin layer. There are no special tentacle-bulbs, but the tentacles 
spread out, clasping the bell-margin, as Haeckel (18) has described 
for Tiara pileata. In the preserved condition no eye-spots could 
be made out. 

The walls of the stomach are very much folded, and pushed out 
into pouches on either side of the radial canals, forming dependent 
diverticula from the angles of the stomach. There are four or 
five pairs of these diverticula. They bear the gonads, and are 
suspended to the radial canal by a double band or mesentery 
(cf. Haeckel, 18). The proboscis is poorly developed. 

The four rather large mouth-lobes are perradial in position, 
scalloped, and finely fringed. 

The gonads and stomach occupy less than the upper half of the 
bell-cavity. The radial canals are spindle-shaped in outline, 
Haeckel’s “lanzettformig.” Throughout thei length they have 


1903. | MEDUS FROM BRITISH COLUMBIA AND ALASKA. 171 


well-marked unbranched lateral diverticula. These diverticula are 
absent on the upper margin of the circular canal, and are thus 
unlike the condition described in Catablema by Haeckel. 

Colour.—Bell bluish throughout ; tentacles, gonads, and stomach 
dark red or purplish. 

Habiiat.—St. Paul Island, Pribyloff Islands, collected by 
Kineaid. 

Discussion.—The large number of tentacles in double rows and 
the opposite position of the diverticula of the stomach, bearing 
gonads, are the reasons for placing this form under Turris. It is 
true that the stomach and gonads are small for Twrris, but the size 
of these organs is variable. Beyond these considerations this form 
seems to resemble more Tiara pileata than any species of Turris. 
It differs from Turris digitalis in the smaller apical process, the 
size of the gonads, the shape of the radial canals, and the length 
of the manubrium ; from 7’. neglecta in the shape of the stomach 
which is not cubical, in the regular opposition of gonad-pouches, 
and in the number of tentacles. The specific characters are :— 
the short conical apex, the large number of tentacles (140), 
arranged in two rows, the spindle-shaped radial canals having 
simple lateral diverticula throughout their length, the five or more 
pairs of gonadial pouches from each of the four stomach-lobes, the 
small size of these and the stomach ; in these respects it approaches 
the condition in Tiara, and, lastly, in its size. Turris digitalis, 
the original Medusa digitale of Fabricius, 1s remarkable for its 
extensive distribution in northern waters; it is. probable that 
Turris neglecta will be found to have a similar distribution. Four 
out of the seven genera of Haeckel’s subfamily Pandeide have 
been reported from arctic waters. 


Til. Marceuip& Haeckel. 
HiprocrEenE Mertens. 


1. Hrerocrene MERTENSI (Haeckel, 18, p. 92). 


Synonym Bougainvillia mertensit Agassiz. 

Specific description.—The bell is nearly spherical, somewhat 
flattened, and contracted at its four meridional points, having 
thus a cubical appearance, 5 mm. by 4mm. The velum is well 
developed. The oral tentacles are situated at the upper end of 
the proboscis—two large, one small, and one rudimentary : this 
last being due possibly to imperfect preservation or an abnor- 
mality of the specimen. There is a marked inequality of the 
tentacles and the tentacle-bulbs. The former are stout, branched 
at the base, each branch dividing dichotomously about six times. 
Two of the tentacle-bulbs are large, horseshoe-shaped, and twice 
as broad as the proboscis. Hach bears twenty-one tentacles 
and as many eye-spots. The other tentacle-bulbs could not be 


172 MESSRS. L. MURBACH AND C. SHEARER ON [June 16, 


clearly distinguished. The radial canals are four in number, 
simple, and opening into the angles of the stomach. The stomach 
is short and broad, a short proboscis connecting it with the lobed 
mouth below. The gonad-masses are interradial. No gastral 
peduncle could be distinguished. 

Habitat.—Victoria Harbour, collected by Shearer. 

Discussion.—The single specimen was so badly preserved as to 
render certain identification almost impossible. It has been placed 
under H. mertensii, the form it seems to resemble most. This 
Medusa was first described by Lesson (24) under the name of 
Cyanea bougainvillii. Brandt, five years later, described it from 
the drawings and manuscripts of Mertens under the name of 
Hippocrene bougainvillii Lesson, changing this again a year later 
to Bougainvillia macloviana. Lesson met with it at the Island of 
Soledad, while Mertens found it at St. Matthei Island, Behring’s 
Strait. Agassiz reports this species from the region of Puget 
Sound, Port Townsend, and the Harbour of San Francisco. 
Hartlaub (20) has suggested that Agassiz is mistaken in identi- 
fying his species with that of Mertens on account of size. Agassiz 
states that his specimens were larger than B. superciliaris, while 
Brandt describes the species as the size of an “ ordinary pea,” 
B. superciliaris measuring some 8 mm. in height ; this would make 
B. mertensti of Agassiz some 9 or 10 mm. in height. Hartlaub (20) 
has recently given an extensive description of the Heligoland 
species of Bougainvillia in the ‘ Meeresfauna von Helgoland.’ 


B. LEPTOMEDUSA. 


I. THAUMANTID# Gegenbaur. 


THAUMANTIAS Eschscholtz. 


1, THAUMANTIAS CELLULARIA Haeckel (18, p. 129). (Pl. XVII. 
figs. 2, 2a, & 2 6.) 


Synonym Laodicea cellularia A. Agassiz. 


Specific description.—The bell is rather flat, 5 to 9 em. broad 
by 2°5 to 3°5 cm. high, somewhat resembling Stawrophora in 
appearance. The tentacles form a fine fringe around the bell- 
margin, being not more than a third of its diameter in length, 
coiled up to their oval spindle-shaped tentacle-bulbs, which are so 
humerous as to almost touch one another. The number of 
tentacles is about 340. In specimens preserved in formalin 
neither ocelli nor cirri ave visible. In proportion to the size of 
the animal the velum is narrow and delicate, being only 5 mm. 
broad. 

The radial canals run from the circular tube of the bell-margin 
to the highest point in the roof of the stomach, where they cross 


1903, ] MEDUS# FROM BRITISH COLUMBIA AND ALASKA. 173 


as ciliated grooves, a condition somewhat like that shown by 
Haeckel (18, pl. iv. fig. 7, for Catablema) (this is not to be 
confused with the “ gastrogenital Kreuz” of Stawrostoma). In 
the preserved condition the walls of the quadrangular stomach are 
so low that the mouth gapes widely. The crenulated, somewhat 
twisted, oral fringes are very characteristic, from one to one and 
a half times the diameter of the stomach in length. The gonads 
are wavy or serpentine bands depending from the radial canals 
throughout almost their entire length. 

Colour.—Bell pale blue, bell-margin and gonads deep violet- 
blue. 

Habitat.—Puget Sound, Victoria Harbour, collected by Shearer ; 
Friday Harbour, collected by Kincaid. 

Mscussion.—The general appearance of this Medusa is very 
like the figure given by Forbes (16, pl. viii. fig. 1 @) for Thau- 
mantias pilosella. The bell is somewhat flatter and the tentacles 
more numerous, the gonads are also less developed. This 
flat condition of the bell is very well portrayed by Brandt’s 
(6) figure of Staurophora mertensti (pl. xxiv. fig. 2), which also 
well represents the number, colour, and appearance of the tentacles. 
There is a delicate shade of blue through the substance of the 
bell, so that when in water they would almost escape observation, 
if it were not for the darker colour of the bell-margin and of the 
tentacle-bulbs. The colour of the bell so closely matches the blue 
colour of the sea-water, that the Jelly-fish become indistinguishable, 
nothing but the dark ring of the bell-margin showing, contracting 
and expanding as the animals swim. 

It will be seen that the shape of the bell is somewhat different 
from that pictured by Agassiz for this species (2, p. 127, 
fig. 195), There is also considerable difference in size. Agassiz’s 
species measured 3 cm. across the bell; the majority of our 
Specimens measured 5 cm., and some fully 8 and 9 cm.—in fact, 
were so large that we had no suitable jars in which to place 
them. Again Agassiz states that the tentacles number about 
a hundred, whilst in the Victoria examples there are considerably 
over thrice that number, this last difference being due to increase 
in size. The number of tentacles in each quadrant of the bell- 
margin is seldom the same, and not necessarily a multiple of 
four, no two quadrants have exactly the same number, the average 
for a quadrant being 84, 

This species made its appearance suddenly in Victoria Harbour 
on July 7th 1900, in great numbers; it was abundant for three 
days, after which few were seen. Agassiz (2) also found it in 
the region of Puget Sound, July 1859. It is remarkable how 
readily foreign particles adhere to the bell-surface of this Medusa, 
little bits of débris, sand, and seaweed sticking to it with great 
tenacity ; many of our specimens were ruined on this account. 
It is difficult to say in the preserved condition whether this is due 


174 MESSRS, L, MURBACH AND C, SHEARER ON | June 16, 


to a sticky secretion on the bell-surface or to mere roughness 
of this surface on which small particles become lodged. Forbes 
(16), in speaking of 7’. pilosella, says of the bell : ““ It is transparent 
and smooth, except on the sides towards the margin, where it is 
as if woolly, being invested with minute epidermic hairs composed 
of fibrous cells.” Perhaps the presence of these foreign bodies 
was due tosome similar condition. The question cannot, however, 
be settled from formalin material, as in this fluid the bell-surface 
takes on a peculiar scaly appearance, no woolly or hairy condition 
ean be made out. Haeckel (18) has called attention to the 
similarity between this species and that reported from Greenland. 
Our specimens differ from the Greenland species in the possession 
of broader oral lobes, the bell- or bottle-shaped tentacle-bulhs, 
and the fact that the gonads are situated throughout the whole 
length of the radial canals. 


II. CAnNoTID# Haeckel. 
Subfamily Ponyvorcuipm A. Agassiz. 


Potyorcuis A. Agassiz. 


1, Potyorcuis mrnuta, sp. nov. (Plate XIX. fig. 3 and Plate 
XXII. fig. 1.) 


Specific description.—The bell is 15mm. high by 12 mm. broad, 
a truncated oval with thick walls. It is broadest above the 
middle of its height, measuring the 12 mm. already given. 

A prominent cone-shaped gelatinous peduncle depends from 
the bell-roof, and to this the stomach, the upper ends of the 
radial canals, and the gonads are attached. The bell is drawn in 
towards the mouth so that its diameter at this point is only 
9mm. The lower truncated edge of the bell is nearly as broad 
as the velum. This is strong and 2°5 mm. in breadth. There 
are eight delicate adradial lines running meridionally in the bell- 
substance. 

There are 55 tentacles—26 large, 20 medium, and 9 rudimentary. 
There are nine more spaces, so that 64 tentacles should be present 
in all. The hollow tentacles are stout and taper to form a rather 
long root at their proximal end. They are slightly swollen just 
at the junction of the root and tentacle proper forming the 
tentacle-bulb. The mature tentacles which are attached to the 
bell-margin by these long roots are carried very much as shown 
in Pl. XTX. fig.3. Small tentacles during growth move outward 
on the lower truncate bell-margin, producing thus the appearance 
of several rows one above the other. On some of the tentacles 
there appear smooth areas free from nematocysts, whereas, as a 
rule, the whole surface of the tentacle is usually covered with 
large and small nematocysts. 

The ocelli are yellowish green in the preserved condition, and 


1903. | MEDUSH FROM BRITISH COLUMBIA AND ALASKA, 175 


situated on an elevation on the outer side of the tentacle-base, 
where the latter leaves the margin of the bell. No otocysts are 
present. The stomach rests with a small angular base on the gela- 
tinous peduncle, receiving at this level the radial canals. It then 
widens into a pouch, becoming again constricted into the proboscis 
which bears the mouth below. In the living condition the digestive 
cavity is circular in outline. In the distended condition it looks 
thin-walled. The mouth has four short lobes which turn out- 
ward. Hach lobe is again lobulated or coarsely toothed. The 
four radial canals are well marked, cylindrical tubes running 
direct in their course. They pass from the angles of the stomach 
upward along the peduncular cone to the highest point of the 
subumbrellar space and then descend directly to the circular 
canal of the bell-margin. Along the whole course of these radial 
canals short lateral diverticula are given off, the ends of the 
canals alone being free from them. Only a relatively small 
number of diverticula on one side of a radial canal are placed 
opposite those on the other side of the canal. None are 
branched or have their ends enlarged in a club-shaped manner. 
On either side of each radial canal there are about 32 of these 
diverticula. They are shorter near the ends of the radial canals, 
where these approach the bell-margin being mere protuberances. 
Gradually towards the middle of the bell they increase in length, 
until they measure about twice the diameter of a radial canal in 
length. 

The gonads are long finger-shaped processes dependent from 
the proximal part of the radial canals, the part most free from 
diverticula, There are eight gonads in each of the four groups, 
the central ones being the longest and reaching to the level of the 
velum. The outer shorter gonads may be mere rudiments. The 
latter cannot be mistaken for diverticula, being thicker and 
hanging free in the subumbrellar cavity. There are a few small 
diverticula on the radial canals over the region where the gonads 
are attached. 

Colour.—Transparent and colourless except the gonads and 
tentacles, which are tinged a pale yellow in preserved material. 

Habitat.— Puget Sound, collected by Kincaid. 

Discussion.—In Agassiz’s figure (2) of P. penicillata only 22 
of the diverticula are on an average arranged opposite one another 
on the sides of the radial canals. It is very doubtful if any 
importance can be attached to the position of these diverticula ; 
certainly their opposite arrangement is unworthy the generic 
importance assigned it by Haeckel (18). In the generic diagnosis, 
p. 149, he says of the radial canals of Polyorchis: “....im 
Distal-Theile mehrere Paar von gegenstiindigen geschlechtslosen 
Fiederasten tragen.” Both Eysenhardt & Chamisso (12) and 
Eschscholtz (11) represent the diverticula on the course of 
the radial canals, in their figures of these Meduse, by cross-lines 
drawn at right angles to the canals; this produces the appearance 


176 MESSRS. L, MURBACH AND C, SHEARER ON [June 16, 


of the greatest regularity in the arrangement of the diverticula, 
exactly opposite one another. Asa matter of fact, this opposite 
character arises only from a crude manner of representation. 
Probably this kind of symmetry is as little marked in them as 
in P. penicillata or P. minuta; for these reasons in future this 
character should be omitted from the generic diagnosis. | Fewkes 
(14) draws the diverticula in his species opposite, yet identifies 
it with P. penicillata of Agassiz. Our specimens approach the 
species of Fewkes more than they do the original descriptions 
given by Agassiz for this same species. Agassiz (2) obtained 
P. penicillata in the region of Puget Sound, and also on the 
coast as far south as the Harbour of San Francisco, where he 
found it very abundant; Fewkes collected his examples at Santa 
Barbara and Santa Cruz on the southern coast of California. 
He does not doubt that these southern examples belong to the 
same species Agassiz described as P. penicillata from a more 
northern range. As already stated, this revised version of P. pent- 
cillata by Fewkes approaches very closely P. minuta ; in fact 
we have only ventured to give it separate specific rank on 
account of size, a feature of no very great importance. We only 
attach importance to it here because our form was evidently 
mature, the great development of the gonads, the number and 
length of tentacles making this almost certain. Yet the height 
of the bell is 15 mm. in P. minuta, as compared with 40 or 50 mm. 
in the P. penicillata of Agassiz and Fewkes. While Fewkes 
identifies his species with that of Agassiz, as already stated, if his 
drawings and descriptions are accurate there would seem to be 
striking differences between the two. Some of these are the 
shape of the bell, which is broader, a less developed condition of 
the diverticula on the ends of the radial canals next the circular 
canal of the bell-margin, the club-shaped even branched ends of 
these diverticula, the position of the gonads on the part of the 
radial canals descending to the stomach, while Agassiz (2) states 
they are “attached at the highest point of the four chymiferous 
tubes.” 

Agassiz figures four gonads in each group, and these reach 
halfway to the velum; Fewkes figures eight, and these reach 
almost to the level of the velum. Although Agassiz only figured 
four gonads, Fewkes says he subsequently found their number to 
be much greater. We should hesitate, however, in emphasising 
these distinctions, for Fewkes, as assistant to Prof. Agassiz, had 
doubtless ample opportunity of referring to the original specimens 
and notes of Agassiz. 

Agassiz considers his species to be the same as that described 
by Eschscholtz (11) under the name of Melicertwm penicillatum, 
p. 106. Eschscholtz gives a very short description and poor 
figure, from which it is hard to determine anything exact. It was 
found by Exschscholtz on the coast of California. Haeckel (18) 
follows Agassiz in arranging this species under P. penicillata, 


1903. ] MEDUSE FROM BRITISH COLUMBIA AND ALASKA, EP 


although he thinks they may possibly be different; in this event 
he proposes that it should be called after Eschscholtz. Blainville 
(4) gives a description and coloured figure of the MJelicertwm 
penicillatum of Eschscholtz, but these are copied directly from 
Eschscholtz without further additions, 

So far as can be judged, the species figured by Eschscholtz was 
not based on immature specimens, considering the number and 
length of the gonads, the tentacles, and the height of bell. From 
the papers of Agassiz and Fewkes we are familiar with the 
young stages of the Pacific species of Polyorchis ; these are quite 
different in essential points from Eschscholtz’s drawing of the form 
which he took off the coast of California (11, fig. 4, pl. viii.). The 
eight long tentacles, four radial and four interradial, are out of all 
comparison with the four rather large tentacles at the ends of 
the radial canals in the young stages figured by Fewkes. It is 
to Agassiz (2) that we owe the present name of Polyorchis, 
Haeckel (18) retaining this name and placing it under the Lepto- 
medusz, family Cannotidee. 

Of the three species already known, by far the most interesting 
is P. campanulatus, originally described as Medusa campanulata, 
by Eysenhardt and Chamisso (12). Here the bell is much lower 
and more conical than in the other species. It is eight-sided, and 
the sides meet in angles. These characters, combined with the 
position and structure of the gonads as found in all Polyorchids, 
areremarkably like whatis found in the Aglauride. For instance, 
the long finger-shaped gonads of Aglantha in position and structure 
are very strikingly similar to the gonads in Polyorchis, although 
they are more numerous. While there are never more than four 
radial canals in Polyorchis, as compared with the eight of the 
Aglauride, no great significance can always be attached to the 
number of radial canals. 

The possession of free “ Horkolbchen” by the Aglauride would 
seem to separate them definitely from Polyorchis, although there 
are Leptomeduse, such as Zaodice, which possess the true endo- 
dermal sense-clubs of the Trachomeduse. The most distinctive 
character between these two groups is the possession by Polyorchis 
of diverticula on the radial canals, but these undergo marked 
change during the growth of the animal. Hardly distinguishable 
in the young, they become apparent as the animal increases in age ; 
this points to their being a recent acquisition in the evolution 
of the race, probably within the limits of this particular group. 

Of the three species of Polyorchis at present known, two are 
from the Pacific, the third from the Adriatic. It has already 
been mentioned that Agassiz (2) found P. penicillata in the 
region of Puget Sound, while Fewkes (14) found it as far 
south as Santa Cruz on the southern coast of California ; it would 
seem to be, therefore, one of the few Medusa-forms common to 
both the northern and southern fauna of the West Coast of 
N. America. 

Proc. Zoou. Soc.—1903, Vor. I. No, XII. 12 


178 MESSRS. L. MURBACH AND C. SHEARER ON - [June 16, 


PROBOSCIDACTYLA Brandt. 


1. ProposcrDACTYLA BREVICIRRATA Haeckel (18, p. 160). 

Synonym P. flavicirrata A. Agassiz. 

Specific description.—The bell is a truncated oval, 7 mm. high 
by 6 mm. broad; the subumbrellar cavity occupies less than half 
this bell, leaving a clear thick mass of jelly forming the roof of 
the dome. Agassiz has well represented this in his figure in the 
N. A. Acalephe (2, fig. 280, p. 173). The velum is quite 
narrow. The tentacles are short, and number in our largest 
specimen 54, being half as long again as the diameter of the 
bell. Small tentacle-bulbs bear the dark ocelli, and young 
tentacle-buds are seen between some of the older ones. Seen 
from the aboral side, an opaque cross marks the position of the 
four lobes of the stomach, on which a portion of the gonads rest. 
The lower portion of the stomach is more cylindrical, ending in 
the tubular much-folded mouth. This sometimes appears as four 
double folds, curved outwards and upwards. The gonads le in 
the interradial folds of the stomach, and pass out along the 
unbranched portion of the radial canals. The radial canals branch 
twice dichotomously, and then somewhat more irregularly, so that 
there is finally a canal for each tentacle. Between the terminal 
branches of the radial canals are blind delicate canals running in 
centripetally from the margin of the bell, reaching halfway up. 
These canals are on the exumbrellar surface, and do not appear to 
be hollow in section; the radial canals are nearer the sub- 
umbrellar surface. Nematocysts are seen scattered in clusters 
along these tubes at varying intervals, so that many masses of 
nematocysts may be found along the course of one tube. 

Colour.—The stomach is a dirty yellow. Agassiz (2) states 
that this Medusa is quite transparent. Whilst this is true of the 
upper part of the bell, the thick yellow mass of the stomach 
renders the lower portion quite opaque. Preservation in formalin 
seems to have caused considerable shrinkage: our measurements 
apply to preserved animals. 

Habitat.—Victoria Harbour, collected by Shearer; Pleasant 
Beach, collected by Kincaid. 

Jnscussion.—Nothing was seen answering to Agassiz’s (2) 
description of the granular covering of the bell, except the patches 
of nematocysts already mentioned scattered here and there along 
the centripetal canals. Haeckel (18) has thrown doubt on the 
presence of nematocysts on these canals; they are so well marked 
in all our examples that it seems strange he should have over- 
looked them. 

This Medusa was first found by Mertens on the coast of 
Kamchatka. Agassiz (2) reports it from the region of Puget 
Sound, but Haeckel considers A gassiz’s species different from 
that of Mertens, naming it P. brevicirrata. These names are 
liable to give rise to some confusion. It is very doubtful if this 


1903. ] MEDUS FROM BRITISH COLUMBIA AND ALASKA, 179 


separation is justifiable, seeing that Brandt seems to have mis- 
understood the structure of fe species. We should hesitate, 
therefore, to base any distinctions on his account. 


Ill. Kucor1p# Gegenbaur. 
Purauipium Leuckart. 


1, PHIALIDIUM LANGUIDUM Haeckel (18, p. 185). 


Synonym Oceania languida A. Agassiz. 

Specific description.—The bell is hemispherical, about 12 mm, 
in diameter by 6 mm. high. It is folded in the peculiar manner 
described by Agassiz (2). 

The velum is narrow and delicate. The tentacles all told 
number sixty, some closely coiled, others straight. The tentacle- 
bulbs are somewhat smaller here than in P. gregaria, the tentacles 
being sharply set off from the bulbs. There are two otocysts 
between each two tentacles. 

The stomach is small and cross-shaped, the arms of the cross 
receiving the radial canals. There is no proboscis. The curled 


i=) 
and fringed oral lobes are about the same length as the arms of 


the stomach cross. The four radial canals bear the gonads on 
their distal half. They are oblong linear bodies, one-third the 
length of the radial canals, attached to their outer half, but not 
re eaching quite to the circular canal of the bell-margin. 

Colo wir. Greenish blue. 


Hebitat.— Victoria Harbour, collected by Shearer. 


2. PHIALIDIUM GREGARIUM Haeckel (18, p. 188). (Plate XX. 
figs. 1 & 1a.) 

Synonym Oceania gregaria A. Agassiz. 

Specific description.—The bell is 12 mm. by 7 mm., and so 
nearly hemispherical, The velum is quite small. Of the tentacles 
and buds destined to develop there are sixty. The tentacle-bulls 
are spherical and relatively large. The otocysts are evenly dis- 
tributed, one or two between consecutive tentacles; they usually 
contain one, sometimes two otoliths. 

The stomach is very small, quadrangular, receiving the delicate 
radial canals at its angles, and ending in the four perradial, 
moderately long, curled, and fringed oral lobes. There is no 

roboscis. The four radial canals run from the angles of the 
stomach to the circular canal, bearing on their distal half or third 
the gonads, which, however, do not reach quite to the circular 
canal. As the gonads are very narrow linear bodies, our speci- 
mens may be immature. This is borne out by the size and 
number of small tentacles. 

Colour.—White, becoming slightly leone on preservation in 


formalin, 
12* 


180 MESSRS. L. MURBACH AND C, SHEARER ON [June 16, 


Habitat. — Puget Sound, collected by Kincaid ; Victoria 
Harbour, collected by Shearer. 

Discussion.—The principal points in which this species ap- 
proaches the Pacific form given by Agassiz and Mayer (8) are the 
large size of the tentacle-bulbs, the distribution of the otocysts and 
their contents. It differs from it in the shape of the simple oral 
lobes, the tentacle-bulbs, and the position of the gonads. Several of 
our specimens seemed to agree more closely with P. variabile than 
with P. gregariwm, apparently the only difference being in the 
number of tentacles. From Claus’s paper (10) it would seem 
that cirri are present in all the Phialidia, and that they are 
usually on the sides of the tentacle-bulbs. 

Haeckel distinguishes three species of Phialidiwm—P. variabile, 
P. languwidum, and P. gregarium. Maas (26) departs from 
Haeckel in retaining the species P. flavidulum, with its larger 
number of otocysts and tentacles, Haeckel placing it under 
P. variabile. Haeckel has arranged some twenty or twenty-five 
names under P. variabile as synonyms; the original descriptions 
of many of these, as Brown has remarked, are far too vague for 
their identification to-day. Brown (8) distinguishes as distinct 
from Haeckel’s species P. buskianwm, P. temporariwm, and 
P. cymbaloideum. The great variability in the members of this 
group renders it especially difficult to determine the value of the 
various species until their hydroid forms are recognized. 

Agassiz (2), in speaking of Oceania languida (P. languidum), 
remarks on the extraordinary attitudes assumed by this Medusa. 
One of these attitudes is given in fig. 102, where the animal is 
rolled up upon itself, the opposite edges of the bell coming 
together. Many of our examples exhibited this peculiar attitude, 
while others were folded in~a three-cornered manner, something 
like the attitude in which Brandt (5) pictures his Stawrophora 
mertensvi (pls. 24 & 25). 

The specimens from Prof. Kineaid’s collection also exhibited 
a oe They were taken from a different part of Puget 

ound, 


IV. Aquorip# Eschscholtz. 
Mesonema Eschscholtz (Haeckel, 18, p. 225). 


1, Mrsonema vicrorta, sp. nov. (Plate XIX. figs. 1, 1a, & 2, 
and Plate XXII. fig. 2.) 


Specitie description.—The bell is hemispherical, 7 em. broad 
by 3°5 cm. high, tapering to a thin flexible margin. A well- 
developed velum is present. Tentacles numerous, over 100, 
shorter than the diameter of the bell. In the same row with 
the tentacles are found numerous small papille, sometimes 
between the tentacles, sometimes below them. Otocysts and ex- 
cretory papille are present. The gastric cone is not pedunculated, 
but is lens-shaped and almost hemispherical. Stomach about the 


1903.] MEDUSH FROM BRITISH COLUMBIA AND ALASKA. 181 


same depth as the oral peduncle, being very wide at the top, 
almost half the diameter of the bell. Mouth much lobed, the 
lobes being narrow, pointed, and finely toothed. There are half 
as many lobes as there are radial canals. There are about a 
hundred radial canals; these run from the peripheral canal inwards 
and upwards to the highest point of the stomach. Hach radial 
canal is covered on its outer surface by a ridge of glandular 
cells. Sometimes these are continued over the stomach and 
oral peduncle down to the lobes of the mouth, giving an appear- 
ance similar to that presented by the ovaries of Orchistoma. The 
two kinds of umbrellar papille are present. 

Colowr.—Organs and bell-margin white, with a very slight trace 
of blue in the larger tentacles. 

Habitat— Victoria Harbour, Esquimalt Harbour, collected by 
Shearer; Pleasant Beach, collected by Kineaid. 

Discussion.—On first examination this Medusa was taken to 
be Orchistoma. That it is an undoubted Alquorid is borne out 
by the presence of marginal vesicles, of which there are one or 
two between successive tentacles, the absence of eye-spots, and 
the presence of excretory papille under the velum. The papille 
which are in the same row as the tentacle-bulbs are young 
tentacles. Brandt describes (6) bodies on the inner side of 
the bell-margin which Haeckel is probably right in considering 
to be excretory papille. Brandt speaks of them as follows 
(p. 8361) :—‘‘ An der innern Seite der Basis des Saumes findet 
sich eine Anzahl kleiner, tassenformiger, an Gestalt der Cupula 
einer Hichel nicht unihnlicher Korperchen.” No mention is 
made as to their relation with the circular canal, and Brandt 
thinks they are the rudiments of a third row of tentacles. 
Evidently Mertens did not observe whether they were inside the 
velum, and perhaps his remarks refer to the young tentacle- 
buds we have described above in WZ. victoria, and not to the true 
excretory papille under the velum. Some slight confusion has 
arisen over the position of these papille. Haeckel (18) and 
Hertwig (21) describe umbrellar papillae, which are outside the 
velum (centrifugal), which are, according to them, blind tentacle- 
bases, or spurs, possessing no openings. Distinct from these 
are the subumbrellar papillae, inside (centripetal) to the velum, 
arranged one opposite each marginal tentacle-spur on the outside ; 
they are connected by an opening at their apex with the circular 
canal of the bell-margin, and have been interpreted as excretory 
in nature. Haeckel (18) expressly states (p. 119) that they are 
inside the velum, and thus in the subumbrellar cavity; yet in 
pl. xi. fig. 13 of the same work he shows them as if they were in 
the same row with the tentacles on the external surface of the 
bell-edge, and thus outside the velum; from his figure it is 
impossible to make out their true position under the velum*. 
Mayer (28) has described centrifugal excretory papille on the 

* Since the above was written, it has been found that Claus (EO, p. 13) makes 
practically the same comment, although his figures 29 and 30 are not clear. 


182 - MESSRS, L, MURBACH AND C. SHEARER ON [June 16, 


tentacle-bulbs of Zygodactyla cyanea, which are not true excretory 
papille in which an opening is present, but probably simple 
tentacle-buds or spurs. Agassiz (2) has described similar structures 
in a number of Atquorids, as Rhegmatodes tenwis, Zygodactyla 
grenltandica, diquorea albida. Possibly in some cases these may 
be the true excretory papille which he has seen but misunder- 
stood, or they may be simple tentacle-spurs, and so entirely 
different structures ; his descriptions and figures do not make this 
clear: if they are excretory papille he does not recognize their 
structure and proper position, Milne-Edwards (30) speaks. of 
tubercles on the bell-margin of Aquorea violacea, “ ayant la forme 
dun petit mamelon” (pl. i. fig. 1c, d), which may be these ex- 
cretory papille, but here, again, their structure is not recognised. 
It is to Metschnikoff (31) that we are indebted for the first 
correct description of these structures ; he was the first to recognise 
their position under the velum, the opening in the apex, and 
their connection with the circular canal. His figures are by far 
the best, although fig. 2, pl. v., is open to the same objection as 
Haeckel’s figure ; fig. 7 is, however, very plain. 

In W. victoria these structures seem to have a very regular 
arrangement, one opposite each tentacle-bulb and one in the 
middle of the space between consecutive tentacles. In Plate XXII. 
fig. 2 is represented a section passing through the bell-margin 
in the plane of one of these papille; the opening in the apex is 
distinct, being connected with the circular canal by a passage 
lined with strong cilia. These papillae do not seem to be present 
in young specimens, only the larger and older examples appear 
to possess them. It is doubtful if their function is purely 
excretory. Brandt (6, pl. v. fig. 4) shows the continuation of 
the radial canals into the mouth-lobes of his Mesonema (Zygo- 
dactyla) ceerulescens, similar to their continuation into the mouth- 
lobes of JZ. victoria, as already described. He also shows a 
peculiar rectangular communication of these canals, on the 
mouth-lobes, which is not present in our species. While col- 
lecting off the inlet of Victoria Harbour early in July, on several 
occasions lens-shaped masses were picked up, they were about 
3 cm. broad; towards the end of the month, when this Medusa 
was obtained, they proved to be the central gastric peduncle from 
which the peripheral parts had been worn and macerated away, 
leaving a hard, smooth, lens-shaped mass. Fewkes has well de- 
scribed these from M. cyanewm as “.... flat, slightly convex 
wbove, rounded convex below.” He regards them as probably 
homologous with the gelatinous peduncle of genera like Lirope, 
Geryonia, and Carmarina. M. victoria frequently contracts the 
bell when disturbed in a characteristic manner, the thin flexible 
margin being folded in, the tentacles almost rolled under the 
velum, the gastric peduncle making a rounded protuberance on 
the upper surface of the bell. Huxley (23, pl. xxxvii. fig. 11) 
shows this attitude in his drawing of Oceania? Although the 


1903.] MEDUS# FROM BRITISH COLUMBIA AND ALASKA. 183 


tentacles are not rolled in and contracted, this Medusa is evidently 
a species of Mesonema. Great difficulty was found in preserving 
our specimens ; they seemed to become very brittle on short pre- 
servation by the ordinary method in formalin, the slightest shaking 
of the bottle in which they were preserved causing them to break 
up into small fragments ; the majority of our jars reached home 
with nothing but a mass of débris at the bottom. This Medusa 
was quite common about Victoria during July, and is evidently 
as abundant on the opposite shore of Puget Sound, being repre- 
sented by numerous examples in Pyof. Kincaid’s collection from 
the vicinity of Port Townsend. When kept in captivity they can 
be readily observed opening the mouth widely right back to the 
commencement of the radial canals and then rapidly closing it 
again, wrapping the oral lobes into a corkscrew-shaped mass, 
as Haeckel (18) has represented the oral lobes in his plate of 
Polycanna fungina (pl. xiv. fig. 4). This may be repeated rapidly 
over and over again. Possessing about an equal number of radial 
canals and tentacles, this species comes under Haeckel’s subgenus 
Mesonemella, but is different from either of his two species 
M. eurystoma and M. cyaneum (Zygodactyla cyanea of Agassiz). 
Nor does it agree with Fewkes’s new species (13), J/. bairdit, 


because in this there are four times as many radial canals as 
tentacles. 


C. TRACHOMEDUS AS. 
I. Petastp@ Haeckel. 


GonionEMus A. Agassiz. 
1. GoNIONEMUS VERTENS A. Agassiz. 


Specific description.—The bell is described by Agassiz as “an 
oblate spheroid cut in two by a plane passing through the north 
and south poles, the plane of intersection containing the circular 
tube.” He also gave other features that will be embodied in this 
account. Preserved specimens are 1°75 cm. tall and 1°50 em. 
broad, being about the same size during life. The bell is consider- 
ably taller than a hemisphere, is rather thin, and tinged a yellowish 
green during life. There is a slight conical depression in the roof 
of the stomach. The velum is well developed and rather broad 
stretching almost halfway across the opening of the subumbrellar 
cavity. The tentacles are twice the longest diameter of the bell 
in length, and look wiry and somewhat heavy for the size of the 
animal. They show the ringed welts of nematocyst well developed, 
standing out very prominently. There are no true tentacle-bulbs, 
the tapering ends of the tentacles being inserted directly in the 
bell-margin; but below their insertion there are rather large, 
oval, brown basal papille. Some distance from the outer end of 


184 MESSRS. L. MURBACH AND €, SHEARER ON [June 16, 


the tentacles there is a little nodule, the glandular sucker. There 
appears to be a otocyst between each pair of tentacles. The 
stomach appears quadrate, but is attached only along the lines of 
the radial canals. The proboscis is of moderate length, bringing 
the fringed manubrium near the velum. The four radial canals 
pass to the highest point in the bell and then dip under the conical 
depression above mentioned, The gonads are a very closely folded 
band hanging from the radial canals, their free border being longer 
than the attached border ; they are thrown into folds (sinusoidally) 
backwards and forwards across the radial canals, like a ruffle or 
rallies 

Colowr.—Gonads deep red, the radial canals as deep brownish 
lines; bell yellowish green. 

Habitat— Gulf of Georgia, collected by Agassiz; Victoria 
Harbour, collected by Shearer ; Matsmets Bay and Puget Sound, 
collected by Kincaid. 

Discussion.—This is the first notice of this Medusa since its 
discovery by A. Agassiz in the waters of Puget Sound some 
forty-three years ago. They were found swimming vigorously in 
groups of threes and fours in the outer part of Victoria Harbour 
at McCauley’s Point, July 14,1900. It is hard to convey an 
idea of the remarkable beauty of this animal as it swims with its 
dark crimson gonads and dense mass of tentacles thrashing the 
water at every contraction of the bell. Agassiz (2) also states 
that he never found them swimming singly, but always in numbers. 
He says :—“ It at once attracted my attention by its peculiar mode 
of moving. I could see these jelly-fishes, with the tentacles 
spread out to the fullest extent, sinking slowly to the bottom, the 
disk turned downward; the moment a blade of kelp touches the 
disk, they stop, bend their tentacles like knees, and remain attached 
to the sea-weed by means of their lasso-cells, which are arranged 
in rings scattered thickly over the surface of the tentacles; after 
remaining attached in this way a moment, with their tentacles 
extended and mouth turned upwards, they suddenly let go their 
hold, turn upside down, contract their tentacles to a third of their 
former length, and begin their upward movement by means of 
short, rapid jerks, given by the sudden expanding and contracting 
of the tentacles as they are violently thrown out from the cavity 
covered by the veil. They keep up this rapid motion until they 
reach the surface of the water; at the instant the upper part of 
the disk touches the top of the water, the Medusa inverts itself, 


* Haeckel (18) understood this condition to represent lateral diverticula in the 
radial canals something like those of Polyorchis. For this reason he has placed it 
near Polyorchis under the Cannotidz, whereas its true relationship seems to be under 
the Trachomeduse, as we have arranged it above, and not under the Leptomeduse 
although it has undoubted characters of this class. It would seem to be similar to 
Meduse, like Laodice, which do not find a natural place under any of Haeckel’s four 
orders. For Brooks (7) has shown that Laodice possesses the true endodermal sense- 
elubs of the Trachyline, despite the many features that rank it with the Leptomedusz 
under the Leptoline. 


—— 


1903.] MEDUSA FROM BRITISH COLUMBIA AND ALASKA. 185 


and sinks, with its tentacles fully expanded, until it reaches the 
bottom, or another piece of sea-weed where it attaches itself, and 
after remaining suspended a little while, repeats the same opera- 
tion.” Agassiz has emphasised this habit of turning over in 
the specific name vertens. This power of attachment is not, 
however, due, as Agassiz states, to the lasso-cells, but to a definite 
structure—an adhesive pad, an enlarged view of which is shown 
in Pl. XXII. fig. 3, which is situated near the ends of the 
tentacles, and acts like a sucker, which is sufticiently strong to 
tear the tentacles without loosening its hold. 

This power of attachment must be of great service to the 
animal, for it prevents its being carried away by any struggling 
animal it may capture. It is quite probable this Medusa often 
captures animals fully as large as itself, as it readily tries to 
digest pieces of meat, almost twice its size, which are dropped 
into the bell-jars in which they are kept during captivity. These 
Medusxe seem remarkably hardy and remain alive in small jars 
of sea-water without change of water for several days. 


2. GONIONEMUS AGASSIZII, Sp. nov. (Plate XXI. figs. 1, 2, & 3, 
and Plate XXII. fig. 3.) 


Specific description.—The bell is 9 by 17 mm., or a little taller 
than a hemisphere. The subumbrellar surface dips down in the 
centre formimg a gastral cone. 

The velum is rather broad, strong, and well developed. 

The tentacles are shorter, more numerous than in any other 
species of this genus described. They number eighty in specimens 
of the above size. ‘They are inserted into the bell-margin at 
different levels, giving the appearance of one row above another. 
In a moderately contracted condition they are not more than 
two-thirds the diameter of the bell in length. They are much 
thicker at the base, and taper more gradually towards the tips, than 
in any of the species already known. Each is provided with 
a small glandular attachment-pad some distance from the end 
(Pl. XXIT. fig. 3). As already remarked, the proximal end of 
the tentacle is slightly enlarged just before entering the margin of 
the bell as in other species. This end of the tentacle is a tapering 
root connected with the circular canal. Just at the outer margin 
of the velum, under the circular canal, is a small ridge or welt of 
ectoderm filled with urticating-organs. It is reddish-brown in 
colour, the same as the papille below. 

There are papille under the circular canal on the bell-margin, 
just below the insertion of the tentacles, containing diverticula of 
this canal. Their outer layer is composed of the same tissue as 
that composing the nettle-ridge, and of a similar brown colour. 
There are not so many otocysts as tentacles, though there is no 
definite regularity. The pigment-spots at the bases of the tentacles 
are not visible in the preserved condition. 


186 MESSRS. L. MURBACH AND C, SHEARER ON [June 16, 


The stomach is much distorted with food, but is of the usual 
quadrangular shape, with oral lobes finely crenulated. The 
four radial canals pass from the circular tube upwards to the 
gelatinous peduncle, under which they pass downwards to the 
stomach ; their crossing lies deeper on this account than the point 
where they enter the stomach (Pl. XXI. fig. 3). They are 
simple. | 

The gonads have the frill-like arrangement on the course of the 
radial canals, from side to side, similar to their arrangement in 
other species of this genus, but they are denser and run closer 
to the circular canal. 

Colour.—This is hard to determine from preserved material, but 
it is doubtless somewhat similar to the bright colours in other 
species of this group. 

Habitat.— Unalaska, Aleutian Is., collected by Kincaid. 

Discussion.—Prof. Kincaid states that this species was collected 
in a small salt lake in the Aleutian Islands, which was doubtless 
connected directly with the sea by some underground passage, as 
its surface rose and fell with the tide. These Meduse were found 
clinging to stones by means of their attachment-pads, and when 
disturbed moved a short distance, then re-attached themselves 
again. This species probably possesses the same swimming-move- 
ments so characteristic of the other species of this genus. 

As already stated, the genus Gonionemus was founded by 
A. Agassiz for G. vertens, which he procured in the Gulf of Georgia, 
July 1859. The peculiar angle formed by the tentacles being 
bent at the adhesive pad or sucker was sufficiently striking to 
suggest the name Gonionemus for the genus. Haeckel (18) 
subsequently, understanding this name to mean “ kneed threads,” 
changed this to Gonynema*, under which name it appears in his 
‘System der Medusen.’ The resemblance of Gonionemus to 
Melicertwm led A. Agassiz to place it with the latter genus in the 
family Melicertide, the elder Agassiz (1, p. 348) placing it under 
the suborder Sertularie, 

Haeckel (18) arranges four of Agassiz’s genera among the 
Trachomeduse, but placed Gonionemus in the family Cannotide, 
subfamily Polyorchine, believing, from Agassiz’s description or 
drawing, that it possessed lateral diverticula on the radial canals 
as already mentioned. 

No further notice was taken of this Medusa until one of the 
present writers, Murbach (32) published a short preliminary 
report of the occurrence of the genus in the Atlantic at Woods 


* As Agassiz derived the name Gonionemus from ywvia and vjpua, Haeckel’s 
assumption that the first part of the name is derived from yébvv is wrong, only the 
last part of his nameisright. Yerkes, ina recent paper (Am. Journ. of Phys. vol. vii. 
p. 181), has changed the name to Gonionema, in this being followed by Perkins 
(Johns Hopkins Univ. Cir., June 1902). Since they correctly derive the first part 
of the name from ywvia, why does this become Gonionema and not Gonianema 


in the full name? In preference to further change we have retained the original 
name. 


1903.] MEDUSA FROM BRITISH COLUMBIA AND ALASKA. 187 


Holl, Mass. Here it was represented by what was thought to be 
a second species. Two years later Dr. Mayer confirmed this 
opinion, adding that Prof. Agassiz pronounced the Woods Holl 
form different from the Pacific form G. vertens, and Mayer (29) 
proposed the name G', murbachi. 

Agassiz and Mayer (3) described a new species, G'. swavensis, 
from Suava Harbour, Fiji Islands. Again, Mayer (28) con- 
cluded that his Cubaia aphrodite was really a species of Gonronemus. 
The development of Gonionemus seems to indicate that it belongs 
to the lower Trachomeduse ; there are also other features that 
put it in the lower Trachomedusz, such as the position of the 
otocysts, gonads, the marginal welt of nettling-organs, and the 
insertion of the tentacles. Until the life-history of one of 
the species is better known it is difficult for the present to 
determine its further position. 

_ Provisionally, Haeckel’s family Petasidee, subfamily Petachnine, 

with four radial canals, otocysts either free on the margin or 
enclosed, tentacles hollow, is best fitted for its reception. It 
might then be placed between the genera Aglawropsis and 
Gossea. 

With the exception of the number and position of the tentacles, 
the above characters are so constant in the four species now 
known that we can look forward to their being found in all true 
members of the genus. The position and insertion of the tentacles 
will vary most. 

Only a few Meduse are recorded, Bathycodon, Pectantis, &e., 10 
which the ends of some or all of the tentacles are provided with 
means for clinging to foreign objects. But these are not of the 
same nature nor in the same position as are the suctorial pads 
of Gonionemus. Recently, Mayer (27) has found a new Medusa, 
in which every fourth tentacle has an adhesive pad _ that 
corresponds somewhat with the position of that in Gonionemus, 
though unlike it in appearance. This Trachomedusa, he thinks, 
is closely related to Gonionemus, and has indicated this in calling 
it Gonionemoides. There is no reason why the presence of these 
pads, if morphologically the same, may not be one of the marks 
of relationship. The fact that the young of Gonionemoides 
geophila have pads on every alternate tentacle, not on every 
fourth as in the adult, may indicate that in the ancestral form 
they were so arranged. In this case Meduse having adhesive 
pads on every alternate tentacle would be more closely related to 
Gonionemoides than to Gonionemus, or possibly would be inter- 
mediate forms such as Gonionemus (Cubaia) aphrodite Mayer, in 
which every other tentacle possesses an adhesive pad. Behaviour, 
where it is well marked or there are special habits as in this 
case, should enter into the characters of the genus or at least the 
species description. The peculiar habit of swimming to the 
surface of the water and turning over to float lazily downward 
is well marked in both G. veriens and G. murbacht, and we may 


188 MESSRS. L. MURBACH AND C. SHEARER ON [June 16, 


expect to find it in the other species when these are closely 
studied. The four species of Gontonemus at present known are 
as follows :— 


1. Gonionemus vertens A. Agassiz (2). 


2. % murbachii Mayer (29). 
3. 5 suavensis Agassiz & Mayer (38). 
4, 9 agassizit, Sp. NOV. 


In general appearance and structure the four species differ as 
follows :— 


Gonionemus vertens has much the tallest bell and heaviest 
tentacles, longest digestive cavity, and oval large marginal 
papille.— Pacific. 

Gonionemus murbachii has the bell low, the velum well 
developed, the tentacles very flexible, the proboscis short.— 
Atlantic. 

Gonionemus suavensis differs from the other species in the 
presence of four green spots on the proximal ends of the radial 
canals, in the absence of subumbrellar papille, and the extent 
and position of the ovaries.— Pacific. 

Gonionemus agassizii has the bell lower than in G’. vertens, yet 
taller than in either of the other two species. It has a larger 
number of tentacles (in fully grown specimens probably over a 
hundred), which are shorter and smaller than in the other species. 
The depression in the roof of the stomach is more marked in this 
species than in the others with the exception of G. swavensis.— 
Pacific. 


Species incerta. (Plate XX. fig. 2.) 


The following description refers to a peculiar form, a single 
individual of which was collected by Prof. Kincaid at the 
St. Paul’s Island, Pribyloff Islands. It is very badly pre- 
served, and on this account no definite identification has been 
attempted. 

Description.—The bell is 18 mm. high by 10 mm. broad at 
its widest part, a little above the region of the velum. The 
general shape and proportions of the bell are represented 
in Pl. XX. fig. 2. The bell is drawn in somewhat towards 
the mouth, and through this the smooth cylindrical proboscis 
extends a short distance. The proboscis bears four well-marked 
cylindrical mouth-lobes, one of these being much larger than 
the others and rolled up on itself, looking like a small contracted 
tentacle. 

Connecting the four radial canals and running in the bell- 
substance are delicate transverse lines represented in the figure. 
Some of these are larger than others, and look almost like small 
connecting canals. The radial canals are thick, and along their 


1903. ] MEDUS FROM BRITISH COLUMBIA AND ALASKA. 189 ° 


7 

course are numerous irregular masses or enlargements, which may 
be gonad masses; preservation is not sufficiently good to make 
certain of this. They are a bright orange-yellow in colour, and 
become more numerous on the upper part of the radial canals 
towards the apex of the bell. The four short tentacles are 
thick and tightly contracted up to their small rounded tentacle- 
Lodo 

Colour.—Radial canals, and the masses along their course, 
bright orange-yellow. Proboscis, tentacles, and tentacle-bulbs 
yellowish brown. 

Habitat.—St. Paul’s Island, Pribyloff Islands, collected by 
Kincaid, 


D. SIPHONOPHORA. 


I. MonorpuHyi1bD@ Claus. 


Muvearxa Bausch. 


1. Mucerma Koount Chun. 
Synonym Diphyes chamissonis Huxley. 


Specific description.—The single specimen obtained is nearer 
M. kochii than any other Monophyid, although not agreeing with 
Chun’s description in all respects. Only one nectocalyx is present 
and there is no sign of another being detached. It is more 
pointed at the apex, and the oil reservoir is larger in proportion 
than those of IW. kochit. The wings of the nectocalyx are almost 
smooth, except that portion below the hydreecium which is wavy 
in outline. The contracted condition of the hydrosoma will not 
warrant a more definite determination. ; 

Habitat.— Puget Sound, collected by Shearer. 

Discussion.—Muggiea kochti is of great interest, as it was 
from the study of this form that Chun (9) found the interesting 
life-history of the Monophyids to consist of three generations, in 
this case of M. kochti, Hudouia eschscholtzii, and Monophyes 
primordialis. 

Chun’s statement (9) that the nectocalyx of Muggicea is 
very similar to the anterior swimming-bell of Diphyes and 
Diphyopsis is confirmed in this case. The position and shape of 
the parts correspond almost precisely with Mayer's figure (28) 
of Diphyes bipartita, sp. nov. (fig. 114). 

In this connection it may be noted that Fewkes (13, p. 834) 
in a footnote records that “specimens of a Diphyes (%) with but 
one nectocalyx are very common in the Bermudas and Tortugas.” 
Mayer also says he found a Diphyopsis which has no posterior 
swimming-bell, and suggests the possibility that “no such 
structure exists.” May not both these species be Muggica? 
The specimen above described differs from WHaeckel’s (19) 


190 MESSRS. L. MURBACH AND C, SHEARER ON ~~ [June 16, 
M. pyramidalis in its shorter hydrcecium, and from his other 
species in the toothing. 


In addition to the foregoing there are a number of specimens, 
not in a condition for definite identification or description. So 
far as can be determined they seem to belong to the following 
species :— 

Sarsia ecimia. ‘Two specimens 
Sarsia rosario. One specimen. 
Atollia bairdii. One specimen. 
Obelia polystyla. One poorly preserved specimen. 


Vanhoeffen (33) gives a good figure of Atollia bairdi, with 
which our specimen seems to closely correspond. 


BIBLIOGRAPHY. 


1. Acasstz, Lovrs.—Contributions to the Nat. Hist. of the 
U.S.A. Vol. iv. 1862. 

9. Acasstz, ALEXANDER.—Illustrated Catalogue of the Mus. of 
Comp. Zool. at Harvard College. No. 2. N. American 
Acalphe. 1865. 

3. Acassiz, A., & Mavzr, A. G.—Acalephs from the Fiji 
Islands. Bull. Mus. of Comp. Zool. at Harvard College, 
WO, xoemnl, WO, Ys ISIE); 

4. Buatnvitte, De, H. M. D.—Manuel d’Actinologie. Paris, 
1834. 

5. Branpr, J. T.—Prodromus Descriptionis Animalium ab 
H. Mertensio observatum. Recueil des Actes del’ Académie 
Impérial des Sciences de St. Pétersbourg, 1833-34, 

Ausfiihrliche Beschreibung d. v. C. H. Mertens a. 
Siener Weltumsegelung beobacht. Schirmq. Memozrs of 
the Imperial Acad. of St. Petersburg, ser. 6, Scien. Nat. 
50) Bp He Kots zoe 

7. Brooks, W. K.—The Sensory Clubs or Cordyli of Laodice. 
Journal of Morphology, vol. x. pp. 287-3804, 1895. 

8. Brown, E. T.— British Hydroids and Meduse. Proc. Zool. 
Soc. Lond. 1896. 

9. Cuun, Cart.—Ueber die Cyklische Entwl. u. d. Verwandt 
d. Siph. Sitz. d. kéng. Prus, Akad. d. Wiss. z. Berlin, 
1882, p. 1155. 

10. Craus, C.—Beitr. z. Ktns. d. Geryonospiden u. Eucopiden- 
Ent. Arbt. d. Zool. Inst. Wien, iv. p. 89, 1882. 

11. Escuscuo.trz, Fr.—System der Acalephen. Berlin, 1829. 

12. Eyspennarpr & CHaAmisso.—De Animalibus quibusdum e 
Classe Vermium Linneana in circumnavigatione Terre, 
auspicant Comite N. Romanzoff, duce Othone de Kotzebue 
annis 1815-18 peracta observatis. Acta Akad. Nat. Curio- 
sorum, vol. x. 1821. 


1903. ] MEDUS FROM BRITISH COLUMBIA AND ALASKA, 191 


13. 


14. 
15. 


16. 
Pe 


Fewxes, J. W.—Report on the Medusze collected by the U.S. 
F. C. Str. ‘ Albatross’ in the Region of the Gulf Stream. 
U.S. F. C. Report, 1884, pp. 927-975. 

. On a few Californian Meduse. Amer. Nat. xxiii. 

Proll | L8So: 

New Invertebrates from the Coast of California. 
Bull. of Essex Inst. xxi. 1889. Also as separate print 
for the author. Boston. 

Forbes, E.—A Monograph of the British Naked-eyed Meduse. 
Ray Soc. London, 1848. 

GronBERG, G.— Die Hydroid-Medusen des arktischen Gebiets. 
Zool. Jarb. Syst. Abt. Bd. ii. p. 451, 1898. 


. Harcxet, E.—System der Medusen. 1879-80. 


Siphonophore. ‘Challenger’ Reports, xxviii. 1888. 


. Harriaus, Cu.—Die Hydromedusen Helgolands. Meeres- 


fauna von Helgoland. 1897. 


. Hertwie, Oscar & Ricuarp.—Nervensystem der Medusen. 


Leipzig, 1878. 


. Hincxs, T.—British Hydroid Zooph. 1868. 
. Huxury, T. H.—On the Anatomy and the Affinities of the 


Family of the Meduse. Phil. Trans. 1849, p. 413. 


. Lesson.—Zoology of the Voyage of the ‘Coquille’ (under 


Duperrey). 1830. 


. Linco, A.— Beit. z. Ktns. d. Hydromedusen. Zool. Anz. 


Bd. xxv. No. 664, 1902. 


. Maas, O.—Reports on the Explorations of the W. Coast of 


Mexico by the U.S. F.C. Str. ‘ Albatross,’ during the Year 
1891. Mem. of Mus. Comp. Zool. vol. xxiii. no. 1, Die 
Medusen, pp. 1-92, 1891. 


. Mayer, A. G.— Description of new and little-known Meduse 


from the W. Atlantic. Bull. Mus. Comp. Zool. vol. xxxvii. 
HO} Le IOO! 
Some Medusz from the Tortugas, Florida. Bull. Mus. 
Comp. Zool. vol. xxxvii. no. 2, 1900. 

. The Variation of a newly-arisen Species of Medusa. 
Sci. Bull. of Brooklyn Inst. Arts & Sci. vol. i. no. 2, 
1901. 


. Mrunze-Epwarps, M. H.—Aequorea violacea. Annales des 


Sciences Naturelles, 2nd ser. t. xvi. p. 196, 1841. 


. Merscunikorr, E.—Meduse and Siphonophore. (Jn Russian.) 


Moscow, 1871. 


. Mursacu, L.—Pr. Note on the Life-History of Gonionemus. 


Journ. of Morph. vol. x1. p. 493, 1895. 


. VANHOEFFEN, H.—Die Akalephen d. Plankton-Expedition. 


1892. 


192 MEDUSM FROM BRITISH COLUMBIA AND ALASKA. [June 16, 


EXPLANATION OF THE PLATES. 


Puate XVII. 


Fig. 1. Codonium apiculum, sp. nov., P. 165. 
2. Thawnantias cellularia, p. 172. 
2a. 5 op oral lobes expanded. 
2b. as 0 oral lobe contracted. 


PruatEe XVIII. 


Fig. 1. Tunis breviconis, sp. nov., p- 170. 
PO ae 5 view of oral surface. 


PuatE XIX. 


Fig. 1. Mesonema victoria, sp. nov., p. 180. 
la. 3 55 portion of bell-margin. 
2. Be 5 oral surface. 

3. Polyorchis minuta, sp. nov., p. 174. 


PLATE XX. 


Fig. 1. Phialidium gregarium, p. 179. 
la. 0 PA crossing of radial canals on stomach-roof. 
2. Species incerta ?, p. 188. 


PratTE XXI. 


Fig. 1. Gonionemus agassizii, sp. nov., p. 185. 
2. 7 es aboral surface. 
3. . * section of bell. 


PLatE XXII. 


Fig. 1. Polyorchis minuta, sp. nov., p.174. Transverse section of a 
gonad showing the continuation of the cavity of the radial 
canal into the gonad. 

2. Mesonema victoria, sp. nov., p. 180. Section of bell-margin. 


cir.can.=cireular canal of bell-margin. 
ect.papil.= excretion papilla. 
ve.=velum. 
3. Gonionemus agassizii, sp. nov., p.185. Attachment pad. 
4, Codonium apiculum, sp. nov., p. 165. Section of bell-margin. 


cir.can.=circular canal. 
oc.=ocellus. 
ten.ba.=base of tentacle. 
ve.=velum. 


5. Codonium apiculum, sp. nov. Section of stomach. 


1903.] THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 193 


November 3, 1903. 
Dr. W. T. Buanrorp, F.R.S., Vice-President, in the Chair. 


The Secretary read the following reports on the additions made 
to the Society’s Menagerie during the months of June, July, 
August, and September, 1903 :-— 

The number of registered additions to the Society’s Menagerie 
during the month of June was 200, of which 38 were by presen- 
tation, 14 by birth, 10 by purchase, 111 were received on deposit 
and 27 in exchange. The number of departures during the same 
period, by death and removals, was 100. 

Amongst the additions attention may be called to a fine pair 
of Grévy’s Zebras (Hquus grevyi), presented by Lt.-Col. J. L. 
Harrington, C.V.O., C.M.Z.S., on June 5th. There are now in 
the Society's Gardens one male and three females of this Zebra. 


The number of registered additions to the Society’s Menagerie 
during the month of July was 98, of which 46 were acquired by 
presentation and 10 by purchase, 39 were received on deposit, 
and 3 were bred in the Menagerie. The number of departures 
during the same period, by death and removals, was 163. 


The number of registered additions to the Society’s Menagerie 
during the month of August was 130. Of these 39 were acquired 
by presentation and 11 by purchase, 1 was born in the Gardens, 
77 were received on deposit and 2 in exchange. The number 
of departures during the same period, by death and removals, 
was 180. 

Amongst the additions attention may be called to :— 

1. Three fine specimens of the Elephantine Tortoise (7estudo 
elephantina), deposited by the Hon. Walter Rothschild, M.P., 
F.Z.S., on August 8th. 

2. An Echidna (“echidna hystrix), deposited by the Hon. 
Walter Rothschild, M.P., F.Z.S., on August 20th. 

3. Four Cyprus Spiny Mice (Acomys nesiotes), presented by 
R. L. N. Michell, Esq., on August 27th. 


The number of registered additions to the Society’s Menagerie 
during the month of September was 167, of which 43 were by 
presentation and 5 by purchase, 4 were born in the Gardens, and 
115 were received on deposit. The number of departures during 
the same period, by death and removals, was 142. 

Amongst the additions attention may be called to :— 

1. Two Masai Ostriches (Struthio camelus, var. massaicus), pre- 
sented by A. Marsden, Esq., on September 8th. 

2. A fine male Chimpanzee (Anthropopithecus troglodytes), 
deposited by the Hon. Walter Rothschild, M.P., F.Z.S., on 
September 14th. 

3. An interesting collection of animals from the Harrar dis- 
trict of Somaliland, presented by William Northrup M°Millan, 

Proc. Zoo. Soc.—1903, Vou. Il. No. XIII. iN} 


194 ON A RHINOCEROS FROM THE WHITE NILE. [ Nov. 3, 


Esq., on September 21st. Amongst these the Abyssinian Duiker 
(Cephalophus abyssinicus) is exhibited for the first time. 

4. Two Wagler’s Pit-Vipers (Lachesis wagleri), presented by 
A. Herbert, Esq., on September 30th. 


Dr. P. L. Sclater, F.R.S., exhibited the front horn of a 
Rhinoceros (text-fig. 21), lately obtained on the White Nile, and 
made the following remarks :— 

“So far as I know, the only specimen of the Square-nosed or 
White Rhinoceros (Rhinoceros simus) obtained anywhere north 


Text-fig. 21. 


(a) Lateral and (4) front views of horn of Rhinoceros from the White Nile. 


of the Zambesi, of which the locality is absolutely certain, is the 
skull of an individual shot by Major A. St.H. Gibbons, F.R.G.S.. 


1903. | ON ABNORMAL CLAWS IN THE CRAB AND LOBSTER. 195 


near Lado, on the White Nile, which, by the ownen’s kind per- 
mission, | exhibited at a meeting of this Society on Dec. 18th, 
1900 (see P. Z. 8. 1900, p. 949). Its identity had been previously 
established by Mr. Thomas, who had recorded its occurrence in 
‘Nature’ of Oct. 19th of that year (‘ Nature,’ vol. Ixii. p. 599). 
This skull is now, I am informed, in the Carnegie Museum at 
Pittsburg, U.S.A. The horn of an example of the same species, 
which I now exhibit, was obtained by my friend Capt. Claude 
Hawker (Commander of the 10th Soudanese Battalion) from the 
Belgian Officers at Lado in the autumn of 1902, and was taken 
from a specimen unquestionably shot in that district, on the left 
bank of the While Nile. The Belgians did not distinguish it 
from the ordinary #. bicornis, and seemed to believe that all the 
Rhinoceroses of that district belong to one species. 

“The present specimen is a front horn of F. simus, or, at any 
rate, of a closely allied form. It agrees very well with the front 
horn of the mounted specimen of 7. simus in the gallery of the 
British Museum, but is rather longer, measuring 31 inches in a 
straight line from the base to the end. The front horn of 2. simus 
may always be distinguished from the corresponding horn of 
R. bicornis by its broad, flattened surface at the base in front, 
the basal front of this horn in AR. bicornis being more or less 
smooth and rounded and projecting in the centre. 

“Capt. Hawker has returned to the Soudan, and will probably 
visit the southernmost station of the Anglo-Egyptian forces at 
Mongalla, 15 miles north of Gondokoro, again this winter. I have 
requested him to obtain further information about this Rhinoceros, 
and have little doubt that he will do so.” 


The Secretary exhibited a series of photographs of the Indian 
Elephant in the act of congress, which had been presented to the 
Society by Mr. H. Slade, of Rangoon, Burma. 


Mr. Henry Scherren, F.Z.S., exhibited some specimens of 
the Edible Crab (Cancer pagurus) and the Lobster (Astacus 
gammarus) showing meristic variations, and made the following 
remarks :— 

‘“‘ Both these variations are in the left chela. That of the Crab 
I received from Mr. Arthur Patterson of Yarmouth, in whose 
name it will be handed over to the British Museum (Natural 
History). A process grows from the lower edge of the palm, 
resembling, though not very closely, the fixed and movable fingers. 
It is, I suggest, a case of a rudimentary extra pair of fingers, 
of which many examples have been figured by Bateson. The 
variation in the Lobster, also in the left chela, is more complex. 
There is a process growing from the upper edge of the mero- 
podite, having three spines round the anterior margin, and a 
soft, articular membrane arising therefrom. Beyond this is a 
three-pointed process—a duplicate carpopodite, which must have 
been movable in the living animal. On the normal meropodite, 


ks 


196 MR. R. I. POCOCK ON BURCHELL’S ZEBRA. [ Nov. 3, 


near the base of the outgrowth, the shell presents an appearance 
not unlike that of a snail-shell which has undergone repair, so 
that this variation would seem to lend support to the theory that 
such duplicate parts may arise from injury. This, however, 
cannot have been the case. The animal would scarcely have 
survived such an injury. The specimen was the property of 
Mr. G. A. Doubleday, who has kindly presented it to the British 
Museum.” 


Mr. R. I. Pocock, F.Z.S., exhibited two photographs (text- 
fig. 22, opposite) of a specimen of Burchell’s Zebra which has been 
preserved for many years in the City Museum at Bristol. The 
photographs were kindly taken by Mr. William Moline, and every 
facility for doing so was afforded by the Secretary and Curator, 
Mr. Bolton, F.R.S.E., with the sanction of the Committee of the 
Bristol Museum. 

The specimen is a small male, probably not quite full-grown, 
standing 44 inches at the withers. Unfortunately, its locality is 
unknown and nothing of its history can now be traced. Its im- 
portance and interest, however, lie in the fact that it belongs to the 
typical race of Burchell’s Zebra, or, as it should be more properly 
called, Burchell’s Quagga (Hquus quagga burchelli), which is either 
extinct as a wild animal or, at all events, verging on extinction. 
Hence it is desirable that the characters of every specimen now 
living in captivity or exhibited in museums should be permanently 
recorded by photography. 

A marked difference between the Bristol example and the 
typical example described and figured by Gray, but now unfor- 
tunately lost, is to be found in the distinctness and distribution of 
the paler, narrower, intermediate stripes. In the specimen sent 
to the British Museum by Burchell these stripes, as attested by 
the figure, were long, sharply defined, and extended without a 
break from the hind-quarters to the head. In the Bristol speci- 
men, on the contrary, they are short and pass from the hind- 
quarters only halfway along to the shoulder. Owing to scarcity 
of material of this rare animal, the exact systematic value of this 
difference is unknown. 


Mr. Pocock also exhibited an example of a species of Votiphilides, 
one of the Geophilomorphous Centipedes. The specimen came 
from Venezuela, and is remarkable for its great length. It 
measures 283 mm. (or nearly 11 inches) long and 9 mm. (or 
about 3 of an inch) broad—that is to say, it is, roughly speaking, 
twice as long as the average-sized specimens of the largest species 
hitherto recorded. 


Mr. Oldfield Thomas, F.R.S., exhibited specimens of three new 
Mammals, two of them representing new genera, which had been 
collected by Mr. A. S. Meek in British New Guinea. Besides 
these new forms Mr. Meek had obtained in the same region 
examples of several very rare species, such as Dorcopsis macleayj, 


99 


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Mounted Burchell’s Zebra in the City Museum, Bristol, 


198 MR. OLDFIELD THOMAS ON NEW [| Nov. 3, 


Phalanger carmelite, Pseudochirus corinne, Dasyurus albopunc- 
tatus, and Leptomys elegans, the last two having been previously 
unrepresented in the National Museum. 

The new forms were described as follows :— 


Hyomys, g. n. (Muride). 


Size very large; form bulky. Fur coarse and harsh. Feet 
proportionally short; pollex with a broad nail. Tail of medium 
length, very coarsely scaled, practically naked. Mamme 0—2=4. 

Skull stout and heavily built. Nasals very broad in front, 
narrowing rapidly behind to a point. Postorbital processes 
present, quite separate from the supraorbital ridges, which, 
though distinct, are not heavily developed. Palatal foramina 
short ; posterior palate cut out to level of front of m.* Bulle 
small, little inflated, each with a raised rim running along its 
inner margin. Paroccipital processes strongly developed. 

Incisors large, equally broad above and below, and of normal 
depth. Molars (see Pl. XXIII. figs. 2a & 2c) very large and heavy, 
their length one quarter the basilar length, and their breadth 
nearly equal to that of the palate between them ; very distinctly 
laminate, though the laminz have got the normal murine cur- 
vature. M.* and m.° each with a well-developed antero-internal 
cusp so jomed with the anterior lamina as to form a distinct 
Y-shaped structure when worn; no antero-external secondary 
cusps. M.° with the posterior overhang very unusually developed. 
Two anterior lower molars with the normal posterior supple- 
mentary cusp very large, so as to form a short extra lamina 
extending from the centre of the tooth to its inner edge. 

Type, Zyomys meeki. 

This genus was not distinguished by any single character of 
marked importance, but the gigantic Rat on which it was based 
could not be referred to any of the known Papuan genera of 
Muride. Its molars, while enormously larger, had the general 
appearance of those of Mus or Uromys, and had nothing of what 
might be called the zigzag character exhibited in Crateromys, 
See and Mallomys, to none of which did it seem specially 
aliled. 


HyoMYs MEEKI, sp.n. (Plate XXIII. figs. 2-2.) 

Fur harsh, general body-hairs about 25 mm. in length, but 
a number of bristles 60 to 70 mm. long intermixed with the 
shorter fur. General colour dark slaty greyish, the hairs grey 
proximally, with black or brown ends; longer bristles dark with 
whitish or buffy ends. Under surface dirty greyish, not sharply 
defined, the bases grey, the ends dirty buffy. Head like 
back ; whiskers very numerous, stiff, black. Ears short, rounded, 
naked ; a small tuft of whitish hairs above their anterior base. 
Limbs uniformly brown throughout ; upper surface of hands and 
feet nearly naked, blackish; palms and soles naked, pads broad 
and fleshy ; fifth hind toe reaching to the end of the first phalanx 


PZ. S 10S Aol WIP YOST, 


Mintern Bros.imp. 


H.Grénvold del.et hth. 


1.ANISOMYS IMITATOR. 
DIN OMNTS, Wie 


1903. | MAMMALS FROM BRITISH NEW GUINEA. 199 


of the fourth; claws of medium strength and curvature. ‘Tail 
about as long as the body without the head, practically naked, the 
two or three minute hairs which project from below the point 
of each scale only from } to 4 the length of a scale; scales very 
large, perhaps the largest among the Muride, only about 5 to 
the centimetre ; arranged diagonally, their points unusually per- 
ceptible to the touch. Proximal third of tail black, terminal 
two-thirds yellow. 

Skull and teeth as described above. Posterior ends of nasals 
just level with those of premaxillary processes. Interorbital 
region narrow, parallel-sided, strongly concave mesially ; its edges 
raised up into vertical ridges which run backwards to the lambdoid 
crests, and have two lateral projections, one at the fronto-parietal 
suture, and the others in the middle of the parietals. Palatal 
foramina shorter than m.’ and m.’? combined, narrow, parallel- 
sided. 

Dimensions of the type, measured on the skin :— 

Head and body 390 mm.; tail 345; hind foot (s. u.) 63; ear 
(wet) 26. 

Skull—ereatest length 74 mm.; basilar length 64:5; greatest 
breadth 41 ; nasals 28 x 12°3; interorbital breadth 9 ; interparietal 
10°5 x 14:5; palate, length from hensilion 37; diastema 23°3 ; 
palatal foramina 8°3 x 4; length of upper molar series 17 ; 
breadth of m.’ 5:7; combined breadth of upper incisors 8°6. 

Hab. Avera, Aroa River, British New Guinea. 

Type. Female. B.M. No. 3.12.1.12. Collected by A. S. Meek. 
One specimen. t 

This huge Rat looked not unlike one of the Indian Bandicoot- 
rats, but had probably no real affinity with them. It was quite 
unlike anything hitherto described from the Papuan region, 
except perhaps Dr. Jentink’s Mus armandvillei of Flores, which 
differed from it, however, in many details. MJallomys rothschildi 
Thos., another large Papuan rat, had molars of quite a different 
pattern. 

Hyomys meeki was no doubt an arboreal animal, as indicated 
by its shortened hind feet, and it was probable that the large 
pointed scales of its tail served a purpose analogous to that of the 
caudal ‘“ climbing-irons” of Anomalurus. 


Anisomys, g. n. (Muride). 


Size large, form less bulky than in Hyomys. Fur coarse. 
Pollex with a broad nail. Tail of medium length, smoothly 
scaled, thinly hairy. Mamme 1—2=6. 

Skull large, stout and heavy. Nasals long, parallel-sided 
behind and but little broadened in front. Supraorbital region 
broad, flat, with heavy edges which are continued backward as 
low, evenly curved ridges to the back of the skull; a vertical 
postorbital projection connected with the main ridge on each side. 
Palatal foramina very small. Palate with raised lateral ridges 
edging it between the foramina and the molars; behind, it extends 


200 MR. OLDFIELD THOMAS ON NEW | Nov. 3, 


some way behind m.’ Bulle but little swollen. Lower jaw 
remarkably high anteriorly, the usual deep hollow in front of the 
molars largely filled up. 

Incisors very peculiar in that while the upper ones are of 
about normal breadth and depth, the lower are quite dispro- 
portionally narrow and deep; in most Rodents the upper and 
lower incisors are of approximately equal transverse dimensions, 
but here the two lower ones combined are of only the same 
breadth in front as a single upper one, while in depth the lower 
teeth exceed the upper by a third, and nearly resemble in shape 
those of Daubentonia. Their roots are carried unusually far up 
at the back of the jaws, so that their basal inflation is at the 
level of the yoke between the coronoid and condylar processes. 
Molars very small, their length less than one-fifth the basilar 
length, and the palate between them about twice their breadth. 
Their lamine directly transverse, with simple raised enamel-edges 
and concave dentine-spaces ; last lamina of m.' and m.* each with 
a small additional internal ring inserted in front of it, the 
homologue of a supplementary cusp; lower molars with four, 
three, and two simple lamine respectively. 

Type, Anisomys inutator. 

This genus seemed to be even less allied to any known one 
than Hyomys, and it could not be said what were its nearest 
relations. Perhaps when young specimens were obtained, so 
that unworn molars could be examined, some light would be shed 
on this problem. In any case the genus might be readily 
distinguished by the peculiar characters of incisors and molars 
above detailed. 


ANISOMYS IMITATOR, sp.n. (Plate XXIII. figs. 1 a1 e.) 


Size and other external characters remarkably like those of 
the large Uromys (U. validus or papuanus), with which it is 
associated. Fur short and coarse; hairs of back about 10-12 mm. 
in length, unmixed with longer piles. General colour above 
coarsely mixed blackish and fawn or buffy, the resulting mixture 
approaching ‘“mummy-brown” of Ridgway. Under surface dull 
buffy white, the hairs slightly darker at their bases. Head 
rather more greyish than back, heavily lined with black. Eyes 
surrounded by indistinct black rings. Ears of medium size, their 
fine hairs blackish. Arms and legs dark grizzled grey, the 
inner sides rather lighter; hands and feet brown, becoming 
whitish at the ends of the digits; claws rather delicate and 
sharply pointed; palms and soles naked, with large smooth pads : 
fifth hind toe reaching nearly to the end of the first phalanx of 
the fourth. Tail fairly long, set with medium-sized scales set in 
alternating rows, and averaging about 9 to the centimetre; very 
thinly hairy, the short whitish hairs becoming rather longer 
towards the tip; its colour dark brown for its basal fourth, the 
remainder yellowish white. 

Skull smooth and rounded; nasals and premaxillary processes 


1903. | MAMMALS FROM BRITISH NEW GUINEA. 201 


of about equal extent; interorbital region broad, scarcely con- 
cave, its edges thickened, but without distinct beading ; palatal 
foramina minute, narrow, of about the same length as m.’; 
anterior palate concave mesially, with swollen ridges laterally ; 
posterior palate extending behind m.* a distance equal to the 
length of m.” Bullee small, smooth. 

Dimensions of the type, measured in skin :— 

Head and body 300 mm.; tail 320; hind foot (s. u.) 60; ear 24. 

Skull—greatest length 68 mm.; basilar length 65; greatest 
breadth 35; nasals 27 x 8°6 ; interorbital breadth 11:3; inter- 
parietal 8°3x14:5; zygomatic plate 9°2; palate, length 33, 
diastema 20; palatal foramina 3°7 x 3:1; length of upper molar 
series 10-2, breadth of m.' 3:1. Upper incisors, combined breadth 
4-7, depth 4:9; lower incisors, combined breadth 2-4, depth 5:3. 

Hab. Avera, Aroa River, British New Guinea. 

Type. Male. B.M. No. 3.12.1.10. Collected by A. 8. Meek. 
Two specimens, male and female. 

So like was this Rat to the large Papuan Uromys, of which 
Mr. Meek also obtained examples, that it was taken for the 
same species until a closer examination showed its many peculiar 
characters. Externally, indeed, it could only be distinguished by 
its rather darker colour and its slightly hairy tail. 

It would be of interest to find out what this Rat fed on to 
account for the peculiarities of its dentition. Perhaps hard- 
shelled nuts, such as cocoanuts, might demand these powerful 
cutting-incisors, while the small and delicate molars would be 
sufficient to deal with the soft contents. 


PERAMELES ORNATA, Sp. N. 


A small species with prominent black markings. 

Essential characters all very much as in P. longicauda Peters 
& Doria *, to whose immediate neighbourhood it is brought by 
all the characters used in the synopses of species, both external 
and cranial, of the ‘Catalogue of Marsupials’. Coloration, 
however, quite unique, for with a pale brown general body- 
colour there is a prominent deep black mesial line running from 
the muzzle down the back to the base of the tail; this line 
commences between the eyes, broadens to about half an inch on 
the crown and nape, narrows on the anterior back, and broadens 
again on the rump. Through each eye there is also a black 
streak, starting at the root of the whiskers, and running to the 
base of the ears; between the median and lateral dark bands 
the head is grizzled whitish. Under surface dull creamy whitish 
throughout, not sharply defined laterally. Rump with an addi- 
tional black line on each side, running downwards parallel to the 
mesial line and passing on to the back of the hind legs. Ears 
of medium length, rounded, dark brown, a small blackish mark 
behind their posterior bases; metatragus short, triangular. Arms 


* Ann. Mus. Genov. xvi. p. 672 (1881). 
+ P. 229 (1888). 


202 PROF. W. B. BENHAM ON AQUATIC | Nov. 3, 


and legs dark brown externally, grizzled whitish internally ; 
upper surface of hands naked, flesh-coloured, of feet pale brown, 
lightening terminally to whitish; soles quite naked, finely 
granulated. Tail long, finely haired, yellow for its terminal two 
inches and along its under surface; the remainder dark brown. 

Skull and teeth agreeing word for word with the description 
of those of P. longicauda given in the ‘ Catalogue of Marsupials.’ 

Dimensions of the type, measured in skin :— 

Head and body 300 mm.; tail 177; hind foot (s. u.) 59; ear 
(wet) 28. 

Skull—basal length 57:2 mm. ; greatest breadth 23°3; nasals 
27 x 5:2; interorbital breadth 12°8; palate, length 37; combined 
length of three anterior molariform teeth 10. 

Hab. Avera, Aroa River, British New Guinea. 

Type. Adult male. B.M. No, 3.12.1.23. Collected by A.S. 
Meek. One specimen. 

In the conspicuous striping of its dorsal surface this handsome 
species differed from all known Bandicoots, though it was 
possible that when dried skins of P. longicauda were examined, 
some indication of a similar pattern of coloration would be found 
to exist in that animal. 


EXPLANATION OF PLATE XXIII. 


Fig. la. Anisomys imitator (p. 200). Lower view of skull, natural size. 
1b. 


6 x5 $ Left upper molar series, 4. 
le. » A sy Upper view of skull. 
1d. ¥ x Pe Front view of incisors. 
: e Right lower molar series, +. 
‘ 


. 33 ” oy) 

2a&26. Hyomys meeki (p. 198). Lower and upper views of skull, natural 
size, 

2c. Hyomys meeki (p. 198). Right lower molar series, 2. 


The following papers were read :— 


1. On some new Species of Aquatic Oligocheta from New 
Zealand. By W. B. Beyuam, D.Sec., M.A., F.Z.S., 
Hon. M.RS. Tasm.; Professor of Biology in the Univer- 
sity of Otago. 

[Received July 24, 1903. ] 
(Plates XXTV.—XXVL.* and Text-figure 23.) 


In the course of a biological survey of the New-Zealand lakes 
undertaken, during the year 1902, by Messrs. K. Lucas and 
Hodgson, of Cambridge, a considerable number of specimens of 
Oligocheta were obtained which were placed in my hands for 
identification. My best thanks are due to Mr. Lucas for his 


* For explanation of the Plates, see pp. 231, 232. 


A SulS OS, voll ie OaIVe 


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AQUATIC OLIGOCHASTES FROM NEW ZEALAND 


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MP Parker lth. Parker & West imp. 
AQUATIC OLIGOCHA TES FROM NEW ZEALAND. 


IPL ZS. IOS, soleil IPL ZO, 


MP Parker lth. , Parker & West imp. 
AQUATIC OLIGOCHA TES FROM NEW ZEALAND. 


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1903. | OLIGOCHETA FROM NEW ZEALAND. 203 


generosity in thus affording me an opportunity, for many years 
to come likely to remain unique, of examining the deep-water 
Oligocheeta of our lakes. 

In view of the extremely interesting character of the terres- 
trial Oligochetes of New Zealand, both from a morphological and 
a z00-geographical aspect, it seemed probable that our deep lakes 
of the South Island would contain equally interesting species ; 
but the result of my investigation, though not wholly without 
interest, is rather disappointing. For, whereas our terrestrial 
fauna includes several endemic genera, like Maoridrilus, Plagio- 
cheta, Neodrilus, Deinodrilus, and Octochetus, our lacustrine genera 
are, with one exception, of an exotic character; and even the “single 
new genus that I have ventured to create, viz. T’aupodrilus, is 
very near akin to a HKuropean worm, Branchiura coccinea of 
Vejdovsky. 

But amongst the new species that are here enumerated, some 
are of considerable interest. Of the genus Phreodrilus Beddard, 
originally founded for a New-Zealand worm, but now extended to 
include certain South-American aquatic species previously placed 
by Beddard in the genus Hesperodrilus, I find two new represen- 
tatives: one of which is ‘“‘ Hesperodrilid” in the nature of the 
male efferent apparatus, and therein agreeing with the Kerguelen- 
Island form and South-American species, rather than with the 
original representative from New Zealand. This genus thus has a 
distribution similar te that of our earthworms belonging to the 
genus Notiodrilus. 

Two other species deserve mention here: Diporocheta aquatica, 
sp. n., and Plutellus lacustris, sp. n., for both of these genera are 
characteristically Australian. 

The genus Diporocheta was founded originally by Beddard for 
D. intermedia from Lake Brunner in this colony ; but it has now 
been extended so as to include a number of Australian species, and 
hitherto Beddard’s species has been the only representative on this 
island. It is worth noting that both our species occur in water, 
whereas the majority of the species are terrestrial. 

As to. Plutellus lacustris, it differs from the rest of the species 
in certain characters, viz., the loss of gizzard, and the absence of 
nephridia in the pregenital segments ; and at first I was inclined 
to form a new genus for it, but these features, in which it 
approaches Pontodrilus (an inhabitant of the sea-shore in various 
parts of the world), appear correlated with an aquatic habit. 

These two genera, Diporocheta and Plutellus, belong to the 
subfamily Megascolecinz, and they are the only representatives 
of this subfamily in New Zealand. It is true that Schmarda 
attributed MNotoscolex (Hypogeon) orthostichon to New Zealand, 
but this appears to have been due to an error—to a lapsus calami. 
He gives as the locality “ Mt. Wellington ”: now, there is no such 
mountain in the district visited by Schmarda in New Zealand, 
but he did visit Mt. Wellington at Hobart, Tasmania; and there 
is no doubt in my mind that he obtained ‘“ HZ. orthostichon” in 


204 PROF. W. B. BENHAM ON AQUATIC (Nov. 3, 


Tasmania, and not in New Zealand, for the genus Wotoscolex is 
common in the former locality, and has never been recognised 
amongst the numerous collections of New-Zealand worms examined 
by Beddard or by myself. 

Again, Baird described “ Megascolex antarcticus” from New 
Zealand—this in all probability should be placed in my genus 
Plagiocheta. With these two doubtful exceptions, then, the only 
representatives in the New-Zealand area of the extensive sub- 
family Megascolecine are :-— 


Diporocheta intermedia Beddard, 
D. chathamensis Benham, 
D. aquatica, sp. n., 

and Plutellus lacustris, sp. n. ; 


and the occurrence of these four species is rather difficult of 
explanation, for the subfamily is characteristic of Australia, and, if 
we except the Malayan Pheretima, is almost confined to Australia 
and Ceylon. 

The remaining species belong to common and probably wide- 
spread aquatic genera, viz., Limnodrilus, Tubifex, Enchytreus, 
Acheta, and Haplotaais. 

Of the first two genera, representatives have already been 
referred to by Beddard as occurring in this colony, but no 
description of the species has been published. Of the Enchytrzeide 
Beddard * has recorded Henlea ventriculosa and Fridericia galba 
(=F. antarctica Bedd.), both of which are terrestrial species living 
in swampy places; while of Haplotaais we already know H. smithi 
Beddard from this country. 

A special interest appears to be connected with some of the 
new species described in this paper, in that they were obtained 
from very considerable depths ; at any rate the soundings given by 
Mr. Lucas for the hauls enumerated below imply that the worms 
were obtained from the bottom, and, moreover, the intestine is 
loaded with mud- and diatom-valves ; although I do not know what 
means, if any, were employed to prevent the entrance of organisms 
as the dredge was pulled up through the water. The absence 
from this collection of the families Naidide, Aolosomatide, and 
Lumbriculide seems to indicate that some such means were em- 
ployed, as these families occur in shallower waters amongst water- 
weeds; and representatives of Naids and Lumbriculids do occur 
in New Zealand, as I have collected them in the neighbourhood 
of Dunedin, but I have not yet worked them out. 


List of Worms obtained by Mr. K. Lucas, and described 
in the present paper. 
Fam. PHREODRILIDZ. 


1. Phreodrilus lacustris, sp. n. 
2. P. mauianus, sp. n. 


* Beddard, Proc. R. Phys. Soc. Edinb. xii. p, 41, 


1903.] 


OLIGOCHATA FROM NEW ZEALAND, 


Fam. TuBIFICID. 


3. Taupodrilus simplex, gen. et sp. n. 
4, Limnodrilus vejdovskyanus, sp. n. 
5. L. lucasi, sp. n. 
6. ZL. sp. ine.* 

7. Tubifex sp. inc.* 


Fam. ENCHYTR#ID&. 


8. Enchytreus simulans, sp. n. 
9. Acheta maorica, sp. n. 


Fam. HAPLorTaxID#. 
10. Haplotawis heterogyne, sp. n. 


Fam. MEGASCOLECID. 
Subfam. MrcAscoLecina. 
11. Diporocheta aquatica, sp. n. 


12. Plutellus lacustris, sp. n. 


The total number of individuals examined is about 150, and 
these were obtained at 27 stations in six lakes; of which two are 
in the South Island, and four in the North Island. 

The following table shows the distribution of the above species 
in these lakes, the depth at which they were obtained, and the 
number of stations at which each species occurred. 


Table showing Distribution of the Worms. 


Name. 


x mauianus ...... 
Limnodrilus vejdovskyanus . 
a ULES 200 080080008 

3 sp. ine. (A) ... 

33 33 (B) O80) 

»  (C) ... 


Tubifex sp. ine. .. 


bE) 9 


32 33 
Taupodrilus simplex ......... 


bby 2 


39 39 
Enchytreus simulans......... 
Acheta maorica ............... 
Haplotaxis heterogyne 
Diporocheta aquatica 
Plutellus lacustris ............ 


No. of | No. of 
Specimens.|Stations. 


4 

16 ; 
1 1 
30t 1 
1 

many ‘ 1 
1 1 
1 1 
1 1 
1 1 
2 1 
1 1 
6 2 
rh 2 
1 1 
7 1 
1 1 
2 1 
2 2 
17t 7 


Lake. 


Wakatipu. 
Manapouri. 
Taupo. 
Waikare. 
Taupo. 
Rotoiti. 
Wakatipu. 
Manapouri. 


Waikaremoana. 


Taupo. 
Rotoiti. 


Waikaremoana. 


Manapouri. 
Taupo. 
Waikare. 
Taupo. 
Manapouri. 
Wakatipu. 
Manapouri. 
Wakatipu. 


_ 160-500 


Depth in 
feet. 


300-1000 
150-500 
150-500 
9 

300-450 
100-228 
1000 
160-500 
200-750 


100-228 
200-750 
10-1000 
150-500 
9 
P 
350 
550 
350-550 
300-1200 


| 
| 


* The specimens being immature are not named. 
+ Together with six immature specimens probably of this species from Waikare- 


moana, 800-840 ft. 


{ Also some cocoons, which appear to belong to this worm. 


206 PROF, W. B. BENHAM ON AQUATIC [ Nov. 3) 


Analysis of the Oligochete Fauna of each of the Lakes. 


Lake Wakatipu, South Island. 

Greatest depth, 1242 feet. 

Number of stations at which worms were obtained, 10. 

Total number of specimens, 37. 

The commonest worm appears to be Plutellus lacustris, which 
was obtained from 7 stations at different parts of the lake, so that 
it is evidently widespread throughout; but it appears to be 
limited to this lake, as none were obtained elsewhere. 

Phreodrilus lacustris also occurred at three stations, and appears 
to be somewhat less widely distributed, though living at the same 
depth. Haplotaxis heterogyne was obtained only at one station, 
and a single undetermined immature specimen of Limnodrilus. 


Lake Manapouri, South Island. 
Greatest depth, 1458 feet. 
Number of stations, 6. 

Number of specimens, 16. 

This, the deepest lake, so far as this survey is concerned, 
contained a greater variety of Oligochetes than any of the other 
lakes, for it yielded 6 genera. 

Phreodrilus lacustris and Taupodrilus simplex account for 
twelve out of the sixteen specimens; the former was obtained once 
only, the latter at two stations. At two other stations Dzporo- 
cheta aquatica and at another Acheta maorica were captured, 
while an immature Limnodrilus completed the list. 


Lake Taupo, North Island. 

Greatest depth, 534 feet. 

Number of stations, 4. 

Total number of specimens, 32. 

Taupodrilus simplea and Limnodrilus lucasit account for more 
than two-thirds of the total, the former occurring in two hauls, the 
latter in one only. With the former, Phreodrilus mauianus was 
obtained, while at another spot Lnchytreus simulans occurred, and 
an undetermined species of 7'ubifex completes the list. 


Lake Rotoiti, North Island. 

Greatest depth, 228 feet. 

Number of stations, 2. 

Number of specimens, many. 

This lake is characterised by Limnodrilus luweasi, of which 3 or 
4 dozen were obtained at one spot; while at the other station 
immature specimens of Z'wbifex occurred. 


Lake Waikaremoana, North Island. 
Greatest depth, 846 feet. 
Number of stations, 3. 
Number of specimens, 16. 
Unfortunately these were very poorly preseryed, much broken 


1903. | OLIGOCH &TA FROM NEW ZEALAND. 207 


and immature specimens of Z'uwbifew sp. and of Limnodrilus, 
amongst which some appear to be L. vejdovskyanus. 


Lake Waikare, North Island. 

Greatest depth 9 feet. Only one haul was taken, which yielded 
numerous specimens of Limnodrilus vejdovskyanus and one 
individual of Tawpodrilus simplex. 


Tt will thus be seen that the lakes in the North and South Island 
respectively differ considerably in their Oligochetes, so far as 
research has yet gone. 

The two species of Phreodrilus are very distinct, one confined to 
the north, and the other evidently common in the southern lakes. 

Of Taupodrilus, the northern and southern representatives may 
be distinct, though I have included them in the same species, as 
the southern material was not in sufficiently good condition to 
permit me fully to investigate the details of the reproductive 
organ. 

Unfortunately the southern representatives of Limnodrilus are 
immature, but they appear to differ in the form of their chete 
from each of the northern species, L. vejdovskyanus from Waikare 
and L. lucasi from Taupo. 


PHREODRILUS LACUSTRIS, Sp. n.* 


A very narrow worm in which the ventral chet are in couples 
of two kinds: each couple consists of (a) a simple hook-like bristle 
and (6) a hook with a very minute tooth in the back. These 
cheetz measure 0°06 mm. in length, 

The ventral cheetze are absent on segments xii., xiii., though on 
the latter segment they are replaced by special copulatory chete. 
Dorsal chetz solitary, capilliform, beginning on segment 111. 

The clitellum covers the hinder part of segment xii. and the 
whole of xiii. 

The male pores are in line with the ventral chet at the hinder 
margin of segment Xil. 

The oviducal pores are in the same line, at the boundary of seg- 
ments xii./xill. 

The spermathecal pores are in line with the ventral chete at 
the anterior margin of segment xiii. 

Copulatory chete, a pair of couples; each pair in a special ovoid 
glandular follicle just behind the spermathecal pore. 

The preclitellar nephridium appears to extend through segments 
vil. to x. 

No sperm-sacs ; but loose developing spermatozoa in segments 
viii. to xii. 

Spermiducal gland of large size, fillmg segment xii., slightly 
convoluted, receiving the sperm-duct at its short, narrow neck, 
where it enters a large protrusible penis, enclosed in a muscular 


* A detailed account of this species has been sent to the Editor of the Quart. 
Journ. Micros. Sci. 


208 PROF. W. B. BENHAM ON AQUATIC [ Nov. 3, 
penial sac. There is no atrial sac, and the atrium itself is not 
longer than the penis. In its general arrangement it resembles 
the apparatus in Hesperodrilus albus of Beddard. a 

The spermathece (one pair) extend through segments xiil. to xv. ; 
the pore leads into a very slightly dilated muscular duct which 
soon becomes narrow and is much arched dorsally; on passing 
through the septum xiii./xiv. the duct, still narrow, becomes glan- 
dular, and then opens into the ampulla, which occupies the hinder 
part of segment xiv. and the whole of segment xv. 

Dimensions. 20 mm.x4+mm. 75 segments. 

Localities. Lakes Manapouri and Wakatipu, South Island of 
New Zealand. 

Remarks.—This worm agrees closely with those South- American 
worms for which the genus Hesperodrilus was founded by Mr. 
Beddard* ; but Dr. Michaelsen t has shown good reason for merging 
this genus with Phreodrilus owing to the discovery, in Kerguelen, 
of a worm which in certain respects presents the characters of 
both the genera. 

The discovery in New Zealand of two new species, this and the 
following, belonging to the section of the genus hitherto found in 
the Falkland Islands, South America, and Kerguelen, is a most 
interesting additional fact in our knowledge of the geographical 
distribution of the Southern Oligocheeta. 


PHREODRILUS MAUIANUS 4, sp. n. 


This new species is founded on a single immature individual, 
which, however, differs from any hitherto described. 

The ventral chet are, as usual, of two kinds, one of each in 
each bundle, viz.—(a@) a simple hook-shaped, single-pointed bristle, 
and (6) a similar bristle with a very distinct tooth on its upper, 
convex, surface. These chete measure 0:15 mm., and are thus 
much longer than those of the species just described. 

The dorsal chet are capilliform, solitary, and commence in 
segment 1il. 

The esophagus is narrow up to segment vi., where it dilates, 
and is then constricted by the following septa. In the middle of 
segment ix. the gut presents a slight constriction, and the epithe- 
lium suddenly changes in its character—the cesophagus passing 
suddenly into the intestine. The dorsal vessel lies free of the 
gut in segment x. and forwards; a supra-intestinal vessel is 
recognisable in segments vil. to xv. An enlarged commissural 
vessel exists In segment x., and a contorted, swollen, heart-like 
organ in the following segment (xi.), which appears to be connected 


* Beddard, Ann. Mag. Nat. Hist. (ser. 6) xiii. p. 206; & Ergeb. Hamb. Magalhaen. 
Sammelreise, 1896—“ Naid. Tubificid. u. Terricolen,” p. 9. 

+ Michaelsen, Oligoch. d. deutsch. Tiefsee Exped. 1902. 

{ The specific name “ mauianus,” in which the syllable aw has the sound of ow in 
cow, refers to the mythical Hercules of the Pacific, known to the Maoris as Maui. 
The North Island of New Zealand owes its origin to Maui, who, while fishing from a 
boat at sea, hauled up the land at the end of his fishing-line. Hence the original 
Maori name for this island was “Te ika a Maui’”’—the fish of Maui. 


1903.] OLIGOCHETA FROM NEW ZEALAND. 209 


with the supra-intestinal vessel. As I have studied the specimen 
only as an entire object, I cannot say anything as to the structure 
of this organ, but it seems to resemble in its external features 
the “blood-gland” which Beddard describes as occurring in seg- 
ments xil., xiii. in P. subterraneus. 

Dimensions. 15 mm. X¥2? mm. It is much stouter than 
P. lacustris, and consists of 70 segments. 

Locality. Lake Taupo, North Island, New Zealand. 


TAUPODRILUS, gen. nov. 
TAUPODRILUS SIMPLEX, sp.n. (Plate XXIV. figs. 1-9.) 


Of this worm I found about a dozen individuals, most of which 
are only the anterior ends, and a few are entire. They are all much 
coiled, rendering it a matter of some difficulty to make sagittal 
sections. 

The following account is based on the study of three individuals, 
more or less sexually mature, stained and mounted in Canada 
‘balsam, and a series of longitudinal sections through the anterior 
segments, and transverse sections of the middle region, in addition, 
of course, to the usual glycerine and potash preparations. 

The prostomium is conical and relatively long. 

Chetce,—The dorsal bundles consist of capilliform (at any rate 
in the anterior segments), accompanied by bifurcate cheetee 
(Pl. XXIV. fig. 3), with a few delicate intermediate teeth. 
Such chetze may be termed “multidentate” or “ ctenate,” and 
occur also in T'ubifex rivulorum. 

The ventral bundle consists almost wholly of crochet-shaped 
cheetee, with intermediate teeth. In the dorsal bundle there are 
one or two capilliform and usually three or four ctenate chet 
(in one individual the former were only present on segments V., Vi., 
vii.; in another they did not begin till the fourth segment, and in 
one specimen I was unable to detect them), and typically they 
occur only on the first 12 to 15 chetigerous segments. 

In the “ctenate” set the chief teeth or prongs are scarcely 
curved (Pl. XXIV. figs. 4, 4a), and the number of intermediate 
teeth varies from two or three to several; usually they all separate, 
but in a few instances a delicate striated membrane unites the chief 
teeth as in “ Psammoryctes.” The dorsal chetze of segment 11. 
are smaller than the rest. 

In the ventral bundle there are from 4 to 6 chet in the 
anterior segments, diminishing to 3 or 4in the mid-body. The 
chief teeth are curved as in normal forked bristles or crochets ; 
the upper tooth is less stout than the lower, and over the greater 
part of the body the two teeth are of the same length, but in the 
anterior segments the upper tooth is slightly the longer (Pl. XXTYV. 
figs. 5, 6). Generally, all the crochets in a bundle have inter- 
mediate teeth, which lie in a different plane from the main teeth ; 
but in the anterior segments a minority in each bundle are normal 
crochets. 


Proc. Zoou. Soc.—1903, Vou. Il. No. XIV. 14 


210 PROF. W. B. BENHAM ON AQUATIC [Nov. 3, 


The clitellum envelops the body, and extends over three seg- 
ments, from the middle of x. to the middle of xii. (Pl. XXIV. 
fig. 1). 

The male pores are near the posterior margin of segment xi. and 
lie outside the line of ventral chet, which in this segment are 
modified to form a bundle of copulatory bristles. 

The oviducal pore is at the anterior margin of segment XiL., 
practically intersegmental, and further laterad than the male 

ores. 

: The spermathecal pores are close to the anterior boundary of 
segment X. 


Internal Anatomy. 


The pharynx, in segment ii., is provided with the usual dorsal, 
ciliated pouch, whence radiating muscles pass to the body-wall. 
The csophagus is quite narrow; the chloragogen cells begm in 
segment vi.; the gut suddenly enlarges in the tenth segment, 
and food occurs there consisting, amongst other things, of diatom- 
valves; this intestinal region is thereafter wide, but septally 
constricted. 

The Vascular System.—The dorsal vessel is distinct throughout 
the body ; it is connected with the ventral vessel by undulating 
commissurals in each of the segments il. to x., while the dorsal 
vessel bifureates in the first segment, and each branch passes 
forwards into the prostomium, bends downwards and backwards to 
unite to form the ventral vessel in segment 11. 

Of the commissural vessels, those in segments vil. and vill. are 
slightly larger and less extended than the rest; they, however, 
are not dilated to form “hearts” such as occur in Limnodrilus 
and other Tubificids (Pl. XXIV. fig. 2). A pair of vessels passes 
backwards on the sperm-sacs to segment xvii., but as to the exact 
origin of them, or whether there is a second pair supplying the 
ovisacs (as in Lranchiwra coccinea), I am unable to ascertain. 
Nor can I state whether any of the commissurals connect with 
the supra-intestinal vessel which is present in at least part of this 
region. I have not been able to detect any integumental vessels, 
either in sections, or in the glycerine and potash preparations. I 
do not think they exist. 

The Reproductive System (Pl. XXIV. fig. 8).—The testes, ovary, 
and oviducts are in the positions usual in this family. 

There are two pairs of sperm-sacs, one in segment ix., the other 
extending through segments xi. to xvii. and constricted by each 
septum through which it passes, while segment x. is filled with loose 
masses of developing spermatozoa, not enclosed in a special sae. 
Large ova, contained in an ovisac, occupy segments xvi., XVii., 
XVill., in one specimen; but in another, in which the sperm-sacs 
are not so extended, the ovisac is not so far back. 

_ The condition of the male efferent apparatus is the most 
interesting feature of this new genus. 

The sperm-funnel, on the hinder wall of segment xi., is flat and 


1903.] OLIGOCHETA FROM NEW ZEALAND. 211 


moderately extensive. The sperm-duct is comparatively thick; it 
passes below the ovary nearly directly backwards to the hinder 
septum of the twelfth segment; there it makes a single loop round 
the neck of the atrium, and then runs up it to enter its distal 
extremity. 

This atrium (or spermiducal gland—for it is both) is a large 
pyzriform or sausage-shaped sac, with a capacious cavity; its 
broader end is directed upwards, its narrower end is suddenly 
constricted to form an extremely short and narrow duct, just long 
enough to pass through the body-wall. In its microscopic structure 
the atrium presents what is probably to be regarded as an archaic 
character; at any rate, it is simpler than that of any Tubificid 
hitherto described. The epithelium consists of a single layer of 
tall, glandular, and much vacuolated cells, outside which is a thin 
peritoneal membrane with flat nuclei (Pl. XXIV. fig. 9). There is 
apparently no muscular coat, except for a few circular fibres near 
its lower end where it is about to penetrate the body-wall. 

In the figure (fig. 9, which is drawn under a camera, and 
represents as accurately as may be the arrangement of the 
nuclei and vacuolation of the cytoplasm) two kinds of nuclei are 
to be seen in the epithelium: most of them are circular and 
situated towards the outer surface of this epithelium; others, 
however, are oval, elongated in a direction vertical to the surface, 
these lie nearer the inner ends of the cells. These latter suggest 
a columnar epithelium distinct from the gland-cells; but I have 
been unable to detect any cell-boundaries corresponding to these 
oval nuclei, which, moreover, are not arranged in close array to 
suggest an epithelium. It may very well be, however, that some 
of the epithelial cells are short and have not become glandular. 

The sperm-duct opens into this atrium at its apex, 7.e. at the 
point furthest from the external opening. In this feature it bears 
a nearer resemblance to Branchiura coccinea than to B. sowerbyt ; 
but even in the former species the duct enters the side, rather 
than the apex, of the nearly spherical atrium. 

The copulatory chete form a bundle (varying from only 3 to 
as many as 7 or 8) of simple bristles, lying in a pit or depression 
near the male pore. Viewed from the side in a mounted speci- 
men, the two organs appear coincident; but sections show the 
true relation. Hach of these chet, one of which is figured 
(Pl. XXIV. fig. 7), is a nearly straight rod, thicker than the 
other cheetz, terminating in a point, which is not terminal, but 
directed to one side. ; 

The spermatheca is globular, with a well-marked duct about 
half as long as the diameter of the ampulla. The neck of the 
latter is provided with a ring of elongated glandular cells, forming 
a prominent, valve-like structure (in longitudinal section). The 
duct is lined by cubical gland-cells, and is provided with a 
muscular coat. There are no spermatophores. 

Dimensions. Length of an entire individual 15 mm.; diameter 


2mm. Number of segments 70. 


OD) PROF. W. B. BENHAM ON AQUATIC [ Nov. 3, 


Locality. Lake Taupo, North Island, New Zealand. Also one 
individual from Lake Waikare, and some immature and imperfect 
individuals from Lake Manapouri, in the South Island, belong to 
this genus, but Iam unable to say whether they are identical 
with the above. 

Remarks.—This new genus appears to be most nearly allied to the 
genus Branchiura Bedd. * as extended by Michaelsen to include 
“ Tlyodrilus coccineus” Vejdovsky. But it will not fit into this 
genus, chiefly on account of the structure of the atrium. 

In B. coccinea the spherical atrium is lined by a layer of 
ciliated cells, outside which is a layer of vesicular cells, corre- 
sponding to the compact “ prostate” (Cement-driise) of Zubifea, &e. 
Apart from this fact, the general arrangement of the male 
efferent apparatus is like that of the new genus: the thick sperm- 
duct, and the short atrial-duct, with muscles at its base, forming 
a slightly protuberant papilla. 

With this species, too, B. coccinea agrees in the presence of 
copulatory cheetee on segment xi., but in the form of these there is 
considerable difference : for in B. coccinea, according to fig. 13 d, d, 


on pl. iv. of Stole’s memoir f, they are of two kinds, both sigmoid 
and crochets; while in the new species they are of neither of 
these two forms, but simple pointed, straight rods, with a nodal 
swelling. There are other points of difference, as in the undulating, 
instead of simple, form of the commissural vessels, in the position 
of the spermathecal pore, and the presence, in Taupodrilus, of a 
distinct muscular duct. 

From B. sowerbyi there are more numerous differences, apart 
from the presence in this species of the gills, 

So far as the atrium is concerned, the new genus exhibits 
points of resemblance to Clitellio arenariust, in which a thick 
and relatively short sperm-duct enters the apex of an elongated, 
dilated, glandular “atrium,” which appears rather as a swelling in 
the course of the duct. Of its minute structure we have no 
account; but in the total absence of a ‘‘ prostate” there is a close 
similarity to the new genus. But in all other points there seems 
to be no close affinity between the two, for there are no capilli- 
form bristles in the dorsal bundles; the chetz being “only 
forked ”—both Beddard and Michaelsen give this as a generic 
character. Further, two pairs of dilated hearts exist In segments 
<HAltlgg, ID 

It is difficult to separate generic from specific characters, but 
probably we may regard as in the former category : 


(a) the presence of fan-shaped cheetze dorsally ; 
(5) presence of copulatory cheetee on segment Xi. ; 
(c) character of the atrium. 


The structure of the atrium of B. coccinea has hitherto been 
regarded as the simplest amongst the Tubificidee (Beddard, Monog. 


* Beddard, Quart. Journ. Micr. Sci. 1892, xxxiii. p. 326. 


_+ Stole, Abhandl. Béhm. Ges. 1888. 
+ Of, Beddard, P.Z.S. 1888, p. 491. 


1903.] OLIGOCHATA FROM NEW ZEALAND. 213 


p- 233), and indeed agrees with that of certain of the Naidide, 
such as Stylaria lacustris *, though I should place Clitelio at a 
lower grade even than B. coccinea, for in it apparently, and at 
any rate in the new genus, the atrium is still simpler, and 
approaches that found in Vais elinguwis, or in Dero, or in Cheto- 
gaster. In the first named, Stole + shows the atrium as a 
dilatation of the sperm-duct, from which it is not distinctly 
marked off; its lining appears to consist of low, cubical cells 
(? glandular), outside which is a layer of flat, peritoneal cells. 

In the atrium of Chetogaster, as described and figured by 
Vejdovsky t, the glandular. lining and flat epithelial cells are 
distinctly shown; while Beddard § thus describes the organ in 
Dero:—“The atria are lined by a columnar epithelium, but I 
could observe no layer of cells covering this organ externally 
and forming the structure which has sometimes been termed 
prostate.” 

In these forms, and in Taupodrilus, the atrium, indeed, is in 
its most simple form, as a distinct and definite organ—definitely 
marked off from the sperm-duct, but still, evidently, a dilatation 
of its distal extremity. 


LIMNODRILUS VEJDOVSKYANUS, sp.n. (Plate XXV. figs. 10-17.) 


From Lake Waikare I received about two dozen stout, greyish 
worms, in which the cuticle is much wrinkled, the body-wall thick, 
and the segments distinctly bi- and tri-annulate. 

The prostomium is conical, with a rather acute point; it is 
relatively long, and exceeds the length of the first segment. 

The chete are in the usual four bundles: 4 and 5 per bundle; 
even 6 in the bundles of segments ii. to vi.; but throughout the 
greater part of the body there are 4 ventrally and 3 dorsally. 

The dorsal cheetee are slightly smaller in all dimensions than 
the ventral, but in form they are similar (Pl. X XV. figs. 10, 11). 
The upper tooth is very distinctly longer and less robust, and 
has a sharper point than the lower tooth. 

The ventral cheetz are absent on segment x1. 

The clitellum covers segments xi. and xii., and does not 
encroach at all on segment x. The genital pores have the usual 
position. 

Internal Anatomy. 


This was studied both in entire specimens—stained in alum- 
cochineal, and mounted in Canada balsam—and in serial transverse 
and longitudinal sections. 

The alimentary canal is without a distinct muscular pharnyx ; 
the buecal region passes through segment i. into the commence- 
ment of segment ii.; its wall is here folded and provided with a 
few retractor muscles. Following this is the beginning of the 


* Vej dovsky, Syst. u. Morph. d. Oligoch. 1884, pl. iv. fig. 10. 
+ Stole, SB. Bohm. Ges. 1887, p. 228, fig. 7. 

~ Vejdovsky, Syst. u. Morph. d. Oligoch. 1884, pl. v. fig. 7. 
§ Beddard, P. Z.S. 1889, p. 444. 


214 PROF. W. B. BENHAM ON AQUATIC [Nov. 3, 


esophagus, in the hinder part of segment i1.; it thickens out 
considerably, right and left, in segment u1., forming a somewhat 
quadrangular organ, looking—in an entire specimen—lke a 
pharynx. The walls of this organ are comparatively thin, some- 
what folded laterally, and lined by tall, columnar, ciliated epi- 
thelium*, which is surrounded by a very feebly developed 
muscular layer; the floor is raised into a ridge in the median 
line. The structure recalls that described by Nasse for 7’ubifex tT. 
In segment iv. the esophagus, retaining the same structure, 
diminishes in size vertically, but is still rather wide laterally. 

The chloragogen cells—which are quite pale—begin in seg- 
ment Vi. 

In the eighth segment (PI. XXV. fig. 12) the ceesophagus passes 
into the intestine. The former region reaches into the first half of 
the segment, then suddenly, at about the middle of the segment, 
dilates to about three times its former size. The epithelium, 
anteriorly high, is there quite low, and the cells contain blackish 
granules. The chloragogen granules have been dissolved out in 
the sections, but these dark intestinal granules remain. 

The intestine is greatly dilated in each segment, and constricted 
by the successive septa. 

The Vascular System.—In the cesophageal region the dorsal 
vessel is free from the wall of the gut, as is the case in Oligocheta 
generally. In the eighth segment it presents an arrangement 
which appears to be, in its details, unique (Pl. XXV. fig. 12). 
In the anterior (cesophageal) moiety of segment viii. the vessel 
becomes very muscular, and at the junction of cesophagus with 
intestine the dorsal vessel bifurcates; each branch, one on either 
side, passes downwards and backwards, along the line of union of 
cesophagus with intestine, to join the ventral vessel. This con- 
necting vessel adheres closely to the wall of the gut, lying in a 
furrow therein, and has a very thick muscular wall (Pl. XXV. 
fig. 13); it is, however, not specially dilated in the way that the 
“lateral heart” of Zubifex is, and, further, it is connected with 
the dorsal vessel, and not with the supra-intestinal as is the case 


in other Tubificids [vide Stole; Beddard, Monog. p. 240]. 

Behind the eighth segment, the dorsal vessel comes to lie below 
the peritoneal cells, above the gut-wall, in the usual way. 
Anteriorly to the eighth segment there is a ‘“supra-intestinal ” 
vessel adhering to the cesophageal roof; it passes forwards through 
several segments, but I did not ascertain how far it reaches. 

In the intestinal region the vascular network on the gut-wall 
is connected with the dorsal vessel; while in the cesophageal 
region it is, as usual, connected with the supra-intestinal vessel. 

Commissural vessels, undulating along the inner surface of the 
body-wall for the whole length of the segment, occur in segments 


* This arrangement reminds one of the condition found in Molosoma, and 
described in detail by Vejdovsky in his ‘Syst. u. Morph. d. Oligoch.’ p. 101, but in 
that low form there is no preceding buccal region. 

+ Nasse, ‘Beit. z. Anat. d. Tubificiden,’ 1889. 


1903. ] OLIGOCHETA FROM NEW ZEALAND. 215 


ii. to vil. They are all narrow, of about the same diameter, and 
put the dorsal and ventral vessels into communication at the 
hinder part of each of these segments. It appears to me that in 
this region the dorsal and supra-intestinal vessels are connected at 
the septum, and that it is at this point of fusion that the commis- 
sural vessels start. This union does not occur in Lophocheta or 
Bothrioneuron, in which the vascular system has been so well 


figured by Stole. 

I cannot detect any integumental network in spite of repeated 
examination of various individuals, in different media. 

The nephridia are not enclosed in vesicular cells. 

The brain is very slightly notched posteriorly. 

The Reproductive System.—The spermiducal funnel is large and 
flat, and has the usual position on the hinder wall of segment x. 
The sperm-duct takes a much undulated course, pushing the 
septum xi./xii. backwards. The duct gradually widens in this 
region to form the atrium, which receives a quite small lobulated 
“‘spermiducal gland” (prostate, Cement-driise); it then passes 
forwards to become the penis. 

The muscles of the penis are spirally disposed, as in some other 
species. 

The chitinous penial tube is of considerable length, about 10 
times as long as the basal diameter, and when the worm is com- 
pressed it extends through segments x1. and xii., reaching nearly 
to the posterior end of the latter. The free extremity is suddenly 
expanded to form what appears in side view to be a thin flat 

late, the margin of which may be even slightly reflexed 
(Pl. XXYV, figs. 14, 15). 

The penis is almost straight, its slight curvature being possibly 
due to compression, as it is not constantly identical in amount ; 
but there is no sudden bend as in the species next to be described. 
T could detect no “valvular apparatus” at the aperture of the 

enis. 
: The spermatheca appears to be variable in form ; in one entire 
individual it was spirally coiled, so that the ampulla—an elon- 
gated ovoid—formed the larger coil, and the narrower duct an 
inner, smaller coil; while in another case the ampulla was more 
globular (Pl. XXYV. figs. 16, 17). The duct is about half the 
length of the ampulla. 

I observed no spermatophores. 

Dimensions. Length 20-25 mm.; diameter ? to 1 mm. The 
number of segments in one specimen was 66 + a regenerated 
tail of 33 very short ones; and in another 75 + 40 very small 
segments. 

Localities. Lakes Waikare and Waikaremoana, North Island, 
New Zealand. 

Remarks.—This species, so far at least as the general form of 
the penial tube is concerned, is most nearly allied to L. clapare- 
dianus Ratzel; but the latter species is of much greater size, 
viz., from 50-80 mm., and its chetz are from 5 to 10 per bundle ; 


216 PROF. W. B. BENHAM ON AQUATIC [ Nov. 5c 


but how far the number is liable to variation is a subject that has 
received but little attention. In my specimens they are pretty 
constantly 4 ventrally and 3 dorsally, except the very anterior 
ones. At any rate, Vejdovsky’s figure (pl. vill. fig. 22) gives a 
form very different from that of the new species; for the upper 
tooth is not pointed, but blunt, and much larger than the lower 
tooth, the proportions being very different from those in my 
species. 

The presence ofa single pair of hearts in this and the following 
species, which, at any rate in L. vejdovskyanus, have a peculiar 
form and arrangement, would appear to demand the creation of a 
new genus for this worm ; for Michaelsen, in ‘ Das Tierreich,’ gives 
two pairs of hearts in segments vill. and ix. as a character of the 
genus Limnodrilus. But in the character of the cheete, and still 
more in the chitinous penial tube, this species agrees so precisely 
with Limnodrilus, that I deem this procedure unnecessary. 


LIMNODRILUS LUCASI, sp.n. (Plate XXYV. figs. 18-22.) 

A slender worm with thick body-wall; the segments are not 
annulated. 

The prostomiwm is short and rounded; the peristomium is as 
long as segment il. 

The chate are 5 in each bundle in the most anterior segments 
(ii., lii., iv., v.), then diminish to 4; and later (in segment x. and 
posteriorly) to 3 in each bundle. 

The dorsal and ventral cheetz on segment ii. are rather shorter 
than on the other segments. All the cheetee are alike; the prongs 
or teeth are nearly equal in length, but the lower or proximal 
tooth is slightly the stouter, and in the posterior segments it is a 
trifle longer than the lower prong. But even in one and the same 
bundle the relative length of the two prongs exhibits various 
proportions. 

Ventral chetz are present on segment xi. even in the adult. 

The clitellum occupies two segments, extending from 3x. 
to $ xu. 

Internal Anatomy. 


The pharynx extends through segments ii. and iii.; the chlora- 
gogen granules are dark brown and commence in segment v. 

A large heart exists in segment vili., but I have not been able 
to make out, in the entire individuals, the exact relations of this 
organ. The blood has accumulated in the vessels at the posterior 
end of the body, and the vessels are empty anteriorly. I did not 
investigate this species by means of sections. The usual undu- 
lating commissural vessels are present in the anterior segments, 
but I find no integumental vessels. 

The dorsal blood-vessel shifts from its proper position in the 
intestinal region, and takes up a lateral position as in branchiura 
sowerbyt (which, according to Beddard, is the only instance 
amongst the Turbificide of this arrangement). 


1903.] - OLIGOCHETA FROM NEW ZEALAND, 217 | 


Sperm-sacs occupy segments xil., xiii, and eggs lie free in 
segments Xlv., XV. 

The chitinous penis is about ten times as long as the breadth of 
its base (Pl. XXV. fig. 18); it is distinctly bent at a point just 
below its outer end, and terminates in an asymmetrical, some- 
what trumpet-mouthed expansion, which is apparently imperfect 
on one side, as the chitin here becomes very thin (Pl. XXV. 
figs. 19, 20); there appears to be a “‘valve-like” arrangement, 
somewhat like that figured by Vejdovsky for L. claparedianus (pl. x1. 
figs. 7,8). The muscles surrounding the penis are spirally wound. 

The spermatheca (Pl. XXV. fig. 21) has an irregularly pyriform 
ampulla, connected to the pore by a narrow neck passing into a 
short muscular duct, which is rather wider in the middle of its 
course than at either end. The duct is much shorter than in the 
previous species. The spermatophores (Pl. X XV. fig. 22), which 
I observed in one instance, are dumbbell-shaped—~. e., an oval 
constriction round its shorter diameter. 

Dimensions. Length 15-35 mm.; diameter ~ mm. or 3 mm. 
With 60 to 80 segments. 

Localities. Lakes RotoitiandTaupo, North Island, New Zealand. 
Obtained in considerable numbers at both places. 

Remarks.—This worm is much more slender than L. vejdousky- 
anus, and, like it, differs from the majority of species of Limnodrilus 
in possessing no integumental vessels, so far as can be made out in 
preserved specimens. As the blood was distinct enough in the 
intestinal network, it seems unlikely that I overlooked the vessels 
on the body-wall, or going thereto. It appears to approach 
L. dugest Rybka*, from Mexico, from the diagnosis given by 
Michaelsen; but I have not access to the original paper in which 
it is described. 


LIMNODRILUS sp. inc. 


From Lakes Wakatipa, Manapouri, and Waikaremoana some 
immature specimens were obtained to which I will not give a 
specific name, but which differ from either of the preceding species. 

The cheetze are in bundles of 4 or 5 anteriorly, but soon decrease 
to a couple: or in some instances (the single individual from the 
southern lake) the maximum, anteriorly, is two per bundle, and 
posteriorly, both dorsally and ventrally, a single cheta. But in 
both cases the form of the crochet is the same; the upper prong 
is much larger than the lower, indeed as much as twice the length, 
and is much more slender and more elegantly curved, as it seems, 
than in the preceding species ; the lower tooth is slightly stouter 
than the upper. 

A more important difference from the preceding species is the 
possession of two pairs of swollen hearts in segments vill. and ix. 

In one case, although only the rudiments of the generative 
organs are present, the epidermis is slightly, but definitely, thick- 
ened on segments 3 X., X1., 3 XU. 

* Rybka, Mém. Soc. Zool. France, xi. p. 380. 


218 PROF. W. B. BENHAM ON AQUATIC [ Nov. 3, 


Remarks.—In 1889 (P. Z. 8.) Mr. Beddard recorded the occur- 
rence in New Zealand of ‘“‘ Limnodrilus sp. ine.,” but gave no 
details as to its anatomy; but in 1892 (P. Z. 8. p. 354) he states 
that the New-Zealand Limnodrilus possesses two pairs of greatly 
dilated hearts in segments viii., ix. In his Monograph, p. 247, 
he repeats this; and both on this page and p. 230 he refers 
to this worm as “ L. novezelandie.” Possibly this is a MS. name 
and crept into the Monograph unintentionally; but it is regrettable 
that no further details have hitherto been published, and as his 
specimens are stated to be immature, it is probable that we shall 
never know what “ LZ. novezelandiew” is: it is a name that has to 
disappear. It is quite likely, of course, that the present “species” 
is identical with Beddard’s, and I hope to obtain more material 
before giving it any name. 

In view of the general distinctness between the lacustrine worms 
of the Northern and Southern lakes, it is possible that two species 
are included here; but in the immature condition I detect no 
peculiarity sufficient to differentiate them. 


Tn looking up the literature dealing with the genus Limnodrilus 
and other Tubificids, I have been struck with the paucity of 
information, on many anatomical points, about the common Huro- 
pean species. It seems to me desirable to have some information 
as to the amount and degree of variation that may occur in the 
form of the chete of Limnodrilus, so as to be able to ascertain 
how far the relative size of the two prongs is a reliable specific 
character. 

Another point that requires attention is the extent of the 
clitellum in different species of Tubificids, for in Michaelsen’s 
and in Beddard’s Monographs little attention 1s paid to this point ; 
and since in the Harthworms it is of value in identification, it 
seems likely that here, too, it would have a certain, but perhaps 
more limited, value. 

It is only in the case of those species that have been examined 
in recent years that this point has been determined. In neither 
monograph do I find a statement as to its extent, for instance, in 
the common Huropean species of Limnodrilus, and, indeed, in very 
few members of the family. Beddard (p. 85) in a tabular state- 
ment showing the position of the genital organ in the various 
families of the Oligocheta, writes under the heading “ clitellum ”:— 


COMM pou Ooo aotenshcba tae NOW 


But in the discussion on the characters of this family, further on 
in the volume, it is not stated, either explicitly or implicitly, 
whether the comma between the two numerals stands for the 
word “and” or “or”; and no details as to the point in question 
are to be found in the account of the genera or species of the 
family. But from the few records that we have, it is evident that 
in the family Tubificide the clitellam is not limited to one or even 
to both of these segments, and it is more extensive and variable in 


1903. ] OLIGOCHATA FROM NEW ZEALAND. 219 


its extent than would be supposed from the above scant statement. 
For instance, I have picked out from Michaelsen’s systematic 
summary of the family, in ‘Das Tierreich,’ the following definite 
statements, and these are the only ones that I can find :— 


Branchiura sowerbyt  .......0..0.0..0+ x LOX: Beddard *. 
6 1 jeaood 

Tubifex (Heterocheta) costatus ...... —x.to -xii. Benham 7. 
T’. (Psamm tes) velutinus ...... er 

(ESCTAD ROY BIES ) ees 4x. to Xil. Randolph ¢. 
LUM Cece) GINO Sen none odcnecaee ate eee é ‘ 
LE ( SPTOSPETMUG)) FOTO... vo. ocr ane sce- UO Klis 
Tovbojew blamcharde «vis... s. sesso sees: (xe) xvandibar, 
Rhizodrilus (Vermiculus) pilosus...... x. tos Xiv. Goodrich §. 

Tee 5 

RMU CLOUSIT AAG. eae Ne Set cele tates | 3x.to—xill, F. Smith |. 
Bothrioneuron WMericanunr .........00. Sly NOU Beddard 4. 
(QUiiiGLED CHRELUTTAOOIS dosncseecbon0snaasdooone 160) SU, 
LTCC. CUD hagedeicessscect8saca0 60 TEX LON Sexi  eEveavpeniineys 


I have not at my command the earlier literature on the subject ; 
but it is a curious fact that in neither of these monographs is 
there a record as to the position of the clitellum in such common 
European species as Zubifex rivulorum, Limnodrilus clapa- 
redianus, &e. 


TUBIFEX sp. inc. 


From Lakes Taupo and Rotoiti some immature worms were 
obtained which appear to belong to this genus. 

The dorsal bundle contains 2 to 4 long capilliform chetze together 
with 2 to 4 ‘‘ ctenates”; the ventral bundle 4 crochets anteriorly, 
dwindling to 2 posteriorly ; the two prongs of about equal length, 
but the lower is rather stouter than the upper. In one instance 
the capilliforms occur only in the first few segments, in other 
cases (from Rotoiti) they are present throughout the body. Only 
one cheeta exists in the ventral bundles of segment x., and none 
at all in segment xi. Commissural blood-vessels are present in 
segments 11. to xi., those of the last two segments being very 
long; while in segment viii. is a large, much dilated “ heart.” 

Though these characters are insufficient to permit me to give a 
name to the species, it appears to be different from 7’. rivulorum, 
which species has been recorded by Beddard (1889) from New 
Zealand. 


ENCHYTREZUS SIMULANS, sp. n. (Plate XXYV. fig. 25 & XXVI. 
figs. 26-28.) 
Seven short and relatively stout worms were amongst those 


* Beddard, Quart. Journ. Micr. Sci. xxxiii. p. 325 

t Benham, Quart. Journ. Micr. Sci. xxxiii. p. 188. 
Randolph, Jen. Zeit. xxvii. 

£ Goodrich, Quart. Journ. Micr. Sci. xxxvii. p. 253. 

|| Smith, Bull. Illinois Lab. v. p. 244. 

q Beddard, Hrgeb. Hamb. Magalhaen. Sammelreise, p. 7. 

** Watai, Annot. Zool. Jap. ili. p. 5 


220 PROF. W. B. BENHAM ON AQUATIC [ Nov. 3, 


collected in Lake Taupo, and clearly belong to the genus 
Enchytreus. 

The prostomiwm is short, rounded, and somewhat conical; the 
anterior segments are well marked, though the body-wall is thin 
and transparent. 

The chete are in four bundles, of (usually) three in each bundle ; 
they are straight rods with blunt ends; all are alike in form 
and size. The three chetee in each bundle are arranged fanwise, 
i. é., the middle one upright, and the other two making equal 
angles with it on either side. Occasionally, in the anterior ventral 
bundles, a fourth cheta was noted. 

The clitellum covers segments xii., Xili., and part of xiv. 

The male pores, on segment xil., are in depressions on either side 
and in line with the ventral series of chete, which, however, are 
absent in this segment. 


Internal Anatomy. 


The brain is convex posteriorly. Peptonephridia are absent 
from the back of the pharynx. 

The ordinary nephridium has a relatively long and narrow pre- 
septal region (Pl. XX VI. fig. 26); the postseptal region is about 
twice the length of this, and is distinctly marked off from it. 
It consists of an irregularly pear-shaped “ body,” which tapers off 
posteriorly to form a long narrow duct, set nearly at right angles 
to the “‘body,” but slightly inclined forwards. 

The spermiducal funnel is particularly large (Pl. XX V1. fig. 28), 
about four times as long as its breadth; it is thick, and bent in 
more than a U-shaped curve (perhaps due to changes during 
preservation), for it appears S-shaped in longitudinal section. The 
entrance is narrow, but there are no columnar cells at its margin, 
which is not reflexed. The wall of the whole funnel consists of 
long glandular cells, which in their distal moities are filled 
with granules. 

The sperm-duct is short, confined to its segment, and coiled in 
a close and regular zigzag; it opens in a depression which results 
from the contraction of several radiating muscles in this region, 
near a group of gland-cells which open through the epidermis on 
the outer side of the male pore. 

The spermatheca (Pl. XXVI. fig. 27), which communicates with 
the cesophagus, lies in the usual segment. The ampulla is nearly 
spherical; the duct is distinctly marked off from it, is about half 
the length of the ampulla and much narrower than it. 

Gland-cells are present around the duct throughout its length ; 
that is, the epithelium consists of tall cells with clear contents; 
the greater part of each cell projects beyond the muscular wall of 
the duct, and these portions form a continuous extra-muscular 
layer (Pl. XXVI. fig. 28). The actual lining of the duet appears 
to be formed by a protoplasmic sheet, in which I detect neither 
cell-boundaries nor nuclei; and this sheet appears to result from 
the fusion of the internal ends of these “glandular” cells. The 


1903.] OLIGOCH ETA FROM NEW ZEALAND. 221 


condition, indeed, is similar to that figured by Michaelsen for 
“ H, mobii” (= ZL. albidus). 

Dimensions. Length 15 mm.; diameter 2mm. Segments 
about 58. 

Locality. Lake Taupo, North Island, New Zealand. 

Remarks.—lt seemed likely, from a preliminary examination, 
that this species might be 4. albidus Henle, which has a very 
wide distribution ; butin the details of the more important organs 
there appear to be sufficient differences to permit the bestowal of a 
new name. 

The figures of the male apparatus given by Hisen* and by 
Michaelsen+ show a distinct everted lip to the spermiducal 
funnel; the sperm-duct is not so compactly coiled in a zigzag, 
while it may reach as far back as the eighteenth segment. 

The nephridia of HZ. albidus also appear to differ from these 
organs in our species § ; while the absence of a peptonephridium 
in our New-Zealand worm appears to mark it off from #. albidus. 

In possessing only 3 chet per bundle, it resembles #. hyalinus 
Eisen, and H. adriaticus Vejd. From the latter it is distin- 
guished by the form and proportions of the three regions of the 
nephridium. With the former, however, it agrees very closely 
in the structure of the male efferent apparatus §, but the sperma- 
theca in that species has an “ atrium-like dilatation” on its duct, 
which is absent from the new species. The form of the nephridium 
is also a point of agreement. But the fact that #. hyalinus occurs 
in Novaya Zemlya seems to exclude the possibility of its imtro- 
duction into Lake Taupo. 


AcHm@TA mAoRIcA, sp.n. (Plate XXV. figs. 23, 24.) 


A single, but fortunately a mature specimen of this small worm 
was obtained at Station 18 in Lake Manapouri. 

Tt was stained in alum-cochineal and mounted entire, and its 
anatomy studied as far as possible. It was then unmounted, and 
the anterior half was cut into a series of longitudinal sections, 
and the rest of the body into transverse sections. Owing to the 
flattening to which it had previously been subjected, the former 
series was not very satisfactory, especially as the sections were 
a good deal torn by the diatom-valves and dirt in the intestine. 
Nevertheless the anatomy was sufficiently studied for systematic 
purposes. 

The prostomium is short, rounded, and provided with the usual 
terminal pore. 

There are no chetigerous sacs, nor could I detect any “ chloro- 
phyll-glands.” 

The clitellum appears to include only segment xil., encroaching 
put slightly into the hinder region of segment Xi. 


# Hisen, Svenska Ak. Handl. (n. ser.) xv. p. 25, pl. ix. fig. 18, pl. x. fig. 20. 
+ Michaelsen, Untersuch. u. H. mobi, 1886, p. 1, pl. iii. fig. 9. 

+ Goodrich, Quart. Journ. Micr. Sci. xxxix. p. 51, pl. v. fig. 2 (nephridium). 
§ Hisen, loc. cit. pl. x. fig. 20. 


222 PROF. W. B. BENHAM ON AQUATIC [Nov. 3, 


T could not detect any ‘great cells,” such as occur in the clitellum 


of some species of this genus. 
The male pores are paired and situated at about the middle of 


segment Xil. 
The spermathecal pores, paired, lateral, lie at the anterior 


boundary of segment v. 
Internal Anatomy. 


There are septal glands on the hinder septa of segments Iv., V., V1., 
and these septa are rather stouter than the rest. 

Segments vii., viii. (as indicated by their ganglia, for the external 
boundaries are difficult to detect in the transparent worm) are 
much shorter than their neighbours ; the tenth and following are 
almost twice the length of either of these two. 

The dorsal vessel arises in segment x.; but I fail to discover any 
swellings on its course. 

The sperm-funnel occupies about half of segment xi. ; it is urn- 
shaped, without an everted margin (Pl. XXV. fig. 23), and its 
length is about equal to twice its breadth. 'The sperm-duct, less 
than twice the length of funnel, takes a nearly straight course 
backwards to about the middle of segment x1i1., when it bends down- 
wards almost at right angles; it is dilated after passing through 
the septum xi./xii., but it soon narrows again, and the external 
opening is surrounded by a small lens-shaped mass of glandular 
cells (spermiducal gland), which it perforates at about the centre. 

Loose masses of developing spermatozoa occur in the body-cavity 
of segments ix. and x., and a few even in segment viil. 

The spermatheca is much elongated (Pl. XXV. fig. 24); its aper- 
ture (surrounded by a group of gland-cells) leads by a short canal 
into an ovoid dilated sac lying in segment v., thence a narrow canal 
passing through segments vi., vil. begins to enlarge after passing 
through the septum vii./vili. to form a large “ampulla” Jying in 
segments ix. and x., which is constricted at about its middle, 
The ampulla contains bunches of ripe spermatozoa. 

The ovary and duct have the usual positions, and the body-cavity 
of segment xii. is fully occupied by a couple (or more) of large eggs, 
which distend the body. 

Dimensions. Length 4 mm.; breadth very small. Number of 
segments 22, with an anal segment. 

Locality. Lake Manapouri, South Island, New Zealand. 

Remarks.—This species agrees with A. (Anacheta) cameranotr 
Cognetti* in its smaller size and in the total absence of cheetigerous 
follicles, but in nothing else; for its spermatheca somewhat resembles 
that figured by Vejdovsky (Syst. u. Morph. pl. vii. fig. 22) and 
labelled A. etsenti, but which Beddard + suggests really belongs to 
A. bohemica. But in the point of origin of the dorsal vessel it 
differs from each of these species. 

Having only a single preserved specimen, I am unable to give 


* Cognetti, Boll. Mus. Zool. Anat. Comp. Torino, xiv. 1899, no. 354. 
+ Beddard, ‘ Monograph,’ p. 356. 


1903. | OLIGOCHATA FROM NEW ZEALAND. 223 


any details as to lymph-corpuscles, nephridia, and certain other 
anatomical features. 


HAPLOTAXIS HETEROGYNE, sp. n.* 


The prostomium is long, but not annulated. 

The chete are four per segment, isolated, one dorsal and one 
ventral on each side; the form agrees with that found in H. gordt- 
cides. The ventral cheta is from two to three times the length of 
the dorsal, but less difference exists in the anterior segments. 
The dorsal chetze occur throughout the body. 

The clitellum surrounds segments xi. to 4 xiv.; it is complete 
and thickest laterally. 

The genital pores were not detected externally ; but the ducts, 
traced in sections, meet the body-wall at points indicated below. 
There are two pairs of male ducts which reach the epidermis in 
segments Xi., xii. anteriorly to the ventral chete. There is a 
single pair of oviducal pores on segment xiii. laterad of the line 
of ventral chete, but rather nearer the anterior margin of the 
segment than these; but they are much further back in their 
segment than the male pores are. 

‘Two pairs of spermathecal pores lie at the anterior margins of 
segments Viil., 1x. 

“The alimentary y canal is remarkable for possessing a strongly 
muscular gizzard in segment iv.; it is quite different from 

a pharynx, which is here absent. A similar, but more extensive, 
Bae has been recorded by Michaelsen for H. gordioides. 

The first pair of nephridia lie in seamen x.; they are rather 
smaller than the postovarian nephridia. These organs are absent 
in segments Xi., Xll., xXi1., but reappear in segment xiv. et seqq. 

The nephridial canal perforates a string of large cells having 
distinct boundaries and highly vacuolated cytoplasm. Hach 
nephridium reaches the epidermis close to the ventral cheta, 
through a short “ duct” formed of a highly granular syncytium. 

As in the case of the sperm-ducts, I was unable to detect an 
actual perforation or opening through the epidermis. 

Reproductive System (text-fig. 23, p. 224).—Two pairs of testes 
and sperm-funnels occur in the usual positions in segments x., Xi. 
The sperm-duct leaves the large, thick, flattened sperm-funnel at 
its ventral edge, as Beddard { found was the case in H. smithi. 
But the sperm-duct in that species has the usual structure, 
z. é. is surrounded by a definite epithelium. 

In the present species the sperm-duct perforates a series of cells 
the boundaries of which are not distinguishable, and has an undu- 
lating course in this syncytium, which extends up to the level of the 
lateral line. The duct passes upwards to this level, then bends upon 
itself, still within the syncytium, and reaches the body-wall in front 


* A detailed and illustrated account of this worm has been sent to the Editor of 
the Quart. Journ. Micr. Sci. 

+ Michaelsen, Zool. Jahrb. (System.) xii. p. 105. 

~ Beddard, Ann. & Mag. Nat. Hist. ser..6, i. p. 389 (1888). 


294 PROF. W. B. BENHAM ON AQUATIC [Nov. 3, 


of the ventral cheta. A transverse section through the sperm- 
duct has all the appearance of a section across a nephridium : we 
see, not an epithelium surrounding the lumen, but a perforated 
cell, with rarely more than one nucleus in the plane of the section, 


Text-fig. 23. 


S RS 
p.s.---BS 
N 
S 
. _Od. 
XIN N 
O.S--8 
SS ia | ss @ ---- ----- --Ne f. 
N P 
S Ra --Ne. 
S N 
SS 
XIV N 
S S 
S 2 N 
S N 
~ = BS 
xv «Of N 
N BS 
'S S 
S 


Plan of genital organs in Haplotawis heterogyne, composed from study of entire 
worm and sections. The gonads are omitted on the right side m order to 
show the full course of the sperm-ducts, each of which leaves its funnel at its 
ventral edge; it has a course similar to that of a nephridium, and, like the 


latter, traverses a cord of cells. 
We}, the first nephridium; Ve.?, the second; We.f., nephridial funnel; O.d., 


oviduct; O.s., ovisac; Ov., ovary; Sp.d., sperm-duct; Sp.f., sperm-funnel; Sp.s., 
sperm-sac; Spth., spermatheca ; ¢., testis. 


1903. | OLIGOCH ETA FROM NEW ZEALAND, 225 
sometimes none. The general disposition of the perforated syn- 
cytium is very similar to a nephridium, and the canal takes a 
course similar to the latter tube, but is not quite so undulating. 
The canal bears cilia throughout its whole extent. 

So similar is this sperm-duct to a nephridium in this worm, 
that the differences are only perceptible when the organs are 
examined under a very high magnification (such as 74, homogeneous 
immersion). But that this nephridium-like tube does actually serve 
as a sperm-duct, is demonstrated by the presence of sperms within 
the canal, and the fact that in the case of one funnel I noted 
spermatozoa entering the mouth of the canal. Though I traced 
the ducts to the body-wall, I was quite unable to detect the pore 
in either sperm-duct ; nor was Michaelsen successful in finding 
the actual aperture in H. gordioides. Of the two individuals 
obtained by me one was fully mature; the segments x., xi. were 
filled with ripe and developing sperms; and the spermathecez were 
also filled with them ; hence copulation had recently occurred. The 
other individual was quite immature, although the three pairs of 
gonads and the ducts were present ; in this individual, Jikewise, 
there are no nephridia (unless the sperm-ducts are nephridia) in 
segments Xi., Xll., X11. 

It appears to me that in this worm the nephridia do act as 
sperm-ducts, as was suggested by the earlier students of Haplotazis. 
There are two median sperm-sacs, in the form of simple pouches, 
formed by the septa x./xi., x1./xil., which are pushed backwards 
above the gut so as to reach into segments xil., xii. respectively. 
Each of the sperm-sacs is filled with developing and ripe 
spermatozoa. 

There is but a single pair of ovaries and oviducts: the former 
in segment xii.; the latter opens in about the middle of segment 
xiii. The oviduct is a wide tube surrounded by a ciliated epi- 
thelium, and opens by a wide funnel into segment xii. This duct 
is present in the immature specimen, and can be seen traversing 
the anterior half of segment xiii. 

The presence of a single pair of female organs marks off this 
species from the two other known representatives of the genus; 
and in this respect our species resembles Pelodrilus, an allied 
genus, originally founded by Beddard * for a New-Zealand worm, 
P. violaceus ; but the discovery by Michaelsent of P. ignatovi 
from Central Asia, in which the two pairs of sperm-ducts oper 
on independent segments, forms a passage to Haplotaxis. But 
in all other characters this new species agrees with the diagnosis 
of Haplotaxis and differs from that of Pelodrilus. 

A single median ovisac, filled with eggs, passes backwards 
through segment xiil, 

In segments Xi., Xil., xiii. are paired copulatory glands, similar 
to those present in some Enchytreids ; those of segment xii, open 
laterally, near the ventral cheete ; the other two pairs open below 

* Beddard, Trans. R. 8. Edinb. xxxvi. p. 292. 
+ Michaelsen, Verh. naturw. Ver. Hamburg, 1903. 


Proc. Zoou. Soc.—1903, Vou. 11. No. XV. 15 


226 PROF. W. B. BENHAM ON AQUATIC [Nov. 3, 


the nerve-cord. Each gland consists of a bundle of long club- 
shaped cells. a é 
Two pairs of globular spermathece lie in the anterior moiety 
of segments viii. and 1x., and open laterally, at the anterior margin 
of these segments. 
Dimensions. About 20 mm. x 3mm. With about 60 segments. 
Locality. Lake Wakatipu, South Island of New Zealand ; from 
a depth of 550 feet. : 


DrroRoCHATA AQUATICA, sp.n. (Plate X XVI. figs. 29-31.) 


Two individuals were obtained, of which one was entire and 
well preserved; the other, broken, soft, and almost useless for 
study. The former was cut into sections. 

Tt is a short and relatively stout worm. 

The prostomiwm is small and prolobie. 

The chete are about 28 per segment, 2. e. 14 on each side, and 
the gaps aa=ab, 2x=13 yz; thus the midventral “gap” is 
practically absent. 

The clitellum is not fully developed in either individual; it 
appears to cover segments xiv., xv., xvi. and the dorsal region of 
the 13th. 

The male pores, on the 18th segment, are distinctly visible under 
a lens, but are not on papille ; they are rather widely separated, so 
as to lie, when seen from below, on the “edge” of the ventral 
surface; they are on a level with the chetal gapsd/e. There are no 
cheetze visible between these pores. 

The oviducal pores are paired, but close together on a pale oval 
area, in line with cheeta a. 

The spermathecal pores are not visible externally, but are on 
the anterior margins of segments viil., ix. I did not note their 
position relative to the cheete. 


The nephridial pores are situated about halfway up the body- 
wall. 


Internal Anatomy. 


There is no gizzard; the cesophageal wall is vascular and 
folded as it passes through segments x., xi.,xli. It then diminishes 
in diameter and becomes thin-walled in the next two segments, 
dilating in the 15th, to attain the full diameter of the intestine 
in the 16th segment. There is a minute typhlosole in the form 
of a small, low, rounded ridge, deeply separated from the rest 
of the lining by a furrow on each side (Pl. XXVI. fig. 29); 
its epithelium differs from that of the general lining of the 
intestine in that it consists only of ciliated columnar cells 
(Pl. XX VI. fig. 30); whereas in the rest of the epithelium two 
kinds of cells are distinctly recognisable (Pl. XXVI. fig. 31), 
namely, (a) long, narrow, ciliated cells, and (6) gland-cells of 


considerable size, whose bases project into the blood-sinus sur- 
rounding the gut-wall. 


1903.] OLIGOCHATA FROM NEW ZEALAND. 297 


Of the vascular system I noted that the last heart is in 
segment Xiil. 

There is a peculiar ‘ glandular organ” below the dorsal vessel 
in the intestinal region. 

The nephridia are of large size, and visible through the trans- 
parent wall in the entire “ uncleared”” worm. 

The Reproductive System.—The two pairs of testes, the ovaries, 
and oviducts have the normal position. 

There are two pairs of sperm-sacs, in segments ix. and xi.., 
while the two intervening segments are filled with loose masses of 
developing sperms. 

The spermiducal gland is visible through the transparent wall ; 
it extends through segments xix. to xxiii.; it has the usual form 
—long, cylindrical, undulating ; its curved duct is confined to the 
eighteenth segment. 

There are no penial cheetee. 

The spermathecz, in segments vili., ix., are globular sacs, quite 
filling their segments, with a short duct, into which opens a small 
ovoid diverticulum. 

Dimensions. Length 38 mm.; diameter 1? mm., reaching 2 mm. 
anteriorly. Segments 67. - 

Locality. Lake Manapouri, in the South Island of New Zealand, 
from depths of 350 to 500 ft. 

Remarks.—tIn the absence of a gizzard this new species differs 
from all the known species of Diporocheta, and therein agrees 
with Perionyx, from which, however, it differs in all the characters 
that differentiate the two genera. 

No doubt the absence of gizzard is related to its aquatic habit. 
Of the numerous species of this characteristically Australian 
genus, only three have so few spermathece as the present one— 
viz., D. pellucida Bourne *, from India, D. moroea Spencer 7, 
from Tasmania, both of which are of much greater size and have 
more cheetze per segment ; and thirdly, D. scolecoidea Spencer ¢, also 
from Tasmania, which, although of smaller size than the new 
species, has many more chet ; and in all other important respects 
these three differ from the present form. 

It is a fact of some interest that the first New-Zealand species 
to be recorded, viz. D. intermedia Beddard §, was obtained from 
Lake Brunner ; in it the gizzard is present, but of small size: 
D. chathamensis Benham ||, in which the gizzard is also small, 
came from a peat-swamp: while another species, the description 
of which is not yet published, has no gizzard recognisable on dis- 
section ; this is from the Otira gorge, not far from Lake Brunner. 
Thus, of four known New-Zealand species of this genus, two are 
aquatic. This may have some bearing on the sparse record of 


* Bourne, Quart. Journ. Micr. Sci. xxxvi. p. 18. 
+ Spencer, Proc. R. Soc. Victoria, vii. p. 49. 
+ Spencer, Joc. cit. p. 61. 
§ Beddard, Quart. Journ. Micr. Sci. xxx. p. 467. 
|| Benham, Trans. N. Z. Instit. xxxiii. p. 1384, 
5 


228 PROF. W. B. BENHAM ON AQUATIC [ Nov. 3, 


occurrence in this country of a genus so abundant in Australia, 
of which the Earthworm fauna is so entirely distinct, for our 
aquatic worms have, as yet, been scarcely touched. 


PLUTELLUS LACUSTRIS, sp. n. (Plate XX VI. figs. 32-40.) 


Six hauls in Lake Wakatipu brought up specimens of a worm 
which in some respects resembles Pontodrilus. 

The collection consists of 24 individuals, some few fully mature ; 
others of full size, but without a clitellum; and still more quite 
young. One haul (No. 5) contains a number of cocoons, which 
from their size probably belong to this species. 

The worms, as preserved, are robust in form, greyish in colour, 
but the body-wall behind the clitellum is translucent. Those that 
are well preserved have much annulated segments: three and 
four annuli anteriorly, but more in the posterior segments. 
The anterior segments are very short. 

The prostomium is relatively long, equal in length to the first 
two segments; it 1s narrow, only half the breadth of the peri- 
stomum, into which it is not embedded. It is prolobic. 

The chet, eight per segment, are isolated; their formula is 
approximately thus: aa = 2.ab; ab =3bc; be =cd; dd = about 
2aa. The spacing differs somewhat in the anterior and posterior 
regions ; ada is greater and 6c is rather less anteriorly than in the 
greater part of the body (fig. 32). 

The cheetze are of the usual form (Pl. XXVI. figs. 33, 34), 
but are ornamented with a number of extremely fine crescent- 
shaped marks near the distal extremity; such as occur in the 
genus LRhinodrilus and others, including Pontodrilus bermud- 
ensis Bedd. and P. insularis Rosa. 

The cheetze, both dorsal and ventral, of segments ii., iii., iv. are 
smaller than those of the body generally, attaining only half the 
length of the latter; but the increase is gradual, as the following 
measurements show :— 


Ventral cheeta of segment 11. measures ‘015 mm. 
39 2? Vs 2? "0225 3” 
” or) BE, ” 03 ” 
9?) 9 XV1l. ? 05 2) 


The ventral cheetze (a and 6) areabsent on segment xviii., being 
here replaced by a couple of copulatory bristles on each side in 
line with cheta 6. 

The clitellum, in the most fully-developed individuals, 7. e. in 
three out of five clitelliferous worms, extends from 3 xiii. to end 
of xvi. (7. e. 35 segments), the hinder boundary being, in each case, 
very well marked ; it completely encircles the body. 

The male genital pores on segment xviii. are in the line of 
cheeta 6; they are quite minute, and are not indicated by any 
papille ; but the ventral region of this segment differs in appear- 
ance from the rest, and is seen to be glandular in transverse 
sections. In one case this area was somewhat depressed, by the 


1903.] OLIGOCH ETA FROM NEW ZEALAND. 229 


contractions of internal muscles, but the margins were not promi- 
nent nor specially glandular. 

Certain “tubercula pubertatis” are present in the mature 
worms, in the form of one or more median, oval, glandular pads, 
with a definite, rather raised margin and a central depression. 
All my specimens present one such tubercle in segment xvii. ; 
it is prechetal, but not intersegmental in position. In one case 
(a) this alone is present. In other individuals a postchetally- 
situated tubercle also occurs in one or more of the segments 
following the male pores, viz. in (6) it is on segment xix., in (c) 
on Xxil., in (d) on xix. and xx. Thus the maximum number, so 
far as my material enables me to ascertain, is three median 
tubercles. 

The oviducal pores appear as small white spots just in front of 
cheeta @ on each side of segment xiv., and this is confirmed by the 
study of sections. 

The spermathecal pores are four pairs, in line with cheta a, at 
the anterior margin of segments Vi., Vil., viil., and ix. respectively. 

The nephridiopores are in line with cheta 6. 

The dorsal pores commence in the third (or perhaps the second) 
segment, 


Internal Anatomy. 


The septa forming the hinder walls of segments viii., ix., x., xi. 
are a good deal thicker than their immediate neighbours. 

The dorsal blood-vessel is single. I did not detect a supra- 
intestinal vessel. 

There are three pairs of hearts, in segments x., xi., xii. Of these, 
the last only has the usual form and arrangement as a semi- 
circular, dilated commissural vessel passing freely from the dorsal 
to the ventral vessel (Pl. XX VI. fig. 39). 

But the other two pairs present a condition which appears to 
be unique: they are, for the greater part of their extent, adherent 
to the cesophageal wall; indeed, they appear in sections to be 
rather dilated, circularly disposed channels of the general peri- 
cesophageal plexus. Hach becomes free only at its ventral end; 
at the same time it becomes gradually reduced in diameter, and 
soon becomes very narrow where it joins the ventral vessel 
(Pl. XXVI. fig. 40). Anteriorly to the tenth segment I find 
narrow commissurals of the usual character. 

The alimentary canal is provided with a pharynx of the usual 
form ; there is no gizzard nor cesophageal gland. 

The cesophagus and intestine contain abundance of food, 
amongst which the valves of diatoms are present in considerable 
numbers. The cesophagus is a good deal dilated segmentally, and 
passes into the wider intestine in segment xviii., where the 
character of the epithelium suddenly changes from a high to a 
shorter type of cell. 

The worm is meganephric, with large funnels; but nephridia 
are absent in the anterior segments; the first one oceurring in 


feo) 


230 PROF. W. B. BENHAM ON AQUATIC [ Nov. 3, 


the 15th (or in another case the 16th) segment—a point of 
resemblance to Pontodrilus. 

The Reproductive System.—The two pairs of testes and their 
funnels occupy the usual position in segments x., xi. 

The spermiducal funnels are large and prominent, but are very 
little folded. The ducts of each side unite in segment xili., and 
the single duct thus formed opens into the neck of the spermi1- 
ducal gland at the junction of the latter with its muscular duct. 

The gland is tubular, undulating, and occupies segments xvii. 
and xviii.; its muscular duct is short, narrow, and curved, opening 
to the exterior in segment xviii. by a very small pore. ‘The 
structure of the gland is similar to that normal in the subfamily ; 
the lumen is provided with an epithelial lining of short columnar 
cells, between which pass the necks of long, club-shaped gland- 
cells in groups; the layer of gland-cells is of considerable depth 
and is covered by peritoneum ; no muscles are present till the 
duct is reached. 

In connection with each of the male pores is a couple of long, 
delicate copulatory chet (accompanied by one or two reserves on 
each side). Each cheeta is about twice the length of a locomotor 
bristle (Pl. XX VI. figs. 35, 36) and half its width; it is nearly 
straight, and its terminal region presents a few lateral notches— 
which seem to correspond to the crescent-shaped furrows in the 
normal cheete. 

There are two pairs of sperm-sacs occupying segments ix. and 
xi., and the intervening segment is filled with loose masses of 
developing spermatozoa. 

The ovaries and their ducts occupy the usual position. Each 
ovary is of considerable size and appears, in sections, nearly to 
fill the segment ; strings of rather large ova, connected by very 
delicate threads, even extend upwards above the gut. An ovisac 
on each side contains eggs in which I note astropheres and 
spindles in two or three cases. 

There are four pairs of spermathecew in segments Vi., Vil., 
vill., ix.; each consists of a relatively large ovoid ampulla and a 
short, distinct, muscular duct, into which opens a small globular 
diverticulum filled with spermatozoa. 

Dimensions. Length 35 to 40 mm,; diameter 1; mm. Segments 
85 to 90. 

Locality. Lake Wakatipu, South Island, New Zealand. 

Remarks.—This new species differs from the rest of the genus 
Plutellus, as enlarged by Michaelsen to include several species of 
Cryptodrilus and of Megascolides, in two characters, viz., in the 
absence of a gizzard, and in the absence of nephridia from the 
fourteen anterior segments. In both these points it agrees with 
Pontodrilus (a sea-shore Oligochete), and they appear to be related 
to an aquatic habit. But in other points the agreement of this 
species is with the genus Plutellus. 

This is the first time that a representative of this genus (on the 
whole an Australian form) has been met with in New Zealand. 


1903. 


] OLIGOCH HLA FROM NEW ZEALAND. Dail 


Letters used on the Figures. 


am., ampulla of spermatheca. L.m., longitudinal muscles. 
at., atrium (spermiducal gland). nef., nephridial funrei. 
at.ep., atrial epithelium. neé.0., nephridiopore. 
b/., blood-sinus in wall of intestine. o.d., oviduct. 
b.w., body-wall. es., esophagus.  @s.’, esophageal 
c., ciliated epithelial cell. opening of spermatheca. 
e.ep., celomic epithelial nucleus. ov., Ovary. 
chl., chloragogen cells. s., septum. 
e.m., circular muscles. s.gl., septal gland. 
com., commissural blood-vessel. s.i.v., Supra-intestinal blood-vessel. 
cop., copulatory chetw, or sac con- sp.d., sperm-duct. 
taining them. spf., funnel of sperm-duct. 
d., duct of spermatheca. sp.gl., spermiducal gland. 
d.v., dorsal blood-vessel. sp.s., Sperm-sac. 
., gland-cells in intestinal epi- | spth., spermatheca. 
thelium. t., testis. 
gl. gland surrounding spermathecal ty., typhlosole. _ ty.v., typhlosolar 
ore. blood-vessel. 
hé., lateral heart. »., ventral chete. 
int., intestine. v.v., ventral blood-vessel, 
EXPLANATION OF THE PLATES. 
Puate XXIV. 


Taupodrilus simplex, p. 209. 

Fig. 1. View of the ventral surface of the genital region of 

extent of clitellum, the position of the genital pores, 
copulatory chet. 

2. Side view of five pregenital segments as seen in a transparent, stained 

specimen. (X 120.) To show certain features of the vascular system. 

3. A bundle of dorsal chete. 

4. One of the ctenates from a dorsal bundle, 

(X 700.) 

4a. The tip of another dorsal cheta, 
prongs. (X 700.) 

_ A cheta from a ventral bundle. (X 700.) 

. The tip of another ventral cheta with intermediate denticle. 

’ One of the bunch of copulatory chetz. (x 700.) 

_ Side view of the genital organs as seen in a bisected specimen ; somewhat 
diagrammatised. a, point of entrance of sperm-duct into atrium. 

. A portion of the wall of the atrium from a longitudinal section of the same. 
(x 500. The details studied with a; homogeneous immersion of Leitz.) 
The wall is seen to consist of a single layer of much vacuolated cells, the 
boundaries of which are distinctly seen in the lower half of the figure, 
where the cells were cut vertically. The round nuclei near the outside 
belong to the gland-cells; the oval nuclei towards the lumen seem to belong 
to compressed columnar cells, whose limits are not distinguishable. 

The sperm-duct is cut through twice as it winds up outside the wall, 
and again (at d’) where it is perforating the wall. 


the body, showing 
and the protruded 


with intermediate denticles. 


showing the membrane joining the two 


(Xx 700.) 


We} MID 


PuaTE XXV. 


Limnodrilus vejdovskyanus, p. 213. 


Fig. 10. Tip of a dorsal cheta. (X 700.) 
11. Tip of a ventral cheta. (x 700.) 

" \ side view of two segments of the body to illustrate the peculiar arrange 
ment of the “heart” in segment viii., where it closely embraces the gut 
around the line of union of esophagus with intestine. The anterior 
portion of the dorsal vessel is in outline, as is also the commissural vessel 
of segment vii. The posterior portion of the dorsal vessel and the supra- 
intestinal vessel are in black, as are also the intestinal vessels. 

13. An obliquely transverse section of the gut in segment Viil. 

heart lies in the furrow between the cesophagus and intestine. 

14, The penial tube. (Camera, X 120.) 


The lateral 


Fig. 15. 


Fig. 26. 
. A spermatheca—isolated ; the entire duct is covered with gland-cells 


37 
38 
39 


MR. OLDFIELD THOMAS ON THE MAMMALS [ Nov. 3, 


The distal extremity of the penial tube. (Camera, X 700.) 
. A spermatheca, as seen in an entire individual. 
. A spermatheca, isolated and uncoiled. 
Figs. 18 to 22 refer to Limnodrilus lucasi, p. 216. 
. Entire penial tube. (X 120.) Note the distinct bend near the distal 
extremity. 
. The distal extremity of a penial tube. (X 700.) 
. Another view of the same. (X 700.) 
. A spermatheca. 


2. A spermatophore, as seen within a spermatheca. 


. The sperm-duct of Acheta maorica from an entire specimen, p. 221. 

. A spermatheca of Acheta maorica as seen in an entire specimen, p. 221. 

. A transverse section of the spermathecal duct of Enchytreus simulans, 
showing the coating of glandular cells outside the muscular wall; their 
necks passing through this, and uniting to form a protoplasmic lining (a) 
round the lumen, p. 219. 


Prate XXVI. 
Enchytreus simulans, p. 219. 


A nephridium—isolated from a bisected specimen. 


(Cf, Pl. XXV. fig. 25.) 


. A male duct ; isolated. 


Figs. 29-31 refer to Diporocheta aquatica, p. 226. 


. A transverse section of the intestine (X 80), showing the small typhlosole 


(ty.); the dorsal vessel is surrounded by a structure (2). : 


. The typhlosole much more highly magnified. (Camera, X 700.) 
. Part of the lateral wall of the intestine (Camera, X 700), showing the twa 


kinds of cell forming the epithelium ; the base ot the gland-cells dip into 
the blood-sinus. 
Figs. 32-40 refer to Plutellus lacustris, p. 228. 


. Outline of a transverse section of the body showing the spacing of the 


chetz. (From a Camera drawing, X 40.) 
. A locomotor cheta. (Camera, X 120.) 
. The tip ofa locomotor cheta. (Camera, X 700.) 


. A copulatory cheta. (x 120.) 


. The tip of a copulatory cheta. (x 700.) 

. A spermatheca. 

. A spermiducal gland. 

. A somewhat diagrammatic sketch of a transverse section through the gut 
in segment xii., showing the lateral heart. : 

. A similar section through segment x. or xi., showing the peculiar condition 
of the “heart ” in these two segments. 


2. On the Mammals collected by Mr, A, Robert at Chapada *, 
Matto Grosso (Perey Sladen Expedition to Central 
Brazil). By Otpryeip THomas, F.R.S. 


By 


[Received July 23, 1903. } 
(Plate XXVII, +) 
the generosity of Mrs. Percy Sladen, Mr. Alphonse Robert, 


who had already done such good work in Sio Paulo and Parana, 


Was e 


nabled to make, during the latter half of 1902, a collecting 


expedition for the benefit of the National Museum to Matto 
Grosso, Central Brazil, a region in respect to which the Museum 


* Chapada heing a word of Portuguese origin (signifying a plateau) has its accent 


on the 


second a; Cuyabd and other names of Guarani derivation on the final a. 


+ Foy explanation of the Plate, see p. 244, 


UNUBLONYP IGS: SHINO) 
‘dunt soag wre qu SEMEL AE) WSN) arses ie 


WOOO Tel IIE! SOS SZ al 


1903. | OF THE PERCY SLADEN EXPEDITION. 233 


collections had hitherto been extremely deficient. Carried out as 
it was with all his accustomed energy and courage, Mr. Robert’s 
expedition was completely successful in spite of the many difficul- 
ties in his way, and of his being unaccompanied by any European 
helper. The collection, of which the present paper gives an 
account, is an astonishing one for him to have been able to obtain 
and prepare with his own hands, in so bad a climate, especially as 
so many of the mammals are of considerable size. Collections of 
mice are much more easily made than of dogs, peccaries, coati- 
mondis, monkeys, &c. ; and the long series of the different species 
here enumerated speaks volumes for Mr. Robert’s working 
qualities. 

The specimens are all prepared in modern fashion, with flesh 
measurements and separate skulls, and there are besides a number 
of skeletons, so that, bearing in mind the inaccessibility of the 
locality, the collection may be looked upon as one of the most 
valuable that the National collection has received for many years. 
The thanks of all zoologists are therefore due both to Mrs. Sladen 
for her generous help, and to Mr. Robert for the admirable way 
in which he carried out his instructions. 

Mr. Robert started from Santos on the 15th April, 1902, and 
after passing through the usual difficulties and delays of quaran- 
tine &c. incidental to entering Argentina, proceeded via Buenos 
Ayres, Asuncion, and Cuyaba, Matto Grosso, to Santa Anna de 
Chapada, a village situated at an altitude of about 800 m., on 
the Serra do Chapada, some thirty miles N.E. of Cuyaba, arriving 
there on the 17th June, and staying there collecting until the 
29th November, when he came down the river again with the 
results of his labours. 

Mr. Robert spent most of his time collecting mammals, but he 
also obtained a considerable number of birds, some reptiles *, 
mollusca t, insects, &c., all of which have been presented to the 
National Museum by Mrs. Sladen. 

The present is the third considerable collection of mammals 
that has been made in the district of Cuyaba. The first was that 
formed by Natterer in 1825-1829, and described in conjunction 
with his other Brazilian mammals by Wagner and Pelzeln. The 
second was that of the “ American Naturalist Exploring Expedi- 
tion” of 1882, under the leadership of Mr. H. H. Smith, and was 
made at the very village, Sta. Anna de Chapada, where Mr. Robert 
worked, The mammals were described by Prof. E. D. Cope? in 
combination with those obtained at Sao Joao, Rio Grande do Sul, 
where the expedition stayed before passing on to Chapada. 

No other important series from single localities have ever been 
made nearer to the present region than those from the Rio Jordao 
(Robert), Lagoa Santa (Lund and Reinhardt), Paraguay (Azara, 
Rengger, and Foster), and those obtained along the eastern side 
of the Andean chain by P. O. Simons. 


* Cf. Boulonger, supra, p. 69 ; Smith, supra, p. 70. 
f Am. Nat. xxiii. p. 128 (1889). 


234 MR. OLDFIELD THOMAS ON THE MAMMALS [ Nov. 3, 


Cuyaba is on the river level, and the country towards Chapada 
is all at the same low altitude until the abrupt rise of the plateau 
edge is reached, comparatively close to Santa Anna. 

Mr. Robert has discovered a considerable number of new and 
interesting species, of which the most notable is the Wild Dog 
(Pl. XXVII.), which I have named in honour of the late Mr. Percy 
Sladen. 


1, Cepus AZARZ Rengg. 

Gr 10225 TOA 1083," 1085, 1093) ss io vile 
1196, 1197, 1198, 1208. 

©. 1045, 1082, 1086, 1094, 1104, 1105, 1152, 1175, 1176. 

In the absence of any certainty that Spix’s name Cebus libidi- 
nosus (type locality, Rio San Francisco) really belongs to this 
monkey, I use that of Rengger, which antedates Geoftroy’s 
C. elegans by many years. 

This fine series emphasizes the difficulty of working out 
members of the present genus, for there is considerable variation 
both in the general colour, in the relative distribution of black 
and yellow on the head, and in the length and development of 
the temporal tufts. 


2. Aotus AzARa& Humb. 


Nyctipithecus azarce auct. 


@. 1150. 28 September, 1902. 


’ 3. CALLITHRIX* MELANURA Geoff. 
So WSO, 
Oe ey 
4, Hisriorus vELAtTuUS Geoff. 
OPS 6: 


5. MIcRONYCTERIS MEGALOTIS Gray 
©. 1014, 1015. 


6. HEMIDERMA PERSPICILLATUM Linn. 


g. 1063. 
2. 1012, 1013. 


7. LONCHOGLOSSA CAUDIFERA Geoff. 


g. 1000, 1001, 1002. 
2. 999, 1111. 


8. ARTIBEUS PLANIROSTRIS Spix. 
3. 1089. 


* For the use of Callithrix instead of Hapale, see Ann. Mag. N. H. (7) xii. 
p. 455 (1908). 


bo 
qo: 
it 


1903. ] OF THE PERCY SLADEN EXPEDITION. 


9. VAMPYROPS LINEATUS Geoff. 


g. 1091, 1092, 1099, 1100, 1101, 1200. 
2. 1090, 1096, 1097, 1098. 


Quite similar to topotypes from Paraguay. 


10. FELIS wIEDII Schinz. 
Flat skin, native made. 


11. Canis cANcRIVoRUS Desmn. 

oe 11395 0184: 

OPO} 

12. CANIS SLADENI, sp. n. (Plate XX VII.) 

g. 1080. 11 August, 1902. Zype. (B.M. No. 3.7.7.40.) 


Allied to C. vetulus, but larger, with longer skull, greyer in 
colour and with black feet. An additional upper molar present 
on each side in the type. 

Size rather larger than in C. vetulus. Fur close and uniform, 
not intermixed with long black piles; long hairs of back about 
33-35 mm. in length. General colour above clear grey suffused 
with the buffy of the underfur, the net result being nearest to 
Ridgway’s ‘smoke-grey.” Individually the longer hairs are 
light for their basal and black for their terminal halves, with a 
white subterminal ring about a quarter of an inch from their 
tip. Underfur brownish plumbeous basally, buffy clay-colour 
terminally. Under surface little paler, dull brownish, lighter in the 
axille and groins. Muzzie and chin blackish. Crown like back, 
but the light ends to the underfur almost obsolete. Back of ears 
brown, grizzled with clay-colour; longer hairs of inner surface 
creamy ; a large postauricular patch yellowish clay-colour. Fore- 
arms and thighs like body; hands and feet, both above and below, 
blackish brown. Tail brownish grey, broadly washed with black 
at the end and over the caudal gland. 

Skull, as compared with that of C. vetulus, markedly larger 
throughout ; interorbital region longer and narrower, the part 
behind the postorbital processes specially lengthened ; frontals 
smooth and swollen ; palatal foramina very long; palate ending 
opposite to middle of m.?; mesopterygoid fossa long, the distance 
from the hinder edge of the palate to the concavity below the 
hamular processes 22 mm.; front edge of presphenoid 6 mm. 
behind the palatal edge. Penis-bone of the usual canine canoe- 
shape, but much less strongly keeled than in the one example 
of C. vetulus available. 

Teeth of the general type of those of C. vetulus; p.* short and 
rounded ; a supplementary and probably abnormal m.’* present on 
each side, longer on the left than on the right. 

Dimensions of the type, measured in the flesh :— 

Head and body 650 mm.; tail 310; hind foot (s. u.) 130; 
ear 76. 


236 MR. OLDFIELD THOMAS ON THE MAMMALS [ Nov. 3, 


od 


Skull—greatest length 122 mm. ; basal length 112; zygomatic 
breadth 63:5; nasals, length mesially 38; interorbital breadth 19 ; 
tip to tip of postorbital processes 26; intertemporal breadth 17:8 ; 
breadth of brain-case 43°5; palatal foramina (singly) 8 x 2:5; 
palate length 56; length of os penis 47. 

Teeth—p.', length on outer border 8:2; m.' 8x95; m. 64x 
8:6; combined length of m.’ and m.’14°5. P.,, length 64; m., 10, 
m., 6°7, m., 4°4. 

This Dog differs from C. vetulus, to which it is no doubt allied, 
in its darker and greyer general colour, its blackish feet, and 
larger skull. The possession by the only specimen of a third pair 
of upper molars is probably an individual abnormality, but oceur- 
ring here in one of the most primitive types of Canide it presents 
a very interesting and suggestive case of atavism. For this 
reason alone further examples of this animal will be welcomed 
with interest, to see how frequently the extra teeth are present. 

I have named this distinct animal in honour of the late 
Mr. Percy Sladen, whose valuable zoological work is known to all 
naturalists, whose death was lamented by a wide circle of friends, 
and whose widow has chosen the most useful and suitable way of 
perpetuating his memory by bearing the expense of Mr. Robert’s 
expedition to Central Brazil. 


13. Canis vetuLUs Lund. 

3. 1108. 

©, IIists). 

Quite alike externally, these two specimens differ surprisingly 
in the size of their molars, the female having the larger. Further 
specimens will be needed before this difference can be explained, 
but it may be noted that the larger-toothed specimen agrees 
closely with typical C. vetulus, whilst that with the smaller teeth 
has an equal resemblance to the wrongly localized type of Gray’s 
“ Lycalopex fulvicaudus, var. 1. chiloensis.” * 


14. SprotHos venaticus Lund. 

Org NAT 

The Bush-dog is seldom captured by collectors, and this excel- 
lent specimen is a valuable accession to the Museum. 

15. GALERA BARBARA L. 

g. 1064, 1117. 

2. 1065. 

16. Nasua nasua L. | 


3d. 1043, 1054, 1056, 1057, 1069, 1087, 1164. 
2 LOSS, LOMO) LOM W129) 1136, 13 7e le IO ae: 
1157, 1163. 


Among this fine series of eighteen Coatis it might be said that 


* P.Z.S. 1868, p. 511; Cat. Carn. B. M. p. 198 (1869). 


1903. OF THE PERCY SLADEN EXPEDITION, 237 


no two can be found that are exactly alike, the set ranging in 
coleur from uniform red to nearly uniform black, and varying as 
much in details as in general tone. The skulls also vary very 
considerably both in size and in the development of their bulle, 
the extremes appearing at first sight to be quite different species, 
but being connected together by every gradation. 


17. ScruRus LANGSDORFFI Brandt. 


3. 1088, 1154, 1177, 1182, 1195. 
©. 1081, 1124, 1155. 


A valuable series representing the true S. langsdorffi, which 
name has been applied by various authors to many different 
members of the group. 


18. RHIPIDOMYS ROBERTI, sp. n. 
6. 1023, 1140. 


A Rhipidomys intermediate in size of skull between the large 
species of the &. macrurus-latimanus group, and the small ones 
allied to R. pheotis ; most nearly related to the latter. 

Fur of medium length ; hairs of back about 7-8 mm. in length. 
General colour abeve dull fulvous or “ clay-colour,” finely lined 
with black. Sides clearer and more buffy, bordered below by a 
clear ochraceous-buff line edging the pure sharply-defined white of 
the belly. Face like back, rather less heavily lined. Cheeks 
buffy, like sides. Eyes without darker rings. Hars of medium 
length, brown, not strongly contrasting with the general colour. 
Outer side ef arms and legs like flanks, ner pure white on the 
fore limb down to the wrists, but on the hind limb only halfway 
down to lower leg, the fulvous passing round the ankles; hands 
and feet dull whitish. Tail fairly long, rather less hairy than 
usual in this genus, its basal inch furry all round and coloured 
like the bedy ; the remainder uniformly brown. 

Skull shaped most like that of A. benevoleus, although much 
larger. Supraorbital ridges fairly well developed, evenly divergent 
backwards. Outer plate of anteorbital foramen scarcely developed 
anteriorly. Palatal foramina of medium length, narrow and 
pointed anteriorly, broadly open posteriorly ; not reaching back 
to the level of m.' Interpterygoid fossa of normal shape, with 
rounded anterior border. 

Dimensions of the type, measured in flesh :— 

Head and body 110 mm.; tail 145; hind foot (s. u.) 25, 
(c. u.) 26-7; ear 16. 

Skull—greatest length 32 mm.; basilar length 24°5 ; greatest 
breadth 16; length of nasals 11; interorbital breadth 5:6; breadth 
of brain-case 14; length of palate 13°6; diastema 8 ; palatal 
foramina 5 x 3°1; length of upper molar series 4°8. 

Type. Adult $. B.M. No. 3.7.7.67. Original number 1023. 
Collected 6 July, 1902. 

Although Mr. Robert’s trunk measurement of the type of this 
species is slightly less than that given by Mr. Simons for R. pheotis, 


238 MR. OLDFIELD THOMAS ON THE MAMMALS [ Nov. 3, 


its skull is very markedly larger, and its teeth even more so. 
That is, however, the species with which it seems most nearly 
allied. 

The presence of Rhipidomys robert, like that of MNeacomys, 
gives evidence of the affinity of the Cuyaba region of Matto Grosso 
with the eastern slopes of the Andes, from which all the previous 
members of this group have hitherto come. 

In naming this distinct species after Mr. Robert I wish to make 
very special acknowledgment of the admirable work he has been 
doing in South America during the last three years. His collec- 
tions, perfectly prepared in spite of all the difficulties incidental 
to a damp tropical climate and uncivilised surroundings, have 
revolutionised the material available in the British Museum for 
the study of the Mammals of Southern Brazil. Numbers of 
species, previously known to us only by the original descriptions 
or by a few unmeasured, discoloured, or otherwise imperfect 
specimens, are now represented by series of perfect skins with all 
the details incidental to good modern work. And again, the 
present collection, formed in a still more difficult country, links up 
in the centre his own Eastern Brazilian material with that 
obtained along the edges of the Andean chain by the late 
Mr. Perry O. Simons. 


19. NECTOMYS SQUAMIPES MATTENSIS, subsp. n. 


g. 1005, 1019, 1025, 1037, 1050, 1203. 
2. 1031, 1033, 1048, 1109, 1131. 


External characters as in true squamipes and in garleppi ; 
the latter also appears to grade into squamipes. 

Skull rather shorter and more rounded, less slender and 
elongated than in true sqguamipes. Nasals as usual narrow and 
elongate. Supraorbital edges with a marked raised bead, more 
developed than in squamipes, much more than in garleppi. Inter- 
parietal small, narrow antero-posteriorly, its anterior edge 
generally directly transverse. Palatal foramina long, widely 
open, with a slight angular constriction at the junction of their 
anterior and middle thirds. Opening of posterior nares wide, 
the edge of the palate squarely transverse, the median pterygoid 
fossa broader anteriorly than posteriorly, its breadth in front 
greater than the length of m.’ Lateral pterygoid fosse propor- 
tionally narrow and sharply pointedanteriorly. (In WV. squwamipes 
the median fossa is narrow anteriorly, broadening backwards, 
with a rounded anterior border.) 

Dimensions of the type, measured in the flesh :— 

Head and body 200 mm.; tail 200; hind foot (s. u.) 47 (extremes 
44_47), (c.u.) 50; ear 24. 

Skull—greatest length 43 mm.; basilar length 34°6; zygomatic 
breadth 22°2; interorbital breadth 7; interparietal 3-4 x 11-5; 
palate length 20°4; palatal foramina 7:5x3:2; breadth of 
posterior palatal fossa anteriorly 3°8; length of upper molar 
series 6°3. 


1903. ] OF THE PERCY SLADEN EXPEDITION. 239 


Type. Adult male. B.M. No.3.7.7.71. Original number 1050. 
Collected 30 July, 1902. 

The difference in the structure of the pterygoid fosse, perfectly 
uniform throughout the series, seems to make it necessary to 
give a special name to the Matto Grosso Water-rat; but as lower 
down the same river, in Paraguay, specimens agreeing with the 
true WV. sguanupes occur, I only distinguish it as a subspecies. 

It may be noted as a curiosity that a very large proportion of 
the skulls of WVectomys, even when quite old, have their median 
interparietal suture open, a characteristic very rare in mammals 
generally. 


20. ORYZOMYS LATICEPS Lund. 


3g. 1003, 1007, 1008, 1020, 1052, 1053, 1062. 

@. 1004, 1006, 1010, 1016, 1026, 1049. 

These specimens appear to be certainly identical with those 
obtained by Mr. Robert on the Rio Jordaéo*, 8. Minas Geraes, 
not far from the typical locality of the species. 


21. NEACOMYS SPINOSUS AMGNUS, subsp. n. 


g. 1061, 1077, 1188. 
©. 1036, 1051. 


Size averaging slightly larger than in the true Peruvian 
NV. spinosus, and tail rather longer. General characters very 
much as in that animal, in spite of the great distance between 
the localities. Fur shorter and thinner than in spinosus, especially 
on the under surface. Colour above bright ochraceous, liberally 
lined with black along the back, but the dorsal area nevertheless 
not nearly so dark as in spinosus. Sides clearer fulvous, the 
yellowish line along their lower edge not so sharply defined as in 
spinosus. Belly and inner sides of limbs white, the hairs white 
to their roots; line of demarcation not very sharply defined. 
Face rather greyer than body. Ears uniformly brown. Arms 
and legs like sides, or rather duller; hands and feet dull whitish. 

Skull as usual varying a good deal in size and shape, but in a 
general way that of subsp. amenus is a little larger than 
that of spinosus, with more rounded brain-case, less developed 
supraorbital ridges, and slightly larger palatal foramina, but 
none of these points can be regarded as really distinctive. 

Dimensions of the type, measured in the flesh :— 

Head and body 90 mm. (extremes 80-90); tail 102 (102-112); 
hind foot (s. u.) 23 (22-24), (c. u.) 24:5; ear 16 (14-16). 

Skull—greatest length 24-6 mm.; basilar length 18-1; zygomatic 
breadth 12°7; interorbital breadth 4:3; palatal foramina 4; 
length of upper molar series 3:1. 

Type. Old male. B.M. No. 3.7.7.84. Original number 1077. 
Collected 11 August, 1902. 

The occurrence of a WNeacomys in this region was quite 


* Of. Ann. Mag. N. H. (7) viii. p. 530 (1901). 


240 MR. OLDFIELD THOMAS ON THE MAMMALS [ Nov. 3, 


unexpected, as also is the close resemblance it bears to the typical 
N. spinosus of Peru, from which it differs chiefly by its brighter, 
clearer colour. 


22. PROECHIMYS LONGICAUDATUS Rengg. 

3. 1011, 1024, 1027, 1034, 1046, 1047, 1114, 1190, 1197, 
1201, 1204. 

2. 1009, 1021, 1035, 1078, 1113, 1185, 1187, 1194. 

Rengger’s type was obtained on the 21st parallel of latitude, 
therefore not far south of Corumba. It would be interesting to 
get absolute topotypes of it, but the Chapada series no doubt 
belongs to the same species. It is nearly allied to my P. boli- 
vianus, which differs from it by the cranial characteristics given 
in the description of the latter. 


93. CoENDOU BRANDTI Jent. 


3. 1075, 1167, 1183. 

These specimens may be considered to represent typically 
Dr. Jentink’s species, for Brandt’s figures, on which the name 
was founded, were from specimens in the Langsdorff collection, 
and as Sciurus langsdorfi came from Matto Grosso, the Porcupine 
may very likely have done so as well. The skull exactly agrees 
with these original figures. 

C. brandti is certainly very closely allied to Gray’s C. boliviensis, 
but the skull differs in certain details, as to the constancy or 
otherwise of which no opinion can be expressed until a much 
larger series has been examined. 


24, CoENDOU CENTRALIS, sp. n. 

g 1147. 22 September, 1902. Type. (B.M. No. 3.7.7.102.) 

A member of the group for which Dr. Jentink has used the 
name of prehensilis. The skull less swollen than in C. brandi. 

Size about the same as in C. brandti; relative development of 
spines and hair as in that species. General colour rather darker, 
owing to the greater extension of the black on the spines, but 
there is no very material difference in this respect. Belly 
browner. ‘Tail decidedly darker, especially along the middle line 
below. 

Skull of the same less swollen type as is figured by Cuvier * 
and considered by Jentink to be prehensilis. Nasals comparatively 
long and narrow, parallel-sided. Palatal foramina very small as 
compared with those of a Rio Jordaéo (Sao Paulo) specimen of 
C. prehensilis. Teeth smaller than those of C. brandtt. 

Dimensions of the type, measured in the flesh :— 

Head and body 480 mm.; tail 530; hind foot (s. u.) 80, (c. u.) 
90, from back of hallucal climbing pad (s. u.) 66. 

Skull—greatest length 94 mm.; basilar length 82; greatest 
breadth 53; nasals 32 x 25; least interorbital breadth 35; greatest 


* Mém. Mus. ix. vl. 20, fig, 3 (1822). 


1903.] OF THE PERCY SLADEN EXPEDITION. 241 


breadth across postorbital projections 46; greatest height from 
palate 45; palate length 42; diastema 26:2; palatal foramina 
5°4x 4:2; length of upper molar series 19-6. 

Type. No. 1147 as above. Killed 22 September, 1902. 

It seems curious that two Porcupines of the prehensilis group 
should occur at Chapada, but, as pointed out by Dr. Jentink, the 
skull-differences between brandti and the less swollen headed 
animal are too great to be due to individual variation. hat 
C. centralis is not the true C. prehensilis is shown by its differences 
from a specimen obtained in Sao Paulo, this latter locality being 
far closer to the region whence Marcgrave described his “‘Cuandu,” 
the original prototype of Linneeus’s C’. prehensilis. 


25. Dasyprocta AZAR Licht. 


3. 1209. 


I can find no tangible difference between examples of this 
wide-ranging form from Chapada, from Paraguay, and from Sao 
Paulo. But in each locality the specimens differ considerably in 
colour inter se. 

I may take this opportunity to point out how very different 
from all other Agoutis the long-tailed form, D. acouchy, is, and 
to suggest that it ought to form a special genus. This might 
be called Myoprocta ; and its main distinguishing characters would 
be the well-developed tail and conspicuously smaller teeth. 


26. AcoutTiI PAcA L. 
SE LOG OMS. 


27, SYLVILAGUS MINENSIS CHAPAD#, subsp. n. 


g. 1059, 1076, 1107, 1119, 1121, 1122, 1123, 1168, 1206. 
2. 1042, 1066, 1153, 1165, 1173. 


Closely similar in all characters to the true S. minensis of 
Southern Minas Geraes, but the colour throughout perceptibly 
lighter. The back, which has the even lining of S. minensis and 
not the coarse mottling of S. paraguensis, is several shades lighter 
in tone, the sides and rump are a light greyish, and the cheeks 
are less blackened. Terminal half of eara lighter brown. Nape- 
patch nearer “ochraceous buff,’ though duller, while that of 
minensis approaches ‘“ tawny ” or “‘ tawny ochraceous.” 

In other respects, in size, proportions, length of ears, and 
characters of skull, I can find no difference between the Chapada 
Hare and the typical S. minensis obtained by Mr. Robert on the 
Rio Jordao. 

Dimensions of the type, measured in the flesh :— 

Head and body 355 mm.: hind foot (s.u.) 76 ; ear-opening 60. 

Skull—greatest length 74 mm. ; basilar length 59. 

Type. Female. B.M. No. 3.7.7.116. Original number 1066. 
Collected 4 August, 1902. 

This is evidently a pale form of the Hare which is found over 
most of Southern Brazil, the true brasiliensis being, as shown in 

Proc. Zoou. Soo.—1903, Vou. Il. No. XVI. 16 


242 MR. OLDFIELD THOMAS ON THE MAMMALS [ Nov. 3, 


1901 *, a smaller animal occurring near Rio Janeiro. It is 
possible that the more coarsely mottled S. paraguensis will also 
prove to grade into the present form. 

Baby specimens of minensis, chapade, and paraguensis are as 
readily distinguishable from each other as are the adults. 


28. Tayassu ALBIROsTRIS Ill. 
9. 1144, with two feetuses 1145 and 1146. 


99. Tayassu TAJACU L. 


3. 1120, 1126, 1169. 
@ | OA, 


30. MazAMA RUFA F. Cuv. 


65 NOS). 
Yg. go. 1044. 


31. MyRMECOPHAGA TRIDACTYLA Linn. 


3. 1128, 1180. 

OPS HAE TTA), 

The reasons for using the above name are given in ‘ American 
Naturalist,’ xxxv. p. 143 (1901). 


32. TAMANDUA TETRADACTYLA Linn. 


g. 1038, 1103, 1106, 1112, 1125, 1149. 
2. 1017, 1058, 1084, 1181. 


33. Dasypus GiILvIPEs Licht. 


Tatow poyou Azara, Quadr. Paraguay, il. p. 142 (1801). 
Dasypus gilvipes Il. Abh. Ak. Berl. 1811, p. 108 (pub. 1815) 
(nomen nudum). 


Dasypus gilvipes Licht. Abh. Ak. Berl. 1815, p. 215 (pub. 
18). 


Dasypus encoubert Desm. Mamm. 11. p. 370 (1822). 
Go Whats), JUG, UGS, 
©. 1072. 


Under the name of Dasypus sexcinctus L. both the North and 
South Brazilian forms of this group have commonly been con- 
founded, but since they differ conspicuously in size and are clearly 
different, it 1s necessary to consider which species should bear 
Linneus’s name. 

Acting on the general principle that the first reference to 
Linneus’s own works should be taken as a guide in identifying 
his species, we get in this case a reference to the “Mus. Ad. Fr.” 
(p. 7), and the specimen mentioned in that work would be the 
type. That specimen is still in the Stockholm Museum, and Prof. 
F. A. Smitt has been kind enough to inform me that its cephalic 
shield is 77 mm. in length. This corresponds closely with the 


* Ann. Mag. N. H. (7) viii. p.635. 


1903. ] OF THE PERCY SLADEN EXPEDITION. 243 


size of examples from Para, while those from South Brazil and 
Matto Grosso agree with Paraguayan specimens in having this 
dimension about 105 mm. JD. sexcinctus will therefore stand for 
the N. Brazilian form, and D. gilvipes, based on Azara’s Tatou 


poyou, for the large southern one. 


34, Tatu NOVEMCINCTUS L. 


3. 1032, 1040, 1079, 1102, 1133, 1148. 
@. 1039, 1202. 


35, TATU MEGALOLEPIS Cope. 


@. 1115. 

As a topotype of Cope’s species, hitherto unrepresented in the 
Museum collection, this Armadillo forms a valuable accession. 
The species is perhaps not so widely separated from 7’. septem- 
cinctus as its describer supposed, for this specimen has 45-47 
scales in the movable bands, and the Museum contains examples 
of septemcinctus from Rio Grande do Sul with only 54, so that 
the gap between the two, stated by Cope to be from 43) to) 5,38 
thereby considerably reduced. 

In size 7. megalolepis 1s decidedly smaller than the older 
known species, the skull of Mr. Robert’s example being only 
63 mm. in length. 


36. DIDELPHIS PARAGUAYENSIS Oken. 


3. 998. 

@. 1207. 

Practically topotypes of Wagner’s D. pecilotis, which was 
collected at Cuyaba. 


37. MARMOSA CONSTANTIA, Sp. 0. 

g. 1110. 27 August, 1902. Type. (B.M. No. Bollol ol 0a) 

Size rather larger than in I. cimerea. Fur close and fine, 
shorter and less woolly than in M. cinerea; hairs of back about 
9 mm. in length. General colour above paler than usual in this 
group, nearly matching Ridgway’s “ isabella,” or slightly greyer. 
On the sides the colour gradually becomes more buffy, passing 
gradually into the “ buff-yellow” of the under surface, the chin, 
chest, centre of belly, inner side of limbs, and inguinal region 
being all of this latter colour. On the sides of the belly the hairs 
are slaty at base, but on the rest of the under surface they are 
yellow to their roots. Face pale buffy grey ; black orbital rings 
present, but narrow, ill-defined, and not prolonged forwards on 
to the sides of the nose. Cheeks to base of ear dull bufty yellow. 
Ears large, naked, greyish brown. Outer side of fore and hind 
limbs dull buffy isabella, like the sides. Tail rather shorter than 
usual in this group, its basal furry portion about 13 inches in 
length, brownish isabella above, yellowish below; naked portion 
about half brown and half white, the junction of the two colours 


mottled and irregular. 
16* 


244 MESSRS. C. J. GAHAN AND G.J. ARROW ON THE __[ Noy. 3, 


Skull stout and strong, with broadly spread zygomatic arches, 
expanded nasals, well developed postorbital processes, and broad 
interorbital region. Palatal vacuitier about normal, opposite the 
first three molariform teeth. 

Middle upper premolar about equal to the posterior; middle 
lower one rather larger than that behind it. Molars large, as in 
M. cinerea, markedly larger than in M. regina. 

Dimensions of the type, an old male, measured in the flesh :-— 

Head and body 180 mm. ; tail 195 ; hind foot (s. u.) 27; ear 31. 

Skull—greatest length 45:8 mm.; basal length 4271; zygomatic 
breadth 26:6; nasals, length 20°7, greatest breadth 6:8, least 
breadth 3-2; interorbital breadth 8:2; breadth across postorbital 
processes 1071; breadth of brain-case 16; palate length 25; 
length of three anterior molariform teeth 7°8. 

Marmosa constantie is larger, paler, and has a shorter tail 
than MM. cinerea, its nearest ally, with which it shares the 
characteristic shape of skull and nasals. The Bogota J/. regina 
Thos. has a narrower skull, less expanded nasals, and unicolor 
tail. 

I have named this pretty species in honour of the donor of the 
present most valuable accession to the National Collection, in 
recognition of her enlightened method of commemorating her late 
husband’s memory. 


EXPLANATION OF PLATE XXVII. 


Canis sladent, sp. n., p. 230. 


3. List of the Coleoptera collected by Mr. A. Robert at 
Chapada, Matto Grosso (Percy Sladen Expedition to 
Central Brazil). By C. J. Ganan, M.A., and G. J. 


ARROW. 
[Received August 18, 1903.] 


(Plate XX VIII.*) 


In this collection the Lamellicornia, Longicornia, and Phyto- 
phaga are especially well represented ; but, since our knowledge of 
the coleopterous fauna of Brazil is mainly confined to more 
northern regions, the whole collection, which contains specimens 
of 175 species, forms a valuable contribution to the materials avail- 
able for the study of the distribution of South American insects. 
New species of Lamellicornia, Rhynchophora, Heteromera, Longi- 
cornia, and Phytophaga are described, and there are in addition 
others which, for want of sufficient specimens or other reasons, 
it is not advisable at present to describe. The earlier groups, as 
far as and including the Rhynchophora, have been dealt with by 
Mr. Arrow, while for the remainder Mr. Gahan is responsible. 


* For explanation of the Plate, see p. 258. 


PZS.1903,vol I. PLAXVII 


André & Sleigh? 


COLEOPTERA from CENTRAL BRAZIL. 


a 


o 

i 
aT 

in 
2 


1903. ] COLEOPTERA OF THE PERCY SLADEN EXPEDITION. 245 


All the Coleoptera were collected by Mr. Robert in the month 
of November at an altitude of 800 metres. 

In an Appendix a few new forms contained in a collection 
from Paraguay, simultaneously received from Mr. W. Foster, have 
been described. 

CICINDELIDE. 
Odontochila discrepans Horn. 
Prepusa distigma De}. 
CARABIDA. 
Brachygnathus sp. near imperialis Chaud. 


GYRINIDA. 
Enhydrus tibialis Rég. 
STAPHYLINIDZE. 
Staphylinus sp. 
HIsSTERIDE. 
Homalodes brasilianus Mars. 


NIvTIDULIDe. 

Colastus latus Murray.—Murray described this species from 
Mexican specimens, and gave the name C’. tonsws to an indi- 
vidual from Brazil, but I am unable to distinguish them and 

_ believe all will be found to represent a single widely-distributed 
species. 
DERMESTID&. 


Dermestes vulpinus ¥. 


ByRRID&. 
Chelonarium ornatum Perty. 


ERoryLip&. 
Morphordes sp. 


CoccrNnELLID&. 
LEpilachna cacica Guer. 
Poria sp. 
PASSALID. 
Neleus punctiger Perch. 


CopRID2. 


Ontherus suleator F. 

O. sp. 

Canthidium sladeni, sp. n. 
Canthon histrio Lep. & Serv. 

CO. chalybeus Blanch. 

C. edentulus Har. 

C’. sp. 

Deltochilum fuscocupreun Bates, 


MESSRS. C. J. GAHAN AND G.J. ARROW ON THE  [Nov. 3, 


Deltochilum cupricolle Luc. 
Pinotus roberti, sp. i. 
P. fissus Har. 

P. nisus Oliv. 

P. carbonarius Mann. 
P. subeneus Lap. 
Phaneus mimas Linn. 
P. palemo Lap. 

P. paleno Blanch. 

P. kirbyi Vigors. 
Gromphas inernis Har. 


MELOLONTHIDE. 
Astena 2 spp. 
Ceraspis cornuta Blanch. 
Lsonychus griseus Mann. (2). 
Clavipalpus tenwis, sp. n. 
Liogenys sp. 
Lachnosterna fulvipennis Blanch. 


RUTELIDE. 

Anomala 2 spp. 

Macraspis pantochloris Blanch. 

Pelidnota sumptuosa Vigors.—A series of this species 
was obtained showing nearly every shade of metallic 
colouring. 

Pelidnota ceruginosa Linn. 

Leucothyreus pruinosus Perty. 


L. sp. 

DYNASTIDZ. 
Cyclocephala 14-punctata Mann. 
C. sp. 


C. ovulum Bates. 
Bothynus ascanius Kirby. 
B. sp. 

CETONIID. 
Gymnetis liturata Oliv. 
G. sp. 
Kuphoria lurida Fab. 


CURCULIONID&. 
Naupactus perpastus Schon. 
AN, B 0) 
Pantomorus sp. 
Hoplopactus lateralis, sp. n. 
Platyomus transversesignatus Schén. 
P. argyreus L. 
Rhigus latruncularius Perty. 
Homalonotus validus Oliv. 
Hilipus nevulus Mann. 


1903.] | COLEOPTERA OF THE PERCY SLADEN EXPEDITION. 247 


Hilipus validus Pase. 

HT, sp. 

Rhinochenus reichei Boh. 
f. stigma L., var. (2). 

&. 2 spp. 
Cryptorhynchus 3 spp. 
Celosternus stwrio, sp. n. 
Centrinus 4 spp. 
Copturus sp. 

Belopeus niger, sp. n. 
Rhynchophorus palmarum L. 


HLATERID 2, 
Chalcolepidius limbatus Esch. 
Pyrophorus noctilucus Linn. 
P. ganus Herbst. 
Psp. 
Monocrepidius sp. 


LAMPYRIDA. 
Lucidota sp. 


BostRYCHID&. 
Bostrychopsis uncinata Germ. 


TENEBRIONIDE. 
Epitragus sp. 
Nyctobates variolosa Kab. 
NV. gigas Linn. 
Zophobas opacus Sahlb. 
Calymmus cucullatus Pasc. 
Scotinus (?) sp. 
Uloma retusa Fab. 
U. sp. 
Camaria levipennis (De}j.). 
Strongylium azureum Gervo. 
S. sp. 
Hoploptera sp. 

-CISTELIDE. 


Prostenus cyanipes Lucas. 

P. sladeni, sp. n. 

Lystronychus ceruleus Sol., var. 
L. latipennis Maklin. 


MeEoip 2. 


Epicauta atomaria Germ. 
L. strigata Gyll. 

Hi. sp. 

ff. sp. 

Lf. sp. 

Tetraonyx sexguttatus Oliv. 


48 


MESSRS. C. J. GAHAN AND G. J. ARROW ON THE 


PRIONIDS. 


Mallodon spinibarbe Linn. 
Anacanthus costatus Serv. 


CERAMBYCIDA. 
Achryson surinamum Linn. 


Hammaticherus lacordairet Gahan. 


Butherium corvinum Germ. 
NXestia suturalis Perty. 

X. polita Waterh. 
Spherion sladent, sp. 1. 
Trichophorus electus, sp. 1. 
Octoplon flavopictum Perty. 
Callichroma suturale Fab. 
Lophonocerus barbicornis Linn. 
Dorcacerus barbatus Fab. 
Oxymerus basalis Dalm. 
Rhachidion obesum Newm. 
Megaderus stigma Linn. 


LAMIUID. 
Hypsioma sp. 
H. fasciata Thoms. 
Oncideres sladent, sp. 0. 
Steirastoma stellio Pasc., var. 
Acanthoderes lateralis Bates, var. 
(dopeza pogonocheroides Serv. 


BRUCHID, 
Caryoborus sp. 


CLYTHRIDZ. 
Luryscopa proxima Lac. 
Megalostomis obesa Lac. 
Tellena varians Lac. 
Urodera familiaris Lac. 


CHLAMYDIDA. 


Chlamys hieroglyphica Koll. 
C. smaragdina Klug. 

C. cistella Germ. 
Poropleura cuprea Klug. 


CRYPTOCEPHALID. 
Griburius pretextatus Suftr. 
Metallactus kollara Suffr., var. 
M. sp. 
LAMPROSOMIDA. 


Lamprosoma chameleon Lac. 
L. viride Lac. 


[Nov. 3, 


1903.] COLEOPTERA OF THE PERCY SLADEN EXPEDITION. 249 


EuMmoLrPip&. 
Lamprospherus thoracicus Baly. 
L. scintillaris Baly. 

Iphimeis fulvipes Baly. 

Agbalus sp. 

Colaspis 14-costata Lefév. 

C. flavipes Oliv., var. 

C. sp. 

C. sp. 

Chalcophana viridipennis Germ. 
ELumolpus surinamensis Linn. 
LE. clavipalpus Chap. 


CHRYSOMELID. 


Doryphora flavozonata Blanch., var. 
Deuterocampta 12-maculata Stal. 

D. musicalis Stal. 

D. sladene, sp. n. 


HALtTIcIDz. 


Orimissa piceicollis Jac. 
Gdionychis opima Germ. 
@. nigropunctata Clark. 
@. 12-notata Clark. 


GALERUCIDE. 
Galerucella sp. 


HIspip2. 


Alurnus thoracicus Perty. 
A, nigripes, Guér. 


CassIDIDA. 
Tauroma sp. 
Dolichotoma strigata Hoff. 
Batonota gregaria Bohem. 
B. ensifer Bohem. 


CANTHIDIUM SLADENI, sp.n. (Plate XXVIII. fig. 1.) 


C. decorato valde affine, sed plerwmque paullo minor, profundius 
sculpturatum aliterque coloratum. Susco-viride, capite pro- 
thoraceque lete viridibus, elytrorum dimidio basali rufo-cupreo ; 
capite sat fortiter punctato, medio leviter tuberculato, clypeo 
bidentato ; prothorace crebre punctato, lateraliter fere regu- 
lariter arcuato, postice medio angulato, subtiliter marginato ; 
elytris profunde punctato-striatis, interstitiis conveais ; pygidio 
minute punctato. 

Long. 7-8'5 mm. 

The elytral interstices of this insect are more convex than in 

any other species of the large genus Canthidiwm known to me. 


250 MESSRS. C. J. GAHAN AND G.J. ARROW ON THE _ [ Nov. 3, 


It can only be compared with C. decoratwm Perty, these two 
being the only forms with parti-coloured elytra, at least in the 
normal condition. In @. decoratwm, however, the elytra are not 
coppery, and the posterior half is blue, while in the new species 
it is green and is bounded anteriorly by a backwardly-curved 
line. In C. decoratwm the line of demarcation is angular, and 
its projection is in the opposite direction, so that it almost 
touches the base of the elytra at the suture. This line is less 
distinct in C. sladeni, owing to the deeper coppery-red colour of 
the anterior half of the elytra and their more deeply-channelled 
surface. The new species appears to be slightly smaller on the 
average than C. decoratwm, judging from the series of five speci- 
mens of each in the British Museum. 


PINOTUS ROBERTI, sp. n. 


P. carolino atqgue eremite valde affinis, sed plerumque paullo 
major ; crassus, niger, postice plagis griseis decoratus ; capite 
utriusque sexus ut im P. eremita carinato, maris carine 
angulis paullo magis productis, thoracis lobo mediano minus 
quadrato, leviter sulcato, sulci fundo absque stria, twbherculo 
laterali minuto acuto, haud cariniforn. 

Long. 27-32 num. 

fab. Brazil: Chapada, Para. 

This is one of the small group of species in which the elytral 
strie are posteriorly dilated and filled with a greyish earthy 
secretion. It is very closely related to P. eremita Har., of 
Venezuela and Colombia, but the average size is rather larger 
and the thorax of the male shows slight differences of form. The 
median lobe is more rounded, its lateral angles being hardly 
traceable, and the longitudinal depression down its centre has no 
definite stria at the bottom. Above the lateral fossa of the 
thorax there is a short acute tubercle, which is represented in 
P. eremita by an elongate crest. I am unable to compare the 
female of P. roberti with that of P. eremita, the five specimens 
of the latter in the British Museum being all males; but the 
sexes of the new species differ extremely little. The head of the 
female is more elongate and pointed, and there is a broad crest 
very similar to that of the male but placed farther back. The 
centre part of the prothorax is only slightly less prominent. 
There is generally a trace of a median stria, and the tubercles 
above the lateral fossee are not acute. The elytral furrows which 
contain the remarkable earthy deposit are larger in the female 
than in the male in this and the other species characterised by 
this peculiarity. 

Mr. Robert found nine specimens of this species, among which 
one only isa male. A second male from Para was confused with 
our series of P. eremita, so that the species is a fairly widely 
distributed one. Its very close similarity to the more northern 


form is no doubt the reason for its having hitherto escaped 
detection. 


1903.] COLEOPTERA OF THE PERCY SLADEN EXPEDITION. 251 


CLAVIPALPUS TENUIS, sp.n. (Plate XXVIII. fig. 2.) 


Hlongatus, convexus, rufo-castuneus ; capite parvo, crebre punc- 
tato, clypeo parabolico, margine leviter reflexo ; prothorace 
polito, profunde wregulariter punctato, lateribus valde angu- 
latis, angulis anticis et posticis obtusis; scutello utrinque 
leviter punctato; elytris haud regulariter striato-punctatis, 
interstitiis minutius haud crebre punctulatis, foemine inter- 
stitio secundo lato grosse punctato-rugoso ; pygidio convexo, 
crebre punctato, linea media levi; corpore subtus lateribusque 
fulvo-pilosis, segmento abdominali 5° maximo; antennis 
10-articulatis, maris clava longitudine ad stipitem equali ; 
tibiis anticis 3-dentatis, unguibus omnibus fissis, maris tarsis 
antics et posticis quam tibiis plus duplo longioribus. 

Long. 22-13 mm. . 

This is a more slender insect than any other species of Clavi- 
palpus known to me, but conforms in all essential particulars to 
the generic type. It is smooth and shining on the upper surface, 
with rows of deep but rather irregular punctures upon the elytra. 
The head is small and the clypeus regularly rounded, the pro- 
thorax very convex and much narrowed in front and behind, and 
the elytra long and of approximately equal width throughout 
their length. The antenne are ten-jointed, but the articulation 
between the 5th and 6th joints is less distinct in the female than 
in the male. The former sex appears in this genus to be less 
easily found than the male, and I believe has not hitherto been 
described. The enlargement of the terminal joint of the maxillary 
palpus, from which the genus has received its name, occurs in the 
male only, but in the present species this joint is of normal size 
in both sexes. In all the species the 5th ventral segment is very 
large in the female, and the distinction drawn by Lacordaire 
between the Macrodactylides and Clavipalpides (viz. the absence 
of this enlargement in the latter) is therefore misleading. The 
segment is enlarged in both sexes, but especially in the female. 


HOoPLOPACTUS LATERALIS, sp.n. (Plate XXVIII. fig. 3.) 


ELlongato-ovatus, fuscus, squamis griseis vel sulphureo-cinereis, 
equaliter sed haud dense obtectus, margine laterali sparse 
viridi-squamoso, squamis marginem intus attingentibus densius 
aggregatis, viridi-argenteis, corporis subtus squamis viridibus 
et roseo-argenieis, abdomine medio denudato; capite lato, 
rostro breve, antice vie angustato; prothorace transverso, 
lateribus equaliter arcuatis ; elytris punctato-striatis, margine 
obscuriore wregulart, paullo ante apicem subito dilatato ; 
pedibus rufo-fuscis, griseo-setosis, squamis pallide viridibus 
interspersis, femoribus magis fuscis, anticis intus spino acuto 
post medium armatis. 

Long. (rostro excl.) 8-9 mm. 


This is the sixth species so far assigned to the genus, and is 
easily recognisable by the dark external margin of the elytra 


252 MESSRS. C. J. GAHAN AND G. J. ARROW ON THE [ Nov. 3, 


expanding just before the apex, and clearly defined by the massing 
together and brighter coloration of the scales bordering it 
internally. The head and thorax are rather broader than in 
H. injucundus Schén., the rostrum shorter and the thorax as 
wide in front as behind. 

A dozen specimens were collected. 


CG@LOSTERNUS STURIO, sp. n. (Plate XXVIII. fig. 4.) 


Crassus, niger, argillaceo-squamutlosus, subtus squamulis pallidi- 
oribus vestitus, prothoracis maculis magnis inferioribus late- 
ralibus maculaque parva dorsali albo-flavis, hac sagittiformi 
ad marginis postici medium posita ; elytris nodosis, squamulis 
argillaceis aggregatis, squamas magnas quasi-piscinas for- 
mantibus, videlicet suturalibus 5, quarum 2 internalibus acute 
tuberculatis, lateralibusque 4, setis minutissimis albidis ubique 
rare sparsutis squamulisque albidis ad margines lateralia 
irregulariter aggregatis; capite postice profunde sulcato, 
utringue transverse carinato ; prothorace medio acute longi- 
tudinaliter carinato, antice valde constricto, postice lateribus 
valde arcuatis et crenatis, angulis obsoletis ; femoribus omnibus 
medio fortiter dentatis. 

Long. (rostro excl.) 10 mm. 


This curious Weevil is remarkable for the manner in which the 
earthy-coloured scales of the elytra are aggregated into large areas 
resembling the scales of a Ganoid fish, and separated by a very 
fine network of black scales. These areas are elevated in the 
centre, the four median ones adjoining the suture very strongly 
and acutely. The posterior part of the head is furrowed and 
transversely elevated on each side of the furrow, and the centre 
of the pronotum is longitudinally elevated. The only other 
species with which this can be compared is C. exornatus Boh., 
which is smaller, narrower, and more brightly coloured. In this 
species also the scales upon the elytra tend to form isolated 
patches separated by darker intervals. In C. sturio, however, 
the patches have spread over the entire surface and are divided 
only by extremely narrow lines. 

The unique type-specimen is probably a male. 


BELOPQUS NIGER, sp.n. (Plate XXVIII. fig. 5.) 


Ellipticus, niger, nitidus, elytris velutinis, rostro tenue, ad pro- 
thoracis et elytrorum longitudinem cequali ; prothorace nitido, 
haud dense punctato, absque sulcis, antice constricto, lateribus 
ab ante medium fere parallellis ; elytris opacis, velutinis, 
profunde striatis, interstitiis seriatim punctatis, singulo postice 
arcuate truncato; pygidio velutino et fulvo-setoso, grosse 
punctato ; corpore subtus polito, sierni lateribus abdominisque 
segmento ultimo punctatis. 

Long. (rostro eacl.) 10 mm. 

Hab. Brazil: Chapada, Para. 

In all structural features this insect is similar to the only 


1903.] | COLEOPTERA OF THE PERCY SLADEN EXPEDITION. 253 


hitherto described species of its genus, B. carmelitus Gyll. It is 
larger, however, and has a still more slender rostrum, which in a 
specimen brought by H. W. Bates from Para is as long as the 
prothorax and elytra together. In the Chapada specimen it is 
slightly shorter. This may be the other sex, but I think most 
probably both are females. The polished surface of this species, 
clothed only upon the elytra with a velvety tomentum, and the 
uniform black colour are quite sufficient to distinguish it. 


PROSTENUS SLADENI, sp. n. (Plate XXVIII. fig. 6.) 


Capite pronotoque viridi-cyaneis, capitis fronte inter antennas 
dense fortiterque, clypeo parcius, punctato ; prothorace quam 
latitudine basis vin breviore, ante mediwm latiore, supra 
omnino fortiter reticulatimque punctato ; elytris eneo-cupras- 
centibus, nitidis, subovatis, valde convexis, fortiter striato- 
punctatis, interstitiis inter series punctorum punctis minoribus 
piligeris, humeris utrisque tuberculo conico munitis ; corpore 
subtus pedibusque cyaneis, nitidis, his longis, femoribus modice 
clavatis subtiliter parce punctatis, tabs densius punctatis ; 
antennis medium elytrorum paullo superantibus, articulis 
quinque ultimis sat valde dilatatis, articulo 3° quam 4° paullo 
longiore. 

Long. 12, lat. 44 mm. 

Hab. Central Brazil. 

Head and prothorax bluish green above, elytra bronzy green 
and nitid. Head closely and rather strongly punctured between 
the antenne, the clypeus being as strongly but less closely 
punctured. Antenne reaching beyond the middle of the elytra, 
with the last four or five joints rather strongly dilated and the 
3rd joint perceptibly longer than the 4th. Pronotum widest a 
little before the middle, very slightly narrowed anteriorly, but 
distinctly narrowed towards the base, where its width is about 
equal to its length along the middle; its surface strongly punc- 
tured all over, the punctures, many of which are setigerous, 
forming a reticulation. EHlytra strongly punctured in rows, with 
smaller setigerous punctures on the intervals between the rows, 
each with a small conical tubercle or prominence at the shoulder. 


SPHERION SLADENI, sp. n. (Plate XXVIII. fig. 7.) 


2. Nigrum ; elytris eneo-viridescentibus pube grisea densa et 
setis rigidis sparsis obtectis; prothorace lateraliter obtuse 
tuberculato, disco utrinque leviter tuberculato, medio crebre 
punctulato, plaga parva postico-mediana polita ; elytris sat 
dense punctatis ; antennis quam corpore paullo longioribus, 
articulis 3°-6"™ apice via perspicue spinosis. 

Long. 14, lat. 33 mm. 

Hab. Central Brazil. 

Black, with the exception of the elytra which have a somewhat 

brassy-green tint; the whole rather densely covered with a fine 
greenish-grey pubescence, with here and there some erect fulvous 


254 MESSRS. C.J. GAHAN AND G.J. ARROW ON THE [ Nov. 3, 


sete. The prothorax is obtusely tubercled at the middle of each 
side, and has two feeble tubercles on each side of the disk; the 
latter is finely and very closely punctulate, but has a small smooth 
shining space in its median posterior part. The elytra are dis- 
tinctly but not very closely punctured, and each has a short 
exterior spine at the apex and the sutural angle obtuse. The 
acetabula of the middle coxee extend outwards to the epimera, the 
present species agreeing in this respect with S. eyanipenne Serv., 
the type of the genus. 


TRICHOPHORUS ELECTUS (Dej. MS.), sp. n. (Plate XXVIII. 
fig. 8.) 

Fusco-brunneus ; elytris, pedibus, abdomine et antennarwm 
articulo primo plus minusve rufescentibus ; pronoto lineis 
duabus interruptis flavo-pilosis ; scutello flavo ; elytris utrisque 
maculis tribus flavo-pilosis—prima ante medium prope suturam, 
secunda et tertia obliquiter positis inter medium et partem 
tertiam apicalem. 

Long. 15-23, lat. 4-6 mm. 

Hab. Central and South Brazil. 

This species greatly resembles 7’. interrogationis Blanch., but 
has no pubescent spots on the vertex of the head, and the two 
posterior spots on each elytron are quite separate from each other. 
It differs also in that the anterior part of the interrupted yellow 
or white line on each side of the pronotum lies farther back from 
the anterior margin, especially in the female, in which it forms a 
small round spot. The antenne of the female are a little longer 
than the body, those of the male very much longer. The species 
has for a long time been represented in the British Museum 
Collection under the MS. name I have adopted, and has also been 
known as 7’. 7-signatus Chev. MS. 


ONCIDERES SLADENI, sp.n. (Plate XXVIII. fig. 9.) 


O. fasciatee (Lucas) affinis, sed elytris haud fasciatis, prope basin 
densius granulato-punctatis, vittis nigris infra-ocularibus 
capitis latioribus. 

Long. 20-28, lat. 8-104 mm. 

Hab. Central and South Brazil: 8. Anna da Chapada, Rio 

Grande. 

This species is the Oncideres maculosa of Dejean’s Catalogue 
(nec Redtenbacher), and appears to be the Oncideres amputator of 
Thomson (nec Fab.). It belongs to the section of the genus in 
which the thorax is slightly wider behind than in front, and in 
which the frontal processes of the male are directed forwards and 
downwards. It is closely allied to O. fasciata Lucas, but easily 
distinguishable from it by the absence of fascie from the elytra, 
the latter being entirely covered with sparse fulvous spots with 
much smaller spots on the intervals between them. The basal 
margin of the elytra is fulvous, the scutellum grey ; the black 
bands on the head below the eyes are wider than in the allied 


1903.] | COLEOPTERA OF THE PERCY SLADEN EXPEDITION. 255 


species, each being almost equal in width to the eye itself. The 
front coxze of the male have a more or less obtuse tubercle at the 
distal end. 


DEUTEROCAMPTA SLADEN, sp.n. (Plate XXVIII. fig. 10.) 


Testacea; elytris nigro-fuscis, utrisque margine basalt et vittis 
sex (quarum prima brevissimc) aureo-flavis ; pronoto medio 
parce subtiliterque, ad latera fortiter densiusque, punctato ; 
articulo ultimo tarsorum apice subtus bidentato. 

Long. 8, lat. 5 mm. 

Hab. Central Brazil: S. Anna da Chapada. 

Brownish testaceous; elytra blackish brown, each with the 
basal margin and six longitudinal bands of a golden-yellow colour, 
the innermost band being very short and narrow; the 2nd and 6th 
bands unite at the apex, just in front of which they are jomed by 
the 5th; the 3rd and 4th meet a little way in front of the apex, 
and a short common band unites them with the 5th; distinct 
rows of punctures mark the boundaries between the alternating 
yellow and dark brown bands. Antenne with the first three or 
four joints testaceous, the rest black ; the last five being dilated to 
form a club. Pronotum sparsely and finely punctured in the 
middle, strongly and more closely at the sides, near which it is 
slightly depressed. 

Tn structural characters this species comes close to D. musicalis 
Stal, though easily distinguishable from it by the markings of the 
elytra, which are suggestive rather of the genus Zygogramma 
than of Deuterocampta. 


APPENDIX. 


Descriptions of « few new Species of Coleoptera from Sapucay, 
Paraguay. By Giusurr J. ARRow. 


The following species were contained in a small but very inter- 
esting collection of insects sent by Mr. W. Foster from Sapucay, 
in the (entomologically) little-known State of Paraguay. 


APTERODEMA PARAGUAYENSIS, Sp. Nn. 


Ovata, convexa, fusco-rufa, elytrorum lateribus maculaque 
suturali postice attenuata’ vage fuscioribus ; capite crebre 
punctato, clypeo magno, margine semicirculart, oculis parwes ; 
prothorace transverso, persprcue punciato, lateribus amntoce 
arcuatis, postice parallellis, angulis posticis fere acutis ; 
scutello parce punctato, lateribus valde curvatis ; elytris 
convexis, ubique irregulariter grosse punctatis, stria suturale 
aliisque indistinctis lateribus fortiter sed parce setosis, ab 
humeris ad post medium leviter divergentibus deinde regu- 
lariter arcuatis, angulis sutwralibus fere rectis; pygidio 
detecto, parce ac minute punctato ; corpore subtus fere nudo 
sed femoribus longe brumneo-setoso, segmentis abdominalibus 


256 MESSRS. C, J. GAHAN AND G.J. ARROW ON THE _[ Noy. 3, 


libris, equalibus, tibiis anticis brevibus, validissime bidentatis, 
tarsis omnibus longis, unguiculis integris ; alis imperfectis. 

Long. 10 mm. 

There are three specimens of this very anomalous little beetle, 
which seems to form a second species of M. Fairmaire’s genus 
Apterodema, although it is not altogether impossible that it may 
have no affinity with it, since the genus was constituted from a 
single specimen in such a condition, as the author says, that 
neither mouth nor antenne could be seen. Enough is said, how- 
ever, to show that in declaring its close relationship to Liogenys, 
one of the Macrodactylides, M. Fairmaire has entirely mistaken 
its systematic position, all the characteristic features of the 
Macrodactylides being absent. The specimens before me, which, 
like M. Fairmaire’s specimen, are females, represent a very 
isolated type, but can only be placed in the Sericoides. The 
elytra are not fused together as in Apterodema acuticollis Fairm., 
but the wings are partially atrophied. There are six free and 
equal abdominal segments. The labrum is free, subvertical and 
emarginate, and the labium consolidated, broad, and _ slightly 
concave in front. The antenne are nine-jointed and the claws 
simple. 

A. paraguayensis is rather larger than the Patagonian insect, 
of a dark red colour, with the margins of the elytra vaguely 
darker, the dark sutural margins forming a broad band at the 
scutellum, which they enclose, but narrowing behind. The pro- 
thorax is broad behind, and the posterior angles slightly produced 
as in A. acuticollis. The only clothing consists of stiff sete along 
the sides and upon the legs. The sculpture is quite different to 
that described by M. Fairmaire. 

I may advantageously take this opportunity of making an 
observation upon an allied South American genus. Lacordaire 
and Germain many years ago pointed out the probability that 
the genus Accia of Curtis, hitherto known only from a single 
female specimen in the British Museum, may prove to be in- 
separable from Maypa Blanchard. Having lately been able to 
examine a male, I can now definitely confirm this supposition. 
The genus should be called Sericoides, into which must be merged 
Accia, Maypa, and Macrosoma. 


MACRASPIS BRASILIENSIS, sp. n. 


Niger, via nitidus, supra ubique creberrime coriaceo-punciatus ; 
prothoracis lateribus subtiliter rugosis ; scutelli medio longi- 
tudinaliter leviter lineato-depresso ; pygidio ubique transverse 
strigoso ; abdominis medio vix punctato, nitido, processu 
mesosternalt recto, haud fortiter clavato; tibiis intermediis 
subtus fere rectis. Q elytrorum lateribus medio haud expla- 
natis. 

Long. 20-24 mm. 


Hab. Brazil: Ega, &e.; Paraguay: Sapucay. 


1903.] COLEOPTERA OF THE PERCY SLADEN EXPEDITION. 257 


Although, owing to the confusion which has prevailed as to 
the not easily distinguishable black species of Jd/acraspis, this 
species has remained without a name, it is a well-known Brazilian 
insect and evidently ranges over a large tract of country. The 
specimens from Kga, referred by Mr. Waterhouse to M. dichrous, 
belong to this species. Of J/. dichrous Mann. the British Museum 
contains as yet only a single female specimen, exactly agreeing 
with that figured by Mannerheim. This species is barely dis- 
tinguishable except in colour from JZ. cribratus Waterh., but 
may be easily distinguished from J. brasiliensis when the 
specific characters have been recognised. The latter differs from 
both in having no lateral expansion of the elytra in the female. 
The male has the last abdominal segment rather less strongly 
trilobed, the middle lobe being broader and less produced ; and in 
both sexes the rows of large setigerous punctures near the margins 
of the segments have almost disappeared. The sculpture of the 
upper surface is almost the same as in I. cribrata, but generally 
rather finer, and there is a very slight but constantly visible linear 
impression along the middle of the scutellum, which serves to 
distinguish this species from all the rest. . 


CYCLOCEPHALA PARAGUAYENSIS, Sp. Nn. 


Fusco-rufa, vertice nigro, elytris testaceis ; capite grosse punc- 
tato, medio leviter longitudinaliter sulcato, clypeo subquadrato ; 
prothorace scutelloque sat fortiter punctatis ; elytris annu- 
lariter punctatis ; pygidio crebre subtiliter punctatis. 3 wun- 
guiculorum anticorum internalium ramis haud valde diver- 
gentibus. 

Long. 12 mm 

This is one of the closely similar species forming the melano- 

cephala-group. Like C. laminata Burm., it is distinguished from 
the other members of that group by the anterior claws of the 
male, in which the two branches of the inner member are not 
strongly divergent. From C. laminata it differs by the thicker 
puncturation of the thorax and scutellum and the densely and 
finely punctured pygidium, which, however, is not rugose as in 
C. melanocephala, dimidiata, and vincentie. There is no appreci- 
able difference between the two sexes in the sculpture of the 
pygidium. 


NAUPACTUS TUBERCULATUS, Sp. Nn. 


Niger, pallide brunneo-pubescens, capitis prothoracisque linea 
alba media e squamis rotundatis composita,; scutello aldo- 
squamoso ; elytris costatis, interstitiis squamis minutissimis 
rufescentibus instructis ; rostro paulo lato, lateribus parallelis ; 
prothorace (3) quam latitudinem longiore, post medium 
lateraliter valde spinoso, disco leviter 6-tuberculato, tuberculis 
4 medio approximatis, 2 pone marginem anticam aliisque 
minus evidentibus prope angulos, omnibus nudis nitidis ; 
singulo elytro costa suturali aliisque duabus nudis, nitidis 

Proc. Zoot. Soc.—1903, Vou. Il. No. XVII. Wi 


258 MR. F. A. HERON AND SIR G. F. HAMPSON ON THE [ Nov. 3, 


ornato, interstitiis squamulis minutis rufescentibus vestitis, 
linea albo-squamosa ultra costam lateralem, hwmeris Sortiter 
extus productis, tibiis anticis intus dentibus numerosis validis 
armatis ; corpore subtus brunneo-vestito, abdominis medio 
depresso, nudo. 

Long. (rostr. excl.) 12°5 mm. 

This species is closely related to WV. nodicollis and perpastus of 
Boheman (the former has no affinity with the species of which it 
stands as a synonym in the Munich Catalogue, that insect, lewco- 
gaster Perty, belonging to the genus Pantomorus). The lateral 
spines of the new form are longer than in the other two, and 
those of the thorax are placed farther back. The thorax is pro- 
portionally longer, and the smooth elevated spots on its disc are 
peculiar to this species, as is the single sharply elevated discoidal 
costa of the elytron. This costa does not quite reach either the 
base or apex. The pinkish colour of the scales upon the disc of 
the elytra is perhaps not constant. We have only a single male 
specimen. 


EXPLANATION OF PLATE XXVIII. 


Fig. 1. Canthidium sladeni Arrow, p. 249. 

. Clavipalpus tenuis Arrow, p. 251. 

. Hoplopactus lateralis Arrow, p. 251. 

. Celosternus sturio Arrow, p. 252. 

. Belopeus niger Arrow, p. 252. 
Prostenus sladeni Gahan, p. 253. 
Spherion sladeni Gahan, p. 253. 

. Trichophorus electus Gahan, p. 254, 

. Oncideres sladeni Gahan, p. 254. 

. Deuterocampta sladene Gahan, p. 255. 


SO OTH Mw ro 


= 


4. On the Lepidoptera collected at Chapada, Matto Grosso, 
by Mr. A. Robert (Perey Sladen Expedition to Central 
Brazil). By F, A. Hieron and Sir Guorce F. Hampson, 


Bart., F.Z.S. 
[Received October 21, 1903.1 


LEPIDOPTERA PAPILIONINA, 
By F. A. Huron. 


LEMONIIDA, 
STALACHTIS PHLEGETONIA Perty. ©. 


Acrea phlegetonia Perty, Del. Anim. Art. p. 153, t. 30. ff. 2,26 
(1830-34). 
LYc@HNIDA, 
Lyczna casstus Cramer. ¢ ¢. 
Papilio cassius Cramer, Pap. Ex. i. t. 23, C, D (1775). 


Belongs to a section differing from Syntarucus in the absence 
of a tail to the hind wings. 


1903. ] LEPIDOPTERA OF THE PERCY SLADEN EXPEDITION, 259 


HESPERIIDE. 

HELIOPETES sp. ¢. 

This form, which belongs to a species very closely allied to 
HT, arsalie Linn. (Pap. arsalte Linn. Mus. Ulr. p. 245, 1764), is 
represented by a single badly damaged specimen which differs 
from the usual form of arsalte in its smaller size, less pointed 
wings, and the extreme reduction of the dark scaling at the bases 
of both wings and along the inner margin of the hind wing above, 
and below in the absence of dark and orange scales on the fore- 
wing costa and along the first median and internal veins of the 
hind wing. 


LEPIDOPTERA PHALAIN A. 
By Sir Grorce F. Hampson, Bart., F.Z.8. 


Nocruipa, 

ACRONYCTINE, 
EXMARGINEA GAMMoPHORA Guen. Noct. ii. p. 289 (1852). 
Ieee 


Nocruina. 
GLYMPIS HABITALIS W]k. xvi. 39 (1859). 
Wes 
GLYMPIS sp. 
1 Oe 


Erepus AGARISTA Cram. Pap. Exot. ii. p. 112, pl. 170, A, B 
(1775). 
29% 
Hypsip&. 


Pericopis sAcriricA Hibn. Zutr. ex. Schmett. ii. p. 21, 
ff. 473-4 (1825). 


POR 


GEOMETRIDA. 
BoARMIAN A. 

Merocavusta vinosa Dogn. Lép. Loja, p. 62, pl. vi. f. 17 (1891). 
MEQ 
OXYDIA sp. 
1 3g in bad condition. 

LiMAcoDID#. 
Susica QuADRATA WIk. v, 1132 (1855). 
2d. 
AsBoLiA vitLosiPes Wlk, xxxii, 555 (1865). 
Won 


ie 


260 ON THE CYPRIOTE SPINY MOUSE. [ Nov. 3, 


CASTNIADA. 
CasTNIA DEDALUS Cram. Pap. Exot. i. pl. 1, A, B (1775). 
RO 


PYRALID&. 
PHYCITIN A. 


ELASMOPALPUS PYRRHOCHRELLUS Rag. Nouv. Gen. p. 23 (1888) ; 
id. Rom. Mém. ix. p. 429, pl. xvii. f. 13. 


Wks 


SACCOPLEURA ? sp. 
1 $ without palpi. 


CHRYSAUGINE. 


PYRAUSTINA. 
GLYPHODES FLEGIA Cram. Pap. Exot. u1. p. 66, pl. 140, D (1775). 
Ue. 

TINEID&. 
1 ¢ belonging to a genus allied to Yortricopsis Newm., from 
Australia. 

HEPIALIDA. 

Dawaca METELLUS Druce, P. Z.8. 1889, p. 509, pl. 43. f. 2. 


3 ¢. Antenne bipectinate ; claspers asymmetrical, the right 
clasper large, the left minute. 


DALACA SLADENI, Sp. 0. 


g. Antenne minutely serrate; claspers symmetrical. 

Head, thorax, and abdomen rufous mixed with brown. Fore 
wing rufous, with numerous pale strie and reticulations, the 
costal area, the medial area below the cell, and the area between 
veins 3 and 6 suffused with dark brown; the pale strie forming 
an oblique ivregularly sinuous line from costa before middle to 
inner margin beyond middle, where it is met by an oblique slightly 
waved line from costa before apex ; a curved whitish mark com- 
posed of two small spots at origin of veins 5 and 6. Hind wing 
red-brown, the costa chequered ochreous and dark brown. 

Hab. Central Brazil, Chapada (A. Robert), 1 3 type. Hap. 
34 millim. 


5. Note on the Cypriote Spiny Mouse. By P. CHaumers 
Mircuett, M.A., D.Se., Secretary of the Society. 


[Received September 10, 1903.] 


Early in the summer of 1903, Mr. Roland L. N. Michell, 
British Commissioner at Limassol in Cyprus, wrote to me through 
Sir A. K. Rollit, M.P., F.Z.S., respecting a large spiny mouse, 
small numbers of which occurred among the rats sent in by the 
villagers in connection with the protective measures against 
plague. I asked him to send home skins and skulls, or, if possible, 
living specimens. He very kindly brought home a number of 


1903.] ON THE MUSCLES OF THE UNGULATA. 261 


living examples, four of which arrived at the Gardens in August. 
They turned out to be specimens of Acomys nesiotes, a new species 
described by Miss Bate* in June of this year from specimens 
brought by her from Cyprus and presented to the Natural History 
Museum, where Mr. Thomas very kindly showed them to me. 
Miss Bate obtained her specimens from the Kerynia Hills, not 
far from the village of Dikomo. She added: “ I never met with 
or heard of it in other parts of the island, though probably it 
occurs at any rate over the whole of the Mesorcea, or central 
plain, and the southern slopes of the Kerynia Hills.” 

From information given me by Mr. Michell it is clear that the 
Cypriote Spiny Mouse has a range in Cyprus much wider than 
that attributed to it by Miss Bate, and probably extending to the 
whole of the island. The south-western portion of the island is 
hilly, the range separating it from the central plain running up 
to over 5000 feet. Some of Mr. Michell’s specimens came from 
the village of Ayios Konstantiiios, at an elevation of over 4000 feet ; 
others were from various regions in the vine-clad hills, and others 
again from the low coast-area near Limassol. The mice are very 
timid and seldom seen by the villagers. They live in deep holes, 
and are reported to do much damage to the caroub trees. They 
are not known in towns. 

It happened, curiously enough, that the four specimens in the 
Gardens and all those obtained by Mr. Michell were devoid of 
tails. This corroborates the remarks of Miss Bate on the brittle- 
ness of the vertebre in these animals. The mice bred while in 
captivity, producing two at a birth. The young had normal tails, 
but lost them in a very short time. 


6. On the Muscles of the Ungulata. By Brrrram ©. A. 
Winnie, Se.D., M.D., M.A., F.R.S., Professor of 
Anatomy in the University of Birmingham, and F. G. 
Parsons, F.R.C.S., F.Z.S., F.L.8., Lecturer on Human 
and Comparative Anatomy at St. Thomas’s Hospital, 
late Hunterian Professor in the Royal College of Sur- 
geons, England. 


Part I1.—Musctizs oF THE HIND-LIMB AND TRUNK. 
[Received September 8, 1903. ] 
(Text-figures 24-27.) 

The first part of this paper, which dealt with the muscles of 
the head, neck, and fore-limb, was read before the Society on 
December 17, 1901, and published in the ‘ Proceedings’ in April 
1902 (see P. Z. 8. 1901, vol. ii. pp. 656-704), and we now wish to 


give an account of the muscles of the hind-limb and trunk, 
together with some general observations on the Order. 


* Ann. & Mag. Nat. Hist. (7) xi. p. 565 (1908).. 


262 MESSRS. B. C. A. WINDLE AND F. G. PARSONS ON [ Nov. 3, 


List of Animals. 
Division A, UNGULATA VERA. 
Suborder ARTIODACTYLA. 


Family HrprororaAMID&. 


. Hippopotamus amphibius. Gratiolet et Alix (IIL). 
Humphry (1V.). 
Cuvier et Laurillard (1.). 


99 73 


Co bo et 


99 99 
Family Sumpz, 

4. Sus scrofa (Domestic Pig). Lesbres (V.). 

D. Chauveau (IT.). 

Geter i ies A: Bronn (VL.). 

7 Cuvier et Laurillard (1.). 

8 Macalister (unpublished). 


OMe eh Le. a Meckel (VII.). 
LOM ee (Waldabear): Cuvier et Laurillard (1.). 
11. ,, porcus (Red River-Hog). 
12.. Babirusa. Vrolik (VIII). 


13. Dicotyles (Peccary). Cuvier et Laurillard (1.). 
14, 9) ? 


Family CAMELIDA. 


15. Camelus dromedarius. Chauveau (I1.). 


16. e eH Lesbres (V.). 
Wile es aa Walton (IX.). 
18. ee a Meckel (VII.). 
L$). F i P. Thompson. 


Family TRAGULIDE. 


20, Tragulus javanicws (Chevrotain). Kinberg (X.). 


? 9 
22. 3 kanchil. 
23. 3 stanleyanus. Macalister (unpublished). 


24. Dorcatheriwm (Water-Chevrotain). Chatin (X1.). 


Family Cervip#. 


25. Cervus axis. Macalister (unpublished). 

26. 4, manchuricus. 

27, Cervulus muntjac. 

28. Cariacus rufus (Brocket), 

: as mexicanus (Brocket), 

30. Alces machlis (Elk), Watson & Young (XXX. & XXXII). 
31. Moschus moschiferus ** (Musk-Deer). Bell (XIV.). 


* By a regrettable error this animal was described as the Musk-Ox (Ovibos mos- 
chatus) m the first part of this paper (P.Z.S. 1901, vol. ii. p. 657). 


56. 
57. 
58. 
59. 
60. 
61. 
62. 


63. 
64. 
65. 
66. 
67. 
68. 
69. 


. Ovis aries. Lesbres (V.). 


THE MUSCLES OF THE UNGULATA. 263 


Family GIRAFFIDA. 


. Camelopardus giraffa. Joly et Lavocat (XIT.). 


» 55 Murie (XXXII.). 
” ” Owen (XIIT.). 


Family Bovip2. 


. Bos taurus. Chauveau (IT.). 


» 9 Lesbres (V.). 

* + Meckel (VII.). 

” % Bronn (V1.). 

” 30 R.C.S. Museum (C. 216). 
” ” sy (C. 220). 
0 oe Chauveau (I1.). 

99 5 Bronn (VI). 

9 50 Meckel (VII.). 

0 a R.C.S. Museum. 


5 ,, steatopyga (Fat-tailed Sheep). 
os », (Syrian Sheep). 
»» musimon (Mouflon). 


. Capra hircus. Lesbres (V.). 


9 Pe Chauveau (IT.). 
9 9° Haughton CXaVE): 
R.C.S. Museum. 


Cephalophus grimmi (Duiker-bok). 
. Tragelaphus scriptus (Harnessed Antelope). 


Suborder PERISSODACTYLA. 


Family TAPIRIDH. 


Tapirus americanus. Turner (XVI.). 


5 “ De Longchamps (X VIIT.). 
Ke “ Cuvier et Laurillard (1.). 
a - Vrolik (VIII.). 

. us Lesbres (Y.). 


a ecm. Murie (XVII.). 


Family Equip. 
Equus caballus. Chauveau (1I.). 


i 53 Bronn (V1I.). 

Ms Pr Lesbres (V.). 

i ES Cuvier et Laurillard (1.). 
. th Meckel (VIT.). 

» asinus. Cuvier et Laurillard (I.). 


s “s Steel (XIX.). 


264 MESSRS. B. C. A. WINDLE AND F.G. PARSONS ON __| Nov. 3, 


Family RHINOCEROTIDA. 
70. Rhinoceros sumatrensis, Beddard & Treves (XX.). 


ale in ? sp. Haughton (XXI.). 
72. is 5 Owen (XXIT.). 
U3 e ES Macalister (XXIII.). 


Division B. SUBUNGULATA. 


Family Procaviup2&. 


George (X XVI). 
Meckel (VIT.). 


bP) 3) 


74. Procavia dorsalis (Hyrvax). Mivart & Murie (XXIYV.). 
UB 8 capensis. 

76. 5 ¢sp. Brandt (XXV.). 

77. 

78. 


7 2? 


Family ELEPHANTIDE. 
79. Elephas indicus. Anderson (XX VII.). 


80. i ms Mayer (XXVIII). 

81. a i Miall & Greenwood (X XIX.). 
82. x a Watson (XXX.). 

83. a 5, Young (XXX1.). 

84, An if Cuvier et Laurillard (1.). 

85. 59 Paterson & Dun (unpublished). 


86. »  Gfricanus, R.C.S. Museum. 


Muscles of the Gluteal Region. 


Ectogluteus (Gluteus maximus).—In the Ungulata the muscular 
sheet formed by the ectogluteus, femoro-coccygeus, and biceps is 
less separable than in any other order we know, and the difticulty 
of deciding how much of it is to be called ectogluteus is added to 
by the fact that the femoro-coccygeus or agitator caude has quite 
lately been recognised as a separate muscle, and so appears only 
in papers of recent date. The origin of the muscle is in all cases 
from the spines of the sacral and anterior caudal vertebre, and 
sometimes from the iliac crest ; the most interesting thing about 
its Insertion is that among the Artiodactyla, in which there is no 
third trochanter, it ends entirely in the fascia of the outer side 
of the thigh and has no bony insertion into the femur. This is 
the case in the Hippopotamide (1, 3), Suide (4, 5, 8, 11, 13, 14), 
Camelide (15), Tragulide (22), Cervide (26, 27, 29, 30), and 
Bovidee (35, 36, 38, 39, 41, 42, 43, 47, 50, 51,52, 54,55). In the 
Perissodactyla, in which a large third trochanter is present, the 
ectogluteus is inserted into it, though it did not seem to us that 
the muscle in the odd-toed Ungulates was usually better developed 
than in those with an even number of toes. This insertion into 
the third trochanter is recorded in the Tapiride (58, 61, 62), 
Equide (63, 65), and Rhinocerotide (71). Among the Subungu- 
lata the muscle was certainly inserted entirely into the fascia in 


1903.] THE MUSCLES OF THE UNGULATA. 265 


our specimen of Hyrax (75) (see text-fig. 24), and in that of 
George (77), but we think that the other authors have included 
the femoro-coccygeus in their descriptions. In the Elephantide, 
Miall and Greenwood (XXIX.) notice a strong attachment to 
the middle fifth of the femur, and this is confirmed by Paterson 
and Dun. We are therefore able to generalise by saying that 


Text-fig. 24. 


Wy 


Yy 


UT Y 

\ (s M, i W Ak 
NTC 
\\ AH 


} 


i 


———— 


———— 
——= 


——— 


e 
—————— 
—— 


=—— 


—_ 


y 


Muscles of outer side of Thigh of Hyrax. 


T.F.F. Tensor fascize femoris. Bi. Biceps. 
E.G. Ectogluteus. 8.T. Semitendinosus. 
A.C. Agitator caude (femoro- | S.M. Semimembranosus. 
coccygeus). | 


in the Artiodactyla and the Procaviide the ectogluteus is inserted 
into fascia only, while in the Perissodactyla and the Elephantidee 
there is a definite femoral insertion. The nerve-supply in the 
Antelope, Sheep, and Pig is from a definite inferior gluteal nerve 


266 MESSRS. B. C. A. WINDLE AND F..G. PARSONS ON [ Nov. 3, 


which comes off the sciatic outside the pelvis and below the point 
of origin of the superior gluteal. 

Femoro-coccygeus (Agitator cawde).—This muscle can be made 
out readily enough in most Ungulates by anyone who is familar 
with it and who appreciates the fact that it is the part of the 
great muscular sheet which lies between the ectogluteus and 
biceps. The origin is from the posterior sacral and anterior caudal 
spines, and the insertion chiefly into the outer side of the patella ; 
above and below this it blends with the fascia (see text-fig. 24). 
As this muscle was not known by most of the older writers on 
myology, its description is often included with that of the ecto- 
gluteus or biceps. We believe that it is present in every Ungulate, 
and that its attachments are very constant. It is supplied by 
the same nerve as the ectogluteus. 

Tensor fascie femoris.—As the femoro-coccygeus connects the 
ectogluteus and biceps, so the muscle under consideration lies 
between the ectogluteus and sartorius though it is often separated 
by a good interval from the former. It rises from the iliac crest, 
which of course is a short structure, and spreads out into a fan- 
shaped muscular mass which usually becomes lost in the fascia lata 
about the middle of the thigh. In the Elephant (79, 81, 84), 
however, it runs two-thirds of the way down, while in the Tapir 
(61, 62) it is fleshy as low as the outer side of the patella. Pater- 
son and Dun have noticed in the Elephant (85), and Murie in the 
Giraffe (33), that the fascia lata on the outer side of the thigh is 
elastic. Its nerve-supply is the superior gluteal. 

Mesogluteus (Gluteus medius)—This muscle, as is usual in 
mammals, is generally larger than the ectogluteus; it rises from 
the greater part of the gluteal surface of the ilium as far as the 
margin of the great sciatic notch, as well as from the fascia cover- 
ing the muscle. At its origin it is probably inseparably blended 
with the pyriformis, at all events the latter can never be traced 
to its human origin from the ventral side of the sacrum. The 
insertion of the mesogluteus is into the great trochanter on its 
outer and often also on its posterior surface. It is supplied by 
the superior gluteal nerve in the Pig, Sheep, Antelope, and Horse, 
but in the Harnessed Antelope (55) we also found a small twig 
passing to it from the inferior gluteal. In the Perissodactyla the 
muscle seems specially large, and its origin creeps along the fascia 
lata covering it to the sheath of the erector spine and the sacro- 
iliac and sacro-sciatic ligaments. Meckel (VII.) says that in the 
Horse it is twelve times as large as the ectogluteus, and it is also 
very large in the Tapir. In the Pig and Hyrax it is relatively 
much smaller, and Meckel describes it in the latter animal as 
being absolutely smaller than the ectogluteus. In our specimen 
of Hyrax (75) we do not feel justified in going as far as this, but 
the muscle was certainly very thin in the middle. 

Entogluteus (Gluteus minimus).—This is also a large muscle, 
but its exact attachments are difficult to define owing to the fact 
that it is often closely connected with the gluteus ventralis. We 


1903. THE MUSCLES OF THE UNGULATA. 267 


believe that these two muscles are always present in Ungulates, 
and that the line of separation can be determined, if carefully 
looked for, by the branch from the superior gluteal nerve to the 
tensor fascie femoris passing through it. The origin is from the 
ventral part of the gluteal surface of the ilium near that portion 
of the bone usually described by veterinary anatomists as the 
neck, while its insertion is into the front and outer side of the 
great trochanter. In the Horse, Lesbres (V.) says that the ento- 
and meso-glutei can only be distinguished from one another at 
their insertions. The nerve-supply seems always to be the 
superior gluteal. 

Gluteus ventralis (Gluteus quartus ; Scansorius).—This muscle, 
when it is distinct, as it generally is in the families of Camelide, 
Girafiide, and Bovide, rises from the ventral border of the ilium 
and is concealed by the overlapping entogluteus, and is inserted 
into the anterior surface of the great trochanter. In Ungulates 
belonging to other families than those just mentioned, it is often 
difficult to separate the muscle from the entogluteus unless the 
nerve to the tensor fasciee femoris is looked for. The nerve-supply 
is the superior gluteal. 

Gluteus profundus (Gluteus quintus ; Ilio-capsularis).—It is 
quite certain that this is not a constant and well-defined muscle 
in Ungulates, though it is occasionally found. Lesbres (V.) 
describes the ‘abductor trochitereen” in the Horse as very 
distinct, and says that it lies over the superior surface of the 
capsule of the hip, rises from the supracotyloid crest, and is in- 
serted into the anterior part of the internal surface of the great 
trochanter which is known to veterinary anatomists as the con- 
vexity of the trochanter. He regards it as the serial homologue 
of the infraspinatus secundus (see Part I. of this paper, P. Z.8. 
1901, vol. i1. p. 687), but it answers very well to our idea of the 
gluteus profundus. In addition to this, he describes another muscle 
in the Horse, rising from the ilium close to the origin of the rectus 
and inserted into the anterior surface of the femur close to the 
head; this he considers is the homologue of the ilio-capsularis of 
the Carnivora. For practical purposes it does not seem to us that 
these two muscles require separate names; they are probably two 
slips of the deepest delamination of the gluteal mass lying in 
contact with the capsule of the hip. In the Harnessed Antelope 
(55) we found a gluteus profundus, and Kinberg (X.) describes it 
in the Chevrotain under the name of M. tenuis femoris. In the 
other Ungulates the muscle is not as a rule delaminated from the 
entogluteal sheet. The nerve-supply in the Harnessed Antelope 
is from the sacral plexus just below the origin of the superior 
gluteal nerve. 

Pyriformis.—We have already stated that the mesogluteus is 
usually fused more or less completely with the pyriformis, but in 
some cases, é. g. the Duiker-bok (54) and Harnessed Antelope (55), 
the entogluteus is more closely blended with it than is the meso- 
gluteus; it must, however, be remembered that in the Ungulates 


268 MESSRS. B. C. A. WINDLE AND F.G. PARSONS ON [ Nov. 3, 


the delamination of the gluteal mass is not nearly so complete as 
it isin Man. In the Pig (4, 5, 8, 9, 11, 14), Ox (35, 36, 38), and 
Sheep (41, 42, 43, 46, 47) the pyriformis is so completely fused 
with the meso- or entogluteus that no trace of it can be found, 
Tn no case does it come out of the great sciatic foramen, for this 
is closed by a membrane, and when the muscle is distinct it rises 
from the surface of this. The nerve-supply in the Duiker-bok 
and Harnessed Antelope is the superior gluteal. 

Obturator internus.—This muscle in the Suide (4, 8, 11, 14), 
Tragulide (20, 21), Giraffide (32-34), Bovide (36, 39, 41, 46, 48, 
50, 53, 54, 55), and Procaviide (74, 75) is small, and more nearly 
resembles a gemellus than the usual mammalian obturator in- 
ternus: the reason for this is that the obturator tertius usurps 
the normal origin of the muscle, and the internus has to arise 
from the body of the ischium at the lesser sciatic notch, thus 
reverting to what is probably the original and generalised origin. 
Of its arrangement in the Cervide we are not sure, but Watson 
and Young describe it in the Elk (XXX. & XXXI.)as having the 
usual origin from the inner side of the obturator membrane. In 
the Perissodactyla and Elephantide the muscle rises from the 
inner side of the true pelvis as usual. In all cases it is inserted 
into the bottom of the trochanteric (digital) fossa, and it is 
supplied by a branch of the sacral plexus. 

Gemelli.mIn those animals just mentioned in which the obtu- 
rator internus rises at the lesser sacro-sciatic notch, the gemelli are 
fused with it; in the others only an inferior gemellus is as a rule 
present. In the Horse, however (63, 65), three gemelli are often 
met with, the third rising a little ventral and caudal to the 
inferior, between it and the obturator externus. In the Elk (30) 
the two gemelli are joined. 

Obturator externus and Obturator tertius (see text-figs. 25 & 26).— 
Mivart and Murie nearly forty years ago described a muscle In 
Hyrax (74) passing through the obturator foramen and being 
inserted into the trochanteric fossa with the obturator externus ; 
this they named the obturator tertius. Later Murie found the same 
muscle in the Giraffe (33),and Lesbres(V.) has evidently noticed the 
same thing, for he says that, in the Horse, Pig, and Rumimants, part 
of the obturator internus rises from the pelvis above the obturator 
vessels as high up as the ilium, while the other part passes through 
the obturator foramen. We have carefully studied this muscle 
in the Chevrotain, Ox, Sheep, Goat, Pig, Duiker-bok, Harnessed 
Antelope, and Hyrax, and are convinced that the following is the 
explanation of it. The obturator tertius is really a part of the 
obturator externus, which has made its way through the obturator 
foramen from outside, pushing the obturator membrane in front 
of it. In doing this it has stolen a good deal of the usual origin 
of the obturator internus and pushed that muscle out of the pelvis, 
so that it rises, as we have shown, from the margin of the lesser 
sciatic notch. There is, therefore, the curious paradox of the 
obturator externus rising inside the pelvis, while the internus 


1903. ] THE MUSCLES OF THE UNGULATA. 269 


rises outside. Our reasons for coming to this conclusion are, 
firstly, that in dissecting away the obturator externus to expose, 
as we thought, the obturator tertius we found that the two 
muscles were practically one, and that no division was possible 
between them; secondly, that the obturator membrane can be 
clearly seen on the pelvic surface of the so-called obturator tertius ; 
and, thirdly, that the part of the muscle which passes through 
the obturator foramen is supplied by the obturator nerve, and not 
by a branch from the sacral plexus. This last fact is perhaps the 
one on which we are inclined to lay most stress. We regret that 
in many of the animals which passed through our hands we did 
not pay enough attention to this muscle, and it was only when 


—-z 


\ To LESS8 TROGHR 


\ro BEHIND LESS TROCHR 
= QUA DE VES 


Hip-region of Sheep, showing double Quadratus femoris muscle. 


G.MED. Gluteus medius. O.I. Obturator internus. 
G.min. Gluteus minimus. O.E. Obturator externus. 


working through Hyrax with the aid of Mivart and Murie’s 
paper that we first noticed it. We have seen enough, however, 
to feel sure that it is present in the Suide, Tragulide, Giraffide, 
Bovide, and Procaviide. We expect, from what we know of 
other muscles, that it will also be found in the Hippopotamide, 
Camelide, and Cervide, if carefully and specially looked for, but 
the various writers on these families were, like ourselves, evidently 
ignorant of its existence. With regard to the Perissodactyla, we 
have satisfied ourselves, through the kindness of the authorities at 
the Royal Veterinary College, that no muscle passes through the 


270 MESSRS. B. C. A. WINDLE AND F.G. PARSONS ON [ Nov. 3, 


obturator foramen in the Horse, and this in spite of Lesbre’s. 
assertion to the contrary. In the Tapiridee, too (61), we failed to 
find it, and so did Murie (X VII.), who certainly would have been 
on the look out for it since he was the original describer. It is 
not described by any author in the Rhinocerotide, but further 
observation is necessary here. In the Elephant no mention is 
made of it, though Miall and Greenwood (X XIX.) and Paterson 
and Dun describe the obturators somewhat carefully. With 
further opportunity for research we believe that it will be found 
that the obturator externus passes through the foramen in the 
Artiodactyla and Procaviidee, but that it does not do so in the 
Perissodactyla and Elephantide. 

Quadratus femoris.—This muscle varies a good deal in size in 
different Ungulates, but it is usually present. Owing to the 
horizontal position of the pelvis, it passes obliquely from the tuber 
ischii to the back of the upper part of the shaft of the femur so 
as to form an X with the tendon of the obturator externus; it is 
usually somewhat constricted at its Insertion, so that the name 
quadratus femoris used in human anatomy does not very aptly 
describe it. It is especially large in the Hyrax (74, 75), and in 
one Sheep (46) (see text-fig. 25) it was found to be double, but this 
does not seem to be always the case. Among the Perissodactyla 
it is not well developed ; in the Horse (63) it is quite small, and 
Meckel (VIT.) says absent altogether. In the Tapir (61, 62) it is 
also small, while in the Rhinoceros (71) Haughton failed to find 
it*. In all the other animals on our list the muscle was present, 
and in the Duiker-bok, Sheep, and Harnessed Antelope was 
supplied by a branch from the sacral plexus as in Man. 


Muscles of the Posterior Femoral Region. 


Senumembranosus and Presemimembranosus.—These muscles 
have the usual mammalian attachments; they are always fused at 
their origin from the tuber ischii; and it is characteristic of 
ungulates that this fusion continues much lower down the thigh 
than it usually does in other mammals. This is the case in the 
Hippopotamus (1), Pig (4, 11, 14), Chevrotain (21, 22), Deer (26, 
29, 30), Ox (36, 40), Sheep (41, 46, 47), Goat (50, 52), Antelope 
(54, 55), Tapir (58, 61), Rhinoceros (71), Hyrax (74, 75), and 
Elephant (81, 84, 85). Meckel (VII.) and Lesbres (V.) point out 
that in the ruminants the origin is entirely ischial, while the 
Horse has a caudal origin besides; we think that we are justified 
in broadening this generalisation and in saying that in the Artio- 
dactyla there is only an ischial head, while in the Perissodactyla 
and Subungulata ischial and caudal heads are found (see text- 
figs. 24 & 26, S.M.). 

The semimembranosus has the usual mammalian insertion by a 
rounded tendon deep to the internal lateral ligament: in the 


is The dissection of this animal seems to have been carried out mmder great 
difficulties, so that the muscle may haye been overlooked. 


1903. ] THE MUSCLES OF THE UNGULATA, 271 


Elephant (81, 85), however, an expansion from this insertion is 
found running down to the inner side of the ankle. According 
to some authors the semimembranosus may occasionally be absent 
altogether: this was the case in Kinberg’s Chevrotain (20), 
Chatin’s Water-Chevrotain (24) *, and Lesbre’s Horse (65). The 
presemimembranosus is usually larger than the semimembranosus, 
and is inserted by fleshy fibres into the lower part of the femur 
just above the internal condyle; besides its fusion with the sem1- 
membranosus it is very intimately connected with the adductor 
mass and so has escaped notice by some observers. The nerve- 
supply in the Pig, Antelope, Sheep, and Hyrax is from the nerve 
to the hamstrings, a large branch which comes off the internal 
popliteal element of the great sciatic. 

Semitendinosus.—This muscle, like the last, may rise by one or 
two heads, of which the one from the tuber ischii is always present. 
A caudal head is rarely found in the Artiodactyla, Chatin’s Water- 
Chevrotain (24) and the Pig (4, 6) are the only exceptions we 
know. In the other suborders, the Horse (63, 64, 65), Tapir (58, 
62), and Hyrax (74, 75, 78) have two heads (see text-figs. 24 & 
26, 8.T.). The insertion is into the second quarter of the 
internal surface of the shaft of the tibia as a rule, and from 
this a small expansion is sent off to help the biceps and gracilis 
in reinforcing and providing a sheath for the tendo Achillis, 
and so acting on the calcaneal tuberosity. These expansions 
from the inner and outer hamstrings are seen at their best 
in the Ungulata; but they are so common among the Mammalia 
generally t, that they probably have a phylogenetic history to 
account for their presence, as well as a physiological advantage 
to account for their special development in Ungulates. In all 
probability their past history is that they are remnants of the 
longitudinal muscle-fibres passing uninterruptedly from the 
pelvic girdle to the foot, such as are found among the Reptilia, 
while the explanation of the fact that they are so well developed 
in Ungulates must be looked for in the mode of life of these 
animals. Among all the orders of mammals there is not one 
which is more distinctly cursorial in the habits of its species; 
with the exception of the Tree-Hyrax there are no arboreal types ;> 
the Hippopotamus and Water-Chevrotain are the only ones which 
are partially aquatic, while there is no example of a fossorial 
Ungulate. For these reasons, any modification which we meet 
with in the limb-muscles of Ungulates as a group, will probably be 
explained by its being an advantage to them in running. ‘The 
expansions of the hamstrings to the calcaneum act as plantar 
flexors of the ankle-joint, while the hamstrings themselves are 
flexors of the knee. In watching the sequence of movements of 


* In our specimens of Chevrotain (21, 22) the muscle was undoubtedly present, 
though so closely connected with the presemimembranosus that it might easily be 
overlooked without careful dissection of the insertion. 

+ The fascial expansions from the semitendinosus and gracilis in Man will at once 
occur to the human anatomist. 


272, MESSRS. B. C. A. WINDLE AND F.G. PARSONS ON [ Nov. 3, 


the hind-limb in walking, one notices that the knee is extended 
and the ankle plantar flexed in order to lengthen the limb and so 
pull and push the body of the animal over the spot on which the 
foot is planted, while at the same time the knee is slightly flexed. 
It is probable that the expansions from the hamstrings effect the 
plantar flexion of the ankle by pulling up the calcaneal tuberosity, 
at the same time that they are causing slight flexion of the knee. 
An interesting and, so far as we know, unaccounted for structure 
in the semitendinosus of Man, is the fibrous streak running across 
the muscular belly; this is seldom seen in the lower mammals, 
and we have not noticed it in any Ungulate, although we have 
carefully looked for it. Macalister, however, found it distinctly 
in the Muntjac (27). The nerve-supply of the semitendinosus is 
the special nerve to the hamstrings from the internal popliteal 
element of the great sciatic (see text-fig. 26, p. 274). 

Biceps (Flexor cruris lateralis)—It has already been pointed 
out how closely the biceps is connected with the femoro-coccygeus 
in the Ungulata, and how constantly the latter muscle is inserted 
into the side of the patella; we shall therefore describe as biceps 
only that part of the superficial lateral plane of muscle which is 
inserted below the patella. As the muscle is so inseparable from 
its neighbour the femoro-coccygeus (agitator caudze), it is very 
difficult to say whether any of its fibres really rise from the caudal 
vertebre ; we are, however, inclined to agree with Lesbres that 
in the Ungulates the biceps has only an origin from the tuber 
ischii (see text-fig. 24, Bi.). Its insertion is into the fascia of the 
upper half or more of the outer side of the leg, while from its 
lower border a strong fibrous process passes down to the calcaneal 
tuberosity, helping the gracilis and semitendinosus to ensheath 
the tendo Achillis. With the exception of relative size, there is 
hardly any variety in the biceps of different Ungulates ; it perhaps 
reaches its greatest development in the Tapir (58, 61, 62). 

Bicipiti accessorius (Tenuissimus).—This in most Ungulates is 
either suppressed or completely blended with the biceps. The 
only record we can find of it is that it 1s figured in Cuvier and 
Laurillard’s Elephant (84), and may possibly be the deep part of 
‘the biceps of the Tapir described by Murie (X VIL). 


Muscles of the Internal Femoral Region. 


Pectineus.—This muscle is often difficult to distinguish from 
the adductor longus, especially as the latter may or may not be 
delaminated from the adductor mass. The origin is fairly constant, 
coming from some part of the ilio-pectineal line, while the insertion 
seems to range all over the shaft of the femur. In the Hippopo- 
tamus (1) Gratiolet describes three parts, pubic, symphysial, and 
ischial ; of these the latter two are evidently adductor longus and 
quadratus femoris. In the Suide (6, 7, 11, 14) the muscle rises 
from the whole ilio-pectineal line and is inserted into the middle 
or distal part of the femur; it is comparatively small in this 


1903.) THE MUSCLES OF THE UNGULATA. 273 


family. In the Chevrotain, Kinberg (X.) says that it is inserted 
into the trochanteric fossa, but the origin is the same as in the 
Suide. Among the Cervide we have records of it in Cervus (26), 
Cariacus (28), and Alces (30), and in these it passes to the middle 
third of the femur. Of the Giraffide we have no definite records. 
Among the Bovide it seems to be variable. In the Ox, Chauveau 
(II.) notices that it is in two bundles, and so it is in the College 
of Surgeons’ specimen (39); Lesbres (V.) says that this is true for 
all ruminants, and we are not surprised to meet with this deserip- 
tion since we have found it double in other orders of mammals ; 
still in our specimens of Sheep (46, 47, 49), Duiker-bok (54), and 
Harnessed Antelope (55) we looked in vain for a division. In 
these three animals the origin was instructive: in the Sheep (46) 
it rose from the whole of the ilio-pectineal line and was inserted 
into the upper two-thirds of the femur; in the Fat-tailed Sheep 
(47) and Duiker-bok (54) it rose only from the angle of the pubis 
and went to the upper half of the femur; while in the Harnessed 
Antelope (55) it rose from the outer end of the ilio-pectineal line 
close to the psoas and was inserted into the femur just below the 
lesser trochanter. In the Tapiride (61, 62) the origin is from the 
pelvie brim as usual, but the insertion is very high up behind the 
lesser trochanter or even, according to Murie, into the trochanteric 
fossa. In the Equide (63, 65, 67) the muscle is well marked and 
rises either from the ligamentum teres, which in this animal comes 
out of the cotyloid notch and runs forward to the hypogastric 
region, or splits to allow that ligament to pass through its origin ; 
it is inserted into the middle third of the femur. In the Rhino- 
ceros (71) it is inserted into the junction of the middle and lower 
thirds of the femur by a rounded tendon. Among the Subungu- 
lata, in the Hyrax (see text-fig. 26, p. 274) (74, 75, 78) and 
Elephant (81, 84) it has the usual origin from the anterior ramus 
of the pubis, while the insertion is into the middle third of the 
femur. 

The nerve-supply may, as in Man, be anterior crural, obturator, 
or both. In the Duiker-bok, Sheep, and Hyrax it was supplied 
by the obturator alone, in the Harnessed Antelope by the anterior 
erural alone, while in the Peccary it received twigs from both 
nerves. 

Adductor mass.—Many of the dissectors of Ungulates in the 
past have, we believe, been stimulated by their knowledge of 
human anatomy to artificially divide up this mass in an unnecessary 
manner. In the animals which we have dissected we have noticed 
that the nerve to the gracilis from the obturator forms a convenient 
indication of the interval between the adductor longus and the 
rest of the mass. When this interval is clearly marked an ad- 
ductor longus may fairly be described, but unless this is the case 
there seems to us little object in separating one part of the mass 
from the rest. Among the Artiodactyla there seems more difficulty 
in distinguishing an adductor longus from the rest of the mass 
than there is in the Perissodactyla or Subungulata, though it is 


Proc. Zoot. Soc.—1903, Vou. II. No. XVIII. 18 


274 MESSRS. B. C. A. WINDLE AND F. G. PARSONS ON __[ Nov. 3, 


always an arbitrary proceeding to settle whether a muscle is 
divisible or not. In the following animals no natural division 
was made out:—Hippopotamus (1), Pig (4, 11), Peccary (14), 
Chevrotain (20, 21, 22), Deer (25, 26), Hlk (30), Ox (35), Sheep 
(42, 46), and Duiker-bok (54). A fairly distinct adductor longus, on 
the other hand, was found in the Giraffe (33), Harnessed Antelope 
(55), Tapir (62), Horse (63, 65, 67), Rhinoceros (70), Hyrax (74, 
75, 78) (see text-fig. 26), and Elephant (81, 85). We have not 
been able to satisfy ourselves of a single instance of the presence 
of an adductor brevis, for the muscle which is called by this name 
in the veterinary books is clearly the homologue of the human 
adductor longus. The adductor longus, when it is distinct, rises 
close to the anterior part of the symphysis, while the insertion is 
into the shaft of the femur, usually in its middle third. The rest 
of the adductor mass rises from the whole length of the sabpubic 
ramus and is inserted into the lower half or two-thirds of the 
femur. The whole of the adductor mass is always supplied by 
the obturator nerve. 


Text-fig. 26. 


NERVE 70 H.S 


Muscles of inner side of Thigh of Hyrax. 


P.s. Psoas. P.S.M. Presemimembranosus. 
Prcr. Pectineus. Q.F. Quadratus femoris. 
O.E. Obturator externus. C.F. Femoro-coccygeus. 
A.M. Adductor mass. Bi. Biceps. 
O.N. Obturator nerve. 8.T. Semitendinosus. 
Gr. Gracilis. I.L.Lig. Internal lateral ligament. 
Gt.Sc.N. Great sciatic nerve. H.S. Ham-string. 


S.M. Semimembranosus. 


Gracilis (Adductor cruris)—This is a very constant muscle 
among the Ungulata; it rises as usual from the whole length of 
the symphysis and runs down as a single broad sheet to the usual 
insertion at the upper part of the inner surface of the tibia. Its 
tendon of insertion is generally more or less directly fused with. 


1903. ] THE MUSCLES OF THE UNGULATA. 275 


that of the sartorius, while from its lower margin it gives off an 
expansion to the inner side of the tendo Achillis which blends 
with similar expansions of the biceps and semitendinosus to form 
the sheath for that tendon. This sheath has already been dis- 
cussed under the head of the semitendinosus. The muscle seems 
so constant in the various families that it is needless to repeat 
each individual record, especially as there are thirty-four of them 
fairly evenly distributed over the order; it will be sufticient to 
point out that according to Murie (XX XIT.) the muscle is absent 
in the Giraffe, while Chauveau (II.) says that in the Camel its 
origin is bifid, the anterior branch being the smaller. Meckel 
(VII.), on the other hand, says that there are four heads to the 
gracilis in the Camel, so that it is clear that further knowledge 
is needed on this point. The nerve-supply in every animal we 
dissected was the obturator. 


Muscles of the Anterior Femoral Region. 


Sartorius (Llio-tibialis)—The origin of this muscle is usually 
not so high as the anterior superior spine of the ilium, more often 
it comes from the iliac fascia covering the insertion of the psoas 
parvus as well as from Poupart’s ligament close by. In the Ox 
(35, 36, 37), Sheep (41, 42, 46, 48), and Goat (50) it has another 
origin from the pubis just internal to the femoral vessels, but the 
two origins soon join so that the vessels pierce the conjoined 
origin. The insertion is into the upper part of the tibia by 
means of the fascia of the thigh; sometimes the fleshy fibres 
reach as far as, or even beyond, the knee, but more often they are 
lost in the fascia lata or join the tendon of the gracilis before 
the knee is reached. In the Perissodactyla the sartorius seems 
rather better developed than in the rest of the Ungulata, at all 
events it remains fleshy below the knee in the Horse (63), Tapir 
(58, 59, 60, 61), and Rhinoceros (71). Sometimes the muscle is 
absent, this was the case in Hyrax (74, 75); and we have no 
doubt that what Meckel describes in this animal (78) as sartorius 
is really tensor fasciee femoris. It was also absent in Anderson’s 
Hlephant (79), and Cuvier and Laurillard (84) do not figure it, 
though it is definitely described by Miall and Greenwood (81) as 
well as by Paterson and Dun (85). On comparing this muscle in 
the Ungulata with its condition in other orders, it will be seen 
that it is better developed than in many of them, and may cover 
in the femoral vessels, making a definite Hunter’s canal. Its 
nerve-supply in the Pig, Sheep, Duiker-bok, Harnessed Antelope, 
and Elephant is the anterior crural, the special branch to it 
passing over (superficial to) the femoral vessels a little below 
Poupart’s ligament. 

The Quadriceps extensor cruris has only a few points of special 
interest so far as our observations go. The rectus, as is usual in 
mammals, always has a reflected head, and often a straight head 
too, though the two may be continuous and so only one is 
described. We have so far not met witha single ungulate in 

13% 


276 MESSRS. B. C. A. WINDLE AND F.G. PARSONS ON _[ Noy. 3, 


which by cutting the rectus and turning its origin up, evidence 
of the two heads could not be found. Lesbres (V.) says that in 
the Pig, Sheep, and Goat the reflected head is absent; we have, 
however, paid special attention to the point in the Pig (11, 14) 
and Sheep (46, 47), and are convinced that both heads are really 
there. With regard to the crureus in Ungulates, we have often 
noticed that it is more distinct than in many other orders of 
mammals, and it rises from all three surfaces of the femur. Some- 
times, as in the Ox (40) and Sheep (41), it is divided more or 
less completely into an inner and an outer portion. The vastus 
externus is usually larger than the internus, but in the Tapir 
(61, 62) both Murie and ourselves were struck by the enormous 
size of the vastus internus. The nerve-supply of the quadriceps 
is always the anterior crural. 


Anterior Tibial Region. 


Tibialis anticus—This muscle usually has a femoral origin 
from the front of the external condyle, rising with the extensor 
longus digitorum, as well as a tibial origin from a small part of 
the upper and front portion of the tibia. 

In the Hippopotamus (3) the two parts are quite separate, the 
tibial portion being inserted into the base of the second (index) 
metatarsal, while the condylar part forms an arch in front of the 
ankle which binds down the extensor tendons, the two horns of 
the arch being attached to the bases of the index and minimus 
metatarsals. In the Suide (4, 5, 11, 14) the two parts are also 
distinct, but the fibrous arch is not present, and they are both 
inserted into the index metatarsal and middle cuneiform bones. 
In all the other Artiodactyla, viz. Chevrotain (22, also the “‘ Napu 
Deer” described by Haughton XV., which is the Javan Chevrotain), 
Virginian Deer, Nylghau, Elk (30), Brocket Deer (29), Sheep 
(47, 48), Ox (35, 40), Goat (50, 52), and Antelope (54, 55), the 
arrangement is practically identical with that of the Duiker-bok 
(54), the variations being very slight. In this animal (54) the 
femoral head is the larger, and opposite the ankle is pierced by 
the tibial tendon, it is then inserted into the inner side of the 
dorsum of the base of the large metatarsal bone, which is common 
to the medius and annularis digits. The tibial tendon, which 
pierces the one just described, runs to the ventral part of the 
inner side of the base of the same metatarsal bone. In the 
Perissodactyla the same femoral and tibial origins are found in 
the Tapiride (61, 62) and Equide (63, 64, 65), and the same 
perforation occurs in the Horse, though we did not notice it in 
the Tapir (61). As to the Rhinoceros we have not sufficient data 
to form an opinion. In the Subungulata the femoral origin is 
absent; this is the case in both the Hyrax (74, 75) and the 
Elephant (80, 81, 84, 85). In the Hyrax its insertion is into the 
second metatarsal, while in the Elephant it is usually inserted 


into the first as well. It is always supplied by the anterior 
tibial nerve. 


1903. ] THE MUSCLES OF THE UNGULATA. 200 


Extensor proprius hallucis.—TVhis muscle is often described in 
Ungulates, but, we think, without any real reason. As the 
hallux and often the fibula are suppressed, the: muscle called 
extensor proprius hallucis is a slip of the extensor longus 
digitorum passing to the index or medius digit. Unless the 
authors who use the name have good grounds for thinking that 
this is really a displaced muscle of the hallux, it seems a needless 
complication to retain the name in the Ungulata. 

Extensor longus digitorwum.—tiIn the Hippopotamide (1, 3) and 
Suide (4, 7, 10, 11, 12, 13, 14) this muscle is inserted into all 
four toes, and its origin is from the front of the external condyle 
of the femur with the tibialis anticus. Meckel (VII.) denies 
that it has a femoral origin in the Pig, but his was clearly an 
exceptional case. In the other Artiodactyla the origin is always 
femoral and the insertion into both toes. An interesting point 
about this muscle in the Ox (40), Sheep (46), Duiker-bok (54), 
Harnessed Antelope (55), Chevrotain (20), and Camel (17) is 
that there is an insertion into the middle phalanx of the medius 
digit as well as into the terminal of the medius and annularis. 
A more careful examination of the muscle in the other Artio- 
dactyles would possibly show that this is a constant arrangement, 
In the Tapiride (59, 61, 62) and Rhinocerotide the origin is 
femoral and the insertion into all three toes. In the Equide 
(63, 65) the origin is also femoral and the insertion into the 
middle and terminal phalanges of the single digit (medius). 
Among the Subungulata the origin is femoral in Hyrax (74, 75), 
though Meckel (VII.) says that it is tibial; its insertion is into 
all the digits. In the Elephant (81, 84, 85) the origin is certainly 
tibial and the insertion into the 2nd, 3rd, 4th and 5th digits. 
Both Paterson and Dun (85) and Miall and Greenwood (81) notice 
that insertions are given to some of the proximal as well as the 
distal phalanges, though which digits have slips to these phalanges 
varies in different individuals as well as on opposite sides of the 
same individual. The nerve-supply is the anterior crural. 
Summing up, it may be said that the origin of this muscle in the 
Ungulata is always femoral except in the Elephant. 

Extensor brevis digitorwm—This muscle is always present and 
has the usual origin from the front of the upper surface of the 
caleaneum. In the Hippopotamus (1, 3) it is inserted into 
the index, medius, and annularis digits. In the Suide (4, 5, 10, 
11, 14) it usually goes to the medius and annularis only, but it 
may send a slip which joins the tendon of the extensor longus 
digitorum on the dorsum of the foot. 

In the Tapiride (61, 62) the muscle is very large and is 
inserted into all three toes. Murie (X VII.) notices that the slip 
to the medius is inserted into the proximal phalanx. In the 
Equide (63, 65) there is only one insertion into the extensor longus 
as in the Ruminants. In the Rhinoceros, Haughton (XX1I.) 
only found a tendon for the medius toe. Among the Subungulata, 
Hyrax may have three tendons for the proximal phalanges 


278 MESSRS. B. C. A. WINDLE AND F.G. PARSONS ON [ Nov. 9, 


according to Mivart and Murie (74), or only two for the hallux 
and index (Meckel, VII.). In our specimen (74) we found 
tendons for the hallux and index, but they were inserted into 
the proximal phalanges. In the Elephant (85), Paterson and Dun 
notice that the muscle rises not only from the caleaneum but 
from the navicular, base of the metatarsal of the medius, and the 
dorsal ligaments of the tarsus; it is inserted into all three 
phalanges of the medius, the slip to the terminal phalanx joining 
the extensor longus to that digit ; on the right side it is inserted 
into the annularis as well. In Miall and Greenwood’s (81) and 
Cuvier and Laurillard’s (84) specimens details are wanting, but the 
muscle only goes to the medius in the former and to the medius 
and annularis in the latter. Like the extensor longus this 
muscle is remarkable for often having an insertion into the 
proximal or middle phalanges. It is always supplied by the 
anterior tibial nerve, 


Peroneal Region. 


Peroneus longus.—This muscle in the Hippopotamide (1, 3) and 
Suide (4, 6, 8, 11, 13, 14) rises from the external tuberosity of 
the tibia and the head of the fibula, though sometimes the latter 
origin may be absent; it passes behind the external malleolus 
and across the sole as usual to be inserted into the innermost 
cuneiform. In the other Artiodactyla its apparent origin is always 
from the external tuberosity of the tibia, but in our Harnessed 
Antelope (55) we found that some of its fibres could easily be 
traced upward to the condyle of the femur, forming an oblique 
band of fibrous tissue on the outer side of the long external 
lateral ligament of the knee. In a former paper (‘Joints of 
Mammals,” Journ. of Anat. vol. xxiv. p. 307) one of us has drawn 
attention to the constant twisting of the fibres of this ligament, 
and we have now very little doubt that its superficial layer which 
runs downward and forward is really the remains of the origin of 
the peroneus longus from the femur. Although the Harnessed 
Antelope shows this in the most marked way, it can be made out 
by careful dissection in most Ungulates. The insertion in these 
Artiodactyla is chiefly into the under side of the base of the great 
cannon-bone. 

Among the Perissodactyla the muscle is present in the 
Tapiridee (58, 61) and Rhinocerotide, and its femoral origin can 
easily be traced. In the Equide (63, 64, 65) there is no peroneus 
longus. In the Subungulata it rises entirely from the head of 
the fibula in Hyrax (74, 75) and is inserted into the base of the 
index metatarsal. In the Elephant, Paterson and Dun made out 
a distinct femoral and tibial origin (85), while the same is clearly 
figured by Cuvier and Laurillard (84). Miall and Greenwood, 
however, failed to notice any femoral origin (XXIX.). The 
insertion, according to Paterson and Dun, whose account is 
by far the most exact and detailed of any we have yet seen 


1903. | THE MUSCLES OF THE UNGULATA. 279 


on EKlephant’s muscles, is into the bases of all the metatarsal 
bones from the fifth to the first. The nerve-supply is from the 
external popliteal in the Duiker-bok, Chevrotain, Sheep, and 
Klephant. 

Peroneus brevis.—In animals such as the Lemur and many of 
the Rodents, where a complete set of peroneals is found, the 
peroneus brevis runs from the fibula to the base of the fifth 
metatarsal bone, while the peronei quarti et quinti digitorum 
pass on to the dorsum of the phalanges. Judging by this 
standard, we have no hesitation in saying that we have as yet 
seen no reason to believe that a peroneus brevis is ever present 
in Ungulates, except, perhaps, in the Elephant. It is described 
by a great many authors, but in every case we have found that 
what they call peroneus brevis is continued on to the dorsum of 
the fourth (annularis) digit, and does not even end in the fourth 
metatarsal bone. When one realises how commonly the fibula and 
the fifth metatarsal are suppressed in this order, one is prepared 
to expect a similar suppression of the peroneus brevis. In 
Paterson and Dun’s Elephant a muscle was found which they 
call peroneus minimus, which rose from the fibula and was 
inserted into the caleaneum and base of the fifth metatarsal; this 
we believe is a peroneus brevis, and the only example of it found 
in the Ungulata. The authors call it peroneus minimus because 
they have fallen into the common mistake of calling the peroneus 
quarti digiti the peroneus brevis. 

Peroneus quarti digiti—TVhis muscle is always present in 
Ungulates. When the fibula is well developed, as in Hyrax and 
the Elephant, it rises from the middle third of that bone, but 
when, as in most Ungulates, there is only the head of the bone 
present, the muscle rises from that and often from the adjacent 
part of the tibia. The tendon usually runs in a groove on the 
outer side of, not behind, the external malleolus, and is inserted 
into the extensor longus digitorum tendon of the annularis. In 
the Horse, however, with the suppression of the annularis, the 
peroneus quarti digiti, which is the only peroneal present, shifts 
its insertion on to the dorsum of the medius. We have records 
of the peroneus quarti digiti under various names in the following 
representative series of animals:—Hippopotamus (1, 3), Pig (4, 
6, 7, 11), Peccary (13, 14), Camel (17), Chevrotain (20, 22, 24), 
Deer (25, 26, 27, 29), Elk (30), Musk-Deer (31), Ox (35, 36, 38, 
40) (in the last-named animal, 40, the tendon was inserted into 
the middle phalanx of the annularis toe), Sheep (41, 42, 43, 47, 
48), Goat (50, 51), Antelope (54, 55), Tapir (58, 62), Horse (63, 
64, 65), Rhinoceros (71), Hyrax (74, 75), and Elephant (81, 85). 
The nerve-supply is always the musculo-cutaneous. 

Peroneus quinti digiti.—This muscle is present in the Hippopo- 
tamus (1, 3) and sometimes in the Pig (4), Musk-Deer (31), and 
Elephant (81, 85). It rises from the upper part of the fibula 
and is inserted into the extensor tendon of the fifth digit. Its 
nerve-supply is the musculo-cutaneous. 


280 MESSRS. B. C. A. WINDLE AND F.G. PARSONS ON [ Nov. 3, 


Posterior Tibial Region. 


Gastroenemius.—The two heads of this muscle rise, as usual, 
from just above the condyles of the femur; the inner is generally 
more extensive in its origin than the external and often extends 
over a considerable portion of the popliteal surface of the femur. 
One of the most notable points in this muscle in the Ungulata is 
the absence of fabellee in its origins, The only member of the 
order in which we have found fabelle is the Hyrax, and in it 
only the outer one was present. The tendo Achillis shows the 
characteristic rope-like twisting previously described by one of us 
(Journ. Anat. vol. xxviii. p. 414) very well, and in some animals, 
e. g. the Elephant (79) and Musk-Deer (31), the tendons remain 
separate as far as the calcaneum. In the Chevrotain, Kinberg 
(X.) describes the outer head as having an anterior and posterior 
origin. In the Hippopotamus, Gratiolet and Alix (III.) say that 
the origins are continued down to the tibia. The expansions 
from the thigh-muscles forming a sheath for the gastrocnemius 
tendon have already been described. The nerve-supply is always 
the internal popliteal. 

Soleus—In the Hippopotamide (1, 3) and Suide (4, 6, 9, 11, 
12, 14) this is either wanting or, more probably, has shifted its 
origin upward to the external condyle of the femur and is fused 
with the external head of the gastrocnemius. In Cuvier and 
Taurillard’s Peccary (13), however, its origin is drawn rising 
from the surface of the peroneus quarti digiti. In the Tragulide 
(20, 22) the muscle is distinct, and in our specimen (22) rose 
from the middle third of the fibula. In the Cervidze (29, 30) it 
rises from the external condyle, but is a distinct, though small, 
muscle. In the Bovide (40, 48, 54, 55) it usually rises from 
the rudimentary head of the fibula and joins the outer head of 
the gastrocnemius. In the Tapiride (61, 62) it is condylar and 
very small. Turner (X VI.) failed to find it at all. In the 
Equide (63, 64, 65) it rises from the external tuberosity of the 
tibia. We have no record of it in the Rhinocerotide. 

In the Procaviide (74, 75) it comes from the head of the fibula 
and is fairly well developed. In the Elephantide (74, 77, 79) it 
is also well developed, rises from the head of the fibula and is 
inserted separately into the caleaneum. 

From the foregoing it will be seen that this muscle in the 
Ungulata is subject to a good deal of variation, but in all except 
the Subungulata is feebly developed. In the Hippopotamide, 
Suid, Cervide, and Tapiride its origin is usually shifted up to 
the external condyle, while in the other families it rises either 
from the fibula or from the adjacent external tuberosity of the 
tibia. Its nerve-supply is the internal popliteal. 

Plantaris—This muscle is always present in the Ungulata. 
In the Hippopotamus (1) it has a very large muscular belly rising 
from above the external condyle of the femur; this tendon in the 
sole has a few fleshy fibres on its deep surface and is the only 


1903. ] THE MUSCLES OF THE UNGULATA. 281 


instance, with the exception of Hyrax, we have met with in the 
whole order of any trace of a fleshy flexor brevis digitorum. 
Slips are given off which act as flexores perforati to all four 
digits. In the Suide (4, 7, 11, 12, 13, 14) the fleshy belly in 
the calf is also fairly large and is wrapped up in the outer head 
of the gastrocnemius; passing round the tuber calcis its tendon 
gains the sole and usually divides into three slips for the index, 
medius, and annularis digits, though occasionally (12) it goes to 
all four digits. In the rest of the Artiodactyla its fleshy belly 
is much smaller, and its tendon, after passing round the pulley- 
like surface on the back of the tuberosity of the caleaneum, enters 
the soleand gives off a loop from its deep surface which surrounds 
the flexores tibialis et fibularis, then the tendon divides for the 
medius and annularis digits, and each slip acts as a flexor per- 
foratus before being inserted into the middle phalanx. In the 
Ox especially the perforation of the tendon takes up a good deal 
of its length, and a section through the perforans and perforatus 
tendons shows the former enclosed by a complete ring of the latter. 

In Dorcatherium (23) Chatin describes tendons to all four 
digits. In the Perissodactyla the muscle rises from the supra- 
condylar fossa and passes to the middle phalanges of all the toes, 
7.e. one in the Horse and three in the Rhinoceros and Tapir. 
In the Subungulata, Hyrax (74, 75) has a fleshy flexor brevis 
developed in the sole. The nerve-supply is always the internal 
popliteal. 

Popliteus.—There is but little of special interest about this 
muscle in the Ungulata; it rises as usual from the outer surface of 
the external condyle and is inserted into the upper quarter or third 
of the posterior surface of the tibia. Asa rule no sesamoid is 
developed in its tendon, though we found one in our Tapir. The 
anterior tibial artery apparently always passes in front of the 
muscle. With regard to the nerve-supply the Duiker-bok (54) 
and Harnessed Antelope (55) have a branch from the internal 
popliteal entering the muscle at its lower border but not curving 
round that border as in Man, in addition there are two or three 
smaller branches entering the posterior surface. 

Flexor fibularis (Flexor longus hallucis of human anatomy).— 
This, except in the Camel and Elephant, is much larger than the 
flexor tibialis, and rises from the posterior surface of the tibia a 
little lower than the latter; it winds round the internal malleolus 
and is soon afterwards joined by the flexor tibialis; it then 
divides into a variable number of tendons which pierce those of the 
flexor brevis digitorum or continuation of the plantaris, and are 
inserted into the terminal phalanges. In the Hippopotamidee, 
Suide, and Dorcatheriwm there are four tendons, in the other 
Artiodactyla only two; in the Rhinoceros, Tapir, and Hyrax 
three, in the Horse one; and in the Elephant the combined 
flexores tibialis et fibularis give tendons to all five digits, though 
the chief contribution is from the tibialis. It is always supplied 
by the posterior tibial nerve, 


282 MESSRS. B. C, A. WINDLE AND F.G. PARSONS ON [ Nov. 3, 


Flexor tibialis (Flexor longus digitorum of human anatomy).— 
This rises from the upper part of the posterior surface of the 
tibia, and from the back of the head of the fibula just below 
the popliteus. In every case of which we have records it joins 
the flexor fibularis in the upper part of the tarsus. Asa rule it 
only sends fibres to the tibial digits, while the flexor fibularis 
supplies them all. In the Elephant and Camel, however, the 
flexor tibialis is the larger tendon of the two. Like the flexor 
fibularis it is supplied by the posterior tibial nerve. 

Lumbricales.—Speaking generally, the Ungulates are not well 
provided with these muscles. In the Hippopotamide, Gratiolet 
(II1.) found one for the medius digit. In the Suide they are 
apparently absent, at least they were so in our specimens (11, 14), 
and we find no mention of them by other writers, nor are they 
drawn in Cuvier and Laurillard’s plates (I.). In the rest of the 
Artiodactyla they are, we believe, always absent. In the Perisso- 
dactyla they are more constant. Chauveau (II.) describes two in 
the Horse, one from each side of the flexor perforans tendon. In 
the Tapir (61, 62) there are three, the outermost being much the 
largest. Among the Subungulata, Hyrax (74, 75) has two, which 
pass round the tibial sides of the medius and annularis digits ; 
in our specimen (75) they were both supplied by the external 
plantar nerve. In the Elephant, Paterson and Dun found only 
one small lumbrical, which was on the tibial side of the medius 
digit and was supplied by the internal plantar nerve. Miall and 
Greenwood (XXIX.), however, describe some more superficial 
ones rising from the plantaris and being inserted into the plantar 
sheaths. 

Titialis posticus.— This muscle, when it is present, rises from 
the upper part of the back of the tibia and joins the flexor 
fibularis in the upper part of the tarsus before that tendon unites 
with the flexor tibialis. We have records of it in the following 
animals: Hippopotamus (3), Suide (4, 7, 12, 13, 14), Camel (15) 
——here it is said to form the chief working part of the flexor 
perforans,—Chevrotain (20), Water-Chevrotain (24), Axis Deer 
(25), Ox (35, 36, 40), Sheep (41, 42), Goat (50, 51), Horse (63, 
65). In the Elephant, according to Paterson and Dun, it rises 
from the tibia, fibula, and interosseous membrane, and is inserted 
into the dorsum of the bases of the second and third metatarsal 
bones as well as slightly into the first. Miall and Greenwood 
(XXITX.)and Anderson (XX VII.) agree with this, though neither 
of them found any insertion into the first metatarsal, but in 
Anderson’s specimen it went to the fourth in addition to the 
second and third. It is certainly absent or quite fused with the 
flexor fibularis in many ungulates; thus Bronn (VI.), Meckel 
(VII.), and Cuvier say that it is absent in the order, and in the 
following animals it could not be made out :—Chevrotain (22), 
Brocket Deer (29), Gazelle, Sheep (47), Duiker-bok (54), Har- 
nessed Antelope (55), Tapir (56, 61, 62), and Hyrax (74, 75). 
Its nerve-supply is the posterior tibial. 


1903.] THE MUSCLES OF THE UNGULATA. 283 


Plantar Region. 


Flexor brevis digitorum pedis.—This rudimentary structure 
has already been described with the plantaris, of which it is the 
continuation into the sole. 

The Accessorius is always absent, the only approach to it is a 
small fleshy bundle seen by Mivart and Murie in Hyrax and by 
ourselves in the Ox (40). 

Deep Plantar Muscles—In the Hippopotamus (1, 3) there is 
an abductor indicis, an abductor minimi digiti, and double-headed 
flexores breves for each toe. 

_ In the Suide there is a greater suppression and evidently con- 
siderable individual variation. Lesbres (V.) describes only a few 
fleshy fibres on the outer and inner side of the foot, while 
Chauveau (II.) says that four interossei are present. In our 
specimen of Red River-Hog (11) we found one interosseus going 
to the tibial side of the medius, one to the fibular side of the 
annularis, and one to each side of the minimus, in addition to 
a feeble abductor minimi digiti. Im the Peccary (14), on the 
other hand, there were only three muscles, that to the tibial side 
of the minimus being absent. In the rest of the Artiodactyla it 
is usual to find only two double-headed flexores breves, each head 
having a sesamoid bone developed in it; there are, however, no 
fleshy fibres in these structures. An exception is the Water- 
Chevrotain, in which Chatin (XI.) describes four interossei, and 
from his figure there appears to be a double-headed fleshy flexor 
brevis to each toe. The increased development is not surprising 
when one remembers how large the lateral digits are in this 
animal. In the Tapir (61) we found a double flexor brevis to 
each toe, and Murie (X VII.) agrees with this but describes in 
addition two superficial muscles to the medius. In the Horse two 
tinterossei (sic) are described by Chauveau (II.) and Cunningham. 

Among the Subungulata the sole-muscles are more numerous ; 
Hyrax (75) has an adductor indicis and annularis on a superficial 
plane and deep to them three double flexores breves, while in 
addition to these there is an abductor indicis and quarti digiti. 
In Mivart and Murie’s specimen all these were present as well as 
an adductor medi on the superficial plane. 

In the Elephant (85), Paterson and Dun found an abductor 
hallucis rismg from the caleaneum, and a “flexor brevis digiti 
secundi,” which comes from the tuber calcis and covers the long 
tendons entering the sole. There is some reason to think that this 
represents the flexor brevis digitorum of Man, save that its 
insertion is into the metatarso-phalangeal capsule instead of into 
the middle phalanx. The minimus hasan abductor and a double- 
headed flexor brevis. The four interosseous spaces are well sup- 
plied with muscles which do not seem to be arranged as double- 
headed flexores breves. Without an accurate diagram it is 
difficult to say whether any of these belong to the more super- 
ficial adductor series of the mammalian foot, or whether they are 


284 MESSRS, B. C. A. WINDLE AND F.G. PARSONS ON [ Nov. 3, 


flexores breves dislocated into plantar and dorsal imterosse1 as in 
Man. The other authorities on the Elephant give us little help, 
and we must reserve the question for future investigation. 


Muscles of the Trunk. 


Serrati dorsales.—The anterior and posterior muscles can 
usually, though not always, be distinguished, and it is probably 
owing to the great difficulty in deciding how much was anterior 
and how much posterior that the great discrepancies in different 
authors’ accounts have occurred. The origins of the two muscles 
are from the thoracic and anterior lumbar spines. The following 
are some of the recorded insertions of the anterior muscle :-— 


Hippopotamus (1) ............ 3. ribs. 
Bios (A) aes n tc asian eee ee a pte  nhas 
Olnensaxounnan (PO) geedoscnadenes DO 6) Vs 
a (22) aschih copes aes 3-12) 4; 
Oc GON Mess insecieiand tous eee Davy ie: 
Sheep (42) wreeccrt aces eeeet HO 4 
roan) loyar® (5D) Wesoscoueensosccode Aen 
Tela (GS OD)" we onbaseesondes 5-14 ,, 
Tehyaeeme (4 WS) US) Vesedes Bee Soe Ae). 
Elephant (84) .................. Hertel ets 
The following are those of the posterior muscle :— 
Hippopotamus (1) ............ 4-15 ribs. 
1 ae (7 ai anaes teea ees Reyes 8-14 ,, 
Olneraxonennn (70) sgabssseseaess. Gr 2s 
(O30) (Gia) Maen saa Nm ke Mat ce 10-13 ,, 
Sheep (AD) casecee nus ces eee 10-13 ,, 
Ambelope (00). seeccenasse teers HORT 3. 
Mlepheamits(S4)) pecceasscre cee 9-22 ,, 


In the Chevrotain (22) and Hyrax (74, 75, 78) the muscle 
formed one single sheet and it was impossible to distinguish 
anterior and posterior parts. 

The nerve-supply is from the posterior primary divisions of the 
thoracic nerves. 

Splenius capitis—This muscle varies a good deal in different 
animals; it rises from several of the anterior thoracic spines, 
but never seems to reach farther back than the 6th. In the 
Giraffe it is entirely absent according to Murie (X XXII.), while 
in the Camel (15, 18) it is so small as easily to escape notice. In 
the Ox (36) and Sheep (44) it is also very small. In the Hippo- 
potamus (1) and Pig (4) it is fairly well marked, and in the 
latter animal Lesbres (V.) found it divided into two portions, 
one being inserted into the occipital crest, the other into the 
mastoid portion of the temporal. In the Elephant, Miall and 
Greenwood describe it (X XIX.) as bilaminar, while in the Horse 
(63, 65) and Hyrax (74, 75) it is well developed. 

Splenius colli.tIn the Artiodactyla this musele, when present, 


1903. ] THE MUSCLES OF THE UNGULATA, 285 


is small and is usually only inserted into the atlas, though in the 
Ox (36) it reaches the axis as well. It is absent in the Camel 
(15, 18) and Giraffe (33), though present in the Chevrotain (20). 

In the Perissodactyla, judging from the Horse (63, 65, 67) and 
Tapir (62), it is specially strong, and in the latter animal is 
inserted into the six anterior cervical transverse processes. Among 
the Subungulata it is present in Hyrax (74, 75), but absent in 
the Elephant (81). 

Erector spine.—The outer portion of this muscle, which in 
Man is composed of the sacro-lumbalis, accessorius, and cervicalis 
ascendens, is not well developed in Ungulates except the Sub- 
ungulata (Hyrax and Elephant); the sacral origin is very feeble 
and is represented only by a delicate fibrous band rising from 
the common origin on the dorsal surface of the sacrum. ‘The 
accessorius 1s the best developed part, and rises from the hinder 
ribs near their angles to be inserted into the same place on the 
anterior ribs. ‘The cervicalis ascendens is represented only by 
one slip going to the transverse process of the 7th cervical 
vertebra. Although these muscles are feeble they are usually 
quite clearly marked off from the longissimus dorsi. In the Horse 
the accessorius and sacro-lumbalis are less well developed than 
in the Artiodactyla, and, according to Chauveau (II.), are in- 
separable from the longissimus, though other authors were easily 
able to distinguish the two. 

The Longissimus dorsi is always well developed and has an 
external series of attachments just internal to the angles of the 
ribs, and an internal to the thoracic transverse processes; this 
internal series of attachments is continued up into the neck as 
the transversalis colli, trachelo-mastoid, and complexus. 

While the spinalis colli is very strong, the spinalis dorsi is less 
well developed but still quite distinguishable. The semispinalis 
is well developed, especially in the neck. 

The Z’ransversalis colli is always present, and is inserted by a 
series of slips into the first five cervical vertebre. 

The Trachelo-mastoid (transversalis capitis) is, as usual, a con- 
tinuation upward of the longissimus dorsi and is inserted into 
the paramastoid process of the temporal bone; it is closely con- 
nected with the transversalis colli. Lesbres (V.) says that it 
divides at its insertion, one part being attached to the skull and 
the other into the transverse process of the atlas. From our 
own dissections we feel sure that Lesbres has included ‘a slip 
of the transversalis colli in his description. In the Chevrotain, 
Kinberg (X.) says the muscle is absent, but we found it in our 
specimen (21). In the Hyrax (75) we noticed that the muscle 
continued to rise in the neck from the articular processes of the 
cervical vertebre. 

Complexus.—This muscle rises from the transverse processes of 
the anterior thoracic vertebre, varying from the 3rd to the 10th, 
as well as from the articular processes of the posterior cervical 
vertebre ; it is inserted into the skull just below the occipital 


286 MESSRS. B. C. A. WINDLE AND F.G. PARSONS ON | Nov. 3; 


erest. In the Hippopotamus (1) and Pig (4) it is very large, and 
in the former reaches back as far as the 10th thoracic vertebra ; 
in the Pig it is divided into a central and lateral bundle, the 
former of which corresponds to the biventer cervicis of other 
mammals although no central tendon is present. In the Camel 
(18), Giraffe (33), and Chevrotain (20) the biventer forms the 
greater part of the muscle and is really a biventer. In the Ox 
(36, 37), Sheep (41, 44), and Antelope (55) the two bundles are 
present, but the inner or biventer does not deserve its name as 
it has no central tendon. In the Horse (63, 67) the two parts 
are present, and four oblique myocommata are found intersecting 
one or both. In the Tapir (55) Murie makes no mention of a 
biventer, though he certainly knew of the existence of such a 
muscle. In our own Tapir unfortunately the deep neck-muscles 
could not be examined. In our specimen of Hyrax (75) the 
muscle was composed of three distinct parallel bundles; of these 
the innermost was continuous with the longissimus dorsi, the 
middle rose from the anterior thoracic transverse processes, while 
the external came from the articular processes of the hinder 
cervical vertebre. All three were inserted together into the 
occipital bone, and a single myocomma was present in each of the 
external and internal bundles though not opposite one another, 
This description differs a good deal from that of Mivart and 
and Murie (X XIV.) in the Hyrax; the arrangement they found 
differs so much from that of any other ungulate that we fancy it 
must have been abnormal. 

The Multifidus spine, Interspinales, and Intertransversales show 
nothing of special importance. The description in Man applies 
to them very well. 

Suboccipital Muscles.—'These have the usual mammalian 
arrangement: the posterior (inferior) oblique is very massive, and 
the rectus capitis dorsalis (posticus) major, as is so frequently the 
case, is a double muscle consisting of an external and an internal 
part; the latter Lesbres (V.) calls rectus capitis posticus medius, 
and he says it is always present in Ungulates. Meckel (VIL), 
however, says that in the Sheep only a single muscle is present, 
and we are so far inclined to agree with him that in our speci- 
men (46) we found the separation a very arbitrary one. 

The Kectus capitis dorsalis (posticus) minor and lateralis are of 
fair size and have the attachments as in Man. 

Triangularis sterni.—This muscle, as Lesbres (V.) remarks, is 
better developed in the Ungulates than in Man. Its origin is, 
as usual, from the caudal portion of the dorsal surface of the 
sternum. In the Chevrotain (20), Duiker-bok (54), and Horse 
(63) it is inserted from the 2nd to the 8th rib, in the Harnessed 
Antelope from the 2nd to the 9th. 

Diaphragm.—This muscle in Ungulates differs in a good many 
respects from that of Man. The costal origins usually come from 
about half the total number of ribs, and where they are inserted 
into the central tendon each digitation can still be distinguished, 


1903.] THE MUSCLES OF THE UNGULATA. 287 


so that the junction between the flesh and the tendon has a 
scalloped appearance. There is only one crus, which rises far 
back in the lumbar region; in the Harnessed Antelope (55) it 
comes from the last lumbar, it runs forward as a long narrow 
tendinous band and becomes fleshy just before the aortic opening ; 
here the main part passes to the right of the aorta and later on 
divides to form the cesophageal sphincter; the part which passes 


Text-fig. 27. 


x 
i 
Wes 
" 
vs SZ 
yy Bry 
yh ROH 
“4 es 
4 , BY, 
Fes “i RY 
cy ti ey 
G f 
: All 
V4 
"| 
Y | 
| 
4 
y | y 
4 44 Y} 
4 WA 
H 
Wy Y Hoh | 
Mth eee | 
94944 


Diaphragm of Harnessed Antelope. (Abdominal surface.) 


A. Crus. F. Costal origin. 

B. Aortic opening. G. Psoas. 

C. Csophageal opening. H. Retractor ultime cost. 
D. Caval opening. I. Last (18th) rib. 


E. Central tendon. 


on the left side of the aorta is much smaller, and in some animals, 
e.g. the Chevrotain (21), is practically absent. There is only one 
arcuate ligament on each side, since the quadratus luamborum is 
quite concealed by the psoas. The caval opening is on the right 
side of the central tendon as usual. In the Horse, Chauveau (I1.) 


288 MESSRS. B. C. A. WINDLE AND F. G. PARSONS ON | Nov. 3, 


= 


says that the costal origins do not interdigitate with the trans- 
versalis. 

Obliquus abdominis externus.—This muscle usually rises from a 
large number of ribs, as the following list shows :— 


Je Choy ooimroaviss (IL) 5) -ecagacosascseasanb one 4-15 
Pye? (hI bageandsosbodstsoposex022 28600056 7-13 
@heyrotaria (2050212) ieee aeeee reece eet 6-13 
Shovereyo) (AES) Snesocacoceeaossocacnucaducd on 6-13 
Harnessed Antelope (55) ............ 5-12 
iDyunilieereal oxalic (G45) | Bebe -cubossdeqoeos ces 6-14 
Nayar (82) eabcodpe occoses: so2neccaped je 4-18 
la lovaste) (Sa) a sarbeaacnenoseconcacdseccnce 418 
leRymaahe (1D) > pepasonseccodasct4oerce sce 5-22 
bilepiamt s(Si)) Weer meee ee ere eeere 3-19 
JM keyoloewaNe (1) \ Gesooosdscoonccocoddndec 3-20 


The insertion into the crest of the ilium is very shght, so that 
Poupart’s ligament is not a well-defined structure as in Man. 
Inthe Horse the external abdominal ring is a large oval structure, 
but in the Ruminants it is a mere slit, the two pillars joining at 
the pubic attachment. The muscle is largely supported by the 
tunica elastica which lies superficial to it and is specially well- 
developed in the Solipeds, Ruminants, and Elephant. 

The Supracostalis is the forward continuation of the external 
oblique; we have little doubt that it is always present in Ungulates, 
although it is so thin and transparent that it requires careful 
looking for, and many authors make no mention of it. It rises 
from the sternum opposite the second and third ribs and passes 
forward and outward, covering the anterior part of the rectus to be 
inserted into the first near the junction of the bone and cartilage. 

Obliquus abdominis internus.—This has a definite origin from 
the iliae crest and lumbar aponeurosis; it is much more fleshy in 
the hinder than in the fore part of the abdomen and is inserted 
into the cartilages of the hinder ribs. Myocommata are frequently 
found extending into it. 

Retractor ultime coste.—Chauveau (II.) describes in the Horse 
a small oblique muscle lying dorsal to the quadratus lumborum, 
rising from the anterior lumbar costal processes and being inserted 
into the posterior border of the last rib; he regards this as part of 
the internal oblique sheet, and we have little doubt that heis right 
since it is entirely in series with the internal intercostals. We 
have found the muscle in the Cervide, Bovide, and Tragulide, and 
believe that it is present in all Ungulates (see text-fig. 27, H., 
p. 287). 

The Vransversalis abdominis requires some little care to 
separate from the internal oblique, but, unlike that muscle, it is 
more fleshy in the anterior part of the abdomen, and is usually 
attached to the deep surfaces of about half the ribs—in the Chevro- 
tain and Harnessed Antelope to the last six, in the Tapir and 
Horse to the last eleven. 


1903. ] THE MUSCLES OF THE UNGULATA. 289 


Rectus ventralis (R. abdominis).—This, as usual, rises from the 
ventral aspect of the pubis near the symphysis and is continued 
forward to the first rib. Ventral to the anterior two or three ribs 
it lies deep to the supracostalis, and here its fleshy fibres usually 
cease and it is continued to the first rib by a thin aponeurosis. 
Many authors describe it as ending at the third, fourth, or fifth 
ribs, but wherever we have been able to repeat their dissections 
we have always been able to make out a delicate continuation of 
the muscle to the first rib. The number of linez transverse varies 
from five to ten but eight or nine isthe commonest number; they 
are, however, not well marked, and Steel (XIX.) has pointed out 
that in the Ass their number varies in individual specimens. 

Pyramidalis.— We have never seen this muscle in Ungulates, 
and our experience agrees with that of Lesbres (V.) and Meckel 
(VII.). Mivart and Murie (X XIV.) were certain of its presence 
in Hyrax, but in our specimen it was undoubtedly absent. 

Allthe preceding ventro-lateral muscles of the belly-wall are 
supplied by the intercostal nerves. 

Quadratus lumborum.—This is a much narrower muscle in the 
Ungulata than in Man, and is attached posteriorly, by a narrow 
tendon, to the sacro-iliac joint or to a tubercle on the ilium close 
by. Anteriorly it is inserted into the lumbar transverse processes, 
and usually into the heads of several of the last ribs. In the 
Hyrax (74) it goes to the posterior twelve ribs, in the Chevrotain 
(20) into five, in the Horse (64, 65), Pig (4), and Elephant (81) 
into two, while in the Bovide its costal insertion seldom extends 
beyond the last rib. 

Psoas magnus.—This rises from the transverse processes of all 
the lumbar vertebre as well as from the sides of the bodies, it often 
also rises from a few of the lower thoracic bodies; its insertion is 
as usual into the lesser trochanter. In the Hippopotamus (1) it 
comes from the last two thoracic as well as the lumbar vertebre. 
In the Pig (9, 11) it is not well developed and only comes from 
the lumbar region. Jn the Elephant (81) its attachment seems to 
be the most extensive, as in that animal it rises as far forward as 
the last four thoracic vertebre as well as the last four. ribs near 
their heads. 

The Psoas parvus was present in every animal of which we have 
records except the Red River-Hog (11), and we have no doubt of 
its being a very constant muscle in Ungulates; it rises from the 
bodies of the last three or four thoracic, “and several of the lumbar 
vertebre ; it is inserted by a ribbon-like tendon into the ilio- 
pectineal eminence. 

The /liacus is a small muscle and is laterally campressed ; it 
rises from the iliac fossa, the ventral sacro-iliac ligament, and the 
margin of the sacrum. Before its insertion it blends with the 
psoas. Maurie, in the Giraffe (XX XII.), noticed that the iliac and 
sacral origins remained separate for some time, and we found the 
same in the Harnessed Antelope (55), though it certainly 1s not 
always the case in Ungulates. 


Proc. Gi, INT INO; DUDS 19 


290 MESSRS. B. C. A. WINDLE AND F.G. PARSONS ON [ Nov. 3, 


Perineal and Caudal Regions. 


Sphincter ani—This muscle has the human attachments in 
male animals, but in females it is continuous with the sphincter 
vagine, forming the sphincter cloace. This is certainly the case 
in the Mare, the Sheep, the Chevrotain, and the Hyrax. 

Transversus perinei.—Thompson (XXXVI.) says that this 
muscle is absent in Ungulates; it is described, however, by Kinberg 
in the Chevrotain (X.) and by Chauveau in the Horse and Bull 
(II.) We have seen it in the Sheep, but we agree with Thompson 
that it is not well marked. 

Bulbo-cavernosus.—As the bulb is a bifid structure in Ungulates, 
the two halves of the muscle are separated by a wide raphe and do 
not unite in the middle line. 

Tschio-cavernosus.—This is always well developed and rises from 
the inner side of the tuber ischii, it then turns forward, wrapping 
round the crus penis or clitoridis. Thompson gives a good illus- 
tration of it in the Camel (XXXVI.). 

The Compressor wrethre surrounds the membranous portion of 
the urethra and consists, as in Man, of an upper and lower bundle ; 
it differs from the human muscle in not being definitely attached 
to the pelvic wall. 

Levator ani.—Thompson (XXX VI.) has shown that the levator 
ani of Man is a compound muscle consisting of two modified tail- 
muscles—the ilio- and pubo-coccygeus supplied by the sacral plexus, 
and an ischio-anal muscle which is a derivative of the sphincter 
cloace and, like that muscle, supplied by the pudic nerve. He 
points out that in the Ungulates only the ischio-anal muscle is 
present, and in this we can agree with him, though Gratiolet (III.) 
describes an ilio-coccygeus in the Hippopotamus and says that it is 
inserted into the anterior three caudal transverse processes. 

Ischio-coccygeus (Coccygeus).—This is well developed in the 
Artiodactyla, and rises from the pelvic surface of the ischial spine 
or the region where that structure would be, as well as from the 
sacro-sciatic ligament. It is inserted into the anterior caudal 
transverse ‘processes. Lesbres (V.) points out that in the Equide 
it is not well developed, and Chauveau (I1.) agrees with this. Joly 
and Lavocat (XII.) deny its presence in the Giraffe, but Murie 
definitely found it in that animal. It is supplied by the sacral 
plexus. 

Sacro-coccygeus superior (Levator caudee; Extensor caudse),—This 
rises from the sacral spines, and is inserted into the dorsal surfaces 
of the caudal vertebre by a series of separate tendons. In those 
Ungulates in which the tail is well developed, a division into 
external and internal layers may be made out as in other mammals. 

Sacro-coccygeus lateralis is the continuation backwards of the 
semispinalis layer of the erector spine. 

Sacro-coccygeus inferior (Depressor caude ; Curvator coccygis).— 
As Lesbres points out (V.), there are external and internal parts of 
this muscle, the latter being slightly developed and reaching only 


1903. ] THE MUSCLES OF THE UNGULATA. 291 


to the 6th or 7th caudal vertebra, while the former runs to the 
tip of the tail. 

The Jnterspinales and IJntertransversales caude are feebly 
developed and diminish as the bones decrease in size. 


ADDENDUM. 


Orbital Muscles.—In addition to the recti and obliqui of human 
anatomy, there is a retractor bulbi or posterior rectus which sur- 
rounds the optic nerve and is inserted into the posterior part of 
the eyeball. This in the Chevrotain (20, 21) is divided into an 
upper and lower part, while in the Horse (63) and Ass (69) the 
division is more complete, for it is divided into four slips corre- 
sponding to the four recti. In the Horse (63) and Ass (69) there is 
also a middle oblique muscle which rises from the inner wall of the 
orbit below the pulley and later on joins the superior oblique; it 
is apparently a part of this muscle which does not pass through 
the pulley. We have looked for this muscle in the Artiodactyla 
but have never seen it, nor do we know whether it occurs in the 
Rhinoceros and Tapir among the Perissodactyla. — 


GENERAL SUMMARY. 


We now propose to consider whether a study of these muscles is 
of any value from a classificatory point of view, and it will perhaps 
be best to collect first those myological points which are common 
to all Ungulates though not common to all mammals. 


Myological Characteristics of the Ungulata. 


1. Almost constant presence of masto-styloideus. 
2. Absence of epitrochleo-anconeus. 
3. Rudimentary condition of pronator radii teres. 
4. Absence of pronator quadratus (except in Tapir). 
5. Position and function of extensor carpi ulnaris as a flexor of 
the carpus. 
6. Absence of supinator brevis. 
7. Insertion of caudo-femoralis into patella. 
8. Semi- and presemimembranosus fused almost to insertion. 
9. Marked expansions from hamstrings to caleaneum forming 
a sheath for the tendo Achillis. 
10. Only an ischial origin for the biceps femoris. 
11. Usual absence of bicipiti accessorius. 
12. Constant presence of straight and reflected heads to rectus 
femoris. 
13. Usual presence of femoral origin of tibialis anticus (Sub- 
ungulata the exception). 
14, Usual presence of femoral origin of extensor longus digitorum 
(Elephant the exception). . 
15. Peroneus longus can be traced up to femur with external 


lateral ligament. 
19* 


292 MESSRS. B. C. A. WINDLE AND F.G. PARSONS ON [ Nov. 3, 


16. Almost constant absence of peroneus brevis. 
17. Absence of accessorius pedis. 

18. Constant presence of trachelo-mastoid. 

19. Absence of pyramidalis. 


We will next consider whether the division of Subungulata 
(comprising the Hyrax and Elephant) has any special points dis- 
tinguishing it from the Ungulata vera. 


Myological Characteristics of Subungulata. 


— 


Semimembranosus and presemimembranosus have ischial and 
caudal heads, but these are also found in the Perissodactyla. 

Sartorius always absent in Hyrax and often in Elephant. 

Tibialis anticus has no femoral origin, 

Lumbricales are fairly well developed, but so they are in the 
Tapir. 

Abdomino-humeralis well developed. 

Palmaris longus present. 

Flexor brevis digitorum mants present. 


aS) 


be 


“IS OU 


It will be seen, therefore, that there are not many points 
common to the Hyrax and Hlephant, but nos. 38 and 7 are 
certainly significant so far as they go, 

Between the Artio- and Perissodactyla there are a good many 
rather striking points of difference. 


Mypological Characteristics of the Artiodactyla. 


1. Ectogluteus is inserted into the fascia of the thigh. 

2. Obturator internus rises outside the pelvis, 

3. Obturator tertius (so-called) is present. 

4. Semi- and presemimembranosus have only an ischial head. 
5. Semitendinosus has usually only an ischial head. 

6. Lumbricales are absent except in the Hippopotamus, 

7. Splenius colli is feeble or absent. 


Mypological Characteristics of the Perissodactyla. 


Kctogluteus is inserted into the third trochanter. 

Obturator internus rises inside the pelvis, 

Obturator tertius absent. 

Semi- and presemimembranosus have ischial and caudal heads. 
Semitendinosus has ischial and caudal heads. 

Lumbricales usually present, 

Splenius colli strongly developed. 


ee 


NTS OVE OO 


Speaking, therefore, from a myological point of view, there is 
good reason for dividing the Ungulata vera into two sharply 


defined groups, but there is little reason for placing the Hyrax 


and Elephant together in a separate division, for, excepting the 


absence of the femoral origin of the tibialis anticus and the 


1903.] THE MUSCLES OF THE UNGULATA. 


presence of the flexor brevis digitorum mantis, they have little in 


common. 


The following are some of the myological characteristics of 
the different families of Ungulata, but they will be better 


appreciated by referring to the table :— 


— 


ONIAKPwrwe 


Mm OO ONS OF to bor 


HIPPCPporTaMIDs#. 


. Posterior belly of digastric often wanting. 
. Sterno-masseteric absent or very feeble. 

. Well-developed rhomboideus capitis. 

. Well-developed sterno-scapularis. 

. Coraco-brachialis entirely absent or only the longus present. 
. Supinator longus sometimes present. 


Peroneus quinti digiti present. 


. Soleus absent. 

. Flexor brevis digitorum pedis fleshy. 
- One lumbrical present. 

. Tibialis posticus present. 


SUIDA. 


. Sterno-facialis is superficial to platysma. 
. Posterior belly of digastric often absent. 
. Hyoideus transversus absent. 

. Sterno-thyroid often double. 


Omo-hyoid may or may not have a posterior belly. 


. Sterno-masseteric absent. 

. Dorso-epitrochlearis from axillary border of scapula. 
. Rhomboideus capitis present. 

. Sterno-scapularis present. 

. Subglenoid muscle occasionally present. 

. Coraco-brachialis medius present. 

. Palmaris longus absent. 

. Supinator longus absent. 

. Extensor profundus digitorum sometimes present. 
. Obturator internus rises outside pelvis. 

. Obturator tertius present. 

. Peroneus quinti digiti present. 

. Soleus absent. 

. Tibialis posticus present. 


CAMELID. 


Sterno-mastoid has an insertion into the angle of the jaw. 


. Rhomboideus capitis absent. 

. Coraco-brachialis medius apparently present. 
. Palmaris longus absent. 

. Supinator longus absent. 

. Peroneus quinti digiti absent. 

. Tibialis posticus very large. 

. Splenius colli absent. 


bo 
ge) 
= 


Se ON aR wn = 


eI 


Uy 


MESSRS. B. C. A. WINDLE AND F.G. PARSONS ON __[ Nov, 3, 


TRAGULIDA. 


Omo-hyoid has no posterior belly. 
Sterno-mastoid has an insertion into fascia over masseter. 
Dorso-epitrochlearis from latissimus dorsi or fascia over infra- 


spinatus. 

Rhomboideus capitis absent. 
Sterno-scapularis small, sometimes absent. 
Coraco-brachialis medius. 

Palmaris longus absent. 

Supinator longus absent. 

Obturator internus rises outside pelvis. 
Obturator tertius present. 

Peroneus quinti digiti absent. 

Tibialis posticus may be present or absent. 


CERVID&. 
Omo-hyoid has no posterior belly. 
Sterno-masseteric (mastoid) into anterior border of masseter. 
Omo-trachelian absent apparently. 
Dorso-epitrochlearis from latissimus dorsi or panniculus. 
Rhomboideus capitis absent. 
Sterno-scapularis small, sometimes absent. 
Coraco-brachialis medius or medius and longus. 
Palmaris longus absent. 
Jumbricales absent. 
Obturator internus and tertius doubtful, probably as in 
Tragulide, Giraftide, and Bovide. 
Peroneus quinti digiti absent except in the Musk- Deer. 
Tibialis posticus present or absent. 


GIRAFFID&. 


Two or more tendinous intersections in the sterno-hyoid. 
Omo-hyoid has no posterior belly. 

Sterno-masseteric (mastoid) into anterior border of masseter. 
Omo-trachelian rises from 6th and 7th cervical vertebrie 
instead of from atlas. 

Rhomboideus capitis absent. 

Sterno-scapularis probably absent. 

Coraco-brachialis medius and longus, 

Palmaris longus absent. 

Supinator longus absent. 

Obturator internus rises outside pelvis. 

Obturator tertius present. 

Peroneus quinti digiti absent. 


. Splenius capitis absent. 
. Splenius colli absent. 


Bovips. 
Omo-hyoid has no posterior belly. 


1903. ] THE MUSCLES OF THE UNGULATA. 295 


2. Sterno-masseteric into anterior border of masseter except in 
Sheep. 
3. Dorso-epitrochlearis small or absent. 
4. Infraspinatus secundus sometimes, though rarely, present. 
5. Rhomboideus capitis absent. 
6. Sterno-scapularis small, often absent. 
7. Coraco-brachialis medius or medius and longus. 
8 
9 
10 


. Palmaris longus absent. 
. Lumbricales absent. 
. Supinator longus absent. 
11. Obturator internus rises outside pelvis. 
12. Obturator tertius present. 
13. Peroneus quinti digiti absent. 
14. Tibialis posticus present or absent. 


TAPIRID®. 


Central tendon of digastric absent. 

Omo-hyoid well developed. 

Sterno-masseteric into angle of jaw. 
Dorso-epitrochlearis from latissimus dorsi. 
Rhomboideus capitis absent. 

Sterno-scapularis comes from three costal cartilages. 
Infraspinatus secundus present. 

Coraco-brachialis medius and longus. 

Palmaris longus absent. 

10. Flexor brevis digitorum manus sometimes present. 
11. Supinator longus present. 

12. Pronator quadratus present. 

13. Obturator internus rises inside pelvis. 

14. Obturator tertius absent. 

15. Peroneus quinti digiti absent. 

16. Tibialis posticus absent. 

17. Splenius colli very strong. 


° 


2 COMI D CUR 09 DD 


EQuiIps. 


Vertical slp of panniculus over shoulder, 
Ship of digastric to ramus of mandibie. 
Sterno-hyoid has a central tendon. 
Oino-hyoid well developed. 

Sterno-masseteric into angle of jaw. 
Dorso-epitrochlearis from axillary border of scapula. 
Rhomboideus capitis absent. 
Sterno-scapularis from three costal cartilages. 
. Subglenoid muscle occasionally present. 
Infraspinatus secundus present. 
Coraco-brachialis brevis and medius. 
Palmaris longus absent. 

Supinator longus absent. 

Obturator internus rises inside pelvis. 


YN 


a 
~ 
° 


— — 
LS) C2) 0) SSD Is 


a 
He Co bo 


18. 


5 CO SS 


FSP CN Se 


SUR oe bo 


MESSRS. B. C. A. WINDLE AND F.G. PARSONS ON [ Nov. 3, 


. Obturator tertius absent. 
. A third gemellus is often present. 


Peroneus quinti digiti absent. 
Peroneus longus absent. 
Tibialis posticus present. 


_ Splenius colli strongly developed. 


RHINOCEROTIDE. 
(Further details of this family are badly needed.) 


Sterno-scapularis present (no details). 
Coraco-brachialis medius or longus. 

Palmaris longus absent. 

Supinator longus present. 

Obturator internus apparently rises inside the pelvis. 
Obturator tertius apparently absent. 

Peroneus quinti digiti absent. 


PROCAVIID. 


Abdomino-humeralis well developed. 

Sterno-facialis absent. 

Zygomaticus absent. 

Masto-styloideus absent. 

Omo-hyoid absent. 

Sterno-masseteric enormous. 

Dorso-epitrochlearis from latissimus dorsi and fascia 
infraspinatus. 

Rhomboideus capitis present. 


. Sterno-scapularis from three costal cartilages. 
. Coraco-brachialis medius, sometimes longus. 


Palmaris longus present. 


. Flexor brevis digitorum manus present. 
. Olecranal head of flexor carpi ulnaris well developed. 


Supinator longus absent. 


. No femoral insertion of ectogluteus. 

. Obturator internus rises outside pelvis. 
. Obturator tertius present. 

. Tibialis anticus has no femoral origin. 

. Peroneus quinti digiti absent. 

. Fabella in outer head of gastrocnemius. 
. Flexor brevis digitorum pedis fleshy. 

. Tibialis posticus absent. 


ELEPHANTIDA. 


Abdomino-humeralis well developed. 
Sterno-facialis absent. 

Sterno-hyoid absent. 

Omo-hyoid absent. ; 

Two sterno-masseteric attachments. 


over 


30. 


31, 


. Sterno-masseteric 


. Sterno-scapularis 


. Tibialis anticus 


fU juve Pp. s4!.] 


. Abdomino-humeralis. .......... 
. Sterno-facialis 
bh ZAVFROMTEN MODIS Gos0a5 000008 sceeseese 


5 De Rate cdoopanneaeacoddcsscsiaseo 
. Masto-styloideus 


b SIKH HONG) Goo cnono5 aap He 5 
. Sterno-thyroid 


5 OAOINVOG! oecasovanacnced sovosons 


. Hyoideus transversus 


. Rhomboideus capitis............ 


. Dorso-epitrochlearis ............ 


. Subglenoid muscle......... 


= Coracosbrachialisiees nears 
ee ealimarissl on ouseeeeer yeeeeeee 


. Olecranal head of pasa 


carpi ulnaris .......... 


. Supinator longus 

. Flexor brevis digitorum mantis 
SS arboriliseseesee sere eee ee 
5 IBYOROFANURNEIUE), co cocoon nsooaecoe ano. 


MObburaorstertius mee eee 


. Origin of Semi- and Presemi- } 


membranosus.............. 


. Lumbricales 
5 SOUS (HEIN). o55 seo consed nson 
Seuibialissposticustasseeeeeeeere 


. Flexor brevis digitorum pedis. 


. Extensor longus feet 


(origin) 


Peroneus quinti digiti 


SPOS COW ccc aanano anapnoonncce 


HIPPOPOTAMIDA. 
2 


) 


TAPIRIDA. 


> 


Posterior belly often ‘No central tendon. Two b 
| 
2 a 
Prolonged to jaw. Strong. C 
Single. O} 
No scapular attachment. Some Scapular attachn 
a 
Present. 7 9 
To post. border of ramus. To angle of jaw. Ne 
Present. 
| 
| Se 
Present. 
I 
> From e ; from lats. dorsi. E 
Absent. Sombq, 

Absent or longus only. Mfedius and longus. ‘Br 
Separate muscle. Feeble. F 
Present or absent. Present. 

Pal 
ometimes present. 
Sarre 
SS ie 
Inserted 
= — 
Q Absent. 
re One ae Present. 
ee a Il 
Femoral. Femoral. 
zy Absent. 


Of 


Present. 


ie 


Present. Very stron; 
1 


1903. | THE MUSCLES OF THE UNGULATA. 297 


6. Dorso-epitrochlearis from latissimus dorsi and axillary border 
of scapula. 
7. Rhomboideus capitis absent, 
8. Sterno-scapularis present. 
9. Subglenoid muscle occasionally present. 
10. Coraco-brachialis medius and longus. 
11. Palmaris longus present. 
12. Flexor sublimis digitorum absent. 
13. Flexor brevis digitorum manus present. 
14. Olecranal head of flexor carpi ulnaris well developed. 
15. Supinator longus present. 
16. Extensor profundus digitorum present to index. 
17. Ectogluteus has a femoral insertion. 
18. Obturator internus rises inside pelvis. 
19. Obturator tertius absent. 
20. Extensor longus digitorum has no femoral origin, 
21. Tibialis anticus has no femoral origin. 
22. Peroneus quinti digiti present. 
23. Tibialis posticus present. 
24. Splenius colli absent. 


The Table facing this page will, we hope, be useful to future 
investigators. It shows at a glance the arrangement of 31 of the 
most interesting muscles in the bodies of Ungulates. So far as we 
know, the muscles which are not included have either the gene- 
ralised mammalian arrangement, or else are specially character- 
istic of all Ungulates, in which case they have already been given 
on p. 291. Another point in which this Table may be valuable is 
that it draws special attention to the gaps in our knowledge, and 
this is often the first step towards filling them up. 


BIBLIOGRAPHY. 
T. Cuvier et Laurintarp.—‘ Planches de Myolog-e.’ 
1849. 
II. Cuavuyeau’s ‘Comparative Anatomy.’ Fleming, 2nd ed., 
1891. 


TIT. Grarroter et Atrix.— Recherches sur lAnat. de 
VHippopotame.’ 1867. 
TV. Humpury.—‘ Observations in Myology.’ 1872. 
V. Lespres.— Myologie Comparée.’ Lyon, 1897. 
VI. Bronn.—Klassen und Ordnungen des Thierreichs. 
Band vi. Abtheilung 5. 
VII. Mecket.—‘ Manuel d’Anatomie comparée,’ tome vi. 
- VIII. Vroum.—‘ Recherches d’ Anat. comp. sur le Babirussa.’ 
Amsterdam, 1844. 
TX. Watton.—‘ The Camel: its Anatomy, Proportions, 
and Paces.’ 1865. 
X. Kiyserc.—‘ De Tragulo Javanico.’ Lunde, 1849, 


To face p. 297.} 


TABLE OF THE MYOLOGY OF THE UNGULATES. 


[Proe, Zool. Soc. 1903, Vol. II. 


PETC RCD: PERIERODACT SUZ: SUBUAG Ure 
- HIPPOPOTAMID®. SUIDA. CAMELID#, TRAGULIDA, CERVID&. GIRAFFID A, BOVIDA. TAPIRIDAS, EQUIDA. RHINOCEROTID&. PROCAVIIDA. ELEPHANTIDA. 
1. Abdomino-humeralis  .. P Feeble. P - Strong. 1. 
i} 
I ¥ ll 
2. Sterno-facialis ? Strong. Feeble if present. Absent. 2. 
ll y ll 
3. Zygomaticus .................. Present. Absent. Present. 3. 
4. Digastric .....--..........-.. Posterior belly often vanting. Two bellies. P Two bellies. No central tendon. Two bellies and an extra ? Fan-like. Central tendon may be 4. 
2 slip. present. 
5. Masto-styloidens Present. Absent. Present. 6. 
= | 
6. Sterno-hyoid ............ Prolonged to jaw. Strong. ? Fused posteriorly. Slight. Many tendinous inter- Slight or absent. Strong. Central tendon. P Strong. Absent. 6. 
ections. 
7. Sterno-thyroid - Single. Otten double. Sas Single. 7. 
8. Omo-hyoid ... _ Noscapular attachment. Sometimesa scapular No scapular attachment. Scapular attachment. " No omo-hyoid. 8. 
attachment. il = II 
9. Hyoideus transversus ... ..... Present. Absent. ? ie ? Present. iD Present. P : 2 ? 9. 
10. Sterno-masseteric ........._ To post. border of ramus. Absent. To angle of jaw. To fascia over To anterior border of To angle of jaw and Usually to anterior To angle of jaw. Nesr angle of jaw. P Very large; to surface Two mandibular 10. 
’ we Dacia masseter. masseter, surface of masseter. border of masseter. of masseter. insertions. 
11. Rhomboideus capitis............ resent. mite 2 Present. Absent. 11. 
ya | 
12. Sterno-scapularis . peresentd Probably absent. Present ‘re feeble. Probably absent. Present or absent. Present. 12, 
| 2 ~ 
13. Dorso-epitrochlearis ......... ? Krom seap. or lats. dorsi. ? From infenspinatas o =F) go unicalis or P Feeble; origin very Strong; from lats. dorsi. From scapula. P From infraspinatus and From scapula or lats. 13. 
7 - : fats, i Fe | " : 
14. Subglenoid muscle............... Absent. Sometimes present. gee gle Not recorded. pees Bondi preeen 14. 
\ } F 7 
15. Coraco-brachialis ............... Absent or longus only. Bodine only. Apparently medius only. Medius only. Medius always, longus ey Medius always, longus — Medins and longus. ie: Brevis and medius. Medius or longus. Medius always, longus  Medins and longus. 15. 
16. Palmaris 1 sorouimes, sometimes. ; sometimes. 
h WORT renscesce coc. L Avent, : Presents 16. 
17. Olecranal head of Flexor . N S I 
carpi ulnaris ..........6.... i SeDAxate no usole! | EEE ° Feeble or absent. Present, Apparently absent. Feeble or absent. Feeble. Feeble or absent. Present. 17. 
18, Supinato peeagecrare resent or absent. | il : 
upinator longus Presen aus Present. | Absent. presen Absent? ipraeant 18. 
19. Flexor brevis digitorum mantis | ote Sometimes present. Ae Present. 19. 
PO PSArtOrius)ssccsecsstesseesvevssseee aan 
Present. } Absent. Usually absent. 20. 
21. Ectogluteus.. Inserted i : ; 5 j 
gluteus: nserted into fascia. - P) Inserted into fascia. Inserted Into Ge trochanter. Inserted into fascia. Inserted into femur. 21. 
22. Obturator tertius ........... Present. : ; 
28, Origin of Semi-andP : Exevent. e Present. Present. ee eee . Probably absent. Present. Absent. 22. 
. Origin of Semi-and Presemi- Sef) aed { 
Membranosus.........6... } Seed Only aie OnE ee Ischial and c ovigin. 23, 
oa Ohe a ae Re Present, Pp eee 24, 
25. Soleus (origin)... Remora. ? Fibular. ; Femoral. e ? Pibular. Femoral. Tibial. ° Fibular. 26 
Present or absent. al 


30. Peronens quinti digiti .. 


Splenius colli 


Present or absent. 


Fibrous, 


Absent. 


Present. 


Femoral ae present. 


Absent. 


XXIV. 
XXV. 
ORW IL, 
XXVIII. 
DOXeValniile 
X XIX, 


XXX, 
XXXT. 
XXXIT. 
XXXII. 


XXXIV 


XXXV. 


XOXOXGV Ale 


. OWEN. 
. Bett.— Proc. Zool. Soc. 1876, p. 184. 

. Haveuron.—Proe. Irish Acad. vol. ix. p. 527. 
. TURNER. 
. MovRieE. 
. De Lonecuamps.—Mém. de la Soc. Linn. de Normandie, 


MR. F. E. BEDDARD ON THE [ Noy. 3, 


. Guatin.—Annales de Sci. Nat. (Zoologie), sér. 5, vol. xv. 
oD 2) ?) 


art. no. 12. 


. Jouy et Lavocat.——Mém. de la Soe. des Sci. Nat. de 


Strasbourg, 1846, vol. 111. 
Trans. Zool. Soc. vols. 11. & 11. 


Proc. Zool. Soc. 1850, p. 102. 
Journ. Anat. vol. v. p. 131. 


vol. viii. 


. STEEL.— Proc. Zool. Soc. 1880, p. 2. 

. BeppARD & TrREvES.—Proe. Zool. Soe. 1889, 

. Haveuton.—Proe. Irish Acad. vol. ix. p. 51 
XXII. 

XXII. 


joe fo 

D. 

OwsEN.—‘ Anatomy of Vertebrates.’ 

Macauister.—‘‘ Flexor Muscles of Vertebrate Limbs,” 
Journ. Anat. vol. vu. p. 283. 

Mivart & Murie.—Proe. Zool. Soc. 1865, p. 329. 

Branpt.—Meém. de |’ Acad. des Sciences de St. Pétersb. 
vol. xiv. 1869. 

GrorcE.—Ann. de Sci. Nat. (Zoologie), sér. 6, vol. i. 
art. no. 9, p. 123. 

ANDERSON.—Journ, of Anat. vol. xvii. (1882) p. 491. 

Mayver.— Verhandl. Leopold.-Carolin. Akad. d. Natur- 
forscher, vol. xxii. p. 1. 

Mratt & GREENWooD.—Studies in Comp. Anat. no. 11. 
(1878); also Journ. Anat. vols. xii., xii. 

Wartson.—Journ. Anat. vol. ix. p. 118. 

Youne.—Journ. Anat. vol. xiv. p. 289. 

Muriz.—Ann. & Mag. Nat. Hist. vol. ix. 1872, p. 177. 

Cuvier.—Lecons d’Anat. Comp. 

WInDLE & Parsons.—Proc. Zool. Soc. 1897, p. 370. 

WeELKER.— Archiv fiir Anat. 1878, p. 20. 

P. THompson.—J. Anat. vol. xxxili. p. 423. 


7. On some Points in the Anatomy, chiefly of the Heart 


and 


Megalobatrachus japonicus. 


Vascular System, of the Japanese Salamander, 
By Frank E. Bepparp, 


M.A., F.RS., F.R.S.H., Prosector to the Society. 


[Received September 29, 1903. ] 
(Text-figures 28-34.) 


The anatomy of the Japanese Salamander has formed the 


subject of more than one memoir of late years, and has been 
furthermore treated of incidentally in a number of papers dealing 
with the anatomy of organs and systems of organs in the 
Amphibia generally*. The dissection of two examples of this 


* Osawa (Mitth. Medicin. Facult. k. Japan. Uniy. Bd. v. no. 4, 1902) gives a 
copious bibliography. 


1903. ] ANATOMY OF THE JAPANESE SALAMANDER. 299 


great Amphibian which died in the Society’s Gardens enables me, 
however, to add a few new facts to what is already known of its 
structure. The larger example measures 39 inches, the smaller 
20 inches. The large Wegalobatrachus was injected, and though 
the injection did not run with equal regularity through all the 
branches of the arterial system, certain regions were filled with 
the gelatine, and allow of the description of quite minute arteries, 
some of which are of importance and apparently have not been 
described. 
§ On certain of the Abdominal Viscera. 


About one-third of the length of the right lung is free from 
any mesenteric attachments, and lies freely over the abdominal 
viscera. The rest of this organ, nearly up to the root, depends by 
two mesenteries which are inserted along the interna] edge of the 
lung where the pulmonary artery runs. One of these mesenteries 
attaches the lung to the liver, the other to the aorta posteriorly, 
and to the cesophagus anteriorly after the origin of the gastric 
artery. Anteriorly the two mesenteries arise separately from the 
lung; posteriorly they arise in common and ultimately become 
fused into one membrane, which at the end of the liver is 
continuous with the mesoarium. 

The tip of the left lung is also free from mesentery, but for a 
shorter space than that of the right lung. One membrane only is 
attached to it, which passes to the csophagus and stomach 
anteriorly, and posteriorly joins the mesogastrium. Posteriorly 
the spleen is suspended in the mesogastrium, and between the 
spleen and the lung is a funnel-shaped depression of the mesentery 
up which passes the gastro-splenic artery. Posteriorly, the mesen- 
tery attaching the stomach to the aorta passes into the mesoarium, 
as has been described on the right side of the body. The liver 1s 
furthermore attached to the esophagus and stomach by a mem- 
brane which corresponds dorsally to the falciform lgament 
ventrally. ‘The spleen is borne upon the outside of the hepato- 
cesophageal mesentery. 

The liver therefore underlies a considerable cavity which is 
largely separate from the rest of the abdominal cavity, the 
walls of which are formed by the various mesenteries attached to 
the liver. Anteriorly this cavity extends beyond the liver, and 
reaches up to the wall of the pericardium; its boundaries are 
shown by the dotted line in the accompanying drawing (text- 
fig. 28, p. 300). Its wall is very strong and tough. Posteriorly 
the mesenteries supporting the ovaries and the oviducts are not 
attached separately to the body-wall. The arrangement is as 
follows :—In the middle line is the mesentery supporting the gut, 
which is low in the rectal region where the gut is straight, and 
is higher anteriorly where the gut has a more sinuous course. 
It ends anteriorly upon the ventral surface of the liver on the 
right side of the gall-bladder, beyond which it does not extend. 
It is parallel to the falciform ligament, which is attached on the 


300 MR. F. E. BEDDARD ON THE [ Nov. 3, 


left side of the gall-bladder. It is also attached on the left side to 
the hepato-cesophageal mesentery, and on the right to the pulmono- 
hepatic. Externally to the intestinal mesentery 1s the mesoarium, 
and externally to this again the sheet supporting the oviduct, 
which is much more pigmented than the other mesenteries. _ 


Text-fig. 28. 


Dissection of the base of the liver and lung of Meaalobatrachus on the right side 
of the body, to show arrangement of divisions of Ccelom. 


L, right lung; Liv., liver; Od., oviduct; P, pericardial cavity. 


The mesenteries of all the abdominally situated viscera, there- 
fore, converge wpon the vena cava and aorta. Quite anteriorly, 


1903. | ANATOMY OF THE JAPANESE SALAMANDER. 301 


however, the oviducal mesentery leaves the common dorsal 
mesentery, and goes to the right and to the left to follow the 
oviduct to its internal aperture. The aorta itself is removed by 
quite a long distance from the dorsal median line, from which the 
dorsal mesentery, over an inch wide in parts, suspends it. It 
should be observed that this membrane is continuous and not 
fenestrated as it is in the frog, where the aorta also does not lie 
close to the backbone. Within this mesentery are suspended the 
intercostal arteries and certain portal veins, both of which are de- 
scribed elsewhere (see p. 312). The halves of this mesentery are 
so loosely attached to each other that the slightest pull brings 
them apart. J am disposed to think that this ready separation 
argues a lymph-space or spaces of considerable extent within the 
eavity of this dorsal mesentery. 

The above description of the mesenteries has been given partly 
to serve for a comparison with Menopoma. The differences 
between the two forms are not, however, great. In the first place, 
the left lung is very much shorter than the right in Menopoma. 
In Megalobatrachus it is slightly longer. Nevertheless, in Meno- 
poma the free tip of the lung to which no mesentery is attached 
is proportionately much longer thanin Megalobatrachus. Finally, 
the left lung of J/enopoma is connected only with the cesophagus. 
This membrane, however, runs for a little distance over the 
mesentery attaching the stomach to the aorta, and is a trace of 
the former greater extension of the lung. The gastro-hepatic 
ligament bears, of course, the gastric and cesophageal branches of 
the portal, which show differences in the two animals. In both, 
the cesophageal portal is quite distinct from the gastric, and enters 
the liver at about the middle of its length. In MWenopoma this 
trunk is formed by the union of two equally sized vessels. In 
Megalobatrachus there is but one. In the larger specimen of this 
genus, moreover, the condition was still more specialised; there 
was no separate cesophageal trunk at all. Furthermore, in 
Menopoma the gastric trunk consisted of two main trunks meeting 
near to their entrance into the liver posteriorly; the anterior of 
these has two branches, the posterior three. In Megalobatrachus 
this vessel is more obviously one trunk with many branches. 
These and other reasons (such as the at least general persistence 
of a pair of gill-slits) lead me to dissent from those who would 
unite the American and Japanese Salamanders into one genus, 
Cryptobranchus. 

A sheet of membrane, quite tough in character, extends from 
the right lobe of the liver to the parietes, and arches over a section 
of the body-cavity. Into the pocket formed at the corner of this 
next to the lateral parietes opens the oviduct. The general 
relations of these various parts are shown in the accompanying 
sketch (text-fig. 28). This cavity is entirely floored by the lung, 
and its boundary near the middle line of the body is the 
mesentery connecting the liver with the lung. On the right side, 
and below the shelter afforded by the transverse and horizontal 


302 MR. F. E. BEDDARD ON THE [ Nov. 3, 


sheet of membrane just referred to, is a foramen in the hepato- 
pulmonary membrane, which, whether normal or abnormal, is 
certainly natural. It is a foramen with a perfectly clear margin 
as seen with a lens, and four to five millimetres across. In the 
drawing a probe is represented inserted in this foramen. On the 
left side of the body the membrane which bears the orifice of the 
oviduct arises also from the edge of the liver anteriorly. 

This state of affairs, which has been described by others, is not 
apparent in the smaller, male, individual. In this Salamander the 
base of the lung is closely attached to the lateral parietes on both 
sides of the body bya tough and very short membrane. There is 
no trace whatever of a fold of membrane arching over this and 
bearing the mouth of the oviduct. Evidently, therefore, the 
existence of this fold is associated with the oviduct, for the smaller 
animal was fully mature. 

The sperm-duct could be traced forward to below the shelter of 
the lung, 7. e. where the latter fills up the space between the liver 
and the parietes. Here the membrane supporting it curves 
round in a semicircle to the right as in the larger individual, but 
there was no trace of the oviduct along the semicircular curve. 
On the other hand, what I take to be the end of the sperm-duct 
lay in the same straight line with the rest of the duct*. I may 
remark, furthermore, that the extent and depth, as shown by 
passing a probe above it, of the semicircular membrane is greater 
proportionately in this male individual. A bristle tipped with 
sealing-wax could be passed forwards as far as to a peint beneath 
the pericardium. This extension forwards of the body-cavity 
then comes to be obliterated in the female, or indeed possibly in 
course of growth, perhaps owing to the pressure of the lung. 

On the left-hand side of the body I found precisely the same 
state of affairs. 

$ The Heart. 


The heart, which has been described by Osawa + more accurately 
than by others, lies in a spacious pericardium which is roughly 
of a circular form, and proved to have the following dimensions : 
greatest diameter 65 mm., greatest length 6°5 mm. The heart 
does not by any means completely fill this pericardium. As to the 
general shape of the heart, the ventricle is rhomboidal, the posterior 
angle of the rhomboid lying a little to the right of the entrance 
of the vena cava inferior. The right-hand angle of the rhomboid 
is tied down by a gubernaculum (text-fig. 29, G), not figured by 
Osawa, which is attached partly to the wall of the pericardium, 
and partly to the right vena cava superior. The gubernaculum » 
is broad and can be seen, by passing a probe beneath it and then 

* The sperm-ducts end anteriorly in a small flattened projecting body only 
attached by one end to the membrane which supports it. I find a precisely similar 
body in the large female, but entirely unconnected with any further rudiment of the 


male-duct. This is referred to on p. 315, in connection with the termination of 
the Arterie comites aorte. 


: + “Beitrage zur Anatomie des Japanischen Riesensalamanders,” Mitth. Medicin. 
Facult. k. Japan. Univ. v. p. 221 (1902). 


1903. | ANATOMY OF THE JAPANESE SALAMANDER. 303 


raising it, to pass over (literally, of course, under) the ductus Cuviert 
before becoming attached to its posterior wall and to the posterior 
wall of the pericardium. In Menopoma, on the other hand, I 
observe that the gubernaculum is smaller and less band-like and 


Text-fig. 29. 


Ventral view of Heart of Wegalobatrachus. 
Au., auricle; G, gubernaculum cordis ; Vici. vena cava inferior. 
(For this drawing and text-fig.28 I am indebted to the kindness of Mr. R. Crawshay.) 


is attached at once to the vein without passing beyond it. More- 
over, in the latter Amphibian as contrasted with Megalobratrachus, 


304 MR. F. E. BEDDARD ON THE [ Nov. 3, 


the vein has a much shorter course within the pericardium, and 
leaves that cavity at its posterior corner on each side, instead of 
passing forward and running for some space within the pericardium 
at an angle with its former course. Hyrtl figures veins in the 

lace of the gubernaculum which he describes as ramifying over 
the heart ; but he describes no coronary artery. On the contrary, 
I find a very distinct coronary artery (see p. 308), but have seen 
no coronary veins. In the interior of the ventricle I could find 
no traces of a subdivision into two cavities. The demarcation 
between the pylangium and the synangium is quite visible exter- 
nally, and plainly to be seen in the accompanying drawing (text- 
fig. 29, p. 303). The entire truncus arteriosus measures 43 mm. 
as against 84 mm., which is the extreme length of the ventricle. 
Of this 43 mm., 28 mm. belong tothe pylangium. The pylangium 
is rather plainly divisible into three divisions, which are subequal. 
The first division, that nearest to the ventricle, seems on a dis- 
section to belong partly to the ventricle and partly to the truncus 
arteriosus. Externally, however, as the drawing shows, it appears 
to be clearly a part of the truncus; its smooth walls and sharp 
demarcation from the ventricle mark it out as a portion of the 
truncus. The synangium does not abut upon the anterior wall of 
the pericardium as in Menobranchus*. It divides while within the 
pericardium into right and left halves, which separately perforate 
the pericardium. The diameter of the synangium is very much 
greater than that of the pylangium, about twice as great, and it 
has an almost square outline. 

The auricle is attached to the ventricle for the greater part of 
the anterior left-hand border. 

It is naturally, on acccount of its thin walls, of somewhat 
irregular outline. It is a little difficult to compare the sizes of 
the auricle and the ventricle on account of their different shape ; 
but it appeared to me that the auricle is slightly, but not very 
distinctly, the larger. Whether the auricle does during life I do 
not know, but it can lie entirely on the left hand of the ventricle 
owing to the large pericardium. The sinus venosus is spacious 
and opens into the auricle not far from the opening of the latter 
into the ventricle. It receives below and approximately in the 
middle line the vena cava inferior. On each side opens into the 
sinus venosus a transverse trunk which is, I presume, partly 
ductus Cuviert and partly superior cardinal. Each transverse 
trunk runs along the whole of the transverse diameter of the 
pericardium and in contact with its posterior wall. 

The auricle is very distinctly divided into two cavities (text- 
fig. 30). The larger, which 1 presume to be the equivalent of the 
right auricle, lies nearest to the ventricle into which it opens. The 
sinus venosus opens into this, and its orifice is shielded on the 
side remote from the ventricle by a flap, which so far completely 
partitions the auricle, since it extends from roof to floor, and has 


* Huxley, “On the Struetme of the Skull and of the Heart of Menobranchus 
lateralis,’ P. Z.S. 1874, p. 186. 


1903.] ANATOMY OF THE JAPANESE SALAMANDER. 305 


no fenestre—at least visible fenestrae. This flap appears to be 
purely membranous. Anteriorly it is continuous with the rest 
of the interauricular septum, which, on the contrary, is entirely 
muscular. It is much fenestrated, and consists below of vertical 
pillars passing from the dorsal to the ventral wall of the auricle. 
There is also, particularly on the side adjoming the fibrous part 
of the septum, a mesh-like disposition of the muscular strands 
such as Huxley has figured in Menobranchus*. The completeness 
of the separation of two auricles, not indicated externally, I may 
repeat, is furthermore shown by the fact that they debouch 
quite independently of each other into the ventricle. The septum 
between the two auricles is almost suggestive of a heart-valve in 
being partly muscular and partly membranous. And it may 
possibly be the case that the septum is physiologically something 
more than a mere partition-wall—and incomplete at that—be- 
tween two neighbouring cavities. It may possibly by its own 


Text-fig. 30. 


Auricular septum of Megalobatrachus, viewed from in front. 
The membranous part is dotted. 


contraction, by the contractions of the muscular part playing 
upon the tendinous sheet, do something to stop or facilitate at 
different periods of the heart’s beat the admixture of arterial 
and venous blood before it reaches the ventricle. I greatly regret 
that, having only one large heart at my disposal, I am unable to 
push this matter any further. 

I do not find myself absolutely in accord with the account of 
the interauricular septum given by Hyrtl or with his figure of the 
same. Hyrtl describes the septum as extremely delicate but 
imperforate, and as arching over the atrio-ventricular orifice. It 
is clear that this is only that part of the septum which I have 
described as thin and imperforate. Hyrtl does not refer to the 
muscular pillars which tie down the rest of the thin, and in parts 


* Loc. cit. pl. 32: fig. 5. 


Proc. Zoo, Soc.—1903, Vou II. No. XX. 20 


306 MR. F. E. BEDDARD ON THE | Nov. 3, 


fenestrated, interauricular partition. Nor is there an indication 
of these structures in his figure. Nevertheless, an inspection of 
the drawing which I herewith submit (text-fig. 30, p. 305) will, 
I think, prove that the septum is not so simple as illustrated by 
Hyrtl*. On the other hand, Dr. Chapmany writes:—“It is 
needless to add that the author found the delicate septum sepa- 
rating the auricles incomplete.” But there is no detailed descrip- 
tion, nor does Osawa figure a perforate septum, though he states 
that it is ‘‘ vielfach durchlocherte.” 

The advantage of a large-sized specimen for investigation is 
clearly brought out in the examination of the structure of the 
heart in this Amphibian. I have been able to compare the large 
individual with a much smaller one, not more than a sixth of its 
size. In this Salamander the ventricle did not show any marked 
traces of the characteristic rhomboidal form which is so apparent 
in the large Amphibian ; its contour was indeed of a general oval 
form, much like the heart as figured by Osawa. Furthermore, 
the pylangium only appeared divisible externally into two 
divisions, of which one, that nearest to the ventricle, was not 
more than half the length of the larger distal portion. It became 
naturally a question of interest to determine which of the three 
divisions so plainly visible in the larger specimen were fused 
together to form the single larger barrel-shaped section. I found 
by an examination of the internal structure of the proximal 
shorter part of the pylangium, that this is to be looked upon as 
the first and ventricle-like portion of the pylangium. 

Pylangiwm.—tThe proximal division of this section has for the 
most part such strongly muscular walls that it might be regarded 
as a portion of the ventricle, were it not for the fact of its marked 
constriction from the ventricle and the smooth character of a 
portion of its internal surface. Atthe end of this smooth portion 
is situated the first row of watch-pocket valves, which appear to 
me to be accurately transverse in direction, but not symmetrical 
with reference to the dorsal and ventral lines of the pylangium or 
in their distances from each other. Furthermore two are equi- 
sized and larger than the third. There is also a fourth in the 
specimen dissected by myself. In comparing the pylangium of 
this large Salamander with that of the smaller example, a fact 
which appears to me to be of some little physiological importance 
comes out. The proportions between the conus arteriosus and 
the ventricle are the same in both individuals, but the heart as 
a whole is of course larger in the larger specimen. The valves, 
apparently, are nearly of the same size in the two specimens, and 
thus do not grow, or grow but slightly. Furthermore, in the 
smaller specimen the valves are so nearly in contact, that at 
the requisite times they must occlude the pylangium and thus 


* Cryptobranchus japonicus: Vindobone, mdccclxy. p. 93, pl. xii. fig. 3. 

+ “Observations on the Japanese Salamander, Cryptobranchus maximus (Schlegel),”’ 
Proc. Acad. Nat. Sci. Philad. 1893, p. 227. 

{ According to Hyrtl and Osawa there may be a fifth. 


1903. ] ANATOMY OF THE JAPANESE SALAMANDER. 307 


perform their function perfectly. On the other hand, this could 
hardly be the case with the big Salamander; and it may be that 
the unequal growth of the various parts of the heart is a direct 
cause of death *. 

Concerning the second row of valves I have nothing to add to 
the account given by Osawa, who corrects previous errors 7. 

Synangium.—A. transverse section through the synangium at 
about its middle point shows that the term ‘“synangium” is 
especially applicable to Megalobatrachus. In Menobranchus this 
region of the heart as figured by Huxley ¢ is divided into four 
cavities only, the subsequent subdivisions of which form the several 
aortic trunks. In Megalobatrachus all the arteries may be said to 
arise separately from the pylangium. The transverse section, in fact, 
shows the cavity of the bulky synangium to be primarily divisible 
into a right and left half. Hach half is again divided by septa into 
cavities, which are those of the four arterial trunks. These are 
placed one above the other in a series of stories, and increase in 
size from below upwards, the lowest and smallest being, of course, 
the Carotid and the largest and most dorsal the Pulmonary. The 
vertically-disposed origin of the arteries contrasts with their 
nearly horizontal position on emergence from the common 
sheath §. 

§ Aortic Arches. 


T donot find that my dissections of the arteries arising from the 
conus arteriosus agree altogether with the figures or the descrip- 
tions of Hyrtl and Osawa, even in important details. As will be 
seen from the drawing submitted herewith (text-fig. 33, p. 311), 
there are four branches, as is correctly shown by Schmidt, 
Goddard, and Van der Hoeven in their memoir ||, and by 
Dr. Chapman | and Prof. Osawa **. Nor are the views which 
Hyrtl gives of sections through the conus arteriosus correct ac- 
cording to my own observations. When the conus is cut through 
just before the arteries diverge, the four trunks are seen to 
lie close together in the relations shown in the accompanying 
drawing (text-fig. 31, p. 308). The largest cavity is that of the 
pulmonary trunk; the two next largest are subequal and belong 
respectively to the two aortic trunks. The smallest is that of the 
carotid. I need say nothing of the exact position which each 
holds in the complex, inasmuch as the facts are accurately shown 
in that drawing. If the drawing be compared with the two 
figures given by Hyrtl and with those of Osawa which illustrate 

* See note in ‘ Nature,’ vol. lxviii. p. 497 (1903). 

+ Loc. cit. 

t+ Loe. cit. pl. xxxii. fig. 6a. 

§ Tt will be seen that my drawing does not agree with that of Osawa (Joc. cit. 
pl. xxxvili., B and C). 

|| “ Aanteekeningen over de Anatomie van den Cryptobranchus japonicus,” Nat. 
Verh. Maatsch. Wetensch. Haarlem, deel 19, 1864. It is curious that this error 
should have been perpetuated in Bronn’s ‘ Thierreich’ by the reproduction of Hyrtl’s 
figures. 

4 Proc. Acad. Nat. Sci. Philad. 1893, p. 227. 

** Toe. cit. pl. xii. figs. 1 & 4. 


20* 


308 MR. F. E. BEDDARD ON THE [| Nov. 3, 


the point, it will be seen that in any case the conditions obtaining 
in the example dissected by those anatomists differ from those 
which characterise the example dissected by myself. Moreover, 
Hyrtl shows the pulmonary artery as arising from the third 
aortic arch. 


Text-fig. 31. 


Transverse section through right branch of Synangium of Megalobatrachus. 


P, pulmonary; C, carotid. Between them the two aortic trunks. 


Coronary Artery.—Before leaving this region of the heart it is 
necessary to call attention to the coronary artery, which has not 
been studied by Osawa. ‘The artery in question lies upon the 
dorsal surface of the conus arteriosus, to which region it is 
limited so far as a naked-eye inspection enables me to judge. It 
is much contorted in its course, a state of affairs which, of course, 
allows of the expansion of the conus without unduly distending 
the artery. The coronary artery is exactly median in position, 
corresponding to the internal septum which divides the synangium 
mto two halves. At its origin, however, which is from the 
externally undivided region of the synangium, it is deflected to 
the right, and is seen to communicate with one (the second) of 
the two aortic arches; its orifice is guarded by a strong valvular 
flap. I could not trace the coronary artery with certainty farther 


1993. | ANATOMY OF THE JAPANESE SALAMANDER. 309 


than the posterior end of the conus arteriosus. It seemed to me 
to pass from it to the surface of the auricle near to the opening 
of the latter into the ventricle. 

In the heart of Menobranchus, as Huxley has shown and as I 
have observed myself, the synangium ends in a single (that is, of 
course, apparently single) trunk, which leaves the pericardium in 
the anterior middle line. In Megalobatrachus, on the other hand, 
as I have already pointed out, the synangium gives rise to two 
aortic trunks, which leave the pericardium separately one on each 
side. There is not, however, by reason of this a vacant space left 
between the anterior end of the synangium and the anterior wall 
of the pericardium. On the contrary, a rounded piece of tissue 
of a different colour and appearance to the synangium fastens it 
to the anterior wall of the pericardium. This piece of tissue was 
dissected, and the dissection reveals that the mass of tissue contains 
a cavity, which runs for a very short way beneath the synangium. 
Physiologically, I imagine this structure acts as a kind of buffer 
to prevent impacts during contraction of the heart against the 
anterior wall of the pericardium, and the fact that the “ buffer ” 
is hollow would aid its efficiency in that capacity. 


Ao 
Junction of vascular arches of Megalobatrachus to form dorsal aorta. 


A, vertebral artery; Ao, dorsal aorta; B, carotid arch ; b, deep carotid ; 
C, first aortic arch; ¢, muscular branch ; D, second aortic arch. 


The Dorsal Aorta is formed by the union of three of the 
“branchial” arches. Hyrtl, however, figures* only two of these 
arches as taking part in its formation, viz. the second and third. 
I shall presently deal more minutely with the course and branches 
of the various arteries which issue from the synangium. But the 
broad outlines of their course will be apparent from the drawin 


* Doe. cit. pl. xi. fig. 47. 


310 MR. F. E. BEDDARD ON THE [ Nov. 3, 


(text-fig. 32, p. 309). It will be seen from that drawing that 
three arches actually unite to form the median dorsal aorta, 
and that all of these trunks are pervious where they join ; 
there is not a question of an imperforate Ductus Botalli. The 
injection has enabled me to ascertain this matter, and, moreover, 
the arteries are so large that a dissection allows of the orifices 
of the several tubes being seen with ease. The second and the 
third of the vascular arches are the first to join each other. The 
common aorta thus formed gives off a thickish branch to the 
muscles dorsally shortly after this junction. A little beyond this, 
j.e. nearer to the middle line, the first, or Carotid, arch unites with 
the common trunk. The aorta of each side of the body then gives 
off a branch which perforates the musculature, and is that vessel 
figured by Hyrtl* as running backwards through the transverse 
processes of the vertebre. I have not, however, followed it 
farther than into the first vertebra which it approaches. A little 
after this the two aorte meet and become the single dorsal aorta. 
This junction takes place at about the level of the anterior end of 
the pericardium. The junction of the several aortic arches does 
not seem to be accurately represented by Osawa. For a con- 
siderable distance the aorta is closely adherent to the middle line 
of the parietes, and indeed has even the appearance of being 
sunk a little below it. It is covered with a sheet of fibrous tissue, 
which has to be dissected away to expose the artery. This sheet 
is apt to be a good deal pigmented. The aorta leaves its close 
contact with the muscular parietes just before the transverse 
membranes which tie down the two oviducts to the parietes, 
which membranes come into contact in the middle lne. At this 
point, then, the aorta ascends on to a sheet of mesentery, and the 
ascent increases as the abdominal region is entered, until finally 
the distance of the aorta from the actual middle line of the 
parietes is more than an inch. 

The four aortic arches increase gradually in size, as might be 
inferred from the sections through the median part of the 
synangium and its two branches. The Carotid is the smallest 
and the Pulmonary the largest. 

The course of the Carotid arch is shown in the accompanying 
drawing (text-fig. 33), which does not entirely agree with the 
figure of Osawa t. When it has freed itself from the synangium, 
it passes along more or less parallel to the next arch. It lies 
anteriorly to the first of the branchial muscles and ultimately 
joins, as has been already stated, the common aortic trunk. At 
the point where it is most curved in its course it gives off a strong 
lingual branch (text-fig. 33, A) which runs forward, This branch 
is also attached to the carotid trunk by a pervious connection 
situated nearer to the root of the carotid, and thus allowing of a 


* Osawa (loc. cit. pl. xxxix., vert., & pl. xlii. fig. a, vert.) represents this artery 
as arising from the afferent carotid arch in one figure and from the efferent second 
arch 1m the other. 

¥ Loc. cit. pl. xxxix. 


4 
; 


1903. ] ANATOMY OF THE JAPANESE SALAMANDER. 311 


“short circuiting” of the blood-stream*. On the opposite side 
of the carotid, just where this transverse connection occurs, a 
small artery is given off which also runs forward. It is not shown 
by Osawa, The lingual branch just referred to runs forward and 
ends in two slender branches ; before this termination it gives off 
a backwardly-running branch, which lies parallel to itself, and then, 
bending to the middle line of the body, runs parallel to the trunk 
of the carotid at its origin. It gives off an equally strong branch 
running forwards. The deep carotid artery (text-fig. 32, p. 309) 
arises from the carotid arch just before the junction of the former 
with the common aortic trunk. Close to the lower bend of the 
carotid, just where it gives off the lingual artery, is a small reddish 
body, served by an artery from the carotid (not shown in text- 
fig. 33), which I take to be a part of the thymus. Another body 
of a similar form lies further back in the neck. 


Heart and Arterial Trunks of Megalobatrachus from the side. 


1-4, vascular arches ; A, anteriorly running branch of carotid ; 
Br, branchial twigs; P, main branch of pulmonary. 


The second arch is partly protected by a branchial muscle, and 
a smaller muscle arches over the third branchial trunk. The 
second arch gives off a small branch running upwards, which 
appears to me to correspond with the small branch supplying 
the thymus in the case of the carotid. This is not figured by 
Osawa, who does figure such a twig arising from a connecting 
branch between this and the next arch. 

The third vascular arch again gives off a similar branch ; but 
this joins a twig from the pulmonary, which will be described 
immediately. 

The fourth or pulmonary artery runs a course which is by no 
means so simple as is figured by Hyrtl or Osawa. It bends back, 

and follows almost exactly the edge of the pericardium, The 


* T found this only upon one side of the body. 


Bul MR. F. E. BEDDARD ON THE [ Nov. 3, 


artery gives off several branches of varying importance as to size. 
The first of these is one of the smallest in size but greatest in 
morphological importance; it arises from where the artery has 
begun very definitely to bend backwards, and is directed upwards 
towards the dorsal side of the body. It is, in fact, the Ductus 
Botalli, the link between this aortic arch and the preceding one. 
The principal branch in point of size arises from the same side 
and a little further on. This closely accompanies the pulmonary 
artery, so closely that it is bound up in the same sheath and had 
to be carefully dissected away. It gives off two branches on the 
dorsal side and one on the ventral side. At the base of the lung 
this vessel abandons its close companionship with the main 
pulmonary trunk, and runs along a different face of the lung. 
This branch is not figured by Osawa. 

The Ductus Botalli, as has been already mentioned, joms a 
branch from the third vascular arch, and the conjoined vessels 
give off a number of twigs to the adjacent musculature, which are 
illustrated in the drawing submitted herewith (text-fig. 33,p.311)*. 
I take it that this series of small vessels, which have been referred 
to as existing in all of the vascular arches, are the arteries which 
originally supplied the branchia. There is a final fact about the 
pulmonary artery, to which I desire to refer. The right pulmonary 
artery gives off a branch which supplies the cesophagus. I could 
find no corresponding branch on the left side, and indeed I do not 
think that one exists. I imagine that the various small branches 
of the pulmonary arch and of its large branch collectively represent 
the cutaneous trunk of the Amphibia generally. 

The Pulmonary vein might easily escape attention, as it is not 
at all apparent on a dissection. It lies in fact directly above the 
vena cava inferior, and being of much smaller dimensions, it 1s 
completely covered by that vein when the animal is dissected 
from the ventral surface. The two pulmonary veins join each 
other just within the pericardium, where they form a tolerably 
wide trunk. The common pulmonary vein increases in diameter 
as it passes along the sinus venosus, and finally opens into the 
smaller division of the auricle. There is thus at least a partial 
separation of arterial and venous blood in the heart. 

Intercostal Arteries and Veins.—The very deep fold of peri- 
toneum which ties down the aorta to the dorsal parietes enables 
me to see clearly the course of the intercostal arteries and of 
certain intercostal veins, which are presumably branches of the 
portal system in the abdominal region. Of the arteries I counted 
seven, four of which are arranged in pairs, while the three 
remaining arteries are not so disposed, They arise from the 
aorta, however, definitely on one side or the other; they are not 
median in origin, Furthermore, the points of entrance into the 
thickness of the parietes are not median; they are on one side or 
the other, and in the case of the paired arteries they enter the 


ge LEROHE. Osawa represents nothing more than a simple transverse Ductus Botalli 
uniting the arteries in question. 


1903.] ANATOMY OF THE JAPANESE SALAMANDER. 313 


parietes side by side in a symmetrically paired fashion. It will 
bé seen that this account does not exactly agree with that of 
Osawa. J saw only four intercostal veins, which are disposed 
singly. Two of them lie at intervals from each other in front of 
the first of the intercostal arteries; then follow two arteries; 
after this the third vein; then a pair of arteries, and finally the 
last of the intercostal veins. 

Arterie comites aorte *.—The unpaired dorsal aorta for some 
little distance before and after the origin of the subclavians is 
accompanied, as shown in the drawing (text-fig. 34, ae: 314), by 
a slender aorta on each side, not distant more hem 2 or 3 milli- 
metres from the aortic trunk. At the origin of the subclavians 
these vessels each give off a branch which accompanies in the 
same way its own subclavian. These two vessels arise from the 
aortic trunk between the union of the two aorte to form the 
dorsal aorta and the origin of the subclavians, but much nearer 
to the latter. Hach of these vessels, immediately after its origin, 
gives off a forwardly-running trunk which accompanies the aorta 
on each side for a little distance, and is lost among the muscles of 
the back. The main branches run posteriorly and end upon the 
transverse membrane which ties the transversely-running portion 
of the oviduct to the parietes. Osawa does not appear to have 
studied these arteries. 

Each arteria comes aorte is reinforced during its course by 
other branches arising from the aorta. That of the left side has 
two such supplementary roots. One springs from the aorta just 
in front of the origin of the subclavian ; it or osses over (really, of 
course, below) the vioelriiem. and runs Rowe a considerable distance 
alongside of the first branch before joining it. A second branch 
springs from the aorta on the posterior side of the subclavian. 
This also runs alongside of the two above-mentioned vessels for a 
considerable distance before joining them. The left-hand vessel 
has only the first of the two accessory branches, which arises 
actually from the posterior margin of the subclavian. About 
halfway between this point and the oviducal mesentery the artery 
divides into two branches, of which the inner spreads out over 
that mesentery, while the outer ends in a projecting reddish body 
which is dealt with later (p. 315 

The anatomy of this region of the body in Megalobatrachus is 
in several respects by no means unsuggestive of the mammalian 
diaphragm, or, to be more accurate, a portion of the diaphragm, 
which is itself a complex structure. The transverse position of 
the membrane supporting the end of the oviduct (found, it must 
be remembered, in both sexes) is the first notable fact to be pointed 
out. Secondly, the heart and a great part of the lungs lie in 
front of this transverse septum, which from its position delimits 
a “chest” not incomparable with the mammalian thorax. The 
arteries, which have just been described as ramifying upon the 

* These arterics must not be confused with the deep-running vertebral arteries 
figured by Hyrtl and just referred to by myself (p. 310). 


314 ON THE ANATOMY OF THE JAPANESE SALAMANDER. [ Nov. 3, 


surface of this membrane, call to mind the superior phrenic 
artery of mammals, Finally, its relations to the oviduct are not 


Text-fig. 34. 


<a \M 
NO si hinttt 
> a FEOY 


Weeds itp 
‘ 


‘ INAS 
\ \ |= 
“ip N ES 
) Ut ttt ting, GSN 
Aes pry, hy Sy 
\ ea ig 
\ SO SRS THN 
{ \\ '§ 
N 


ii j 
ea : aya b my 
as EL spec a ete ALY 


4s 
Lay 
ws 


Ian 
Le 


YY ff 7 


exiled eee IZ 


Dissection illustrating relation of arterie comites aorta of Megalobatrachus 
to “ diaphragm.” 
Ao, aorta; Od, oviduct ; P, arteria comes aorte; Scl, subelavian; V.m, vertical 
membrane (“diaphragm ”’); «, problematic body. 


1903. ] THE SECRETARY ON ADDITIONS TO THE MENAGERIE. BLS) 


so fatal to the hypothesis here dealt with as might at first appear. 
In some mammals, for instance in Galictis barbara, the broad 
ligament continuous with the suspensory ligaments of the gonads 
and their ducts is continued forwards very nearly as far as the 
diaphragm. Furthermore, in some birds the oviduct is in actual 
contact with the oblique septum, which is a structure also to be 
compared with these septa that we are now considering. It must 
not be lost sight of that the transversely-running membrane in 
Megalobatrachus is not a characteristic of the female animal only, 
and serving merely for the support of the oviduct : it is equally 
developed in the male. 

In the figure that I have just referred to a projecting body 
(text-fig. 34) lettered “a” will be noticed. This is brownish in 
colour, more or less oval in form, and depends from the anterior 
side of the oviducal membrane. From its position I take it to 
represent a somewhat similar body which I have already referred 
to as existing in the male Megalobatrachus. I have investigated 
both by means of serial sections. 

In the case of the female the oval body consists of an outer 
muscular layer which is thickened at the solid “ neck” and con- 
tinuous with the membrane, from which it freely depends into the 
body-cavity. Through the “neck” passes a large blood-vessel, a 
branch of the arteria comes aortee. The interior of the body con- 
sists of largish nucleated cells of which I could detect no regular 
arrangement. In the male the structure seems to be the same, 
with more muscular fibres interspersed in the core. I imagine 
that this body is the persistent rudiment of the pronephros, but 
the structural indications are quite inconclusive. If so, it is plain 
that the oviducal funnel of the female can have nothing to do with 
the pronephros. 


November 17, 1903. 
H.G. Tae Duke oF Breprorp, K.G., President, in the Chair. 


The Secretary read the following report on the additions to the 
Society’s Menagerie during the month of October 1903 :— 

The registered additions to the Society’s Menagerie during the 
month of October were 180 in number. Of these 83 were 
acquired by presentation and 3 by purchase, 15 were born in the 
Gardens, 78 were received on deposit and 1 in exchange. The 
total number of departures during the same period, by death and 
removals, was 139. 

Amongst the additions attention may be called to :— 

1. A fine male Chimpanzee (A nthropopithecus troglodytes), from 
the Albert Nyanza, presented by Lt.-Col. David Bruce, F.R.S., 
Oct. 4th. This animal, so far as is known, is the first living 
example received by the Society from Eastern Africa. 

2. Two Scoresby’s Gulls (Leucopheus scoresbir), deposited by 
the Hon. Walter Rothschild, M.P., Oct. 17th. 

3. Two Wharton’s Fruit-Pigeons (Carpophaga wharton?) and a 


316 ON TSETSE-FLIES FROM AFRICA. [N ov. 17, 


Ghristmas-Island Dove (Chalcophaps natalis), from Christmas 
Island, in the Indian Ocean, presented by Capt. A.W. Cole, Oct. 27th. 


Mr. Henry Scherren, F.Z.8., exhibited the front horn of a 
Burchell’s Rhinoceros (Rhinoceros sinus), the largest yet received 
in England from the Soudan. It was the property of Col. B. 
T. Mahon, O.B., D.S.O. (for whom it had been mounted by 
Mr. Rowland Ward), and measured along the anterior curve 
362 in., with a circumference at the base of 192 in. This is only 
three-fifths of the length of the record horn obtained in South 
Africa by Roualeyn Gordon Cumming, which has a length of 
622 in. In addition to the horn exhibited and that belonging to 

‘apt. Hawker (cf. above, p. 194), other horns from the same 
region were in the possession of Major-Gen. Sir F. R. Wingate, 
Hon. Capt. McNaughten, Major Sanders, and Capt. J. G. A. 
Massey. Mr. Scherren stated that this Rhinoceros was reported 
as being fairly numerous on the northern boundary of the Congo 
Free State and in the adjacent parts of the Soudan, and that 
Mr. Rowland Ward had known of these horns coming from that 
district for many years before Major Gibbons secured his specimen 
(cf. P. Z.S. 1900, p. 949). 


Mr. R. I. Pocock, F.Z.S., exhibited a piece of basalt picked up 
between tide-marks on the coast of Victoria, Australia, by 
Mrs. Kenyon, which contained a specimen of the web of the 
Marine Spider Desis kenyone, described in the ‘ Proceedings’ of 
last year (cf. 1902, ii. p. 102). Although broken, the web served 
to illustrate the habit of these marine spiders of spinning a 
closely-woven sheet of silk over a crevice in the rock as a pro- 
tection against the rising tide. 


Mr. Pocock also gave an exposition, illustrated by drawings, of 
a new suggestion as to the white rump-patches of Ungulata, with - 
special reference to the races of Burchell’s Zebra. 


Mr. E. E. Austen exhibited specimens, with pupe, of Glossina 
palpalis Rob.-Desv., the species of Tsetse-fly which is the active 
agent in the transmission of sleeping-sickness in Uganda, by 
conveying Zrypanosoma castellanit Kruse, the parasite which is 
the cause of the disease, from man to man. Examples of four 
other species of Tsetse-flies were also shown for the sake of com- 
parison, and the general characteristics of the genus Glossina, 
which is confined to Africa, were pointed out. Attention was 
drawn to the remarkable mode of reproduction in the genus, 
which renders it impossible to attack the insects in their breeding- 
places, as has been successfully accomplished in many places in the 
ease of mosquitos ; for, instead of laying eggs, Tsetse-flies produce 
but a single larva at a birth, which forthwith crawls away and 
assumes the pupal condition on reaching some sheltered spot. Seven 
species of Tsetse-flies are ab present known, and of these at least 
two, Glossina morsitans Westw. and Gl. pallidipes Austen, are 
probably concerned in the dissemination of ‘* Nagana,” or Tsetse- 


1903. | ON A HAIR-WHORL IN A GAZELLE. Sly 


fly disease, caused by Zrypanosoma brucei, among domestic 
animals. The demonstration of the connection between Glossina 
palpalis and sleeping-sickness in Uganda is due to the recent 
researches of Lieut.-Colonel Bruce, R.A.M.C., of which a detailed 
report is shortly to be presented to the Royal Society. 


Mr. Oldfield Thomas, F.R.S., exhibited on behalf of Mr. de 


Winton a drawing of a female Gazelle (Gazella muscatensis (°)), 


Text-fig. 35. 


Lp TT 1 ANS 
f ie Fe ya 
a ADA, id 


Diagrammatic sketch of skin of Gazelle to show hair-whorl on withers. 


obtained at Sheik Oman near Aden, and presented to the Society 


318 ON SANSKRIT REFERENCES TO THE TIGER. [Nov. 17, 


by the Officers of H.M.S. ‘Cossack,’ which showed a complete and 
perfect whorl of the hair on the withers (text-fig. 35, p. 317). A 
male of the same species obtained at the same time exhibited no 
sign of this whorl, but the hair lay in the direction of the tail from 


the base of the neck. 


Mr. C. Tate Regan read a paper entitled ‘‘ A Revision of the 
Fishes of the Family Zoricariide,” in which nearly 200 species 
were recognised as valid, 35 being described as new to science. 
The types of the species described by Cuvier and Valenciennes 
and by Castlenau, in the Museum at Paris, had been examined. 
The genera were grouped into five subfamilies— Plecostomine, 
Hypoptopomine, Loricariine, Neoplecostomine, and Argiine, the 
last two being regarded as closely related on anatomical grounds, 
although differing in external characters. 

This paper will be published entire in the Society’s ‘Transactions.’ 


The following papers were read :— 


1. On Early Sanskrit References to the Tiger. 
By V. V. Ramanan, M.A., F.Z.8. 


[Received September 14, 1903. ] 


In his paper “On the Original Home of the Tiger” (P.Z.S. 
1903, vol. i. p. 109), Col. C. E. Stewart made some statements 
that I am able to correct. He said that, after enquiry, he could 
find no Sanskrit word for the Tiger. The word Vydghra is the 
Sanskrit term for the tiger, and is common in juvenile books. 
The modern Hindustani word to which Col. Stewart referred is a 
corrupt derivative of the Sanskrit original. 

T have also to note that references to the tiger are frequent in 
Sanskrit literature. Col. Stewart stated that he could find no 
mention of the tiger in Sanskrit works treating of the fighting 
between Rama and Ravana. The Ramdyan of Valmiki is the 
most ancient of the many Sanskrit works that deal with the 
famous “ Lanka” war. In Ramayana there are numerous allu- 
sions to tigers, especially in the chapter where the primeval 
forest, Dandaka, is described. I pass over many similar references 
in other works dealing with the same war, as these are of later 
date. 

In the Mahabharata, a poem which is of the same age as, if 
not more ancient than, the Ramdyan, there are numerous refer- 
ences to the tiger. The legend of the Vydghra-pdda, the “ tiger- 
footed” Rishi, is familiar. 

In the ‘ Vedas,’ the most archaic documents in Hindu literature, 
there are references to the tiger. One quotation, from the 
Tuttiriya Brahmana section, is enough for my purpose :— 
“ Vyaghroyam Agnau charati pravrshthah.” 


1903. ] ON THE ANATOMY OF THE HAMADRYAD SNAKE. 319 


There is not an Indian vernacular, ancient or modern, Dravidian 
or Aryan, without its own name for the tiger. It is interesting 
to note that in South India there are vernaculars of quite indi- 
genous origin that owe no part of their vocabulary to Sanskrit, 
and which, on that account, are supposed to be more ancient than 
Sanskrit. In these there are many synonyms for the tiger, and 
these enter frequently into the composition of the names of South- 
Indian villages. Villages bearing the names ‘“ Tiger-town,” 
“Tiger-village,” ‘Small Tiger-town,” and so forth are quite 
common. 

T shall add no more, as the discussion, if pursued further in 
this direction, would lose its zoological interest and become 
purely etymological. But I shall be pleased to give any zoologist 
more detailed information. I am not competent to say if the 
question of the original home of the tiger, which is one of zoo- 
geography and paleozoology, can be made to rest on a foundation 
of philology. I should like, however, to elicit the opinions of 
Mr. Lydekker and Dr. Blanford, whose labours have made them 
competent judges. 


2. On the Trachea, Lungs, and other Points in the Anatomy 
of the Hamadryad Snake (Ophiophagus bungarus). By 
Frank H. Brepparp, M.A., F.R.S., Prosector to the 
Society. 

[Received November 3, 1903. ] 


(Text-figures 36-40.) 


Although some recent systematists * include the Hamadryad 
within the same genus (aia) as the Hooded Cobra and its 
immediate allies, there are nevertheless a number of anatomical 
features in which the Hamadryad differs. from Waia tripudians, 
and which, therefore, tend to justify its earlier separation as a 
distinct generic type. It differs, for example, in the form of the 
liver. In Ophiophagus, as in other Snakes, the liver has two 
distinct though closely-adjacent lobes, which are divided by the 
vena cava posterior, and of which each is lodged in a separate 
ccelomic compartment 7. These lobes are, however, quite unequal 
in size. The left-hand lobe measures 16 inches in length, the 
larger right-hand lobe 19 inches. The latter, however, extends 
for 73 inches beyond the termination of the former, but it begins 
43 inches after it. The inequality in size between the lobes 
is thus accentuated by their position with reference to each 
other. In the Cobra (Waia tripudians) precisely the reverse con- 
ditions obtain. The left lobe (that nearest to the stomach, when 
the animal is opened and viewed from the ventral surface) is 

* H.g. Boulenger, Cat. Snakes B. M. vol. iii. 1896, p. 386. 

on “On the Subdivision of the Body-cavity in Snakes,’ P. Z.S. 18924 
p. 477. 


320 MR. F. E. BEDDARD ON THE [ Nov. 17, 


considerably the longer; it measures 11 inches; it commences 
3 inches before the right lobe and ends 7 inch behind it. The 
right lobe, therefore, measures 77 inches. It is clear, therefore, 
that the proportionate difference also varies in the two species. 
Gall-bladder—As I have been at some pains to make a careful 
dissection of the complicated series of anastomosing tubes by 
which the bile enters the gut, | venture to include in the present 
communication a description of the same, which is illustrated by 
the drawing (text-fig. 36)*. The two specimens which I have 
examined differ slightly in the exact relations of the tubes of 


Text-fig. 36. 


Gall-bladder of Ophiophagus bungarus. 


D., diverticulum ; G.B., gall-bladder ; H.D., hepatic duct of intestine ; 
Iyv., intestine. 


the rete, but not so materially as to interfere with the general 
truth of the figure. The actual plexus is not very complex, but 
the hepatic duct, as will be seen, enters into it. Ultimately five 


* The drawing was made from the injected gall-bladder, which J caused to be filled 
with coloured gelatine, 


Oe in ee a a 


1903. ] ANATOMY OF THE HAMADRYAD SNAKE. Boil 


bile-ducts emerge from the rete and pursue a long course (long 
as compared with the plexus region) through the pancreas to 
the intestine. Arrived at the intestine they do not debouch 
directly into its lumen by five orifices, but open separately, or 
nearly so, into a thick-walled diverticulum of the gut, which itself 
communicates with the gut by an orifice not wider than one of 
the separate bile-ducts. I have noticed a similar diverticulum, 
which receives both cystic and hepatic ducts, in Westerman’s 
Cassowary. As compared with the Cobra*, which I have also 
examined, the bile-duets of Ophiophagus are more numerous and 
more complex in their intercommunications. The hepatic duct, 
for example, is single, though it communicates with more than 
one of the cystic ducts. The gall-bladder itself has not so long a 
common cystic duct as that which characterises Ophiophagus. I 
do not think that a more minute description of the cystic network 
in the two types is useful at present; but I may observe that 
sketches of the gall-bladder published by other anatomists, and a 
few made by myself, indicate that the general plan of the cystic 
rete is not without use in distinguishing and comparing various 
genera of serpents. 

Again, the Cobra has a testis proportionately longer than that 
of the Hamadryad and differently situated. The anterior testis is 
24 inches long and its anterior border is 11 inches from the liver ; 
its posterior border is a long way in front of the kidney. 

In Ophiophagus bungarus the anterior testis abuts so closely 
upon the anterior kidney that the two organs are only just 
disconnected ; while the spermatic artery, influenced, so to speak, 
by this near approximation, sends a branch to the kidney before 
entering the testis. The anterior end of the testis is, moreover, 
22 inches away from the end of the liver, and it measures only 
1? inches in length though very much plumper than the corre- 
sponding testis of the Cobra. In estimating these facts 1t must be 
borne in mind that the Hamadryad measured 93 inches from tip 
of snout to vent, and the Cobra 51 inches. 

More possibly, but still to my mind not certainly, associated 
with difference of size is the fact that the Hamadryad possesses 
six gastric arteries as against three in the case of the Cobra. I 
recognise—it may be explained—as gastric arteries those vessels. 
which arise from the aorta after the posterior end of the liver and 
supply the stomach only. 

In the Ophidia generally there are two Carotid arteries, both of 
which spring near together from the right aortic arch. The left- 
hand artery is, I believe, always the largest, and extends right up 
the neck as far as the head, giving off branches along its course 
to the gullet and windpipe. The second and smaller of the two 
Carotids is, as a rule, of much smaller calibre, and in the distribu- 
tion of its twigs is hardly more than a thyroidean artery. In 
Ophiophagus bungarus these conditions obtain, and the artery in 
question does not extend up the neck very much beyond the 


* Bronn’s ‘Thierreich,’ Rept., Bd. vi. pt. mi. pl. exxx. fig. 2. 


Proc. Zoou. Soc.—1903, Vou. Il. No. XX. 21 


322 MR. F. E, BEDDARD ON THE [ Nov. 17, 


thyroid gland. On the other hand, in the Cobra the conditions 
are quite different. The right Carotid, although perhaps not 
more than one-half of the diameter of the left, is quite as fully 
developed as regards the length of its course, and runs right up 
to the head and lies closely side by side with its fellow artery. 
It is rather curious that both arteries plunge into the thickness 
of the tissues of the head, not on opposite sides as might be 
expected, but close together on the same (left) side. 

Another difference in the situs viscerwm is indicated by the 
following measurements :—In Ophiophagus the origin of the 
superior mesenteric artery is 73 inches from the mandibular 
symphysis and 214 from the cloacal aperture. The corresponding 
measurements in the Cobra which | examined were 35 and 
15 inches respectively. There is thus in the Hamadryad a 
shortening of the ‘‘abdominal” region of the body as compared 
with the Cobra, and a consequent crowding of the viscera therein 
contained. This is also expressed by the nearness of the kidney 
to the testis already referred to. In spite of this the lung of the 
Hamadryad has a longer vascular, and therefore respiratory, 
region than the Cobra. The measurements are as follows :—Iln 
the Cobra the lung ceases to be vascular 18 inches from the snout, 
in the Hamadryad 38 inches. Were the proportions the same, the 
length of the Hamadryad from snout to vent would be 1074 inches ; 
it is, as a matter of fact, 93 inches. It follows, therefore, that the 
vascular part of the lung in this snake is proportionately longer 
than in the Cobra. 

It seems to me to be plain that these various characters are, at 
least, more striking as a means of separating the two snakes than 
those used by the original describer of ‘‘ Hamadryas hannah” * 
or by those who have succeeded him. 

But a more remarkable character still distinguishes this snake 
from its ally, the Indian Cobra. In the latter snake, as is well 
and has been long known t, the tracheal rings are incomplete 
dorsally as cartilaginous rings, the interspaces being merely mem- 
branous. The membrane, however, except at the beginning of the 
trachea, has a diameter fully as great as that of the cartilaginous 
tract ; it is, moreover, tough, longitudinally creased, and obviously 
dilatable. 

In Ophiophagus the tracheal rings are similarly incomplete from 
the very first, but the membranous interval is less tough and 
strong (though the snake is considerably larger), and very much 
less in relative and even in actual diameter. So far, therefore, 
the differences are but slight. A very remarkable difference 
is Illustrated in the accompanying drawings (text-figs. 37, 38, 
pp. 323, 324). At intervals this dorsal tracheal membrane is 


* Cantor, ‘‘Sketch of an undescribed Hooded Serpent, &c.,” Asiat. Researches, 
vol. xix. 1836, p. 87, and also P. Z.S. 1838, p. 73, where it is described as “ Hama- 
dryas ophiophagus,” a reference omitted by Mr. Boulenger in his Catalogue, who 
quotes only, though so far correctly, P. Z.S. 1839, p. 32. 

a Wate “Legons d’Anatomie Comparée réd, par Duvernoy,’ ed. 2, Paris, vol. yii. 

40, p. 93. 


ANATOMY OF THE HAMADRYAD SNAKE, 


1903.] 


Dissection of neck of Ophiophagus. 


B, intercostal artery ; S, air-sacs; T, trachea. 


324 MR. F, E. BEDDARD ON THE | Nov. 17, 


perforated by regular oval foramina, which commence two and a 
half inches from the Jarynx and continue to within half an inch 
of the lung. These apertures vary somewhat in size, 8 mm. being 
one extreme and 3 or 4 mm. the other. They may be really 
divided into two series. Twenty-six or twenty-seven (and it 
should be said that this series comprises the largest foramina) 
lie at approximately equidistant imtervals, corresponding either 
exactly or very nearly exactly to the middle of a large mem- 
branous sac, which is plainly to be regarded as a hernia-like 
outpushing of the lining membrane of the trachea. This series 


Text-fig. 38. 


eee 
> 
4 


iw) 


BEE: 


Wainy 


ITNT 


sayy yes 


A more detailed view of two air-sacs and dividing septum in Ophiophagus. 


A, intercostal vein; B, intercostal artery (both resting upon septum which divides 
two adjacent air-sacs); C, carotid; T, trachea, with two perforations ; 
V.A., vertebral artery; V.V., vertebral vein. 


of sacs surrounds the organs of the neck and practically obliterates 
the celom. ‘The sacs are of about the same dimensions, an inch 
or so in antero-posterior diameter. So closely are adjacent sacs 
pressed together that the bounding walls anteriorly and posteriorly 
run straight across the neck. The appearance thus produced, 
when the parietes of the neck are cut through deeply enough to 
include the ventral wall of the sacs, would Jead naturally, on a 
superficial view, to the inference that we had here to do with a 


ee 


See ee ee. ee ee 


’ 
: 
h 
[ 
} 


1903. ] ANATOMY OF THE HAMADRYAD SNAKE, 325 


ceelom divided by metamerically arranged transverse septa like 
that of a typical Annelid. This, however, is obviously not the 
case unless we are to adopt the totally improbable view that 
the trachea communicates directly with the celom. These 26 or 
27 chambers, extending from the chin to the lung, do not, so 
far as I have been able to ascertain, communicate with each 
other. They correspond fairly, though not accurately, with the 
intercostal branches of the anterior vertebral artery, and in the 
following way :—Those arteries arise at approximately equidistant 
intervals and bury themselves in the thickness of the parietes in 
the dorsal middle line. Along their course they are supported 
by the membranous partitions in question, which adhere to them 
from their origin to their disappearance in the parietes. Only 
four of the total series of over thirty of these arteries were not 
so supported, and only two “septa” were devoid of an arterial 
companion. On the opposite side of the body the vertebral veins 
have a similar relation to the “septa.” In addition to the 


Text-fig. 39. 


iB A- 


\ 
. Ay NK 


A portion of trachea from dorsal surface of Ophiophagus, showing, A, perforation 
leading into air-sac, and, B, circular thinning of tracheal membrane. 


principal tracheal foramina there are smaller ones which le 
between the larger and just between two adjacent air-sacs. The 
arrangement of these, however, is not regular. I could only 
count four or five of them. I imagine that they correspond to 
smaller intercalary sacs, of which there was distinct evidence, 
lying between the larger ones. Finally here and there, and also 
lying between the primary sacs, were regularly rounded thinnings 
of the tracheal membrane (text-fig. 39, B) which may be looked 
upon as—if the expression be allowed—imperforate foramina. I 
could detect no evidence of intercalary sacs in connection with 
them ; but they are ready, as it were, to form such sacs. 

In a second example of this Snake, also a male, I examined the 
air-sacs of the neck, which I found to agree in all essentials with 
those of the first specimen, There are, however, rather fewer 


326 MR. F. F. BEDDARD ON THE [ Nov. 17, 


of the compartments (about 22), which fact, coupled with the 
smaller dimensions of the snake, bears out, to a certain extent, 
my belief that the smaller intercalary sacs grow with the increase 


in size of the snake. 
Text-fig. 40. 


‘> 
fas 


Ending of trachea in lung of Ophiophagus. A, foramina leading into air-sacs ; 
B, terminal air-sac; C, eutrance to rudimentary lung; D, lung. 


The accompanying sketch (text-fig. 40) shows the relation of 
the last of these air-sacs to the lung. It will be observed that in 
Ophiophagus as in Naia there is a rudimentary lung as well as 
the fully developed one, and that the bronchus enters the fully 
developed lung about half an inch posteriorly to the commence- 
ment of the latter. The rings of the bronchus do not extend 
down the lung beyond the origin of the bronchus to the rudi- 
mentary lung, but a narrow white seam is to be observed which 
is developed for some distance along the lung. The median thin 
fibrous wall of the trachea, of course, is in contact with the lung- 
tissue; but there is no transitional area between the two. The 
dark red vascular lung begins abruptly. The last orifice from 
the trachea is shown in the figure, and beyond it will be seen a 
flap with a crescentic edge which covers over a short sac which 
is the last of the series of neck-sacs and in which the trachea 
may be said to terminate. The wall of this terminal sac is 
shown in the drawing to be perforated by a foramen, which, I 
admit, may be artifact. I am unable to be positive about the 
matter. If it is not, then it must be a branch of the terminal 
sac. It must be further observed that the lung can be seen 
through the terminal sac which thus lies ventrally of that organ. 
I have satisfied myself that there is no-communication between 
the terminal air-sac and the lung, except, of course, indirectly 
through the trachea. 

So far as I can ascertain, there is nothing precisely similar to 
this known in other Snakes. Many anatomists, however, have 
described a continuation of the lung-tissue wp the trachea, i.e. 
towards the head*. This may be merely an extension of the 

* For instance, Meckel, Anat. Comp. (French translation), Paris, vol. ii. 1838 ; 
Schlegel, ‘Essai sur la Physionomie des Serpents,’ Amsterdam, 1837; Siebold & 


Stannius, Handbuch der Zootomie, 2° Buch, Berlin, 1856 (2nd ed.) ; Milne-Edwards, 
Le¢ons sur la Physiologie et ’ Anatomie comparée,’ t. ii., Paris, 1857. 


< 


1903.] ANATOMY OF THE HAMADRYAD SNAKE. 327 


anterior bit of the lung which in Ophiophagus lies above the 
entrance of the bronchus, or it may be an increase and conversion 
into lung-tissue of the median membranous region of the trachea. 
Quite recently the late Prof. E. D. Cope has investigated—but 
apparently only in a preliminary way—the lungs of various 
snakes, some of which were known and others unknown to 
previous observers. In the long list of Ophidia investigated by 
Cope representatives of most groups occur, but he had not the 
opportunity of examining the Hamadryad. The pith of his 
paper*, so far as concerns the subject in hand, is contained in 
the following sentences :—‘“ In the Solenoglypha, without excep- 
tion, this extension of the dorsal ¢ lung is present, and extends to 
the head, and its lumen is continuous with the trachea through- 
out its whole length. The same structure exists in the genera 
Hydrus and Hydrophis; and also in the Peropodous genus 
Ungalia, which differs besides from other Peropoda in having 
but one posttracheal lung. Finally the tracheal lung, as I shall 
call it, is distinct from the true lung in Platyurus and in Cher- 
sydrus. Inthe former of these genera the trachea is not separate 
from the lumen; while in Chersydrus it is distinct. It, however, 
communicates with the cells of which the lung consists in this 
genus by a series of regularly placed foramina on each side.” 

It is a little difficult to draw conclusions from so abbreviated a 
statement of fact ; but I am inclined to suppose that Chersydrus 
(also a Colubrine, though aglyphous) comes nearest to Ophio- 
phagus though obviously differing much in detail. I take it that 
the conditions which characterise Ophiophagus are an extreme 
modification of those apparently found in Chersydrus: that the 
metamerically arranged pouches in the former, with their series 
of single orifices into the trachea, are the remains of what Cope 
terms the ‘“ tracheal lung,” in which the respiratory function has 
been replaced by some other use. In the absence, however, of 
complete details as to Chersydrus and the other genera to which 
Cope refers, it is rash to form a definite opinion. It must be 
borne in mind that there is in Ophiophagus no connection with 
the lung. The only structure among Sauropsida, other than 
serpents, which I can think of as obviously comparable to this 
series of pouches in Ophiophagus, is the tracheal pouch of the 
Emu. That bird in a quite similar fashion shows a slit on the 
trachea where the rings fail to meet in the middle line. It is, 
however, ventral rather than dorsal in position, and there is but 
a single slit. It leads into a thin-walled outgrowth of the lining 


* “On the Lungs of the Ophidia,” Proc. Amer. Phil. Soc. xxxiii. 1894, p.217. The 
facts are also to be found in two Other memoirs, viz. ‘ The Classitication of Suakes,” 
Amer. Nat. xxvill. 1894, p. 831, and ‘“ The Classification of the Ophidia,’ Trans. 
Amer. Phil. Soc. xviii. p. 188. 

+ “Dorsal” as used by Cope is the equivalent of “right” in Mr. G. Butler’s 
paper “On the complete or partial Suppression of the Right Lung in the Amphis- 
benidz and of the Left Lung in Snakes and Snake-like Lizards and Amphibians,” 
P. Z.S. 1895, p. 691. 

t¢ Cf. Murie, “On the Tracheal Pouch of the Emu (Dromeus nove-hollandia).” 
P. Z.S. 1867, p. 405, where previous literature 1s cited. 


328 MR G. A. BOULENGER ON FISHES [ Nov. 17, 


membrane of the trachea exactly as in Ophiophagus. Moreover, 
a kind of septum appears to divide that pouch into two, so that 
there are, at least, traces of metamerism. There is, however, no 
evidence of any connection in the past between this pouch and 
the lung. 


3. Report on the Fishes collected by Mr. Oscar Neumann 
and Baron Carlo von Hrlanger in Gallaland and 
Southern Ethiopia. By G. A. Bounencnr, F.R.S., 
Wile yaa 


‘Received October 29, 1903. | 
(Plates XXIX.-XXXI*) 


Mr. Oscar Neumann has entrusted to me for study the fishes 
collected on the expedition from Zeila through Southern Ethiopia 
to the Upper Nile in 1900-1901, of which he gave an account 
at the Geographical Society in June 1902, and which is printed 
in the ‘Geographical Journal’ for October 1902, with a map, 
to which I refer the readers for the position of the localities 
mentioned in this report. 

The fishes are all from East of the Nile system and were 
obtained in the following waters :— 

Modjo, Ivaro, and Wabbi Rivers, affluents of the Webi Shebeli ; 
Kassam, Gadschimboda, and Suksuk Rivers, affluents of the 
Hawash; Maki River, flowing into Lake Zwai; Omo River, 
flowing into Lake Rudolf; and Lake Gandjule, south of Lake 
Abaia. 

The following list of 19 species, 4 of which are described as 
new, extensively supplements our scanty knowledge of the 
fishes of that part of Africa, based on the small collections 
previously made by Marquis Antinori, Capt. Bottego, Prince KE. 
Ruspoli, aud Dr. Donaldson Smith, which have been reported 
upon by Dr. Vinciguerra + and Dr, Giinther =. Collections had 
also previously been made in the Kassam and Maki Rivers by 
Mr. E. Degen on his way to Lake Tsana, but the five new species 
discovered by him and described by me$ are, curiously, not 
represented in the collection made in those rivers by Mr. Neumann 
and Baron v. Erlanger. 


MorMYRID. 


1. MormMyrus KANNUME Forsk. 


A single specimen, from the Wabbi River. 


* For explanation of the Plates, see p. 334. 

ole a Genova, xvill. 1883, p. 691, xxxv. 1895, p. 21, xxxvil. 1896-97, 
i P.Z.S. 1896, p. 217. 
§ Ann, & Mag. N. H. (7) x. 1902, p. 421. 


Nin a 


‘INNVNONGN O84V'T 


“ULI TSP t8E19'P 


‘dust soig wiagunyl 


SUNOC Teal WIE” SOGIE SZ al 


‘ITM GONV Td Shaved 


TEBRE © [je Veean9) e 


‘dar soig wsre7uUyy 


YOOC lal WES BOGS Z al 


IPA. O MOS Soll Wl ICO 


Ss 
Ss 


J.Green del.et hth. Mantern Bros.imp. 


1.DISCOGNATHUS MAKIENSIS. 2.DISCOGNATHUS BLANFORDII, 
3.CHILOGLANIS MODJENSIS. 


1903. | FROM GALLALAND AND SOUTHERN ETHIOPIA. 329 


CHARACINID&. 


2. Hyprocyon FoRSKALIL Cuv. 
Several young specimens from Lake Gandjule. 


3. ALESTES AFFINIS Gthr. 
Several specimens, from the Wabbi and Modjo Rivers. 


4. ALESTES MACROLEPIDOTUS Cuy. 
Several specimens from the Omo River. 


5. MICRALESTES ACUTIDENS Peters. 


Several specimens from the Omo River. 
This fish, originally described from Mozambique, has since 
been found in the Ubanghi, in the Nile, and in the Niger. 


CYPRINIDS. 


6. LABEO NEUMANNI, sp.n. (Plate X XIX.) 


Depth of body 34 to 3? times in total length, length of head 
Al to 42 times. Head I 50 14 as long as br oad ; snout rounded, 
with small horny warts, its length 2 to. a little less than 2 that of 
the head ; eye per fectly lateral, fie diameter 4t to 53 times in tle 
length of ‘the head ; interorbital width 4 length of “head, 21 to 3 
‘shine the diameter of the eye; mouth strongly arched, its width 
2 to % the length of the head; lips with small papillee forming 
transverse plice; lower lip with a fringe of conical papille ; 
rostral flap large, with denticulate edge; a minute barbel in the 
corner of the mouth, hidden under the folds of the mouth. Dorsal 
fin composed of Toe MT rays, a little nearer the end of the 
snout than the root of the caudal, its upper edge concave, the last 
simple ray sometas and as long as or slightly longer than the head. 
Anal with III 5 rays, falcate, . its longest ray nearly as long as the 
head, reaching or nearly reaching the root of the caudal. Pectoral 
subfalciform, as long as or a little longer than the head, not quite 
reaching the ventral, the first ray of which falls below the 4th or 
5th branched ray of the dorsal. Caudal fin deeply emar pinata, 
crescentic when fully spread out. Caudal peduncle 1} to Ls as 


long as deep. Scales finely striated longitudinally, 39- Al) ay 


5 between the lateral line and the base of the ventral, 16 or 18 
round the caudal peduncle. Olive above, whitish beneath ; fins 
greyish (in spirit). 

Five specimens, measuring 140 to 320 millimetres, from the 
Modjo and Wabbi Rivers. 

Distinguished from LZ. gregorii Gthr. by the broader inter- 
orbital space and the more numerous scales. 

Three very small specimens, measuring from 18 to 27 milli- 
metres, from the Wabbi, may, I think, be safely referred to the 
same species. 


330 MR. G. A. BOULENGER ON FISHES [Nov. 17, 


7. LABEO CYLINDRICUS Peters. 


Two specimens, measuring 64 and 150 millimetres respectively, 
from the Modjo River. Diameter of eye 43 and 5 times in length 


of head. D.III9. Sq. 35-37%, 4, 16. 


ey 


8, DiscoGNATHUS MAKIENSIS, sp.n. (Plate XXXI. fig. 1.) 


Body feebly compressed, its depth 5 to 6 times in the total 
length. Head as broad as deep, 14 as long as broad, } to 7 total 
length; snout rounded, projecting beyond the mouth; inter- 
orbital region convex, its width not quite half the length of the 
head; eye lateral, but better visible from above than from below, 
in the middle of the length of the head, its diameter 4 to 53 times 
in the length of the head and twice to twice and a half in the 
interorbital width ; width of mouth not quite + the length of the 
head; upper lip well developed, not fringed ; lower lip forming a 
mental disk which is broader than long; two barbels on each 
side, measuring 2 to 4 the diameter of the eye. Dorsal with 
10 rays, 7 of which are branched, equally distant from the end of 
the snout or the nostrils and from the root of the caudal; first 
branched ray longest, a little longer than the head. Anal II 5, 
first branched ray longest, ? to 2 the length of the head. Pectoral 
as long as or slightly longer than the head, not reaching the ventral, 
which is situated below the middle of the dorsal. Caudal fin 
deeply emarginate, longer than the head. Caudal peduncle 13 to 
1? as long as deep. Scales 38-40 2, 34 or 4 between the lateral 
line and the ventral, 16 round the caudal peduncle. Olive above, 
white beneath. 

Six specimens, measuring from 48 to 80 millimetres. Maki 
River, running into L. Zwai. 


9. DiscoGNaTHUS BLANFoRDII Blgr. (Plate XX XI. fig. 2.) 


Several specimens from the Gadschimboda R., affluent of the 
Hawash. 


10. DiscoGNATHUS QUADRIMACULATUS Riipp. 


Several specimens from the Gadschimboda and Suksuk Rivers. 
Kye in the middle of the length of the head, its diameter 33 
(young) to 5 times in the length of the head; barbels 4 to # the 
diameter of the eye; mental disk small, feebly marked, a little 


longer than broad. Scales 39-42 a 4 between lateral line and 
ventral. The largest specimen is 55 millimetres long. 

Described from Abyssinia by Riippell under the names of 
Gobio quadrimaculatus and G. hirticeps. Young specimens from 
Lake Arsadé, south of Shoa, have been described by Vinciguerra 
as Discognathus chiarinit. 


We are now acquainted with six African species of the genus 


1903. | FROM GALLALAND AND SOUTHERN ETHIOPIA. 331 


Discognathus, which may be distinguished by means of the 
following synopsis :— 


I. Pupil of the eye in the second half of the length 
of the head; barbels not more than half 
diameter of eye; caudal peduncle 14 to 15 as 
long as deep ; scales in lateral line 37 to 40. 


Snout projecting strongly beyond the mouth, the 
width of which is less than half the length of 
the head ; upper lip well developed, with a series 
of papilla forming a fringe .. D. dembeensis Riipp. 
Snout projecting feebly beyond the mouth, the width 
of which equals half the length of the head; 
upper lip scarcely developed, not fringed ......... D. johnstonii Blgr. 
II. Pupil of the eye in the middle, or anterior to 
the middle, of the length of the head. 
1. Mental disk well developed, with free posterior 
border. 
Barbels 2 diameter of eye; eye lateral; dorsal fin 
equally distant from centre or posterior border 
of eye and root of caudal; caudal peduncle as 
long as deep ; ; lat. 1. 35-38 ....... D. vinciguerre Bley, 
Barbels 3 4 to } diameter of eye; eye ‘supero- -lateral ; 
dorsal fin equally distant from eye and root of 
caudal, or a little nearer the latter; caudal 
peduncle 14 to 1} as long as deep; lat. 1 38-87. Dz blanfordii Bley. 
Barbels $ to # diameter of eye; eye lateral; dorsal 
fin equally distant from end of snout or nostrils 
and root of caudal; caudal peduncle 1} to 1$ as 
long as deep; lat. 1 SoM eee .. D. makiensis Blgr. 


. Mental disk very indistinct, or anita to a 
mere pad, without free posterior border ; 
barbels 4 to 2 diameter of eye; dorsal fin 
equally distant from centre or posterior 
border of eye and root of caudal; caudal 
peduncle about 13 as long as deep; lat. 
We B74 icin sscssesessetses Di quadrimaculatus Rupp. 


11. BarBus ERLANGERI, sp. n. (Plate XXX.) 


Depth of body 3 to 31 times in total length, length of head 
33 to 41 times. Snout “gunned. or Paneneinates prominent, its 
length 3 to 34 times in length of head; diameter of eye 33 (young) 
to 5 times in length of head, interorbital width 24 to 3 times ; 
mouth small, inferior, its width + to ¢ length of head; lips well 
developed, the lower continuous across the chin, forming a small, 
rounded median lobe; barbels two on each side, the anterior 

3 the diameter of the eye in the young, 1} in the adult, the 
posterior longer, as long as the eye in the young, twice as long 
as the eye in the adult, the distance between them measuring 
4 (young) to 1 diameter of eye. Dorsal IV 9; last simple ray very 
strong, bony, smooth, its ossified part 5 to % the length of the 
head; the border of the fin feebly seneeanene - the distance of 
the dorsal from the occiput equal to or a little less than its 
distance from the root of the caudal fin. Anal with III 5 rays, 
not reaching, or nearly reaching, the caudal when folded. Pectoral 
fin pointed, as long as or a little shorter than the head, not 
reaching ventral; latter originating slightly in advance of first 


332 MR. G. A. BOULENGER ON FISHES [ Nov. 17, 


ray of dorsal. Caudal fin deeply forked. Caudal nee 1 to 


12 as longas deep. Scales finely striated, 30— 30 fe ee 25 ieee 


Ter line and ventral fin, 12 or 14 round caudal ‘pedunele. 
Silvery, bae sk olive-brown (in spirit). 
14 specimens, measuring from 65 to 270 millimetres. 
Wabbi, Modjo, and Irvaro Rivers. 
A very close ally of B. duchesnw Bler., differing in the shorter 
anterior barbels. 


12. BarBus PALUDINOSUS Peters. 


Several specimens from the Suksuk River. 


13. Barinius NILOTICUS Joannis. 


Several specimens from the Omo River, between Malo and 
Koscha. 


14, Bariurus Loar Bier. 


A single young specimen, same locality as the preceding. 


15. Nropota Borreci Vincig. 


Several specimens from the Wabbi and Modjo Rivers, and from 
the Omo River. 

The genus Veobola is very closely related to Chelethiops, from 
which it differs in the origin of the dorsal fin corresponding or 
nearly corresponding to that of the anal, and in the absence of a 
ventral keel. The snout of Neobola bottegi is shorter and more 
obtuse than that of Chelethiops bibie, and does not at all project 
beyond the mouth; the pectoral fin does not extend or extends 
but very slightly beyond the root of the ventral. The largest 
specimen collected by Mr. Neumann measures 73 millimetres, 


SILURIDA. 


16. CLARIAS ROBECCHIL Vincig. 


Three specimens, Hawash River and its affluent the Kassam 
River. 


17. CHILOGLANIS MODJENSIS, sp. n. (Plate XX XI. fig. 3.) 


Body slightly compressed, its depth 54 times in the total length. 
Head depressed, 1} as long as broad, its length 3 times in ne 
total length. Eye directed upwards, in the second half of the 
head, its ‘diameter 7 times in the length of the head, 13 or twice 
in ibe interorbital width, which is greater than ane distance 
between the eye and the poster 10r nostr il; premaxillary teeth in 
two large contiguous groups, forming 4 on 5 transverse series; a 
group ai 7 to 9 slender mandibular eeu: maxillary barbel + the 


1903. ] FROM GALLALAND AND SOUTHERN ETHIOPIA. 333 


length of the head, longer than the lower labials. Dorsal I 5; 
spine not serrated, not quite 2 the length of the head. Adipose 
fin low, its base measuring about 3 its distance from the rayed 
dorsal. Anal III 6-7. Pectoral spine not serrated, 3 the length 
of the head. Ventral extending to a little beyond the origin of 
the anal. Caudal peduncle twice and a half as long as deep. 
Dark brown above and on the sides, with three yellowish bars, 
the first in front of the dorsal fin, the second above the ventral, 
the third on the caudal peduncle ; lower parts white. 

Total length 40 millim. 

Two specimens from the Modjo River. 

The four species now known of this genus differ in the 
following characters :— 


A. Maxillary barbel much longer than eye. 


C. deckenii Peters.—Depth of body 54 to 61 times in total 
length; interorbital width equal to distance between posterior 
nostril and eye; D. 15; A. I11 6; pectoral spime more than 
= length of head. 

O. modjensis Blgr.—Depth of body 53 times in total length ; 
interorbital width greater than distance between posterior nostril 
and eye; D.15; A. III 6-7; pectoral spine 3 length of head. 

CO. niloticus Blgr.Depth of body 4 to 43 times in total length; 
interorbital width greater than distance between posterior nostril 
and eye; D. 16; A. III 8; pectoral spine # length of head. 


B. Maxillary barbel scarcely longer than eye. 


C. brevibarbis Bler.—Depth of body 6 times in total length ; 
interorbital width greater than distance between posterior nostril 
and eye; D.15; A. IIL7; pectoral spine § length of head. 

The new species may therefore be regarded as intermediate 
between the two previously described, C. deckenii from German 
East Africa, and C. niloticus from the Upper Nile. 


CyYPRINODONTID®. 


18. HapLocuiLus ANTINORI Vincig. 

A single specimen from the Suksuk River. 

The species was only known from Lake Arsade, south of Shoa, 
in the district of the Adda-Gallas, whence the types of the 
Discognathus chiarinit noticed above (p. 330) were also obtained. 


CIGHLID&. 
19. TruaApraA NiILoTICA L. 


Numerous specimens. Lake Gandjule, and Wabbi, Modjo, 
and Suksuk Rivers; also in the hot springs of the Hawash 
River near Filoa. 


304 MR. F, MARTIN DUNCAN’S DEMONSTRATION [Dec. I, 


EXPLANATION OF THE PLATES. 


Pratt XXIX. 


Labeo neumanni, p. 829, reduced }, with upper view of head, reduced 3, and open 
mouth, nat. size. 


Purate XXX. 


Barbus erlangeri, p. 331, reduced 2, with upper and lower views of head, same 
reduction. 


PuatE XXXI. 


Fig. 1. Discognathus makiensis, p. 330, nat. size, with upper (a) and lower (6) 
views of head, X13. 
2. Discognathus blanfordii, p. 330, do., do. 
3. Chiloglanis modjensis, p. 332, nat. size, with upper (a) and lower (6) views 
ot head, X 2. 


December 1, 1903. 
Dr. Henry Woopwarp, F.R.S., Vice-President, in the Chair. 


Mr. F. Martin Duncan exhibited, by means of the Bioscope, a 
series of pictures of Zoological subjects, and made the following 
remarks :— 


For many years past I have kept a permanent record of my 
investigations in various branches of Natural Science, by means 
of the Camera, using in the field various forms of hand and stand 
cameras, and in the laboratory a special photomicrographic outfit. 

My first application of photography as a means of recording 
scientific investigations was in helping my father (the late Pro- 
fessor P. Martin Duncan, F.R.S.) in his works on the Fossil and 
Recent Corals and Echinoderms. Although in many cases a 
single photograph or a series of photographs will, to a greater 
or less degree, demonstrate the appearance or phenomena under 
investigation, I have always felt in zoological work, particularly 
in studying the movements and habits of animal and insect life, 
that ordinary photography left much to be desired. Although a 
long series of photographs will give us an animal in various 
positions, very often the one most characteristic position desired 
is not obtained, and the point, therefore, missed. This is parti- 
cularly the case in dealing with the mode of seeking and capturing 
prey by the Carnivora; and when I brought my photographic 
efforts to bear upon work relating to the habits of wild animals 
in their natural environment, when seeking and seizing their food, 
I found the ordinary series of snap-shots failed lamentably to give 
a graphic idea of the facial expressions and characteristic move- 
ments of the animal. 

Jt was this want which led me primarily to consider the advisa- 
bility of applying Animated Photography to Zoological study. 
As a lecturer and teacher of Zoology and Botany, I felt at once 
that if it were possible to place before my audience or class “living 
pictures” of animal, insect, and vegetable life, I should have a very 


1903. ] OF THE ZOOLOGICAL BIOSCOPE. 335 


important educational factor in my hands. Modern methods of 
teaching all go to prove the importance of teaching through the 
agency of the eye as well as the ear, and it is a matter of common 
knowledge amongst teachers that a lecture demonstrated by a 
graphic series of experiments or pictures is much more vividly 
impressed upon the minds of the students than a simple unillus- 
trated oration. 

As regards the application of Animated Photography to the 
Microscope, I was primarily led to attempt it through my inves- 
tigations into various diseases of plants produced by Bacteria. I 
felt that the ordinary method of preparing bacteria for microscopic 
examination,—to wit: the fixing of the organisms to the cover-glass 
by passing the cover-glass through the flame of a spirit-lamp, and 
then staining (more often than not in heated stain),—gave one 
anything but an accurate idea of the natural appearance of the 
organisms under examination. It appeared to me that, working 
on such lines, one might as well take a sheep out of the meadow, 
roast it whole in front of the fire, dip it into a bath of stain, and 
then hold it up as a true likeness of what a sheep browsing in a 
meadow appears like. Also, in lecturing, I felt that it would be a 
great advantage if I could project on to a screen an “animated 
photograph” of such subjects as the circulation and rotation of 
protoplasm, the circulation of the blood, &e. 

Of course, the great difficulty in seeing living bacteria, with any 
degree of comfort, is the close affinity of their refractive index to 
the media in which they are cultivated. However, after some 
months of experimenting, and trying various optical formule, I 
succeeded in finding a combination of lenses which would permit 
of accurate examination of living, unstained bacteria. The results 
have been very interesting, and I hope on another occasion to have 
the pleasure of bringing them before the notice of the Society. 

Increase of teaching and literary work put a stop for a period 
to my attempting to experiment with Animated Photography in 
my work ; but I was fortunate enough, some little time back, to join 
hands with Mr. Charles Urban (of the Charles Urban Trading Co., 
Ltd.), who has very great experience in Cinematography and is the 
inventor of the Bioscope. The difficulties that had to be overcome 
in applying Cinematography to Natural Science subjects, particu- 
larly zoology, and to the microscope, were many, but have now 
practically been overcome. 

How far I have succeeded in applying Animated Photography 
to Natural Science, as a means of recording and demonstrating 
phenomena, you will now have an opportunity of judging for 
yourselves. I will now project a series of Animated pictures 
showing the life of the Hive-Bee; a comparison of the mode of 
capturing food employed by three reptiles, viz., the Toad, 
Chameleon, and Boa Constrictor ; and also some of my Animated 
Photomicrographs showing the circulation and rotation of proto- 
plasm in Zlodea, the Freshwater Hydra, &c., ke. 


336 ON SPECIMENS OF NAKED RODENTS FROM E. AFRICA, [Dec. 1, 


My. F. E. Beddard, F.R.S., exhibited and made remarks upon 
a portion of the large intestine and the cecum of a Boa (Boa 
constrictor) which had recently died in the Society’s Gardens. 
The walls of the intestine in the neighbourhood of the cecum, 
and of the cecum itself, were thickened and inflamed. The cecum 
was filled with a hard mass consisting of small stones and a number 
of the snake’s own teeth, the presence of which it was thought had 
given rise to the inflammation. 


Mr. Beddard also exhibited, on behalf of Mr. G. A. Doubleday, 
a hairless specimen of the Common Rat (A/ws decumanus) which 
had been caught in a trap at Leyton, Essex, and which agreed in 
its characters with a so-called variety (Jfus nudo-plicatus) of the 
Common Mouse figured in the Society’s ‘ Proceedings’ (1856, p. 38, 
Mamm. pl. xli.). 


Dr. Walter Kidd, F.Z.8., exhibited a drawing of a Beisa 
Antelope (Oryx beisa) showing a reversed area of hair along the 
median line of the back, a character which was found in 
Ruminants only, but not in all of them. 


Mr. Oldfield Thomas, F.R.S8., exhibited an example, the second 
known, of the peculiar little naked rodent described by him in 
1885 * as Heterocephalus phillipsit. ‘This specimen had been 
presented to the National Museum by Dr. A. G. W. Bowen, 
R.N., to whom it was given, at Mogadishu, Italian Somaliland, 
by Dr. Dulio, Governor of the Italian Protectorate. 

Dr. Dulio stated that these animals lived in colonies of from 
50 to 100 in the sandy districts near the coast, that they burrowed 
very rapidly in the loose soil and were for that reason difficult to 
obtain. 

Mr. Thomas said that the chief interest of this specimen lay in 
the fact that, like the type, it had only two cheek-teeth in each 
jaw above and below, while the larger and better-known Hetero- 
cephalus glaber had three. As this important character was thus 
shown to be constant, which could not be assumed from the single 
specimen hitherto available, Mr. Thomas thought that a special 
genus would have to be formed for the two-toothed group, and 
suggested for it the name of Yornarinat. The other differences 
between Yornarina phillipsi and Heterocephalus glaber had been 
more fully detailed in the papers referred to in the footnote £. 

A form of true Heterocephalus occurred also in British East 
Africa, which Mr. Thomas took this opportunity of describing :— 


HETEROCEPHALUS ANSORGEI, sp. n. 
General characters as in H. glaber, but size smaller (see skull- 


* 


P. Z. 8. 1885, p. 612. 
A famous nude by Titian. : 
P. Z. S. 1885, p. 845; Ann. Mus, Genov. (2) xv. p. 3 (1898). 


bee 


1903. ] ON HAIR-WHORLS IN THE OKAPI. 337 


measurements) and the cheek-teeth very much smaller in diameter, 
their transverse and longitudinal diameters subequal. In H. glaber 
the transverse diameter of the middle tooth considerably exceeded 
the longitudinal. Palate ending almost immediately behind the 
last molar. Incisors feebler than in H. glaber, the inner half of 
their anterior surface slightly concave, in correspondence with the 
ill-defined grooves found in this position in H. glaber. General 
shape of lower jaw as in H. glaber, but the teeth equally modified 
with those of the upper. 

External characters, apart from size, apparently quite the same 
as in H. glaber, but the tail has been lost in the single specimen. 

Dimensions :— 

Head and body (approximately) 94 mm.; hind foot 20-5. 

Skull—front of nasals to junction of sagittal and lambdoid crests 
19 mm.; greatest zygomatic breadth 16°8 ; nasals 6°8 x 3-4 ; inter- 
orbital breadth 7:3; intertemporal breadth 5:6; palate, length 
from henselion 10°3; diastema 7; combined length of three cheek- 
teeth 3°2; transverse diameter of middle tooth 1:2. Lower jaw, 
back of angle to front of symphysis 18-6; to back of coronoid 11 ; 
lower tooth-series 3°3. 

Hab. ‘“ Between Ngomeni and Kinani,” Makindu country, 
British East Africa. 

Type. Male. B.M. No. 98.9.25.3. Collected 31 October, 1896, 
and presented by Dr. W. J. Ansorge. 

Dr. Ansorge had noticed that this Heterocephalus “‘ was throwing 
out earth minh its hind feet from a tiny circular hole at the 
bottom of a small crater-shaped mound of red earth.” 


Mr. G. A. Boulenger, F.R.S., exhibited a young hybrid Newt 
(Molge marmorata 3 x M. cristata 2) bred by Dr. Wolterstorff, 
of Magdeburg, in his aquarium, as reported in the ‘ Zoologischer 
Anzeiger,’ Sept. 21, 1903. This specimen agreed in all external 
characters with JM. blasvi de VIsle, of which one of the original 
specimens, from near Nantes, 8. Brittany, forming part of 
M. Lataste’s collection, was also exhibited. 


Prof. KE. Ray Lankester, F.R.S., exhibited two drawings repre- 
senting the arrangement of the hair on the fronto- -par ietal sur face 
of the head of two specimens of Okapi. The drawings are re- 
produced in text-figs. 41 and 42. Text-fig. 41, p. 338, is from the 
subadult female sent over by Sir Harry Johnston, described by 
Prof. Lankester in the Trans. Zool. Soc. vol. xvi. p. 279, and now 
mounted and exhibited in the Natural History Museum. The 
second drawing (text-fig. 42, p. 339) is from a smaller specimen of 
an apparently adult female m the possession of the Hon. Walter 
Rothschild at Tring, by whose kind permission the drawing is 
published. Prof. Lankester made the following remarks :— 

The hair is represented diagrammatically in both cases, the 
arrow-heads corresponding to the free ends of the hairs. 


Proc. Zoou. Soc.—-1903, Vou. Il. No. XXII. 22 


338 _ PROF, E. RAY LANKESTER ON wee, ik 


Text-fig. 41. 


moan WOM yy = LY: 
A NANAN Me 
AN nn ez 
aw \\\ \\ \\ i WZ. va 
SS AN) ANY Wee Wes 
OARAN LSS = ZL, o- 
SAA AWAE Ae 
~ S RYN \SS=7 i pa = 
~ SSS eis : 
Ny eS 
Ss 
SS 


SS 
SSS SSS = 


SS 
UN 
SEQ 
—> =~ 

Ss \\ NX 


ee ape 
HH MON SSESSSSS SSS \ 


Y \ 
ZU NY = 
ee \ oe 
See At i Hl BINS SSS 
SI i HIMWWSN SSS SS SB 
SZ i, SSS 
SEW | (aS Ze 
Zig 
i, Hi VAY / == Zo 
GLEE 
Lujgg 
Gy 
WW 


Yy 
/ 
Mf 


i 
Hy, 
AN 
{\ 
\ 


: 
‘ 


| 


Y 
| 


G44 
ty, 
Yih 

yf 

Ye 

, 

( 

ly 


Ey 
7 


ZS 
Ze 
Ziy 
in 


Za 
4 
EE 


<=, 


Z 
a 


Ss 


| i ARS SGX ‘ZG, jf “Wf Y, 
[IN VANS SSNS I IY 
Hgitin’s Nee SS MY h 


Diagram of the hair-whorls and hair-streams on the fronto-parietal region of the 
head of a female specimen of Okapi in the British Museum. 


Okapia erichsoni Lankester. (Sir Harry Johnston’s specimen, presented to the 
British Museum.) 


b, b, paired bare spots corresponding in position to the horns of the male. Mediad 
of the right-hand spot is a bare patch caused by rubbing; it is not repre- 
sented on the left-hand side of the head. 

_d, single supra-nasal vortex or meeting-point of three hair-streams. 

wv, single spiral whorl on the left frontal region (asymmetrical in position). 

¥, occipital radiating centre. : 


1903. HAIR-WHORLS IN THE OKAPI, 359 


Text-fig. 42. 


QW it WW’ 
WW ZZ 
SN Me 


i QW! 
eS SSS SSS £2 Zp ZEEE SD, 
i ee 
|) Ve ee 
SSE S| SS 
SS ZR yy 
SSS SS 
SS PEEWIIR-CR SS SRR 
EF ”- VW RS SV 
SSS OOOO 
SHH NN 
= iy, ( i INS S 
—— Zw get ly 
Se, GEN|| AV! VX 
=v NY WS 
EN ENE 
: S| HEN ‘i 
EW VRS aN 
SEE AES 
ESS QS 
INS; A NV 
SS) ISNA 
SSS aN IN KA RR 
SS 
SSS AY / 
SSSA 
ASA WA 
SYS SSS \\ Rat (ty WZ 
SSW MZ 
. SS 
i SSA 
a SSSA 
QN SQV \ i 
RN RAS RAM AY | 47, LY 
Ne . SS WW. Vy Fi, 
< INNS Sani Wall hq, / Wifi, 
HR CG 
i ay) ANN SANA Re ws Yi 
NUN I A 
WSIS NRA) 
WIN SSS IRS 
SS ZL Soe aera 
i INN SSS ZZ INNS} i 
WS eS 77/1 
HWINWSS SSSR 411 
"in OSS SSL 4 hi | \ 
UBS See 7 
TSA TAN 
ANN EAN 
EIN TT \ \\ 
TS CTR iN 
YENI) NOMA 
ZL NTS SN 


‘Diagram of hair-whorls and hair-streams on the fronto-parietal region of the head 
of a small adult female specimen of Okapi in Mr. Rothschild’s Museum. 


a, median inter-cornual spiral whorl and meeting-point of hair-streams (not 
present in text-fig. 41). 
b, 6, as in text-fig. 41. 
e, ¢, right and left inter-cornual radiating centres (not present in text-fig. 41). 
y, occipital radiating centre, as in text-fig, 41. : 
22% 


340 ON HAIR-WHORLS IN THE OKAPI. [Decrae 


Tt will be observed that there is considerable difference between 
the arrangement of the hair-whorls, hair-spirals, hair-radiant- 
centres, and hair-streams in the two specimens. We do not yet 
know enough of these curious dispositions of the hair-growth to 
say what value should be attached to one or other difference as 
indicating distinctness of race or species—or whether, indeed, any 
such value can be attached to these structures. 

The publication of this pair of diagrams taken from so interesting 
a creature as the Okapi will, it is hoped, lead to a more extensive 
and complete study of the subject, not only in the various skins 
of Okapi which may be hereafter or have been already received, 
but in large series of other species—such as the Antelopes. 

The differences between the two specimens of Okapi in regard 
to the hair-whorls are best to be appreciated by an examination of 
the diagrams (text-figs. 41, 42, pp. 338, 339) and the explanation 
given below the figures. But it may be briefly pointed out that the 
British Museum specimen (which it will probably be necessary, as 1 
have elsewhere pointed out, to distinguish as Okapia erichsonz) has 
a single median supra-nasal or interorbital vortex or meeting-point 
of three hair-streams (text-fig. 41, d) which is not present in 
Mr. Rothschild’s specimen. Also it has an asymmetrical single 
spiral whorl of hairs on the left frontal region (text-fig. 41, a). 
This also is not present in Mr. Rothschild’s specimen. 

On the other hand, in Mr. Rothschild’s Okapi there is a median 
spiral vortex or meeting-point of hair-streams between the sites 
corresponding to the position of the horns in the male (text-fig. 42, a). 
Also there are, at no great distance from this median spiral, a 
right and a left “radiant centre” (text-fig. 42, ¢,c) from which 
hairs radiate evenly in every direction. Nothing corresponding 
to these is seen in the British Museum Okapia erichsoni. 

The spiral vortex a of the Tring specimen may perhaps be con- 
sidered as represented in some fashion by the one-sidedly placed 
spiral vortex « of Okapia erichsoni, as though a@ had moved 
forward and to the animal’s left. 

In connexion with this apparent asymmetry, I may draw 
attention to the fact that Dr. Forsyth Major has found that the 
horns of the male specimens of Okapi submitted to him by the 
Congo State Museum are invariably asymmetrical. 


Prof. Lankester also exhibited and made remarks upon some 
specimens of Medusze reported to have come from the Victoria 
Nyanza, and sent home by Sir Charles Eliot, K.C.M.G., H.B.M. 
Consul-General and Commissioner in the British East-African 
Protectorate. 


The following papers were read :— 


1903. ] ON THE MAMMALS OF CYPRUS. 341 


1. The Mammals of Cyprus. By Dororay M. A. Barr *. 
| Received October 26, 1903. ] 
(Text-figure 43.) 


1. Introduction. 


A list is given below of all the species of mammals—other than 
domesticated—known up to the present date to occur in Cyprus. 

Hitherto very little has been published with reference to the 
Mammalia of this island. Drs. Unger and Kotschy in ‘ Die 
Insel Cypern’t recorded eight species; in 1879 Dr. Giinther £ 
mentioned four species in a note on a collection of mammals and 
reptiles from Cyprus; and recently the Hare and Spiny Mouse 
have been described as peculiar to this locality. In the collection 
of the British Museum (Natural History) there are also some 
specimens procured by various collectors and presented by the 
late Lord Lilford. 

This present list is chiefly compiled from material obtained 
during a visit of eighteen months to the island in 1901 and 
1902. Two bats, Rhinolophus blasii and Myotis myotis, previously 
recorded from Cyprus, were not included in my collection. 

Compared with the number found in other Mediterranean 
islands, and considering the extent of Cyprus, which contains 
3584 square miles, Mammals are but poorly represented, inasmuch 
as the total number of species does not exceed fifteen. It might 
be thought that the dry and barren condition of the greater part 
of the country is the cause of such lack of variety, but this 
appears unlikely when it is remembered that the character of 
the island has completely altered within historic times. This 
change has been due chiefly to deforesting, which for long con- 
tinued unchecked, with the natural consequence that a large 
proportion of the rainfall, which is small, is now wasted. Parts 
of the central mountain-range still support a somewhat scanty 
growth of pines, ilex, and arbutus, but these have little chance 
of spreading, for most of the seedlings are either burnt up by 
the great summer heat or destroyed by the numerous flocks of 
goats. In former days even the Mesorcea, which stretches from 
the west coast to the east, was densely clothed with trees, whilst 
now it is a bare, dusty plain save for a few months in the year 
during the growth of the cereal crops. 

1 am much indebted to Mr. Oldfield Thomas for his kind 
assistance and for so courteously giving me access to the specimens 
in the Collection of the British Museum (Nat. Hist.). 


Il. List of Species. 
1. Roussetrtus meypriacus E. Geoftr. 


A Fruit-Bat has long been known to occur in Cyprus, though 


* Communicated by OLpFrELD Tuomas, F.Z.S. 
+ Wien, 1865. {t Proc. Zool. Soc. 1879, p. 741. 


342 MISS DOROTHY M. A. BATE ON THE [ Dee. 1, 


yal 


not always considered to be of this species. However, the 
question of its identity has been fully discussed by the late 
Dr. Anderson *, who came to the conclusion that it ought, without 
doubt, to be referred to as Rousettus egyptiacus, which is also 
found in Palestine and Egypt. 

Excessively common in Cyprus, this Bat often does considerable 
damage to the fruit-crops, particularly to the oranges and dates, 
though whilst the latter are ripening they are often enveloped in 
sacks or matting as a protection against the depredations of this 
marauder, During the summer it is said to roost in thick trees, 
in winter taking up its quarters in the roofs of old buildings and 
caves in company with rock-pigeons. Hxtremely noisy, even in 
the daytime and when undisturbed, its shrill note can be heard 
some little distance off. On entering a cave numbers may be 
found hanging head downwards from the roof in closely packed 
bunches ; very restless, every moment one or two detach them- 
selves to fly off squeaking from one group to another. 


2. RHINOLOPHUS FERRUM-EQUINUM Schreb. 

Only one specimen of the Horseshoe Bat was procured, which, 
with the following species, had already been obtained by one of 
the late Lord Lilford’s collectors. 


3. RHINOLOPHUS HIPPOSIDEROS Bechst. 


Common, found in caves. 


4, RHINOLOPHUS BLASIT Peters. 


Four specimens of this Bat were obtained by one of the late 
Lord Lilford’s collectors and are now in the collection of the 
British Museum. I did not meet with it during my stay in the 


island. 


5, PrpisTRELLUS KUHLI (Natt.). 

This seems to be the commonest of the small Bats of Cyprus. 
Its occurrence was recorded by Dr. Giinther in 18797. It is 
unpleasant to handle owing to its heavy, disagreeable odour. In 
June 1901 several were brought to me, one being kept in a box 
for the night. The following morning it was found that two 
young ones had been born; they were perfectly naked, and 
appeared pink owing to the transparency of their skins. The 
mother carried one on either side of her body under the wings. 


6. Myoris myoris (Bechst.). 

This species is included in Unger and Kotschy’s list under the 
name of Vespertilio murinus Schreb., but was not amongst those 
I procured. 


* ‘Zoology of Egypt,’ Mammalia, pp. 87-89 (London, 1902). 
¥* Proc. Zool. Soc. 1879, p. 741. 


1903. | ; MAMMALS OF CYPRUS. 343 


7. MiNIoPrERUS SCHREIBERSI (Natt.). 
I obtained a single specimen of this Bat in a small cave in the 
sea-cliffs at Cape Pyla, Famagusta District. 


8. ERINACEUS AURITUS Gmel. (Text-fig. 43.) 


Three examples were obtained of this Hedgehog, which had 
already been recorded from Cyprus*. These appear to resemble in 
every respect specimens from Egypt in the collection of the British 
Museum. The species is common over the greater part of the 
island, and is said to be eaten by the natives. In June 1902 a 
villager brought me three young ones, which I kept for some 


Text-fig. 43. 


fare 
¥e, 
ery tiyhy, “t 
ye, TP riya, 
bes 
= z 


eae 
Long Ft tage 
Wig” ton t 


on hee 


EHrinaceus auwritus. 


months with an old one found subsequently. They were very 
active, and neither shy nor frightened even when first caught. 
They fed and allowed themselves to be handled without showing 
any signs of fear, unlike our English hedgehog under similar 
circumstances. Amongst themselves they were extremely pug- 
nacious, continually fighting, and at the same time giving vent 
to loud squeals resembling the “ miaoul” of a cat. A favourite 
mode of attack was to seize hold of one of their opponent’s feet, 
thus effectually preventing it from retreating within its protecting 


* ‘Zoology of Egypt,’ Mammalia (Dr. Anderson), p. 158 (1902). 


344 MISS DOROTHY M. A. BATE ON THE ‘[Dee. 1, 


spines. They fed well on bread and milk, hard-boiled eggs, and 
small pieces of raw meat. A Cypriote told me that in summer 
hedgehogs are often found in the vineyards, where they feed on 
the grapes, which they would have no difficulty in reaching, for 
in Cyprus the vines are not trained on poles, but are every year 
eut back, only a short stump being left, so that the grapes may 
often be seen lying on or almost touching the ground. Theaccom- 
panying photograph (text-fig. 43, p. 343) is of a young animal 
taken shortly after its capture. 

Erinaceus europeus is included in the list of mammals given by 
Unger and Kotsehy *, but this name, no doubt, was meant to 
apply to #. auritus, which appears to be the only species of hedge- 
hog found in the island. 


9. CROCIDURA RUSSULA CYPRIA, subsp. n. 


The Shrew obtained undoubtedly belongs to what may be 
termed the C. russula-group. Taking that from Western Europe 
as the typical form, it will be found that races intermediate 
between it and the one from Cyprus occur in different parts of 
the Continent. It is therefore proposed to distinguish the 
Cypriote Shrew by the above subspecific name, denoting the 
habitat of this island race. It differs considerably from typical 
specimens, from which it may be readily recognised by the 
following characters :—Size and general build shghter, approaching . 
that of C. whitakert de Winton, from Morocco* ; skull smaller, tail 
longer, and average length of fur much shorter, this being parti- 
cularly marked in the summer coat. 

The three examples procured are all females—one, caught in 
March, being in winter coat, while the other two, caught in July, 
show the lighter-coloured and shorter summer fur. The following 
measurements (in millimetres) were taken in the flesh, and those 
of a specimen from Liége in the collection of the British Museum 
are given for comparison :— 


Ae, Tail. Hind foot. Ear. 
INOS GGNGO)) ral ener eecueenieln 58 465 11 a 
ING LTC Ors poet genet 60 Al 11 9 
IN@ HIM ( Q Ai77BS)) acdascovoooe 60 A3 12°5 10 
Specimen from Lieége(@)... 79 37 12°5 — 


(Gey WE, ING; YH IIo.) 


Only one skin (No, 111) is accompanied by a skull, though 
several portions of skulls found in the earth of a cave in the 
Kerynia Hills are, in every respect, similar to the recent one. 
This last measures 19 mm. in extreme length and 9 mm. in 
breadth, whilst the corresponding measurements of the skull 
belonging to the specimen from Liége mentioned above are 
21 mm. and 9-5 mm. respectively. 


* Op. cit. 
+ Proc. Zool. Soe. 1897, p. 954. 


1903. ] MAMMALS OF CYPRUS. 345 


The type is now No. 3.12.4.23 of the British Museum Collection. 

The Cypriote Shrew in the colour of its winter coat resembles 
the browner and less speckled specimens from Western Europe, 
although the average length of the hairs is very much less, being 
3-5 mm. as opposed to 5 mm. In summer the hairs are shorter 
still, averaging only 2°5 mm.; also at this season the fur of the 
dorsal region is much lighter and greyer. 

I have been unable to find any previous record of the oceur- 
rence of a Shrew in Cyprus, and it seems likely that it is not 
very plentiful, for I succeeded in trapping only three specimens, 
all of them being caught within a hundred yards of the same 
place. This was close to a stream, flanked by steep banks thickly 
clothed with grass, brambles, and other plants, not far from 
Papho in the south-west of the island. 


10. VuLPEs vuLPEs (Linn.). 


It was expected that the Cypriote Fox would resemble that of 
Syria, which is evidently identical with the V. persica of Blanford *. 
However it is quite distinct, being a small red form very similar 
both in size and coloration, though somewhat yellower dorsally, 
to a specimen from Asia Minor in the National Collection, the 
skull of which was unfortunately not preserved. 

The Cypriote race probably ought to be included in one of the 
subspecies of the typical form, and may perhaps be found to be 
identical with the Fox of Southern Europe (V. melanogaster of 
Bonaparte)?. Its skull agrees in size with that of V. v. egyptiaca. 

Five specimens of this fox were procured, four in thick winter 
coats and one in summer, which latter was shot on Troddos at a 
height of about 6000 feet above sea-level. In this the fur is very 
thin and scanty, and except on the head and limbs has lost almost 
all trace of red or tawny colouring, being brown on the back and 
dull brownish white on the underparts. 

It is commonly found all over Cyprus, and is the only wild 
carnivore known to occur in the island at the present day. It is 
especially plentiful about the cliffs of Cape Pyla, where the natives 
shoot large numbers in winter, selling the skins to be made into 
rugs. 


11. Mus ratrrus Linn. 


This species is represented in Cyprus by the brown form with 
lemon-white underparts (JZ. tectorum, Savi). In spite of the 
reward offered by Government for the destruction of these rats in 
the port-towns (to guard against the introduction of the bubonic 
plague by their means), they are excessively common, being found 
everywhere, both in or near buildings and at a distance from any 
inhabited houses. Mr. Gennadius, Director of Agriculture in 


* Zool. Hast. Persia, 1876, p. 39, pl. 11. 
+ Icon. Fauna Ital. 1832, i. pl. i. 


346 MISS DOROTHY M. A. BATE ON THE | Dee. 1, 
Cyprus, writes * that they cause much damage to the Karoub trees, 
the fruit of which forms one of the most important exports of the 
island. They are very easily trapped, and were especially plentiful 
in the Papho District close to the streams, many of which have 
high banks densely covered with brambles and bushes. In this 
growth, generally at a height of eight or ten feet, they build their 
nests, which are loosely made of leaves and coarse herbage, and 
lined with finer grasses, or else entirely composed of leaves of the 
tall bamboo-like reeds that here and there edge the streams. The 
entrance is at one side of the nest, which is flatly domed like a 
squirrel’s drey, and, from a casual inspection, would appear to be 
nothing more than a bunch of rubbish or dead leaves, which the 
thickness of the vegetation prevented falling to the ground. 

These rats move about without difficulty amongst trees and 
bushes, often at a considerable height above the ground. One 
evening, whilst passing through an orchard, I saw one sitting in a 
clump of brambles about a couple of yards away and busily feeding 
on the blackberries which it held to its mouth with both fore paws 
in an attitude similar to that adopted by a squirrel when eating a 
nut. These rodents also make use of holes in banks and amongst 
rocks, making it seem likely that the nests built in bushes may be 
used only during the warmer months of the year. 

I never met with any examples of the typical black form, but 
was told of some having been seen in a store-house at Larnaka, 
the chief port of the island. 

The Brown Rat from Cyprus has previously been recorded as 
Mus decumanus t and also as MW. alexandrinus, but the latter name 
appears to be strictly applicable only. to the brown form of JZ. rattws 
with grey underparts—a house-haunting race {. 


12. Mus MUSCULUS GENTILIS Brants. 


The specimens of this mouse from Cyprus resemble those from. 
the Soudan in the collection of the British Museum, the fur of 
the underparts being pure white, with the proximal ends of the 
hairs grey in only some examples. 

In the five specimens obtained the tails are long, in each case 
exceeding the length of the head and body, in this respect 
reversing the proportions that obtain in the wild form of 
M. musculus found in Portugal $. This mouse is common in the 
island, being found not only in the open country but also in out- 
houses and buildings near towns, and although it seems most 
likely, I am unable to say whether or not this form also haunts 
the houses in towns. I do not remember seeing any of the darker 
coloured typical W/. musculus. 


* “The Carob Tree,’ by P. Gennadius. Nikosia, Cyprus, 1902. 

+ Unger and Kotschy, op. cit. 

{ See ‘Zoology of Eygpt,’ Mammalia (Anderson), 1902, p. 274. 

§ Mr. Oldfield Thomas, “On a wild-living Mouse of the Mus musculus group in 
Portugal,” Zoologist, vol. xx. 1896, pp. 137-9. 


1903. ] MAMMALS OF CYPRUS. 347 


13. Acomys NESIOTES Bate. 


From the size and general appearance of the adult of this species 
it might be considered an island form of A. dimidiatus, which it 
resembles somewhat closely. On the other hand, the grey 
coloration of the young, which persists until the animal has 
attained almost its full size, points to a possible connection 
between A. nesiotes and A. cahirinus. 

A description of the Cypriote Spiny Mouse has already been 
published*. At that time it had only been procured near Dikomo 
at the foot of the Kerynia Range, but it has since been found 
near Limassol in the south, so that it probably occurs over the 
greater part of the plains and low hills in the island 7. 

The type specimen is now No. 3.12.4.37 of the British Museum. 


14. Lepus cyprius Barr.-Ham. 


Mr. Barrett-Hamilton’s description of the Cypriote Hare was 
founded on three skins—accompanied by two very much damaged 
skulls—presented to the National Collection in 1875 by the late 
Lord Lilford. The specimens procured by me do not agree very 
well with these skins, which are possibly somewhat faded ; therefore 
this further material being now available, it is thought that a few 
additional notes on the species may be of interest. The following 
measurements (in millimetres) were taken in the flesh :— 


Head and body. Tail. Hind foot. Far. 
INTO (isi a bp eeeennase 558 89 133 107°5 
INO» GILG.) saansseocsese D30 82 120 107°5 
IN@s OGG) eacdcosssabe 508 1K 126°5 105 
Notg3 (2) sae 546 89 126-5 1015 


The greatest and basilar lengths (in millimetres) of the two skulls 


obtained are as follows :— 
Greatest length. Basilar length. 


INO» UGG. (CG) eoccabanseee 95 76 
Is 168 (GS) coceseooe ee 90 Tel 


_ In size L. cyprius is apparently similar to L. ereticus, a specimen 

of the latter being 21 inches (532°5 mm.) from the tip of the nose 
to the end of the tail (collector’s measurement). The skulls of 
these two species are also much the same, the total length of that 
of one of the type specimens of Z. creticus being 97°5 mm. 

The newer examples of the Cypriote Hare resemble the Cretan 
species more closely than does the type specimen, particularly in 
showing the light-coloured rump and a slight amount of white on 
the anterior surfaces of the ears, but lack the grizzling on the upper 
lips which is present in all the five skins from Crete in the British 
Museum Collection. 

* Ann. Mag. Nat. Hist. ser. 7, vol. xi. pp. 565-7 (1903). 


+ See P.Z.S. 19038, vol. ii. p. 260. 
+ Ann. Mag. Nat. Hist. ser. 7, vol. xi. pp. 126-7 (1903). 


348 MR. REDCLIFFE N. SALAMAN ON AN [ecaae 


The Cypriote Hare is distributed all over the island and is 
plentiful in many parts, in spite of the numbers that are every 
year shot by the natives. They not infrequently do a considerable 
amount of damage in vineyards, and consequently are looked upon 
as vermin and, subject to certain restrictions, may be killed at any 
time of the year and without a license. 

This hare is included in Unger and Kotschy’s list under the 
name of Lepus timidus. 


15. Ovis opHrion Blyth. 


No list of the mammals of Cyprus would be complete without 
mention being made of the Moufflon, which is still found in the 
forests of the western part of the Troddos Range, this being the 
wildest and least inhabited district in the island. On more than 
one occasion during the summer of 1902 some were seen on Olym- 
pus, the highest point in the island, and probably the most easterly 
to which they now roam. 

During my visit in 1901 and 1902 none were allowed to be shot, 
and at all times a special license is necessary. In spite of this it 
is supposed that many are still killed by the peasants; I myself 
saw several skins and horns, mostly of immature specimens, that 
were undoubtedly illegally come by, to judge by the secrecy with 
which they were brought for inspection. As a rule the horns are 
hidden or thrown away in the forest, where some have subsequently 
been found by the police. 

The measurements of the largest horns of the Cypriote Mouffion 
mentioned in ‘ Records of Big Game’ * are :—Length on front 
curve 25 inches, and circumference at base 8 inches. The owner 
of this trophy is given as Mr, H. Williamson, but it is probable 
that the one referred to is the very handsome mounted specimen 
fae and presented by him to the English Club at Limassol, 

sy prus. 


2. On the Cause of Death of a Polar Bear recently living in 
the Society’s Gardens. By Repcrirre N. SaLaman, 
M.A., M.B. Cantab., F.Z.S., Acting Director of the 
Pathological Institute, London Hospital. 


{Received November 12, 1903. ] 


A fine well-grown male Polar Bear (Ursus maritimus), pre- 
sented to the Society by Mr. Arnold Pike in 1895, died suddenly 
on Noy. 1. There was no history of illness, and the animal had 
been taking its food regularly and was in no way emaciated. A 
preliminary examination was made on Monday, Noy. 2, and I 
completed the post-mortem next day. ; 


* Rowland Ward, 3rd edition, 1899, p. 385. 


1903. ] ANEURYSM IN A POLAR BEAR, 349 


On opening the Thorax an enormous quantity of recent blood 
was found filling both pleure. 

The Lungs were collapsed owing to the large effusion, but 
showed no signs of any disease, nor was there any sign of 
pleurisy. 

It was interesting to note that the lungs showed almost as 
much anthracosis as is found usually in the lungs of human city- 
dwellers. 

The Heart and the Aorta to within a few inches of the 
diaphragm were found to be perfectly normal; however, the 
latter part of the dorsal aorta was found to be bound down by 
recent adhesions to the vertebral column and on its antero-mesial 
surface to be fused to a structure about the size of a hen’s 
egg. On removing the latter six inches of the dorsal aorta and 
the new fibrous structures above described, it was found that the 
aorta was ruptured on its anterior surface, forming a hole (4" in 
diameter) which led into a cavity contained within the above 
fibrous mass. This cavity had a volume equal ,to that of the egg 
of a blackbird; it was filled with greyish and red laminated clot. 
The walls of the cavity consisted therefore of clot and of fibrous 
tissue on the outer, left, and anterior surfaces, whilst on the 
mesial surface the laminated clot came into direct contact with 
the bodies of the vertebra, which were slightly eroded. At its 
posterior extremity this sac had ruptured into the thoracic cavity. 
There was evidence that there had been some slight leakage at 
any rate some hours before the fatal hemorrhage, as there was 
found on the posterior surface of the left lung a blood-stained 
roughened area which corresponded in life to the position of 
the sac. 

We have therefore to deal with an aneurysm of the aorta, itself 
perfectly healthy except for the small rupture which forms the 
aperture into this aneurysmal cavity. No part of the aortic wall 
has shared in the formation of the aneurysm, which therefore 
falls into the group known as “false aneurysms.” 

The etiology of the morbid process remains obscure, and, not- 
withstanding the fact that the esophagus was without obvious 
lesion, it is conceivable that, at some time previous, some sharp 
bone, penetrating the cesophageal wall, may have lacerated the 
aortic wall, and thus been the starting-point of leakage from the 
latter and formation of a “ false aneurysm.” 

I am indebted to Professor M°Fadyean for having kindly 
informed me that aneurysm is, so far as he knows, an unknown 
condition in wild animals and excessively rare in the domestic 
carnivora. 

The aneurysms in horses are examples of dilatation of the 
aorta, and are quite distinct from the aneurysm found in this 
case. 


390 DR. A. G. BUTLER ON THE PLUMAGE (Dewar 


3. On the Development of the Adult Colouring in the Yellow- 
billed Cardinal (Paroaria capitata) from 8S. America. 


By Artur G. Buruer, Ph.D., F.L.S., F.Z.8., &e. 
[Received November 17, 1903. ] 


In the ‘ Avicultural Magazine’ for July 1903 (pp. 294-298) I 
had occasion to point out that Petronia albigularis of Brehm was 
not an immature plumage of P. dentata, but a distinct species, 
specimens of both birds having been in my possession for over five 
years, and, though unaccompanied by a cock of either species, 
building and laying at intervals during the whole time in which 
I owned them. 

In the same Magazine for December 1903 I have a short paper 
with an opposite tendency, dealing with two forms of Paroaria 
which have hitherto been regarded as distinct species. I have 
not, however, entered into details, in that brief account, of the 
method of the growth in colour which gradually established the 
identity of the supposed species: this I now propose to do. 

In July 1893 I imported a fully adult male of Paroaria capitata 
in perfect plumage; unhappily it was subjected to rough treat- 
ment during the journey home from La Plata, and so only 
survived about six months. However, I hada beautiful skin made 
of it, thus enabling me to have something better than a mere 
illustration for future reference. 

This year (September 1903) a consignment of Lesser Cardinals 
was brought into the London bird-market by an Italian, who sold 
his stock to an enthusiastic friend of mine, Mr. J. B. Housden 
of Sydenham, who later brought some of the Cardinals to me as 
‘““Yellow-billed Cardinals” (P. capitata); but, as soon as I saw 
them, I knew that the more advanced examples were unquestion- 
ably the so-called Brown-throated Cardinal (P. cervicalis); some, 
however, were fortunately in the interesting nestling dress. I 
secured four specimens in various stages of colour-growth. 

The young bird at first is somewhat browner above than the 
adult, the white of the under surface less pure; the flanks ash- 
grey; the head and gular streak pale sandy buff excepting the 
region of the so-called “crest” (P. cwcullata is the only species 
with a true crest) which is purplish ashy; the beak ochre-yellow, 
with the tip and the culmen and tomium of the mandible blackish; 
tarsi flesh-pink, slaty-grey in front. 

The complete change from this plumage to the fully adult 
colouring appears to take from nine to ten weeks; and during 
the whole time not a single feather is moulted out, but each 
feather gradually changes from day to day. I am certain of this, 
because I put the youngest bird with one more advanced in a 
large box-cage, so that no feathers could be dropped without the 
certainty of their being seen lying on the sand; moreover, the 


1903. | OF THE YELLOW-BILLED CARDINAL, 351 


change is so extremely slow that, not one but seventy or eighty 
moults would have been required to show all the transitions. 

The first thing which one notices is that the buff of the head 
and throat becomes dulled and deepened (perhaps the ‘“ ochreous- 
brown” described in the Museum Catalogue of Birds, vol. xu. 
p. 812); then little flecks of crimson appear dotted over the 
surface, which increase in extent from day to day until the full 
briliant crimson of the head is attained; but shortly after the 
appearance of these crimson flecks on the chin a small blackish 
spot appears at the extremity of the gular streak, which works 
regularly backwards until it has deepened the distal third of that 
streak (each feather, however, being left reddish at its base); then 
it throws out a flanking line which extends rapidly up each side of 
the beard-like streak, whilst from behind the chin a third central 
line begins to travel in the opposite direction. With the encroach- 
ment of the distal patch and the extension laterally of these three 
lines, the whole of the gular streak becomes black with reddish 
bases to the feathers; simultaneously the flanks become paler and 
the edges of the feathers white, offerimg an indistinctly streaky 
appearance; the back also becomes blacker, but the blackish 
streaking of the mandible and dark tip to the maxilla are 
generally more or less well-defined, though they appear to be 
sometimes lost at a very early stage. 

At this period the bird has therefore assumed the characters 
of Paroaria cervicalis; but, unfortunately for the validity of that 
species, they continue still to develop; the beak becomes wholly 
bright ochreous, the tarsi become wholly flesh-pink, the reddish 
bases disappear from the feathers of the throat, the grey almost 
entirely disappears from the flanks, the upper surface becomes 
quite black; and our P. cervicalis has become adult P. capitata. I 
am afraid the fact is quite indisputable. 

The habitat of the species should therefore stand as— Argentine 
Republic and Paraguay northward to Bolivia and Matto Grosso. 

It is possible, as Dr. Sharpe suggests (Catalogue of Birds, xii. 
p. 814, footnote), that P. gularis may be a further variation of the 
species ; but I should think it more likely that it was distinct. 
The variation in the amount of black in the region of the eye 
may be due to immaturity, as in the black of the throat of 
P. cervicalis, which does not cover the bases of the feathers. It is 
certain that not one of the Cardinals recently imported showed a 
trace of the black fusiform patch over the lores and behind the 
eye of P. gularis: nor did any of them show the slaty maxilla 
illustrated in the Museum Catalogue (vol. xxii. pl. xvi.); but perhaps 
this was an artist’s license, like the raised crests on the crowns of 
figures 2 to 6 of that plate,—birds which are perfectly incapable 
of erecting the feathers on their crowns like P. cucullata. 


352 DR. P. CHALMERS MITCHELL ON [ Dec. 1, 


4. On the Occasional Transformation of Meckel’s Diverti- 
culum in Birds into a Gland. By P. CHaumers 
Mircuety, M.A., D Se., Secretary to the Society. 


| Received December 1, 1903. ] 
(Text-figure 44.) 


Soon after I began to study the varying dispositions of the intes- 
tines in Birds I, like other observers, was struck by the occasional 
presence of a well-marked cecum about the middle of the course 
of the small intestine. This cecum is identical morphologically 
with what is known as Meckel’s diverticulum in the case of Man, 
and is the vestige of the embryonic yolk-sac. Ina communication 
(Trans. Linn. Soc. ser. 2, Zool. vol. vii. pt. 7, pp. 173-275, 1901) 
in which I reviewed the structure of the intestinal tract in many 
hundreds of birds belonging to practically all the families, I was 
able to add considerably to our knowledge of the occurrence of this 
structure and to point out its morphological significance as a 
point of orientation. In mammals, its presence in the adult 
appears to be rare and to be only an individual variation. In 
birds, the curious state of the case is that its presence or absence 
appears not to be an individual variation but to mark coherent 
assemblages. In most cases, when present, it is a true vestige of 
the embryonic condition, an apparently functionless rudiment 
sometimes still containing small granules of yolk. In such cases 
it appears to get smaller with age, but on this point I have not a 
large collection of observations to guide me. In other cases, a more 
peculiar condition occurs—the yolk-sac vestige has, so to speak, 
burst out again into a second life. It has become a glandular 
organ of a highly elaborate nature. I have already stated this fact 
(loc. cit. p. 264) and mentioned that Lénnberg and Jagerskiold 
have drawn attention to it; but as the microscopic structure of this 
new organ has not been described, I propose here to give a short 
account of it. 

The glandular condition of Meckel’s diverticulum is particularly 
well-marked in the Woodcock (Scolopaa rusticula). In that bird 
it is more than half an inch long, thick and very firm, with a 
narrow lumen slightly distended towards the apex and communi- 
cating with the cavity of the gut. The figure (text-fig. 44) shows 
part of a longitudinal section thr ough the diverticulum. The 
epithelium lining the lumen, and continuous with that of the 
intestine, is thrown into a set of deep glandular folds forming a 
branching system that occupies nearly half the wall. In the inter- 
spaces between these tubular glands, numerous patches of lymphatic 
tissue occur. Towards the apex of the gland this secreting layer 
is much thinner and more regular, and there is less lymphatic 
tissue. The rest of the ‘dhticlroese of the wall of the gland is 
composed of longitudinal and circular muscle-fibres, rather irregu- 
larly arranged, and of loose connective-tissue stroma in which 


1903. ] MECKEL’S DIVERTICULUM IN BIRDS. 353 
are embedded very numerous and large masses of lymphatic tissue, 
these latter also being more scanty towards the extremity of the 
gland. Ali the specimens that I have been able to examine were 
obtained from birds that had been dead at least for some hours, 
and the lining epithelium was partly decomposed and mixed 


with the contents of the lumen. 


Text-fig. 44, 


2s 


ei 

= 

~ 

a 

a 

ss = 
eS cd 
o = 
i > 
om 

7 

ae 

ied 


pet? af isa cy, 


* 
ay 
ise! 
= 
Sih 
ke 
a ™ 
yh 
ier 
~ 
= 
. 
ol 
- 
> 
i 
x 


t Depa r 


Sey 
+ 
ees 
§ As Ma See 
Pee Z 
a 4 : 


aan 
a 


marcel 


j 


Meckel’s diverticulum in the Woodcock. Part of a longitudinal section of the wall: 
the lower half of the section shows the intestinal epithelium thrown into 
Peng folds, with patches of lymphatic tissue; the upper part shows 
ongitudinal and circular imuscle-fibres and connective-tissue stroma, with 
islands of lymphatic tissue. 


There can be no doubt from the structure of this organ that it 
is a definite gland, but we are so profoundly ignorant of the 
physiological processes in all animals, except Man and a few 
creatures commonly used in experimental laboratories, that there 
are practically no data for making even a suggestion as to its 
function. It is obviously similar in structure to the paired ceca 
in certain birds, as, for instance, most of the Passeres, where these 
are glandular nipples. It is interesting to note that the glandular 
condition of Meckel’s diverticulum occurs in birds that are other- 
wise specialised and does not appear to occur in birds which in 
other features of their anatomy are archicentric or primitive. 
This supports my inference that the glandular condition is a new 
feature, probably of recent origin in the history of birds, 

Proc, Zoou. Soc.—1898, Vor. Il. No, XXIII. 23 


354 SIR C. ELIOT ON NUDIBRANCHS [ Dec. 1, 


The mode of origin of new organs is a morphological problem to 
which little attention has been given, although it is of great 
biological interest. The most obvious analogies with the present 
case are such changes of function and structure as the trans- 
formation of the lower part of the pharynx into the thyroid gland 
or the metamorphosis of part of a gill-slit into the middle 
chamber of the ear; but I do not remember any case exactly 
similar to the present where the protoblastic structure, having 
fulfilled its function, degenerates, but is replaced by a deutero- 
blastic structure with a new function. It is interestmg to notice 
that the raw materials for this deuteroblastic growth occur im 
many birds, inasmuch as the relic of the yolk-sac, or protoblast, 
lingers throughout life, but it is only in a few cases that the relic 
sprouts again and becomes the new organ. 


5. On some Nudibranchs from East Africa and Zanzibar. 
Part) IM. By Sir C) Bron KCC MG, Me Come 
missioner for the Hast Africa Protectorate, F.Z.S.— 
Dorididz Cryptobranehiate, I. 

[ Received October 16, 1903.] 
(Plates XXXII.-XXXIV.+) 


This paper contains the results of an examination of a number 
of Cryptobranchiate Dorids from Zanzibar and the East Coast 
of Africa referable to Bergh’s subfamilies Archidoridide, Disco- 
doridide, Diaululide, Kentrodoridide, and Platydoridide. They 
inelude the following species :— 


Archidoris Bergh. 


iN africana, sp. N. 

2. 35 minor, Sp. N. 
Staurodoris Bergh. 

oe os depressa, Sp. Ni. 

4, =e calva, sp. n. 
Discodoris Bergh. 

oy . boholiensis Bergh. 

6. x cerulescens Bergh, variegata, subsp. n. 
Peltodoris Bergh. 

ie és angulata, sp. 0. 

8. ie aurea, Sp. n. 


Thordisa Bergh. 
9: » villosa (A. & H.). 
10. 2 stellata, sp. n. 
Iie “3 crosslandi, sp. n. 


* Vor Part II. see P. Z. S. 1908, vol. i. p. 250, 
{ Vor explanatior of the Plates, seep. 385. 


LI. Pl. XXXIL. 


ais 


& 


jpler es 


oN 


aS 


rer 


é 


LATA 


Huth,Lith* London. 


C.Crossland del. 


A. Willey del fig. 5. 


3, Te GROSS ANID 
5.HALGERDA WILLEYI. 


1,2. THORDISA VILLOSA. 
4 TRIPPA MONSONIL. 


P2Z.5.1903, voll Pl XXX. 


Brown pigment. 
a Spiartes. A [A 


RES 
oats ES) ‘ ) 
a) 


C.Crossland del. Huth, Lith? London. 


[Sor Li ORDISA VILEOSA, 43 TCROSSL ANDI. 


ies 
ini 


EGS) UNOS Noli ACORN, 


CiGoasisllemnd Is : een Inte ender 


1,2. HALGERDA WASSINENSIS. 3.SCLERODORIS CORIACEA. 
4 PARASITIC COPAPOD. 5.6.ASTERONOTUS HEMPRICHI. 
7. KENTRODORIS RUBESCENS. 


1903. FROM EAST AFRICA AND ZANZIBAR. 355 


Trippa Bergh. 


Ive »,  areolata (A. & H.). 

Ne » monsont, sp. Nn. 
Fracassa Bergh. 

14. i tuberculosa. 


-Halgerda Bergh. 
15. ss willeyi, sp. n. (From the Loyalty Islands.) 


16. i wasinensis, Sp. D. 
Kentrodoris Bergh. 

ef: 53 rubescens Bergh. 
Platydoris Bergh. 

18. a eurychlamys Bergh. 

1®. 5, scabra (Cuvier). 

20. ce formosa ? (A. & H.). 

21. - elliott? (A. & H.). 

22). $3 pulchra, sp. 0. 

Die ay incerta, Sp. Ni. 

24, i papillata, sp. 0. 
Sclerodoris Eliot, gen. n. 

25. os osseosa (Kelaart). 

26. S tuberculata, sp. 1. 

PAI iy minor, Sp. id. 

28. es rubra, sp. 0. 

29. 59 coriacea, sp. . 
Asteronotus Ehrenberg. 

30. x hemprichi Khrenberg. 


The Oryptobranchiate Dorids are distinguished by haying 
branchix which are completely retractile into a permanent pocket. 
The rhinophores also are retractile into permanent pockets and 
are perfoliate. A radula is always present. In number of 
species they form one of the richest groups of the Nudibranchiata, 
but they show less variety of form than the Doridide Phanero- 
branchiate, and it is not easy to find good generic characteristics. 
The most distinctly marked group is that consisting of the large 
genus Chromodoris (with more than 100 species) and some allied 
genera such as Casella and Ceratosoma. These possess a charae- 
teristic shape, lip-plates, a radula with differentiated teeth, and 
simply pinnate branchie. Well characterised genera are also 
found in the Miamirade (Miamira, Orodoris, Spheerodoris) and 
the Cadlinide (Cadlina, Tyrinna). There remain the five sub- 
families mentioned above, which agree in being usually flat and 
oblong in form and in having a radula without a central tooth, 
consisting of numerous laterals which are generally hamate and 
uniform, though often smaller near the rhachis and degraded or 
denticulate at the outer end of each row. 

Anyone who describes new forms of these Dorids, however 


much he may disclaim any such ambitious task as a revision of 
ORE 2 
av 


356 “SIR C, ELIOT ON NUDIBRANCHS* [ Dec. I, 


the group, is bound to consider the value of the existing generic 
distinctions. The five subfamilies are divided into about 30 
genera all created by the great master of the Nudibranchiata, 
Prof. Bergh, and several of them consisting of a single species. 
He himself observes (on Vhordisa in Semper’s ‘ Reisen’*, xv- 
p- 666) that the definitions of many of these genera seem to be 
too precise, and as new forms are discovered the old divisions are 
found to melt away. The discoverer of » new form often finds 
that it does not accurately fit in to any of the existing genera, 
and must ask himself whether he should create a new genus or 
enlarge the definition. I have httle doubt that in most cases 
the latter is the preferable course. If animals are not divided by 
natural differences, there is no object in emphasising the im- 
portance of minute peculiarities. If Chromodoris is allowed to 
consist of 100 species showing a considerable range of variation, 
including the presence or absence of median teeth, there seems to 
be a want of proportion in splitting up the other Cryptobranchiata 
into so many genera. 

The genus Stawrodoris offers « good example of the difficulty 
of classifymg new forms. Taken by itself, the typical species 
St. verrucosa is remarkably well characterised in both the scien- 
tific and popular sense. Anyone could recognise it at first sight. 
The back is covered with club-like tubercles, the rhinophorial and 
branchial pockets are protected by similar tubercles which act 
as valves, and the branchie are simply pinnate. On the other 
hand, Sé. psewdoverrucosa (von Ihering) has no tubercles on the 
branchial pocket and has bipinnate branchie. Bergh also refers to 
this genus the Doris pustulosa of Abraham, which has bipinnate 
branchie and small, but apparently not valve-like, tubercles. I 
have spectmens from the Indian Ocean which have the dorsal 
surface tuberculate and the following additional characteristics :— 
A. has the rhinophores arising among tubercles and simply pinnate 
branchie, but no tubercles on the branchial pocket (Stawrodoris 
pecten). B. has five pinnate gills, the anterior margin of the foot 
entire, and small tubercles set on the edges of the gill-pocket and 
partly closing it (Stawrodoris calva). C. has tubercles round the 
rhinophores, none on the branchial pocket, and thin bipinnate 
branchie (Archidoris africana). D. has tubercles on the rhino- 
phore pockets but not around the branchial opening, and tri- 
pinnate gills (Archidoris minor). Of these, I think we must admit 
A.. and B. to be Staurodoris, if we accept St. pseudoverrucosa. 
But Archidoris kerguelensis, A. australis, A. rubescens, A. incerta, 
and A. nyctea are all described by Bergh as having tubercles on 
the edge of the branchial and rhinophorial pockets, and must 
come very near the less typical members of Stawrodoris. It is 
hard therefore to say whether C. should be classed as Staurodoris 
or Archidoris. There seems to be a complete series of links 
between the two genera, and, this being so, we must either unite 


The letters S. R. in this paper refer to Prof. Bergh’s “ Malacologische Unter- 
suchungen,” published in ‘Reisen im Archipel der Philippinen,’ von Dr. C. Semper. 


1903.] FROM EAST AFRICA AND ZANZIBAR. 357 


the species in question, or draw an arbitrary dividing line. In 
the latter case, I think we must say that the typical Stawrodoris 
has simply pinnate branchie and valve-like tubercles closing the 
rhinophorial and branchial pockets. One or other of these 
features may be absent or obscure in a species which otherwise 
possesses the generic characteristics ; but when both are absent, 
as in G., the animal, I think, must be referred to Archadoris. 
Again, Staurodoris pseudoverrucosa has the dorsal tubercles some- 
times connected by ridges, and the same phenomenon is found 
in Garstang’s Doris maculata (which appears to be a Stawrodoris), 
and, sporadically, in the true Sé. verrucosa. But, as will be seen 
from a species described below, Halgerda wasinensis, this character 
brings Stauroderis very near to Halgerda. 

It will be well to examine the value of the chief points by 
which the subfamilies and genera under consideration can be 
differentiated. They are as follows :— 

(a) The dorsal surface and general texture. The back is rarely 
quite smooth, as it is in Chromodoris; Halgerda and Asteronotus 
have the skin smooth but raised into ridges or lumps. As a rule, 
the surface is covered with projections which may be either 
minute granulations (Platydoris, Discodoris, &c.), papille (Thor- 
disa, &c.), tubercles (Archidoris, &c.), clavate tubercles or warts 
(Staurodoris), compound tubercles (Zrippa, Fracassa). There is 
sometimes a ridge down the centre of the back. Some genera, 
notably Platydoris, ave exceedingly hard; others, such as 7rippa, 
are so soft as to be almost gelatinous. On the whole, these 
external characters of the skin and texture form a fairly good 
indication of relationship. Platydoris, in which I should be 
disposed to include Hoplodoris, forms a distinct natural group, 
and the warty or tuberculate forms (Archidoris, Stauredoris) also 
hang together *. 

(6) Rhinophores and branchie. Neither the rhinophores 
themselves, nor the pockets into which they are retractile, seem 
to offer good generic characters, though they may often serve to 
distinguish species. But even within a species there may be 
variety: Archidoris tuberculata has the rims of the rhinophore 
pocket sometimes smooth and sometimes tuberculate. 

The branchiz also are disappointing as a means of classification. 
For instance, it does not seem possible to unite Stawrodoris with 
the other genera having simply pinnate branchie (Chromodoris, 
Casella, Ceratosoma, Spherodoris, Halla, Thorunna, Rostanga), 
and these simple branchie often show a tendency to divide at 
the tip and become, strictly speaking, bipinnate. Similarly, we 
cannot bring together bipinnate forms and oppose them to the 
tripinnate. Perhaps the distinction between ample and scanty 
branchiz will prove to be of generic importance. The branchial 
pocket is of various shapes: round, crenulate, or stellate. However, 


* T find it hard to agree with Prof. Bergh’s criticism of his own family Archi- 
doridid that it will prove “ganz unhaltbar und kiinstlich.’ With the exception 
ot Peltodoris the other forms seem to hang well together. 


b) 


308 _ SIR C. ELIOT ON NUDIBRANCHS | Dec. 1, 


id 


it does not seem possible to make any classification according to 
this shape, and to unite, for instance, all the stellate forms. 

In most of the Cryptobranchiata the tentacles appear to be 
digitate, but the Archidoridide have a tendency (not without 
exceptions) towards a flattened and furrowed form. Spherodoris 
has no tentacles, and the same is said of Halla and Hchinodoris. 
No part of preserved Nudibranchs is more liable to distortion 
than the tentacles, which may be variously contracted, flattened, 
or crumpled by the action of the preserving fluid or the pressure 
of the adjacent parts. 

(c) The foot appears to offer good characters; as a rule, but 
not invariably, the forms which have a broad foot and narrow or 
moderate mantle- margin belong in other respects to groups 
different from those which have a narrow foot overhung on every 
side by a wide mantle-margin. The Archidoridide and Disco- 
doridide have both, as a rule, broad feet, but the body of the 
former is plump, and of the latter flattened. The Platydoridide 
have flat bodies and narrow feet. 

In most forms the anterior margin of the foot is deeply grooved, 
and the upper lamina notched so as to form two flaps, which in 
the Kentrodoridide are very ample. Many (though not all) of 
the Archidorididee have the anterior margin simply grooved and 
otherwise entire. In a few genera (7rippa, Halla*, Spherodoris) 
the divided upper lamina is attached to the sides of the head, but 
it does not seem possible to bring together the forms which 
present this peculiarity. 

(d@) In the internal anatomy, the mouth-parts are perhaps the 
most important for classification. It is clear that considerable 
structural differences in other organs ave generally accompanied 
by a difference in the radula. For instance, Acanthodoris and 
Lamellidoris, which bear a strong superficial resemblance to the 
Cryptobranchiata but have no permanent gill-pocket, have also a 
totally different radula, and most of the larger divisions of the 
Nudibranchiata have a characteristic arrangement of teeth. But 
it is hazardous to conclude from this that smadl differences of the 
radula have a generic value. One common variation from the 
ordinary hamate type of radula is the serrulation of the outer 
teeth. This may be present or absent in the same genus (e. g. 
Staurodoris verrucosa and Sé. bicolor), and in some species (Platy- 
doris argo and Halgerda formosa t) is only found in some of the 
vows of teeth. Asa general rule the innermost and outermost 
teeth are smaller: the latter often rudimentary or degraded. 
But it appears that these characters are only of specifie, not of 
generic importance. 


* [I venture to point out that the generic name Hailla is preoccupied by a Poly- 
cheete (Lumbriconereid) worm named by A. Costa in 1844 (cf. Ann. Acad. d. Aspiranti 
Naturalisti Napoli, ii. p. 63 (1844).—C. CrossLanp. 

+ Ina specimen of Halgerda formosa trom the Berlin Museum, given me by the 
courtesy of Prof. Martens, I found at the end of some but not all of the rows small 
rudimentary teeth, some but not all of which bore a few irregular serrulations. This 
1s not quite the same arrangement as observed by Prof. Bergh, 


1903.] FROM EAST AFRICA AND ZANZIBAR. 309 


Another point of importance is the presence or absence of a 
labial armature, that is to say, of a ring or plates on the labial 
cuticle composed of a compact mass of minute hooks or rods, 
There is some reason for dividing the group into those which 
possess and those which do not possess this armature, but still I 
think that an absolute dichotomy of this kind presents difficulties. 
Firstly, a rudimentary labial armature is of some inconvenience 
in classification. Bergh has described such an instance in Platy- 
doris variegata; and the animal described below as Thordisa 
crossiandi has two small areas on the labial cuticle which cannot 
be called plates, but consist of a loose collection of minute rods. 
In Platydoris pulchra the labial cuticle is strengthened with 
similar rods, distributed through its extent, but not collected into 
rings or plates. Secondly, though Discodoris B. is a well-defined 
genus, the same can hardly be said of the family Discodoridide, the 
genera of which mostly agree only in having a labial armature. 
Thus Fracassa and Carminodoris appear to be akin to Trippa, 
Hoplodoris to Platydoris, and Halla to Chromodoris. It would seem 
that the more primitive forms of the Cryptobranchiata are those 
which have a labial armature and some differentiation of the 
teeth, such as occurs in Chromodoris and Cadlina, and that the 
forms with no labial armature and uniform hamate teeth are 
more recent developments. If this be so, it 1s easy to under- 
stand that in many groups a few forms have survived in which 
the labial armature has persisted. Thus Fracassa 1s practically a 
Trippa which has preserved this character, and it appears to me 
that its analogies to Z’rippa are greater than those to Discodoris. 

(c) Considerable use has been made by Prof. Bergh of the 
reproductive organs as a character for classification*, That great 
weight must be attached to important variations in these organs 
—such as the presence of one or two spermathecee—is obvious ; 
put while fully admitting the necessity of examining the internal 
anatomy and the futility of describing only the external characters 
of Nudibranchs, it must also be admitted that it is not con- 
venient to found genera of fairly large animals upon minute 
internal characteristics which can only be discovered by an expert 
microscopist, and by him only in a well-preserved and mature 
specimen. Such a criterion seems desirable only if it is of great 
anatomical importance. But what are the variations presented 
by these organs in the Cryptobranchiata? (1.) Accessory organs 
are sometimes present. These are generally accompanied by 
other characters of systematic importance as in Kentrodoris and 
Asteronotus. (ii.) The male branch of the hermaphrodite system 
sometimes broadens and sometimes does not into a dilatation 
called the prostate. It does not appear that this difference is 
generally accompanied by other characters of importance: ¢. 9. & 


* But see System. d. Nud. Gast. p. 1088. “ Bei dem jetzigen Stande unserer 
Kenntniss .... ist es sehr gewagt eine systematische Gruppirung derselben zu 


versuchen. Es ist es um so mehr als der generische Werth der bei der hier ver- 
suchten Gruppirung dem genital System, hesonders den verschiedenen Conformae 


a 22 


¢ionen seiner Austiihrungsginge, beigelegt ist, als soleher kam sicher gestellt ist. 


369 SIR C. ELIOT ON NUDIBRANCHS | Dec. I, 


prostate is present in both Discodoris and Platydoris, which are 
otherwise so distinct, and is absent in Archidoris but present im 
Anisodoris, which are otherwise identical. (ii1.) In many genera 
is present an armature of the reproductive organs, that is, one 
or many spines, hooks, or plates generally only on the male 
branch. The value of this feature, as estimated by the con- 
comitance of other important characters, varies. In the families 
of Doridopside and Phyllidiadz and in the well-marked genus 
Platydoris of the Cryptobranchiata an armature is, so far as we 
know, characteristic. On the other hand, we find two genera 
like Diaulula and Gargamella apparently identical but for its 
presence or absence. In the Aeolidiade also a hook or spine is 
present or absent in closely allied genera, It appears to me that 
such an armature is not suflicient to constitute generic rank 
without other characters, 

(f) The other internal organs do not offer many features 
which serve our purpose of classification, It does not appear 
that we can unite the forms in which the stomach is enclosed 
in the liver, or in which there is only a single instead of a 
double blood-gland, or in which the nervous system is very 
concentrated. 


As a result of these considerations, I think that the most pro- 
fitable way of classifying new Dorids of the group treated of here, 
is to refer them when possible to the following genera taken in a 
wide sense :—Arechidoris, Discodoris, Thordisa, Trippa, Halgerda, 
Kentrodoris, Platydoris, Asteronotus, and Sclerodoris (gen. n.). 
No doubt, remarkable forms have been and will be found which 
require special genera for their accommodation, but the majority 
of species seem to me to fall under one or other of the divisions 
mentioned. 

Archidoris is distinguished by a fairly plump shape, broad foot, 
and a warty or tuberculate back, No member of this group is 
known to possess a labial armature, and a genital armature is 
rare. The radula generally consists of uniform, simply hamate 
teeth: more rarely some or all are denticulate. I include in 
this genus, in its wide sense at least, Anisodoris, Homoiodoris, 
and Artachea. As mentioned above, though it is easy to define 
the generic characters of a typical Stawrodoris, the genus seems 
to pass into Archidoris by a complete series of connecting links. 

Discodoris.—\ should be inclined to extend this genus go as to 
mean flat, oval animals, not hard, with both foot and mantle- 
brim fairly broad. Back granulate. A genital armature is 
usually absent; a labial armature is either present (Section 
Discodoris) or absent (Section Peltodoris). 

Lhordisa.—Flat, soft animals, much like Discodoris, but with 
the back neither granulate nor warty, but covered with soft 
pointed papille. No member of this group is known to possess 
a well-developed labial armature, though a rudimentary one is 
found in Zh, crosslandi. 


1903. ] FROM EAST AFRICA AND ZANZIBAR, 361 


Trippa.—Soft, and sometimes spongy or gelatinous, The back 
is covered with tubercles which themselves bear smaller promi- 
nences or filaments. A labial armature is occasionally present 
(Section Fracassa). Some species have special glands set round 
the buccal mass, and the sides of the head connected with the 
foot, but it does not appear to me that the absence of these 
characters ought to exclude a form from the group. 

Halgerda.—The texture is entirely smooth and somewhat stiff, 
though ridges may be present, In the known species the branchie 
are scanty, No member of this group is known to possess a labial 
armature, 

Kentrodoris.—Broad, soft, and flat, with the dorsal surface 
minutely granulated. The broad foot is deeply grooved in front, 
and the upper lip, which is notched in the middle, is developed 
into wing-like expansions on each side. The reproductive system 
is sometimes armed, and accessory organs are present. In some 
species, at any rate, the branchie are unusually large. 

Platydoris,— Very ilat forms, of a peculiar hard consistency. 
The back is minutely granulated and rough to the touch. The 
foot is narrow. The branchial pocket is stellate in the known 
forms, A labial armature is rare (Pl. variegata). There is a 
characteristic genital armature of scales bearing hooks, 

Asteronotus,—Of a characteristic leathery consistency. The 
back is quite smooth in texture, but bears lumps and ridges. No 
labial or genital armature. 

Sclerodoris,—This new genus is proposed for certain forms 
which appear to have never come into the hands of Prof. Bergh, 
though I think Alder & Hancock’s Doris osseosa, carinata, apicu- 
lata, and tristis (‘‘ Notes on a Collection of Nudibr. Moll. made in 
India,” Trans. Z. 8. vol. ii. 1864) should be referred to this 
genus. It is characterised by having the same hard texture as 
Platydoris, but the back, instead of being smooth, is marked with 
various ridges and depressions. In the known species there is no 
labial or genital armature. I should wish to bring my Sclerodoris 
under Prof. Bergh’s Dictyodoris, but the generic characters as 
formulated by him do not include the hard texture and raised 
reticulate pattern. 

Of the above-named genera, Asteronotus and Kentrodoris, 
though well characterised and not rare, have not hitherto proved 
numerous In species, 


1, ARCHIDORIS§ AFRICANA, Sp. 1. 


One specimen marked ‘‘ Chuaka, shore.’ 
animal, 

Alcoholic specimen 5 centimetres long, 1:6 high, with a fairly 
uniform breadth of 2°7, plump and not flat. The colour is a 
dirty greyish yellow, with traces of violet. The back is covered 
with tubercles: those in the middle are largest and measure 
4 millimetres across; they decrease in size outwards, and are 
quite small at the mantle-edge. The top of each is lighter, and 


’ No notes as to living 


362 SIR C. ELIOT ON NUDIBRANCHS (Decl, 


was probably of a different colour in life. It is noticeable that 
the tubercles in the middle are all large and not mixed with small 
ones. The mantle-brim is moderately ample, thick, and stiff, and 
bears numerous irregular tubercles on the lower side which are 
probably glandular in character. The rhinophore openings are 
indistinctly bilabiate, not much raised, and bear small tubercles 
on the sides and edges. The rhinophores are large, ample, and 
deeply perfoliate. The branchial pocket is also not much raised, 
indistinctly bilabiate, and at the same time with five irregular 
and not very distinct crenulations. There are tubercles on the 
sides but none on the edges. The branchie are eight, tripinnate, 
but not ample: the two hindermost are smaller than the others. 
The foot is large and broad, with a shallow groove anteriorly and 
a split upper lip. The tentacles, which are set at right angles to 
the head, are unusually large and long (5 millimetres). The 
labial cuticle is black, and corrugated but unarmed. The radula 
consists of 33 rows containing about 60 simple white hamate 
teeth; the innermost are smaller, but the outermost are much 
the same size as the rest. No prostate or genital armature 
was discernible. There was a large purple double blood-gland, 
deeply cleft in both parts so that it seemed to have four divisions. 


2. ARCHIDORIS MINOR, Sp. n. 


One specimen from Wasin. There are no notes on the living 
animal. 

The alcoholic specimen is 2°9 centimetres long, 1:8 broad, and 
9 millimetres high. The colour is rather bright yellow, with 
traces of a darker tint near the mantle-edge. The back is 
covered with flat warts, largest towards the centre, and decreasing 
towards the mantle-edge, but smaller ones are mingled with the 
larger; they show indications of a lighter colour at the top. The 
underside is of a uniform yellow. Round the rhinophore pockets 
are two or three tubercles, which look as if they had been high in 
life. The rhinophores are high, straight, and narrow, strongly 
recalling Bergh’s figure of those of Staurodoris januari (S. R. 
Supp. 1. plate C, fig. 14). The branchial pocket is slightly raised, 
bilabiate, and indistinctly crenulate. Though there are tubercles 
near the edge, these in no sense close over it or act as valves. 
The branchie are eight, tripinnate, but high, thin, and scanty. 
The central papilla, also, is very high and thin. The foot is fairly 
broad, and grooved in front, with the upper lamina notched. 
The tentacles are small and conical. There is no labial armature. 
The radula consists of 30 rows, each containing about 50 long 
hamate teeth on either side of the rhachis; the innermost are 
crowded and smaller, the outermost not much smaller. At the 
side of the base is a groove, terminating in a slight projection at 
the bottom of the hook. 

This specimen has many points of resemblance with Archidoris 
africana, but I am inclined to think that it is specifically distinct, 
for the following reasons :—(1) The prevailing colour is yellow, 


1903. | FROM EAST AFRICA AND ZANZIBAR, 363 


whereas in 4. africana it is violet, both externally and in the 
intestines; (2) the tentacles are small; (3) there are no tubercles 
on the underside of the mantle-edge; (4) the rhinophores and 
their pockets are somewhat different from those of A. africana ; 
(5) so are the teeth. 

It is possible that this is the Doris rusticana of Alder & 
Hancock (‘Notes on a Collection of Nudibranchiate Mollusca 
made in India,” Tr. Z. 8. iii. p. 120), but in view of their statement 
‘“‘No oral tentacles (?); the head with lateral angles; branchial 
plumes five,” identification is not possible. 


3. STAURODORIS DEPRESSA, Sp. 1. 


One specimen from Wasin. No notes as to living animal. 

The alcoholic specimen is 6°3 centimetres long and 4°9 broad. 
The general shape is broad and flat. The thick and fleshy 
mantle-brim is 2 centimetres wide, and the foot consequently 
unusually small compared with the dorsal surface, being only 
2°77 mm. long and about 8 mm. broad. The colour is a uniform 
greyish white, with a slight tinge of violet anteriorly and down 
the middle of the back. The whole upper surface is covered with 
warts, which*are small at the mantle-edge but increase in size 
towards the centre. The top of the larger ones, which measure 
5 millimetres across, is flat and hard, consisting of a mass of 
densely-crowded spicules, and is of a somewhat different shade 
from the rest and in life possibly distinctly coloured. On the 
underside of the mantle-edge are numerous small tubercles of 
glandular appearance. The openings of the rhinophores and 
branchie are tuberculate. The latter orifice is indistinctly stellate 
and also indistinctly bilabiate, but it is not clear what its original 
shape may have been. Both the branchial and rhinophorial 
orifices are closed in the alcoholic specimen. The branchiz are 
six in number, but the hindermost pair are deeply bifid so that 
there appear to be eight. They are mostly bipinnate and rather 
scanty. The foot is grooved and notched in front. The tentacles 
are large, distinct, and somewhat flattened, with rather uncertain 
traces of a groove. There is no labial armature. The radula is 
broad and white, the formula being about 70.0.70X32. The 
teeth are simply hamate and allof much the same size. On some 
of the inner ones I was able to see eight or ten very minute 
denticles on the inside of the hook. This extremely fine serrula- 
tion is difficult to detect, but I expect that it is present on all the 
teeth except the outermost. The stomach is not free, but 1s 
enclosed in the liver. The female reproductive organs are armed 
with small transparent brick-like scales. 

This form offers analogies to both Homoiodoris and Artachwa 
Bergh, particularly the latter, and the thick leathery mantle and 
large warts also remind one of Asteronotus. On the whole I class 
it, though very doubtfully, as Stawrodoris, mainly because the 
openings of the rhinophores and branchie are closed by the 
surrounding tubercles. 


364 SIR C. ELIOT ON NUDIBRANCHS [Deenl; 


4, STAURODORIS CALVA, Sp. 0. 


One specimen from Kokotoni Harbour, Zanzibar; dredged in 
about 5 fathoms. 

The living animal was of a dirty grey colour, but with very 
little pigment at all; the gills and rhinophores sandy, the under 
surface and the smooth band near the rhinophores pinkish. The 
dorsal surface was covered with tubercles, large and small, of 
various sizes, but decreasing towards the mantle-edge, where they 
were minute; small tubercles were set on the edge of the slightly 
raised gill-pocket, which was partly closed by them. The anterior 
portion was prolonged into a nose-like projection, Across it 
extended a smooth pinkish strip, which bore no tubercles. im- 
mediately behind this strip were set the rhinophores. It is 
possible that this singular arrangement may have been an un- 
natural distortion. Some species of Platydoris undoubtedly 
remain fixed in crevices until their shape is altered. Still, the 
present specimen showed no trace of tubercles having been effaced 
on the bare patch, and there is no reason to suspect the character 
except that it is, as I believe, unique among the Doridide. 

The alcoholic specimen, 2 centimetres long and 1-3 broad, is 
much like the living animal. The broad foot has’ the anterior 
margin entire and not grooved. The dorsal tubercles are sur- 
rounded by numerous very distinct spicules arranged in a stellate 
form. The tentacles are small and furrowed. There is no 
labial armature, and the radula consists of 48 rows of simple 
hamate teeth containing about 70 teeth on each side of the 
rhachis. The stomach is large and free. The branchiz are five 
in number and bipimnate. The branchial pocket 1s almost closed 
by the valve-like tubercles on the edge of it. No reproductive 
armature was discovered. 

This form seems referable to Stawrodoris. Though the branchiz 
of this genus are typically only pinnate, they appear to be 


aaa in both S. psewdoverrucosa (v. Ther.) and S. pustulosa 
(Abr.). 


5, DiscoDoriIs BOHOLIENSIS B. 

[S. R. xii. p. 519, xvii. p. 897.] 

Two specimens from Zanzibar. 

The body of the living animal was flat, with a very ample 
flexible mantle. A high, narrow dorsal keel extended from the 
branchial pocket to between the rhinophores. The ground-colour 
was yellowish drab, with a black edging round the wavy mantle- 
edge, which in places extended inwards. The whole back was 
covered with small papille, some brown, some white. The dorsal 
keel was blotched with brown and black. The rhinophores and 
branchie were black. The underside was dirty white, with black 
and brown blotches at the side of the foot. The living animal 
was 23 inches long and 2 broad. The large specimen displayed 
the phenomenon of self-mutilation, The rhinophore openings 


1903. | FROM EAST AFRICA AND ZANZIBAR. 365 


were raised and crenulate, the rhinophores bent backwards. The 
branchial opening is a transverse slit, the two lips almost meet in 
the middle but separate at the sides. The branchie are six, 
tripinnate, the posterior pair bifid. The labial cuticle bears two 
small yellowish plates composed of minute rods. The radula 
formula is about 23 x 40.0.40. The teeth are simply hamate ; the | 
two or three outermost are rudimentary, the innermost are 
smaller and have rather shorter hooks as described by Bergh. 


6. DIscoDORIS CH RULESCENS VARIEGATA, subsp. n. 

[ Bergh, in Semper’s Reisen, xvi. Hiilft 1. p. 805. | 

One specimen from Jembiani, Zanzibar. 

The living animal was about an inch long and of a light slaty 
blue, with many small blackish blotches. The coloration of the 
underside was similar but rather lighter. 

In alcohol the blue parts have become yellow, a remarkable 
change of tint which has also occurred in T’rippa monsona. The 
texture is leathery and stiff but not hard. The whole dorsal 
surface, including the rims of the rhinophore and _gill-pockets 
which are raised, is covered with minute tubercles. The branchial 
pocket is roundish, with a jagged edge and very deep. Within it 
are six tripinnate branchie. The foot is grooved and the upper 
lip deeply cleft. Immediately above this cleft is the mouth, with 
a white, tapering, digitate tentacle on each side. In the upper 
part of the oral tube are two roughly triangular collections of 
minute rods, less definite in outline and consistency than the type 
of armature generally described as labial plates, but sufficiently 
large to warrant us in describing the cuticle as armed. The 
radula consists of 30 rows, containing about 45 white, simply 
hamate teeth on each side of the rhachis. The innermost and 
outermost are somewhat smaller, but not degraded in shape. The 
reproductive system is not armed. The prostate is bent and 
fairly large, but I was not able to discover the peculiar structure 
of the hermaphrodite gland mentioned by Bergh. 

This animal appears to have all the chief characteristics of Bergh’s 
D. cerulescens, and comes from much the same part of the world. 
Mébius’s specimen (from Mauritius) was apparently of a uniform 
bluish white, whereas this one is mottled with darker blotches. 
Hence I describe it as a variety. 


7, PELLODORIS ANGULATA, Sp. 0. 


One specimen from Chuaka. 

The animal has a close superficial resemblance to Thordisa 
villosa, but has six violet-brown spots symmetrically arranged in 
rows of three on each side of the median dorsal line between the 
rhinophores and branchie, and some round chocolate spots on 
the under edge of the mantle near the foot. The violet spots 
seem to be under the surface and visible through the dorsal skin. 
On a closer examination the superficial resemblances disappear : 


366 SIR 0. ELIOT ON NUDIBRANCHS [diee, 1 


the peculiar tubercles of 7. villosa are absent; the general 
texture, though flexible, is not quite soft, but rather stiff; the 
back is minutely granulate but not harsh. The rhinophore 
pockets are raised and have jagged edges. The branchial pocket 
is raised with round smooth edges, turned very distinctly 

outwards. The branchie are six and tripinnate. The anterior 
_ pair are smaller than the others. The most characteristic feature 
of this specimen is, the foot, which has a wide thin margin all 
round, dilated anteriorly into tentacular expansions, similar to 
those found in the Aeolidide, and 3°5 millimetres long. The 
front part seems to be grooved in the middle and the upper 
lamina to be attached to the head on each side. Admitting that 
it is dangerous to speak positively of such characteristics on the 
strength of a possibly distorted alcoholic specimen, I think it 
is clear that the anterior portion of the foot must be expanded in 
a way unique among the hitherto described species of Doridide. 
The tentacles are thin and digitate. No labial plates were 
discernible. The buccal mass was large for the size of the animal. 
The formula for the radula appeared to be about 45.0.45 x 38. 
The teeth are the ordinary simple hamate type; the innermost 
are not smaller; the 3-5 outermost are degraded but are not 
serrulated. The stomach is large and free, laminated internally. 
There seemed to be traces of an inconspicuous armature of 
transparent scales on the glans, but I was not able to satisfactorily 
make out its arrangement. 

The dorsal spots in this specimen seem to resemble those 
described by Bergh in P. mauritiana, but this animal must be 
specifically distinct from that form. 


8. PELTODORIS AUREA, Sp. n. 


Three specimens captured near Wasin. 

The living animal is flat, with an ample mantle which extends 
far beyond the head and tail. The texture is not hard but also not 
flabby ; one specimen is much stiffer than the others. The general 
colour is arich light orange, due to the back being covered with little 
flat orange warts on a sandy ground. At regular intervals round 
the mantle-edge are spots of dull violet ; there is also a spot just 
in front of the gills and one behind the rhinophores. The under- 
side is yellowish with a few brown spots. The dimensions are 
2°2 centimetres length, and 1:4 centimetres breadth. Both the 
rhinophore and gill-pockets are somewhat projecting, but though 
they rise among tubercles they cannot be described as tuberculate. 
The gill-pocket 1s large, somewhat contracted in the middle and 
expanded at the sides, so that the 8 tripinnate gills fall in two 
bundles, right and left. The foot is 1-6 centimetres long and 
only 3 millimetres broad; grooved and notched in front. The 
tentacles are small and button-like. There is no labial armature. 
The radula is small and fragile: it consists of twenty rows, each 
containing about 25 white, simply hamate teeth; the innermost 
and outermost are somewhat, but not conspicuously smaller. The 


1903. ] FROM EAST AFRICA AND ZANZIBAR. 367 


stomach is quite free from the hepatic mass. No armature was 
discernible in the reproductive organs. 

T have some hesitation in classifying this specimen as Peltodoris, 
as the back is not minutely granulated but covered with small 
warts. The shape, however, is not that of Archidoris, and both 
the stiffness and small radula are in favour of the position here 
assigned to the form. 


9. THorpisA vittosa (A. & H.). (Plate XXXII. figs. 1 & 2; 
Plate XX XIII. figs. 1-3.) 


| Alder & Hancock, Trans. Zool. Soc. Lond. vol. 111. (1864) p. 119, 
pl. xxxiil. fig. 1; Bergh, Semper’s Reisen, Heft x1i. (1877) p. 540 ; 
Bergh, Danish Exped. to Siam, Opisthobranchiata, 1902, p. 182.| 

One specimen was dredged in Zanzibar Harbour on a sandy 
bottom with a little Zostera (Pl. XXXII. figs. 1 & 2). The ground- 
colour of the living animal is a translucent yellow, like a bit of 
crystallised fruit. On the ample and transparent mantle-margin 
were blotches of peaty red and of different sizes. Smaller spots 
of the same colour are scattered over the whole body, particularly 
above the visceral mass. The under surface is uniform bright 
yellow with a few brown dots. The whole dorsal surface is 
covered with colourless transparent papille (Pl. XX XIII. fig. 2), 
some simple (especially on the mantle-edge), and some compound 
with two or more filaments. It is also plentifully suppled with 
spicules set in a stellate arrangement, but the general consistency 
is quite soft and not stiff. The rhinophore and _ branchial 
openings are slightly raised and tuberculate, but not stellate. 
The rhinophores are large and slightly bent back; the stalk is 
rather longer than the laminated portion. The branchiz are six 
and mostly only bipinnate, though tripinnate branches also occur 
(Pl. XXXII. fig. 3). They are grey with a brown rhachis. The 
foot is grooved in front but not notched. The tentacles are thin 
and digitate. There is no trace of labial armature. The radula 
consists of about 47 rows of simply hamate teeth, each row con- 
taining 40-50 on either side of the rhachis. They are all of the 
same shape and size, except the outermost five or six, which bear 
from seven to ten long fine hair-like denticles on each side of the 
much reduced central hook. No armature was discoverable in the 
reproductive system. 

The alcoholic specimen is quite flat, and is 2°5 centimetres long 
by 1:6 broad, but the living animal was capable of assuming two 
shapes—one flat with a broad mantle-edge, and one high with 
a much narrower edge (Pl. XX XIII. fig. 1), 

I think this animal may be safely identified with the Doris 
(Thordisa) villosa of A. & H. Bergh seems to think that this 
species is probably identical with his Thordisa maculigera, and 
T share this view, though the formation of the outermost teeth 
is not exactly like either his description or his plate, as the 
denticles are longer and the central hook, though much reduced, 
has not vanished. 


368 SIR C, ELIOT ON NUDIBRANCHS [ Dec. 1, 


10. THORDISA STELLATA, Sp. Nn. 

One specimen from Chuaka. nite 

The living animal was soft, but yet distinctly harsh to the 
touch. ‘The colour isa yellowish grey with small sandy patches 
and also dull chocolate blotches, the latter at the mantle-edge 
and round the visceral mass. The underside is of a greyish white, 
with pronounced chocolate blotches round the foot, and a much 
fainter ring of the same halfway to the mantle-edge. 

The preserved specimen, which is much bent, is 2°8 centimetres 
broad, and would be at least 3-5 centimetres long if straightened 
out. The texture is rather leathery, but the back is covered with 
small soft papille of various sizes and colours, and all simple. 
The rhinophore-openings are slightly raised, closed, and apparently 
crenulate. The branchial pocket is slightly raised, stellate, and 
entirely closed by six lobes. The branchiz are yellow, tripimnate, 
five or six in number according as one much smaller than the 
others is reckoned separately or as an appendage. The rhachis 
is very thick and broad. The foot is grooved and notched in 
front. The tentacles are close together above the mouth and 
somewhat flattened. No labial armature could be found. The 
radula consists of 36 rows, each containing about 70 hamate teeth 
of the ordinary type. The innermost are smaller and the outer- 
most less distinctly formed, but neither rudimentary nor denti- 
culate. No genital armature was discoverable. 

This specimen appears referable to 7ordisa and bears a strong 
resemblance to 7’. villoss, but differs in the more leathery 
consistency, the stellate branchial opening, and the outermost 
teeth of the radula, 


11. THoRDISA CROSSLANDI, sp. n. (Plate XXXII. fig. 3 & 
Plate XX XIII. figs. 4-8.) 

Many specimens of this form were captured at Chuaka, on the 
East Coast of Zanzibar, in 1901-02, 

The animals are large, the measurements of a fine alcoholic 
specimen being, length 12°5 centimetres, breadth 9°1, height 2°5, 
The shape is therefore flat and oval. The coloration is in 
its general effect inconspicuous. The upper surface is sandy 
with blotches of brown irregularly bordered with black. The 
under surface (Pl. XX XIII. fig. 3) is whitish with numerous 
brownish spots and a brownish border. But when the upper 
surface is carefully examined it presents a great variety of shades 
of light and dark brown which cannot be easily described or 
depicted. The back is covered with thick-set pointed papillee, some 
of which are developed into distinct filaments at their extremities, 
The general texture is soft. The openings for the rhinophores and 
branchiz are slightly raised, and may be described as tuberculate 
since they open among tubercles, but they do not appear to be 
provided with special tubercles. The branchial pocket is an 
ivregular oval and not stellate or crenulate. The branchie 
(Pl. XXXIIT. fig, 5) are six in number and tripinnate, The 


1903.] FROM EAST AFRICA AND ZANZIBAR. 369 


rhinophores are bent backwards. The foot is fairly broad: it 
is grooved anteriorly, and the upper lamina is notched and 
developed into fairly ample flaps on each side of the division. 
The oral tentacles are digitate, and white with yellow ends. On 
the white labial cuticle are two small yellow patches, measuring 
1 millimetre in length in the largest specimens. They are 
composed of an irregular collection of rods, similar to those 
which form the labial armature of the Discodorids, but can hardly 
be described as plates since the outline is ill-defined and the 
texture loose. The radula consists of about 45-55 rows, contain- 
ing about 80 teeth, on each side of the naked rhachis, over which 
the innermost teeth close so as to render it invisible. The teeth 
(Pl. XXXITII. fig. 4, a & b) are of the ordinary hamate shape 
and all alike, except that the innermost are distinctly smaller. 
At the outer end of some, but not all the rows, is found a small 
degraded tooth. The stomach is free from the hepatic mass, and 
the lower part is somewhat muscular and laminated internally. 
The genitalia are remarkable for the structure of the glans 
(Pl. XXXITI. figs. 6 & 7), which is long, twisted spirally, and 
provided with two rows of tubercles. The central nervous system 
(Pl. XXXII. fig. 8) is much concentrated, as in Asteronotus, 
and the different ganglia cannot be distinguished. 

All my specimens were found adhering to the underside of stones 
in a manner suggestive of sedentary habits. The animal is able, 
however, to swim well upon occasion with a motion somewhat 
resembling that of a sole. It has also some power of self- 
mutilation, and can cast off portions of the mantle, though it does 
so less readily than some allied forms. The branchiz are very 
sensitive, and retract if the shadow of a hand is allowed to fall on 
them. The dorsal papille are kept in constant motion. 

It is extremely difficult to determine the true affinities of this 
species. It has the general form and soft pointed papille of 
Thordisa, and to that genus I think it had better on the whole be 
referred. But it has also a rudimentary labial armature, a 
concentrated nervous system, and a peculiar conformation of the 
genitalia. In this last point it offers some, but not complete, 
analogies to Phialodoris, in which, however, the back is minutely 
granulated and not covered with papille. 


12. Trippa AREOLATA (A. & H.). 


[A. & H., “Notes on a Collection of Nud. Moll. made in India,” 
Tr. Z. 8. ii. 1864, p. 119.] 

Two specimens, one from Mombasa, the other from Wasin. 

Alder and Hancock’s figure gives a good idea of the living 
animal, but hardly emphasises sufficiently its extraordinary 
resemblance to a piece of old worm-eaten sponge Though con- 
spicuous enough when placed by itself in a basin, it is invisible in 
its natural haunts, among sponges and seaweeds. Both my 
specimens were detected by touch only, not by sight, and I 
suspect that the creature is really common. 

Proc. Zoot. Soc.—1903, Vor. Il. No. XXIV. 24 


370 SIR C. ELIOT ON NUDIBRANCHS [ Dee. 1, 


The living animal was spongy and almost gelatinous in texture. 
The alcoholic specimens though flabby have become considerably 
shrunk and hardened. The larger one (to which all the measure- 
ments given below refer) is 5-7 centimetres in length, 3°8 in breadth, 
and 2°3 in height. Down the centre of the back runs a some- 
what indistinct ridge, on each side of which is a row of five pits, 
with black bottoms. There is one similar pit behind the branchial 
pocket. In the smailer specimen the distribution of the pits 
is different, and it would appear that no particular arrangement 
can be regarded as characteristic of the species. In this specimen 
also the dorsal ridge and a knotty crest between the rhinophores 
are much more distinct than in the larger one, bearing out Alder 
and Hancock’s remark that these features are most conspicuous 
in the young individuals. In both specimens the back is covered 
with irregular tuberculate warts or prominences. ‘The rhino- 
phores project out of tubes which are about 5 millimetres high 
and thickly studded with tubercles, about five being set round 
the edge. The branchial pocket projects about 6 millimetres and 
opens backwards. In the larger specimen it is distinctly bilabiate. 
The upper lip is thickly tuberculate in its whole extent and bears 
three compound tubercles on its edge which close like a valve; 
the lower lip has no tubercles on the edge and is altogether 
smoother than the other. In the smaller specimen the pocket 
opens backwards, but is round and not two-lipped. It is probable 
that the tubercles increase in number and size as the animal 
grows older. The branchie are large and strong, tripinnate, and 
apparently five in number, but so deeply bifid that it would 
hardly be wrong to call them ten. In both specimens the foot is 
deeply grooved and notched in front and the upper lamina united 
to the head below the mouth, an arrangement which differs from 
that seen in Spherodoris (levis), where the mouth seems to be 
between the two lamine. 

The labial cuticle is very strong and much puckered, but no 
armature was discernible. Round the buccal mass, at the 
posterior end of the oral tube, are set a number of glands, of 
which I found ten in one specimen and eight in the other. They 
are mostly three-fingered in shape. The radula consists of only 
23 rows, each containing about 40 teeth on either side of the 
naked rhachis, but looks large and broad on account of the 
unusual size of the teeth, which are simply hamate with yellowish 
bases and colourless hooks. The innermost teeth are very small, 
but gradually increase in size up to the 15th, after which they are 
equal. The two or three outermost are reduced. The stomach is 
small but free. No armature was discernible in the reproductive 
organs. 

I think these specimens are clearly the Doris areolata of 
A. & H., and equally clearly referable to the genus T’rippa, Bgh. 
Probably Doris spongiosa Kelaart (Ann. Mag. Nat. Hist. (3) 
iii, 1859, p. 303) is the same species. Trippa (Phlegmodoris) 
mephitica Beh. is a closely allied form, and I should uot be 
surprised if it even turned out to bea variety of, or identical with, 


1903. | FROM FAST AFRICA AND ZANZIBAR. 371 


this species, for these animals evidently undergo great changes, 
both of shape and colour, in alcohol. 


13. TRIPPA MONSONI, sp.n. (Plate XXXII. fig. 4.) 


One specimen dredged at Chuaka on the East Coast of Zanzibar. 

I made the following description of the living animal :—About 
one inch long and lively in its movements. The mantle ample, 
covering both foot and head completely. The foot narrow, grooved 
in front and slightly notched on the upper lip. The tentacles on 
each side of the mouth white and conical. The whole body is of 
a soft, spongy texture. The back is covered with small purplish- 
white prominent reticulations and also bears white tubercles, 
particularly on the edges of the mantle, branchial and rhinophore 
pockets. Both the tubercles and reticulations bear small papille 
or bristles. Between the reticulations are small purple pits and, 
as a result, the general colour seems purple. Besides these, there 
are four large pits, set symmetrically in a square in the centre of 
the back. They are brownish at the sides and deep purple at the 
bottom. There are five other similar but rather smaller pits, 
three in front of the rhinophores, one behind the gills, and one 
placed quite symmetrically at the side of the upper right-hand 
large pit. A white line runs round the edge of the mantle. The 
rhinophores are yellow and finely perfoliate. The gills are usually 
exposed, though they are completely retractile into a rather small 
pocket. They are six in number, yellowish and tripinnate. The 
under surface of the animal is purplish white; there are no spots 
on the foot, but two rows of purplish blotches on the underside 
of the mantle. 

The alcoholic specimen is 1:1 centimetre long and ‘5 broad. It 
is very spongy, like the living animal, but the colour has changed 
to a uniform light yellow, Only the four central pits remain 
distinct, the others having disappeared. There is no labial 
armature. The radula consists of 28 rows of hamate teeth, 
bearing three very small triangular denticles on the side of the 
hook. The innermost teeth are small and less distinctly formed 
than the others; the three or four outermost are rudimentary, 
and sometimes bifid or trifid. There is no genital armature. 

I think this form may be referred to Z'rippa: the spongy 
texture, the pits, and the small internal teeth all seem character- 
istic. In making the dissection, I unfortunately omitted to search 
for the ptyaline glands, and am now unable to say whether they 
are present or not. 


14. FRacAssA TUBERCULOSA, sp. n. 


One specimen from the East Coast of Zanzibar on the reef. 

The living animal was of a dirty-white colour with dull green 
patches on the centre and edges of the back; the whole upper 
surface was sprinkled with small bright blue dots with chocolate 
borders; also there were several dull yellow spots. The under, 
surface was white with an irregular and indefinite network of dull 
green. The junction of the foot with the under surtace of the mantle 


372 SIR C. ELIOT ON NUDIBRANCHS Bes, 1, 


was marked by a thick line of chocolate with numerous small 
bright blue blotches. The dorsal surface was arched and covered 
with large irregularly shaped tubercles bearing secondary knobs. 

The alcoholic specimen is of a uniform greyish white. It is 
4-7 centimetres long, 2°9 broad, and 1°9 high. The margins of the 
rhinophore-pockets are raised and smooth. The branchial pocket 
is not much raised and in itself forms a fairly regular circle, which 
is, however, somewhat distorted by the surrounding tubercles. 
The margin, however, is not tuberculate as e.g. in Staurodoris. 
The branchie are eight and tripinnate, the anterior pair being 
much larger than the others. The anal papilla is large. The 
spots at the junction of the foot and mantle appear to be glandular. 
The foot is grooved and notched in front. On each side of the 
mouth are two small conical tentacles. There is a labial armature 
of two small yellowish plates composed of minute rods. The short 
but very broad radula consists of only 28 rows, containing about 
65 yellowish teeth on each side of the rhachis. The teeth are 
hamate; the innermost fold over the rhachis: there is an 
accessory denticle in the four or five outermost, and the outermost 
of all are smaller and rudely formed. The reproductive system is 
unarmed; there are ample folds surrounding the orifices and a 
prostate is present. 


15, HaLGERDA WILLEYI, sp.n. (Plate XXXII. fig. 5.) 


One specimen captured by Dr. A. Willey, at Lifu, Loyalty 
Islands, and kindly given by him to me, seems referable to this 
genus. It was accompanied by a drawing (vide Pl. XXXII. fig. 5) 
and this note :—“ Lifu, Sandal B., 3.10.96. Reddish yellow (rich 
ochreous) ribbed Doris. The ribs are ochreous and intervening 
valleys have black linear pigment. Tentacles (7. e. rhinophores) 
white tipped with black girdle. The rest of ground-colour is dull 
greyish black. Cloacal rim a dirty white. Foot orange, produced 
behind. On passing the hand over branchie so as to produce a 
shadow they were retracted.” I have captured an Ophiuroid at 
Zanzibar, the coloration of which exactly resembled Dr. Willey’s 
drawing, so that it is possible that this remarkable pattern may be 
cryptic in certain surroundings. 

The preserved animal is considerably shorter and broader than 
the drawing. The length is 3°1 centimetres and the breadth 1:8. 
The general shape is flat ; the foot long and narrow (2:3 centimetres 
long by ‘5 broad), grooved but not cleft in front. The mantle- 
margin is thin but ample, though a large piece has apparently 
been bitten out behind. The general consistency is tough and 
leathery, but there is no rough feeling as in Platydoris. Though 
smooth to the touch, the back is covered with a series of low ridges 
and valleys arranged in an elaborate pattern, which will be best 
understood from the figure (Pl. XX XII. fig.5). It starts partly from 
the mantle-edge and partly from the median dorsal line, by which it 
is divided into two parts, though there is no raised crest. On the 
dorsal surface the ridges are yellowish brown and the valleys black. 
On the lower surface the coloration is much the same, there being 


1903. ] FROM EAST AFRICA AND ZANZIBAR. 373 


numerous black lines instead of valleys. The tentacles are knob- 
like but large and distinct. The rhinophore-openings are quite 
smooth, whitish yellow, and fairly large; they are flattened, but 
look as if they had once projected. The branchial pocket is fairly 
large, quite smooth, and whitish yellow. The rim is thin but 
projects amply. The direction of the opening is posterior, not 
vertical, The branchise (as shown in the figure) consist really of 
two plumes arising one on each side of the anal papilla, but each 
is split into three subdivisions, so that there appear to be six. 
They are scanty and irregular; mostly bipimnate, but partly tri- 
pinnate. The buccal parts are protruded and are yellowish white 
with black spots. There is no trace of labial armature. The 
radula consists of whitish and simply hamate teeth; the outer- 
most are not denticulate or degraded; the innermost are smaller 
and more crowded than the others. On the left side under the 
mantle is a curious gland-like projection, which is perhaps merely 
a blister caused by alcohol. On the right side in the usual place 
is a large yellow lump with black spots on which are placed the 
genital orifices; they are surrounded by strong folds, but no 
armature was discoverable in the organs themselves. 
I propose to call this species Halgerda willeyt. 


16. HaucGERDA WASINENSIS*, sp.n. (Plate XXXIV. figs. 1 & 2.) 


Three specimens captured by Mr. Crossland at Wasin Island, 
Kast Africa. 

They differ from 7. formosa and H. willeyi in having not only a 
pattern formed of ridges on the back, but also distinct knobs at the 
points where the lines of this pattern join one another. 

In the living animal (Pl. XXXIV. fig. 1) the mantle-edge was 
white, but the ground-colour was a dull red-brown ; numerous 
brown spots of a deeper colour were arranged round the mantle- 
edge and a few scattered over the central dorsal area. Over the 
whole dorsal surface was a raised pattern in brilliant orange. The 
white foot was also spotted with deep reddish brown. This colour 
also appeared on the rhinophores in broad bands alternating with 
white, and in broad lines on the rhachis of the white gills. The 
foot projected behind the mantle in crawling. 

The largest alcoholic specimen is 1-9 centimetres long, 1:2 broad, 
and ‘7 high; the contracted foot is 1:4 long and “4 broad. Traces 
of the original colour remain, but the yellow has mostly disappeared. 
The raised network starts from a central ridge and is developed in 
a pattern composed of roughly triangular spaces. The foot is 
1:2 centimetres long and only 3 millimetres broad; the tail is 
5 millimetres long. The anterior margin of the foot is grooved 
and the upper lip notched. No oral tentacles ave visible, but in 
all the specimens the head is so contracted that it would be unsafe 
to say none exist in life. The rhinophore-openings are set in a 
tubercle and are very difficult to see. The rhinophores are long, 
but only a small part is lamellate. The branchial pocket is quite 


* This name is wrongly spelt “wassinensis” on Plate. 


374 SIR C. ELIOL ON NUDIBRANCHS (Hitec wn 


round, very small, with a thick white rim round it, so that in the 
preserved specimen it resembles a dorsal tubercle. The bipinnate 
branchiz are somewhat scanty and irregular; they are arranged 
as in the diagram (Pl. XXXIV. fig. 2), the three posterior plumes 
being quite small. There is no labial armature. The radula consists 
in one specimen of 18 and in another of 24 rows; in both specimens 
the longest rows contain 26 teeth on each side; the rows bend 
downwards near the rhachis, the ten or twelve innermost teeth 
being smaller and more crowded than the others. All are simply 
hamate except the outermost. These are rudimentary ; sometimes 
they bear three or four long denticles and sometimes seem to be 
split up into small separate rods. There is no trace of any 
armature in the genital apparatus; the organs are small and 
possibly even the largest specimen is immature. 

This form presents resemblances to Stawrodoris, some species of 
which have bipinnate branchie, but the dorsal tubercles are few, 
and none are present round the pockets of the branchie or rhino- 
phores, both of which openings are differently formed from those 
of Stawrodoris. 


17. KENTRODORIS RUBESCENS B. (Plate XXXIV. fig. 7.) 

[ Bergh, 8. R. x. p. 411.] 

Several specimens were obtained at Chuaka in August 1901. 
The distribution was apparently very local and the species was 
found only on this one occasion. The alcoholic specimens have 
unfortunately been lost, but I give a figure drawn from the living 
animal and the following notes :— 

The animal was large (about 17 cm. long and 5 broad), soft and 
almost gelatinous. Its most remarkable feature was the great size 
and elevation of the seven quadripinnate gills, which were retrac- 
tile into a large cup. Their tips were as much as 8 centimetres 
above the level of the back. The mantle projected anteriorly and 
formed an ample loose hood over the head. The ground-colour was 
a light pinkish drab with dull yellow spots and brown blotches 
dorsally, There were also a number of thin chocolate lines running 
more or less longitudinally but often branching laterally. The 
under surface was of a light drab-brown, with similar chocolate 
lines on the foot. The dark colour of the gills was due to a 
multitude of such lines. 


The animals were infested by a number of yellow parasitic 
Copepoda. Aig 


I think this animal is Bergh’s Kentrodoris rubescens, though, as 
the specimens are unfortunately lost, it is hard to be quite sure. 
The huge erect gills are even a more prominent feature here than 
in Semper’s figure (oc, cit. pl. xxxiii. fig. 8). 

18. PLATYDORIS EURYCHLAMys B, 

[Bergh, 8. R. xii. p. 510, Suppl.-Heft i. p. 61, xvi. p. 802.] 

Two specimens from Chuaka. 

According to notes made from the living animal, the texture 
was hard and rough. The larger specimen was 8 centimetres long 


1903. ] FROM EAST AFRICA AND ZANZIBAR. 375 


and 4:5 broad. The dorsal surface was covered with reddish- 
brown granulations, each surrounded by a grey or white ring, and 
there were also in one specimen eight chocolate-coloured blotches 
with white edges, four around the rhinophores and four in front 
of the branchial pocket. The rhinophores were dark brown, the 
gills grey, with a thin white line down each pinna. In the 
smaller specimen (which appeared to be indubitably the same 
species) there were no blotches and the gills were drab-coloured. 
The blotches have also vanished from the alcoholic specimen, 
which is of a dull reddish brown, darker in places owing to 
aggregations of minute black spots. The under surface is of a 
uniform reddish brown. The texture is hard and rough, as usual 
in the genus. The visceral mass is 3°2 centimetres high and 
arched, but the mantle-margin is low and flat. This margin is 
exceedingly ample, measuring as much as 2°6 centimetres at the 
sides, 2°2 behind the tail, and 1-7 before the head, although the 
dimensions of the whole alcoholic specimen are only 6°9 by 6:1. 
The small foot is grooved and notched in front but not very 
deeply. The rhinophore-pocket is slightly raised and crenulate. 
The branchial pocket is entirely closed by six lobes. The anterior 
and posterior lobes are broad flaps, and considerably larger than 
those at the side, which are narrow and pointed. The gills are six, 
tripinnate, not very large or very sensitive. There is no labial 
armature. The radula was injured, but was clearly large and 
composed of closely-packed simply hamate teeth. Perhaps the 
formula may have been about 50x 100.0.100. The innermost 
teeth are smaller than the rest; the outermost irregular in shape. 
The stomach is large and free, thin, and only partly laminated. 
The large double blood-gland is partly anterior and partly pos- 
terior to the central nervous system. ‘There is a genital armature 
of discs and hooks as described by Bergh. 


19. PLATYDORIS SCABRA (Cuv.). 


Three specimens from Wasin. 

The alcoholic specimens have preserved the hue of the living 
animal unusually well. The ground-colour is yellowish white, 
with irregular violet mottlings of varying intensity formed mostly 
by minute rings of the same colour with yellowish-white centres 
or by spots. The underside is a clear yellowish white, with na 
markings except at the sides of the foot which are mottled like 
the back. The branchie are light yellow with grey axes; the 
rhinophores, buccal mass, and viscera all light yellow, and there 
is a thin rim of the same colour round the pockets of the rhino- 
phores and branchie. 

The largest speeimen is 9 centimetres long, 5:2 broad, and 2:2 
high. The visceral mass is somewhat arched and the wide mantle- 
margin undulated. It is 1-9 centimetres broad at the sides, 1°5 in 
front of the head, and 1:2 behind the tail. The foot is small and 
narrow ; it is grooved and notched in front but not very deeply, 
and the upper lip is thick. The branchial pocket has six lobes, as 
in Platydoris eurychlamys, the anterior and posterior lobes being 


376 SIR C. ELIOT ON NUDIBRANCHS © [ Dee. 1, 


larger than the others. The branchie are six, tripinate, very 
ample and delicate. The tufts at the side of the posterior pair are 
almost separate, so that the whole number might be reckoned as 
eight. The rhinophore-pockets are closed by indistinct crenula- 
tions. The oral tentacles are of a fair size, white and conical. 
There is no trace of labialarmature. The radula formula is about 
48 x 60.0.60. The teeth are simply hamate, the innermost smaller, 
the two or three outermost irregular. The penis is armed with 
the hamiferous disks characteristic of the genus apparently set in 
four rows, and the vagina provided with strong folds. 

The animal, both when alive and when preserved, is exceedingly 
hard and rough. It is very sluggish, and I have always found it 
fitted into crevices on the underside of stones, as if it had not 
moved for a long period. 


20. Puatyporis rormosa ? (A. & H.), var. 


[ Alder & Hancock, “ Notes on a Coll. of Nudibr. Moll. made in 
India,” Tr. Z. 8. ii. 1864, p. 116.] 

One specimen from Chuaka. 

The notes on the living animal describe it as bright scarlet, 
shaded in places by minute brown specks, very flat, and rough to 
the touch all over. The end of the mantle had been thrown off, 
probably by self-mutilation, so that the body terminated abruptly 
behind the gill-pocket. 

The alcoholic specimen is much bent, but if straightened out 
would be about 5 centimetres long; the maximum total breadth is 
4 and the maximum width of the mantle-margin 1-4.. The some- 
what projecting rhinophore-pockets have slightly crenulate edges, 
as has also the branchial pocket, which is nearly round. The 
branchiz are six, tripinnate, and rather small. The foot is some- 
what broader than usual in the genus, grooved and notched in 
front. The oral tentacles are large and slightly grooved on the 
outer side. The buccal mass is very Jarge and muscular. There 
is no labial armature but a large dark radula, consisting of simply 
hamate teeth, the innermost smaller, the outermost irregular in 
shape. The formula is about 60.0.60 x40. The genital organs 
are very strong and muscular, the male branch set with numerous 
yellow hooks of the shape usual in the genus: the female branch 
with strong folds and lumps. 

I think this specimen may be referred to Pl. formosa, or at 
least that there is not sufficient ground for creating a new species. 
It is certainly not Pl. arrogans (cruenta), which has scarlet 
blotches but a pattern formed by minute dark lines, not spots. 
Allowing for the variations so common in this genus, the coloration 
corresponds fairly well with Alder and Hancock’s description, and 
the grooved tentacles are a remarkable point in common. The 
chief discrepancy is that the branchial pocket is not distinctly 
lobed or stellate, as in the typical Pl. formosa; but I have noticed 
in many species of Platydoris that though on an average this 


character may be very well marked, it may be indistinct in some 
individuals, 


1903. ] FROM EAST AFRICA AND ZANZIBAR. 377 


21. Puaryporis ELxioTi (2) (A. & H.). 


[Alder & Hancock, “‘ Notes on a Coll. of Nud. Moll. made in 
India,” Tr. Z. 8. iii. 1864, p. 116.] 

One specimen from Wasin, dredged in 10 fathoms. 

The notes on the living animal describe the dorsal surface as 
being on the whole of a reddish brown, very beautifully mottled 
with various shades of sandy colour, the visceral mass being darker 
than the rest. The under surface was white: just outside the edge 
of the foot was a row of dark brown spots, and nearer the mantle- 
edge a bright orange border formed of spots set near together. 

The alcoholic specimen measures 6°4 centimetres in length, 4°4 
in breadth, and 2:4 in height. Like Pl. scabra, it is hard and 
rough. The colour is a mottled pattern of white, a sandy tint, 
and reddish brown. Beneath, the sides of the foot and the 
adjacent parts of the broad mantle, which is 2°2 centimetres wide, 
are thickly spotted with chocolate marks arranged so as to give 
the impression of a continuous band. Seventeen chocolate spots 
are irregularly arranged round the foot ona yellowish ground ; 
then comes a fainter band also composed of chocolate spots ; then 
a yellowish border extending to the mantle-edge. The foot is long 
and narrow, grooved and notched in front but not deeply. The 
rhinophore-pockets are closed by six projections. The branchial 
opening is also six-lobed, the anterior and posterior lobes being 
larger than the others, as in Pl. ewrychlamys and scabra. The 
branchie are six, tripinnate, not very large. The oral tentacles 
are much retracted, white and conical. The buccal mass is large 
and muscular, the labial cuticle very strong but unarmed. The 
radula about 40x 70.0.70; the teeth yellowish, simply hamate, 
the outermost smaller but not much degraded. The stomach is 
large and free, strongly laminated in parts; it appeared to contain 
sand, as well as alimentary matter. The penis is armed with two 
rows of hook-bearing scales of the usual type, but set very close 
together, each fitting into the next; the vagina with lumps but 
no scales. 

I feel somewhat doubtful whether this animal should be called 
Pl. ellioti. Neither my specimen nor those described by A. & H. 
present any very definitely distinguishing characters. But, on the 
other hand, there is no feature of importance which militates 
against the identification, and the colours (which A. & H. record 
as varying) are sufficiently alike. 


22. PLATYDORIS PULCRA, Sp. n. 


Two specimens from the neighbourhood of Wasin, dredged in 
10 fathoms. 

The living animal was of a beautiful orange-red, covered closely 
with minute lighter spots. Round the mantle was a border of dull 
white containing purplish-black spots and small specks in one 
specimen, and in the other dull violet spots. The under surface 
is described in the notes on living specimens as of uniform lighter 
orange, but in the alcoholic specimens there is a rim of faint 


378 SIR C. ELIOT ON NUDIBRANCHS (Dee. 1, 


mottlings round the foot. The rhinophores were dark brown 
with white lamelle ; the branchize sandy-coloured. The animals 
were very stiff and harsh to the touch. 

The larger alcoholic specimen is 3°8 centimetres long, and 
broader behind than in front, the maximum breadth being 2:2. 
The whole dorsal surface is covered with extremely minute 
granulations, which can only be seen under a strong lens. The 
rhinophore-pockets are very slightly raised and crenulate. The 
branchial pocket is stellate, with six not very distinct lobes; the 
branchiz are six, tripinnate, small and deeply retracted. The 
anterior end of the foot grooved and notched as usual; the oral 
tentacles distinct, white and conical. The labial cuticle is 
strengthened by some minute rods, but they are not combined into 
a plate or rim. ‘The radula consists of 36 rows, containing about 
50 teeth on each side of the rhachis. The innermost and outer- 
most teeth are smaller, the two or three outermost degraded but 
not denticulate. The male branch of the reproductive organs is 
armed with colourless scales, bearing erect spines, not hooks. The 
female branch has strong folds. 


23. PLATYDORIS INCERTA, Sp. 0. 


Seven specimens, found on brown sponges at low tide, Prison 
Island, Zanzibar. 

The largest of the living animals was about an inch and a half 
long. They were all very flat in shape, sluggish, strongly adherent, 
and, though smooth, of the stiff coarse texture characteristic of 
the genus. The coloration rendered the animal invisible on the 
sponge, but was somewhat variable. As a rule it was yellowish 
brown, but one specimen was tinged with red and another with 
green. On the dorsal surface were collections of minute sandy 
dots, which in some specimens formed a line down the middle of 
the back. The underside was lighter in colour and without 
markings. The rhinophores were tipped with white, the gills 
brown or sandy. 

The alcoholic specimens are all of a uniform dirty yellow. The 
measurements of the largest are: length 1:6 centimetres, 
breadth 0:9, height 0°4; the foot is 1:5 long and 0°7 broad. The 
rhinophore-pockets are crenulate, in some specimens slightly 
raised, in others closed and almost flat. The rhinophores have an 
unusually long stalk bearing a thick bunch of perfoliations, out of 
which rises a narrow bare tip slightly bent backwards. The 
gill-pocket is rather indistinctly stellate, with about six lobes, and 
contains six somewhat small bipinnate branchise. The foot is 
grooved in front and the upper lamina notched. The tentacles 
are narrow and digitate. There is no trace of labial armature. 
The radula formula is about 35.0.35x60. The teeth are white 
and simply hamate, of a rather thick and clumsy shape; the 
innermost are smaller but formed like the rest, the two or three 
outermost are degraded. The reproductive organs appear to be 
unmature, but both the penis and vagina are armed with trans- 
parent, colourless, brick-like scales, apparently arranged in four 


1903.] FROM EAST AFRICA AND ZANZIBAR. 379 


rows, each containing about ten scales. The central nervous 
system is as usual, with very large and distinct eyes. 

The idea that the specimens are immature is supported by the 
size, which is small for the genus, and perhaps by the fact that a 
good many were found together. They are possibly the young of 
some already described form, and offer certain analogies with 
Pl. vicina, in which, however, only the male reproductive organs 
appear to be armed with scales (Bergh, Semper’s Reisen, Suppl.- 
Heft 1. 1880, p. 63). 


24, PLATYDORIS PAPILLATA, Sp. n. 


Nine specimens from Chuaka. 

The living animals were of varying but somewhat sombre 
coloration, ranging from dark peaty brown to yellowish brown, in 
all cases blotched with grey or black markings, greatly varying in 
extent and pattern. The under side of the ample mantle had a 
whitish border, then a yellowish area covered with minute brown 
dots, and, lastly, chocolate-brown blotches near the foot, sometimes 
few and separate, sometimes united in a band. The foot was 
greyish. The gill-pocket closed as in Asteronotus. The dorsal 
surface was covered with numerous small simple papille, and also 
bore some much larger-branched papille, which may have been 
as much as half a centimetre long. In the living animal they 
looked exactly like: bits of sand. Their number varied greatly 
in different specimens: in some they were numerous, in others 
there were only a few near the mantle-edge. The largest living 
specimen was 11 cm. long and 7 broad. 

The measurements in alcohol are: length 8:1 em., breadth 5:1, 
height 2°8. The mantle-brim very thick and 1:6 wide. The 
texture of all the specimens, particularly on the mantle-brim, 
is very distinctly leathery, but not hard or rough. One spe- 
cimen, with an almost smooth back, presents the appearance of 
Asteronotus, but is clearly distinguished by the presence of a 
few branched papille. Also, in all specimens the back is granu- 
late, and not smooth as in Asteronotus. The rhinophore-openings 
are slightly raised and indistinctly crenulate. The branchial 
pocket can be closed by six lobes which meet over it; they are 
not all of the same size, and vary in different specimens. The 
branchie are six, tripinnate, and large. The foot is fairly broad 
and rather amply developed in front, where it has the usual notch 
and groove. It is, perhaps, as a consequence of this development 
that the oral tentacles, being pressed between the foot and head, 
appear flattened in most of the alcoholic specimens, and in some 
expanded into lobes as in Hexabranchus. There is no trace of 
labial armature. The radula is broad, and consists of from 30 to 
40 rows, containing from 60 to 70 teeth on each side of the 
rhachis. The teeth are long and simply hamate, the two or three 
innermost are somewhat degraded; the two innermost are not 
parallel to the rest of the row, but are set almost at right angles 
to it and project into the large bare rhachis (a somewhat similar 
though less-marked arrangement may be seen in Bergh’s figure 


380 SIR C. ELIOT ON NUDIBRANCHS [ Dec. 1, 


of the radula of Asteronotus bertrana 8S. R. plate Ixviii. fig. 9). 
The male reproductive organs are armed with two rows of hook- 
bearing disks, each disk set at some distance from the next one. 
There appears to be an accessory gland on the female branch like 
the glandula amatoria of Asteronotus, but no hasta was discover- 
able. The central nervous system is much concentrated ; above 
it anteriorly and posteriorly le the two very distinct divisions of 
the blood-gland. 

This species seems in many ways intermediate between Platy- 
doris and Asteronotus. But as it has the characteristic genital 
armature of the former, and as one of the principal characters of 
the latter is that the back is quite smooth and neither granulate 
nor papillous, I have thought it better to refer my specimens to 
Platydoris. 

The chief specific character is no doubt the branched dorsal 
papille. I think it probable that this animal is identical with 
the Doris sordida of Quoy & Gaimard from Mauritius, but as 
neither their description * nor their plate shows the branched 
papillee, identification is impossible. 


25. SCLERODORIS OssEOsA (Kelaart). 


[See Kelaart, ““On new Species of Ceylonese Mollusca,” in 
Annals & Mag. of Nat. Hist. vol. ii. 3rd ser. p. 298, 1859; and 
Alder & Hancock, ‘“ Notes on a Collection of Nudibranchiate 
Mollusca made in India,” Tr. Z. 8. ii. 1864, p. 121.] 


Three specimens from the neighbourhood of Wasin. The notes 
on the living animals are unfortunately not forthcoming. 

The alcoholic specimens vary in colour from pale yellow to 
greyish brown. They are hard and rough to the touch like Platy- 
doris. The largest is 3°8 centimetres long, 2°2 broad, and 1:2 high. 
In all the specimens there is an indistinct dorsal ridge, and the 
back is irregularly reticulate and honeycombed ; but while these 
markings extend over the entire upper surface in two specimens, 
they are confined to patches in the third. In all three there 
is one pit, larger and more conspicuous than the others, and 
surrounded by a protuberance in front of the gills. The mantle- 
brim is wide, and extends about 5 mm. beyond the head and tail. 
The rhinophore-openings are somewhat raised and closed by 
valve-like crenulations. The rhinophores are conical and not 
much bent back; the perfoliations cease before the tip. The 
branchial pocket lies at the end of the dorsal ridge and has raised 
edges; it is directed somewhat backwards and is crenulate. The 
branchie are six or seven, with the stem very large compared with 
the scanty perfoliations, which are bi- and sometimes tripinnate. 
The long and narrow foot is grooved and notched anteriorly. 
The tentacles are small, white, and conical. There is a strong 
bluish labial cuticle without any armature. The radula consists 
of about 40 rows, containing about 45 sunply hamate teeth on 


Unless this feature 1s meant to be included under the observation “ Cette espéce 
aun peu la forme d’une Onchidie.” 


° 


1903. | FROM EAST AFRICA AND ZANZIBAR. 381 


either side of the rhachis; the innermost are smaller, the two or 
three outermost degraded and sometimes bifid. The vestibulum 
genitale, like the rest of the body, is full of rod-shaped spicules, 
but no armature could be discovered in the ducts. 

I think this is undoubtedly the animal described by Kelaart 
(J. c. p. 298) and said to resemble a piece of bone or worm-eaten 
white stone. It is also not unlike Alder and Hancock’s plate 
(. c. xxviii. figs. 9 & 10). But their description does not entirely 
correspond with my specimens, particularly in speaking of the 
branchial pocket as “a cup, the margin of which is scalloped and 
produced into a large lobe in front.” Kelaart, on the other hand, 
says there are four or five branchial plumes which emerge hori- 
zontally from under the posterior termination of the dorsal ridge, 
which is correct. The branchiz seem to be somewhat variable in 
number, and, as is often the case with Dorids, admit of being 
counted in more than one way. 


26. SCLERODORIS TUBERCULATA, Sp. Nn. 


One specimen from Prison Island, Zanzibar Harbour. 

The following notes were made on the living animal :—“ Dark 
brown with sandy spots, exactly like a sponge splashed with sand. 
Underside clear bright brownish red. Branchial pocket crenu- 
late. The middle part of back covered with conical warts, which 
form an irregular keel; smaller warts on mantle-edge. Rhino- 
phores red; branchize eight, voluminous; axes red, tips white. 
Animal alters shape, sometimes rather high, sometimes quite flat 
like Platydoris. Consistency quite hard and rather rough. ‘Two 
depressions with deep black markings as in some species of 
Trippa.” The alcoholic specimen bears a strong general resem- 
blance to Zrippa areolata, but is stiff and spiculose like Platydoris, 
and has nothing of the flabby gelatinous feeling which charac- 
terises 7’. areolata. The back is covered with irregular tubercles, 
simple and compound, of all shapes and sizes, all granulate and 
sometimes connected by ridges so as to form a reticulation. There 
is an indistinct median ridge and two large pits with black 
bottoms, one in front of the branchial pocket and one about half- 
way up on the right-hand side. The general colour is greenish grey. 
The length is 6-5 centimetres, the breadth 4:2, and the height 2:2. 
The oral tentacles are distinct, digitate, and white. The foot is 
grooved and notched in front, the upper lamina being attached 
to the head below the mouth. ‘The labial cuticle is strong and 
puckered, but no armature was discovered nor any ptyaline glands 
as in Z'rippa areolata. The radula consists of 40 rows, containing 
about 50 teeth on each side of the rhachis. These teeth are white 
and simply hamate ; the innermost are smaller than the rest, the 
two or three outermost degraded and often bifid. The stomach is 
free and laminated internally. No genital armature was discernible. 

The rhinophore-pockets are raised and provided with nine valve- 
like tubercles, of which two are much larger than the others. 
The rhinophores are short and thick, with about 50 perfoliations. 
The branchial pocket is entirely closed by ten valve-like tubercles, 


382 SIR C. ELIOT ON NUDIBRANCHS [ Dec. 1, 


some of which are grooved outside and thus appear double. The 
branchize are eight in number, tripinnate, and very much 
retracted. The rhachis is very stout and strong. 


27. SCLERODORIS MINOR, Sp. n. 


One specimen from Chuaka. 

The living animal is described as superficially resembling 
Platydoris papillata, differing only in that the dorsal surface is 
covered with ridges which form a raised reticulate pattern. The 
colour was a uniform greyish brown, with a few patches 
resembling adherent sand. The underside of the mantle was 
warm grey with minute brown spots, the foot dirty orange. The 
gill-pocket did not close completely when the branchize were 
retracted. 

The alcoholic specimen is 2°7 centimetres long, 1-6 broad, and 
1:l high. The texture is rough and leathery, with the peculiar 
feeling common in Platydoris. All the dorsal surface, including 
the reticulations, is covered with minute tubercles. There is a 
rather indistinct median keel, from each side of which extends 
a somewhat irregular reticulate pattern. The rhinophore-openings 
are slightly raised and very slightly crenulate. The branchial 
pocket is also slightly raised, of irregular shape, but not 
crenulate, ridged vertically, and nearly but not quite closed. The 
branchie are eight, the two posterior shorter than the rest; 
the others are tall, thin, and sparse, so that in the alcoholic 
specimen they appear simply pinnate, though they are really bi- 
and sometimes tripinnate. he foot is rather broad; the front 
notched and the upper lamina apparently attached to the head, at 
the side of which are the conical oral tentacles. The snout is 
protruded. ‘There are scattered minute rods in the labial cuticle, 
but they are not combined into plates. The radula consists of 
33 rows of yellow, regular hamate teeth, which do not diminish 
much in size, either at the rhachis or at the end of the rows: 
there are about 45 on the complete rows on each side of the 
rhachis. No genital armature was discernible. 

This specimen bears a strong general resemblance to Sclerodoris 
tuberculata, and may perhaps be a young individual of the same 
species. The radula is, however, not quite the same, there are no 
pits on the back, and the reticulate pattern is more distinct than 
in the larger animal. It is possible that as the animal becomes 
older the tubercles and pits may develop at the expense of the 
pattern. 


28. SCLERODORIS RUBRA, sp. n. 


One specimen from the reef off the East Coast of Zanzibar. 

The living animal bears a most remarkable resemblance to a 
vermilion sponge which is common at Zanzibar. It was not, 
however, found on the sponge, but alone among Zostera near the 
shore, and not in any way concealed. It was picked up under the 
impression that it was a species of sponge. The colour was red, 
with some very natural-looking sandy patches. The texture was 


1903.] FROM EAST AFRICA AND ZANZIBAR, 383 


firm and fleshy, with something of the rough feeling characteristic 
of Platydoris. 

The alcoholic specimen is dirty white in colour with greyish 
blotches. The measurements are: length 5:5 centimetres, breadth 
2°7, height 2°6. It will therefore be seen that the shape is very 
distinctly arched. In the middle of the minutely granulated 
back is an indistinct keel from which extends on either side a low 
fleshy reticulation. Independent of this reticulation, and some- 
times concealing it, are a number of excrescences which, even in thé 
alcoholic specimen, present the most extraordinary resemblance 
to the miscellaneous growths and accretions found on old shells 
and sponges. Some are as much as 4 millimetres high. I en- 
deavoured to pull them off, being sure they could not be part of 
the mantle. They are so, however, and afford an even more 
remarkable case of mimicry than 7’rippa areolata. The edges of 
the rhinophore- and gill-pockets are crenulate, but not raised. 
The eight tripinnate branchize are deeply retracted into the 
bottom of the pocket. The anal papilla is large and has a cleft 
down the posterior side which appears natural. In the alcoholic 
specimen the foot is narrow with the edges turned inwards, but 
it was probably fairly broad in life: it bears a shallow groove in 
front, the upper lip of which is notched. The tentacles are very 
small and digitate. The blood-gland is large, reddish, and double: 
the central nervous system in a reddish capsule and much con- 
centrated. There is no labial armature. The radula, which is 
large and wide, consists of 42 rows, containing about 55 large 
blunt hamate teeth on each side of the rhaehis: the innermost 
are smaller than the others, and one or two of the outermost are 
also smaller and distinctly bifid. The reproductive system is 
unarmed. 


29. SCLERODORIS CORIACEA, sp.n. (Plate XXXIV. figs. 3 & 4.) 


One specimen from a cave near Chuaka on the East Coast of 
Zanzibar. 

The living animal was yellowish brown in colour above and 
light orange underneath. The preserved specimen is 2°4 centi- 
metres long, 1°7 broad, and ‘8 high in the middle of the back, 
the sides of the mantle being very low. The foot is large, 
being 1:9 centimetres long and 1-1 broad: the sides are developed 
into wide and very thin expansions. The anterior margin is 
deeply grooved, but the upper lip is not split though it is indented. 
The rhinophore-openings have somewhat raised and indistinctly 
crenulate edges. The branchial pocket is wide, conspicuous, and 
somewhat two-lipped in shape. The edge is turned outwards and 
is not at all crenulate. The branchie are six and tripinnate. 
The general texture of the animal is leathery, and the whole 
dorsal surface is covered with a distinctly raised but somewhat 
irregular reticulate pattern. Both this pattern and the ground 
surface are granulate. The buccal tentacles are long, thin, and 
pointed. There is no labial armature. The radula consists of 
40 rows, with about 40 teeth on each side of the naked rhachis. 


384 SIR C. ELIOT ON NUDIBRANCHS [Dec. 1, 


The teeth are simply hamate; the innermost are smaller; the 
outermost smaller and imperfect in shape, bifid or irregularly 
serrulate. The stomach is large, muscular, and free from the 
liver. No armature was visible in the reproductive apparatus, 
and the genital orifices were unusually small and inconspicuous. 

On the right side of the liver was found a parasite (PI. XXXIV. 
fig. 4), extending from the fore end halfway down, the head being 
bent downwards round the fore end of the liver. ‘The impression 
of the parasite on the liver was very distinct. 

This form has not the hard feeling characteristic of Sclerodoris, 
but as the back is leathery, reticulate, and granulate, I hardly 
think it advisable to create a new genus for its reception. 


30. AsTERONOTUS HEMPRICHI EKhr. (Plate XXXIV. figs. 5 & 6.) 

[ Ehrenberg, Symbol Physics, Animalia Evertebrata, 1831.] 

This large animal is common under rocks at Zanzibar and in all 
parts of British East Africa, between tides. 

Tt has a characteristic feeling like leather or india-rubber, and 
not hard or rough like Platydoris. The ground-colour varies 
from black to olive or chocolate-brown. The skin is quite smooth, 
but bears a number of irregular lumps, resembling blisters in 
appearance. Towards the edges of the mantle they are smaller 
and somewhat confluent. Down the centre of the back runs a 
more or less elevated ridge, varying much in different specimens, 
All these protuberances are of a much lighter tint than the rest 
of the body. In some specimens there are small white lines round 
the lumps. The large branchiz are white or pinkish. The under- 
side of the mantle is light yellow, with a row of chocolate blotches 
forming a line round the margin of the mantle and another round 
the foot. The coloration shown in the figure (Plate XXXIV. 
fig. 5), though an accurate representation of some specimens, is 
not the commonest. ‘The greater number of individuals are more 
distinctly green. The animals are sluggish and show little 
inclination to move. The branchiz are comparatively insensitive. 
Though completely retractile, they are not hidden when the teeth 
close over the pocket, but can be clearly seen at the bottom of the 
cavity (Pl. XXIV. fig. 6). 

The largest alcoholic specimen measures 8°3 cm. in length by 6, 
and is 2°4 em. high. The foot is 6 in length by 1:5; the anterior 
margin is deeply notched and grooved, the upper flap being very 
ample. The tentacles are large and in two specimens show a 
rudimentary foliation resembling that of Hexabranchus. This 
may be due to artificial compression, but Iam not sure. The lips 
are also ample and protruding, so as to look like a second pair of 
tentacles. The rhinophore-pockets are somewhat raised and 
smooth, sometimes distinctly bilabiate. The branchiz are six, 
ample, and quadripinnate. The margin of the pocket is produced 
into six lobes, which partially close over it. The radula consists 
of about 32 rows, each containing about 45 large yellow teeth on 
either side of the wide, naked rhachis. The teeth are simply 
hamate: the outermost two or three are smaller, and the imner- 


1903.] FROM EAST AFRICA AND ZANZIBAR. 385 


most are set almost at right angles to the rhachis. The stomach — 
is large and free, but thin and not laminated nor muscular. The 
blood-gland is large, and the nervous system is very concentrated. 
The generative system appears to be as described by Bergh, but 
the glandula and hasta amatoria are difficult to see and were 
satisfactorily detected only in one specimen. 

These specimens are, I think, clearly Ehrenberg’s Asteronotus 
hemprichi, from Massaua. He describes it as “ sex-pollicaris, 
oblongus, glaber, vesiculosus, supra fuscus, lineis circulisque niveis 
sparsis, vesicas dorsuales cingentibus, subtus lateritus, pede flavido, 
branchiarum apertura lobulis sex stellatim positis preecludenda. 
.... Branchiarum e dilute laterito seu carneo albicantium 
fasciculus amplus.” 

Prof. Bergh seems inclined to think (S. R. xvii. p. 917) that 
the real species of this genus are not more than three, hemprich, 
mabilla, and cespitosus. The differences between these three 
do not seem to me to be clearly defined, and my numerous 
specimens, which I unhesitatingly refer to one species, present 
connecting links, especially in colour, which make me think that 
the three species are merely varieties of one. 


EXPLANATION OF THE PLATES. 


N.B.—Except in the cases noted, the figures are drawn from the living animal. 


PratE XXXII. 


. Thordisa villosa (p. 367), ventral view. 

Dorsal view of the same. 

Thordisa crosslandi (p. 368), ventral view. The margin of the mantle is 
inturned here and there, showing the mobile papillae which cover the 
dorsal surface. 

4, Trippa monsoni (p. 371), dorsal view, much enlarged. 

5. Halgerda willeyi (p. 372), from a drawing by Dr. Arthur Willey. 


PratE XXXIII. 


Fig. 1. Thordisa villosa (p. 367), head and anterior end. The figure shows an 

extreme elevation of the body, which normally is flat. 

2. Dorsal papilla of the same species, with flexible pigmented end and spicule- 
stiffened base. Also a portion of the mantle-edge magnified. 

3. Branchie of the same. 

4. Thordisa crosslandi (p. 368). Teeth from the radula: a, upstanding; 
b, laid flat. 

5. Gills and anus of the same. A ridge (a) connects the higher part of the 
rhachis with the anal papilla (0). 

6. Dissection of the retracted penis of the same, showing the shape and structure 
of the enclosed glans. 

7. The glans penis of the same is slit open, showing it to be hollow and to 

8 


Fie. 


go 19 


contain a prolongation of the vas deferens which passes to its tip. 
. The central nervous system of the same in its sheathing of connective tissue. 


PratTe XXXIV. 
. Halgerda wasinensis”™ (p. 373), dorsa) view. 
. Diagram of the arrangement of its gill on the rhachis. 
Sclerodoris coriacea (p. 383). Pencil drawing from the preseryed specimen. 
. Degenerate Copepod parasite found in the liver of Sclerodoris (p. 384). 
Asteronotus hemprichi (p. 384). 
. Enlarged view of the gill-opening when the branchie are as completely 
retracted as is possible. 
. Kentrodoris rubescens (p. 374), about half natural size. 


Gg 
C2 OVE Cobo 


i <7 


”? 


* This name is wrongly spelt “ wassinensis” on Plate. 


Proc. Zoon, Soc.—1903, Vou. Il. No. XXV. 25 


386 MR. F. E. BEDDARD ON THE TONGUE (Decal 


6. A Note upon the Tongue and Windpipe of the American 
Vultures, with Remarks on the Interrelations of the 
Genera Sarcorhamphus, Gypagus, and Cathartes. By 
Frank E. Bepparp, M.A., I’.R.S., Prosector to the 


Society. 
: [Received October 15, 1903.] 


(Text-figures 45-48.) 


In a recent communication * which dealt to a slight extent with 
the arrangement of the papille upon the upper surface of the 
tongue in Accipitres, I referred to certain peculiarities in which 
the tongue of Vultwr diverged from that of its allies, though 
agreeing with it in certain broad features that differ from the 
condition in the Falconine group of the Accipitres. J am not for 
the present again concerned with the latter subject, but with a 
more detailed account of the structure of the tongue in Vwltur and 
its comparison with some other forms. 

In an example of Vultur calvus (3), the tongue is 35 mm. in 
length, and thick, of a boat-lke form, the sides being bent 
upwards. There is the usual posterior row of spines, and, in 
addition to these, a lateral row on each side which commences 
quite posteriorly (see text-fig. 45) and extends anteriorly for a 
distance of 11 mm. on the left side and 10 mm. on the right. The 
spines are flattened and backwardly directed, getting blunter in 
form anteriorly ; there are seven on the left side and five on the 
right. In Vultur auricularis the lateral rows of spines are more 
pronounced and somewhat different in form, thus affording, it 
may be remarked in passing, an additional character to distinguish 
the two species. In this bird the tongue measures 39 mm. in 
length, and the lateral rows of spines end at 14 mm. and 13 mm. 
respectively from the posterior end of the organ; as in VJ. calwus, 
the left row is the longest. It will be observed, therefore, that in 
both species the proportions of the spine-bedecked and spineless 
regions of the tongue are much the same. In V. e@uricularis the 
spiny papille are sharper and more numerous. I counted ten of 
them on the left side and eight on the right. An accurate notion 
of their shape and relations will be obtained from an inspection 
of the accompanying drawing (text-fig. 45). The form of the 
tongue, as well as the number of the lateral papille, differentiates 
Vultur from Gypohierax. The latter has a flatter, longer, and so 
far more typically ‘‘ Accipitrine” tongue. It has no traces of the 
characteristic lateral spiny papille, which cannot, therefore, be 
regarded as a “ Vulturine” character, correlated perhaps with 
diet. And though this one instance may not be regarded as 
sufficient of itself to emphasise the suggestion, it is supported by 
converse examples. In Aquila chrysaétus, Nisaétus fasciatus, and 
Gypactus barbatus there ave distinct traces of these same papille 


* “On the Syrinx in the Accipitres, &c.,” P. Z. S. 1903, vol. ii. p. 157. 


1903. ] AND WINDPIPE OF THE AMERICAN VULTURES, 387 


(three or four in number). In the larger species, moreover, the 
tongue is thick and boat-shaped, but not so markedly as in 


Vultur. 


Text-fig. 45. 


Left-hand figure: dorsal view of tongue of Valtwr awricularis. 
Right-hand figure: dorsal view of tongue of Gypagus papa. 


A, lateral spines; B, posterior spines. 


I believe that these facts throw some light upon the extra- 
ordinary tongue of the American Vultures, which differs so much 
from that of other Accipitres that Prof. Fiirbringer rightly uses 
it * as one of the many characters distinguishing these birds from 
the Old World Vultures. The general structure of the tongue in 
the Cathartide has been described: for example by MacGillivray T 
in Cathartes atratus, and by Gadow in Gypagus papa =. In these 
birds and in the Condor the tongue is edged with a regular row 
of backwardly directed papille ‘like the teeth of a saw.” An- 
teriorly these spines are blunter and shorter, and forcibly suggest 
the lamellz on a Duck’s bill. In Gypagus there may be, in addi- 
tion to the lateral spiny papille, a few to the inside of each row ; 


* Untersuch. z. Morph. u. Syst. d. Vogel, Amsterdam, 1888, p. 1304. 

+ In Audubon, ‘Ornith. Biography,’ v. 1839, p. ae 

< Bronn’s Thier-Reich, Végel, pl. XXV ill. fic. 4; this figure is referred in the 
explanation of the plates. to Gyps sp.” Ie eather from the text that Dr. Gadow’s 
description of the tongue was partly drawn from an unpublished memoir ot 
Dr, W. Marshall, and that the drawing belongs to that MS. I fecl sure that there 
is some mistake, and that the tongue figured is that of Gypagus or perhaps 
‘athartes. 


388 MR. F. E, BEDDARD ON THE TONGUE | Dec. 1, 


and there are traces of these, perhaps reduced to one spine, in 
Cathartes. 1 cannot, however, with any confidence distinguish the 
tongues of the two genera. The point, however, which I wish to 
emphasise is that this single inner spine, if there be only one, 
which lies medianly of the lateral fringing row, is the vestige of the 
V-shaped edging of spines which borders the tongue of other 
Accipitres posteriorly. The tongue of the Carthartide, therefore, 
differs from that of other Accipitres in the nearly complete dis- 
appearance of the spines which border the tongue posteriorly, and 
the hypertrophy of the lateral spines which are indicated in some 
Accipitres, particularly in Vultwr. I may remark finally that the 
numerous large glandular orifices (figured by Gadow) which are 
scattered over the tongue in Gypagus and Cathartes are not found 
in Vultwr; nor is the slit of the glottis in the latter edged with 
spines as it is in the Cathartide. j 

In my work* upon the anatomy of Birds, I briefly drew attention 
to the fact that the “‘syrinx” in the American Vultures presents 
two types—one characterising Cathartes, and the other the two 
genera Gypagus and Sarcorhamphus. 1 take the opportunity of 
expanding that account. In remarking that the Cathartide are 
distinguished by their completely closed bronchial rings, I take it’ 
that Prof. Fiirbringer had in view the genus Cathartes only. It 1s 
only Cathartes that figures in the description of the muscular 
anatomy of Birds contained in his great work. In supporting 
Dr. Fiirbringer’s statement, I found it necessary f to limit that 
statement to Cathartes, since in Gypagus and Sarcorhamphus the 
bronchial “rings” are in reality semirings, as in birds with a 
tracheo-bronchialsyrinx. The actual difference, however, between 
Cathartes and the other types is one of degree. In Cathartes the 
bulk of the bronchial cartilages are complete rings, while a few at 
the end (I counted five of these in two specimens selected at random) 
are semirings. In Gypagus, on the other hand, all the bronchial 
cartilages are semirings, In this point Gypagus agrees with 
Sarcorhamphus. A further peculiarity is noteworthy as distinctive 
of the several genera of Cathartide with which I deal in the 
present communication. In Sarcorhamphus 1 have figured ~ the 
membrana tympaniformis as continued upwards along the trachea 
posteriorly, which results, therefore, in the production of tracheal 
semirings. This longitudinal fissure of the tracheal rings exists 
for some distance. In Gypagus precisely the same state of affairs 
is to be found, only it is less extensive than in the Condor. I can 
find no trace of any transverse break in the tracheal rings of 
Cathartes. I have been able to examine four individuals which 
include both sexes. If it occurs, the probabilities are in favour of 
its being exceptional, We have, therefore, in the structure of the 
windpipe in these American Vultures two distinct types—one 
developed in Cathartes, and the other in Sarcorhamphus and 
Gypagus. 


* “The Structure and Classification of Birds,’ London. 1898, 
ii Loe. eit., see also “On the Windpipe and Heart of the Condor,” P. Z.S. 1902, 
vol. 1. p. 399. t P. Z.S. 1902, vol. i., text-fig. 30, p. 241. 


1903. ] AND WINDPIPE OF THE AMERICAN VULTURES. 389 


Interrelationships of the Genera CATHARTES, SARCORHAMPHUS, 

and GYPAGUS. 

The foregoing remarks upon the windpipe lead to the con- 
sideration of the relationships inter se of the American Vultures. 
Garrod pointed out * the agreement of Sarcorhamphus and 
Gypagus in possessing the femoro-caudal, which is absent in 
Cathartes. Pycraft, in an osteological survey of the ‘ Falconi- 
formes” +, has noted characters, some of which lead him to bracket 
together Gypagus and Sarcorhamphus, though others do not 
apparently justify such a juxtaposition. On the whole, it seems 
to me to be clear that the skulls of Sarcorhamphus and Gypagus 
are more like each other than either is to the skull of Cathartes ; 
and I shall attempt to justify this view by calling attention to a 
few minutiz which have not been dwelt upon in this connection. 

The outline of the skull, as seen from the side, is very charac- 
teristic. Cathartes has, relatively speaking, a long low skull with 
a proportionately longer maxillary and premaxillary region ; it 1s 
almost Cormorant-like in aspect. In the other two genera this 
region is deeper and shorter and descends with more of a curve, 
instead of possessing an upper contour-line which is almost a 
straight line. The premaxillary region, in fact the end of the 
beak region, is much more inflated in Gypagus and Sarcorhamphus 
than in Cathartes. The pervious nostrils are of greater length and 
narrower across in Cathartes than in the two other genera which 
agree with each other in this feature; moreover, in Gypagus 
and Sarcorhamphus the conjoined maxillary plates are visible as 
projecting forward beyond the posterior end of the nostril, more 
especially in Gypagus. In Cathartes they are not so visible. On 
the lateral aspect of the skull another curious though small differ- 
ence may be noted, which serves to ally Sarcorhamphus and Gypagus 
and to divide them both from Cathartes. In the last-mentioned 
bird Mr. Pyeraft has correctly figured £ a groove which he terms 
the “nasal hinge.” This appears to separate off the lacrymal (fused 
to the frontal) from the nasal in front. The suture in question 1s 
semicircular and with the convexity anterior. In both Sarco- 
rhamphus and Gypagus the hinge is present but has more the form 
of a half ellipse; moreover, the concavity is forward and it thus 
appears as if a backward process of the nasal were embraced by 
the concave anterior margin of the lacrymal. Mr. Pycraft has 
also figured §, behind the nasal hinge in the same bird, a strong 
ridge which presumably marks the boundary of the lacrymal, and 
is prolonged so as to slightly overhang the orbit. This ridge is: 
quite absent in both Sarcorhamphus and Gypagus, the orbital 
margin of which is here smooth. It must be admitted, however, 
that, in one point exhibited in this particular region of the skull, 
Cathartes is rather intermediate between the Condor at one extreme 
and G'ypagus at the other. This is the backward direction of the: 

* Collected Papers, Loudon, 1881, p. 210. 

+ P. Z.S. 1902, vol. i. p. 277. 

+ Loe. cit. pl. xxxii. fig. 1, nh. 

§ Fig. 10. Wrongly ascribed in the explanation of plates to Serpentarius: 


390 MR. F. E. BEDDARD ON THE TONGUE [ Dec. I 


Text-fig. 46. 


Sarcorhamphus equatorialis. 


Upper figure: insertion of maxillo-jugal (J). : 
Lower figure : mandible of the same, viewed from behind. 


Text-fig. Ah) 


Cathartes atratus. 


Upper figure: insertion of maxillo-jugal (J), 
Lower figure: mandible, viewed from behind. 


1903.] AND WINDPIPE OF THE AMERICAN VULTURES. oon 


descending process of the lacrymal, not so pronounced as in the 
Condor, but more developed than in Gypagus, where it is indeed 
vertical in direction. Another difference which allows of a closer 
association of the Condor with the King Vulture than of either 
with Cathartes, is the condition of the interorbital septum. In 
both Sarcorhamphus and Gypagus there are two large vacuities, 
the larger being in front. In Cathartes the septum is less per- 
forated, the posterior vacuity alone being present—a tiny hole * 
representing the anterior vacuity. Pycraft has called attention to 
a series of stages in the development of the attachment of the 
maxillo-jugal bar. It appears to me that the facts observable tend 
to suggest that Gypagus stands rather apart from both Cathartes 


Text-fig. 48. 


Gypagus papa. 


Upper figure: insertion of maxillo-jugal (J). 
Lower figure: mandible, viewed from behind. 


and Sarcorhamphus. In the former genus (text-fig. 48) the bar 
divides only just before the insertion on to the maxilla. The upper 
lamella is of large size, and, as in the two other genera, fits on to 
a hollowed-out area of the maxilla. In Cathartes (text-fig. 47, 
p- 390) and Sarcorhamphus (text-fig. 46, p. 390) the division of 
the bar occurs much further back. The upper lamella is certainly 
larger than the lower bar, but it is relatively smaller than in 
Gypagus. In Sarcorhamphus, indeed, it is actually of the same 
breadth as in Gypagus, though the skull of the Condor is twice 
the size of that of the King Vulture. Furthermore, the forward 


* This seems to be not always present. It is not figured by Pyeraft. 


392 ON THE TONGUE ETC. OF THE AMERICAN vuLtuRES. [Dec. 1. 


edge of the upper lamella, which forms the joint, is neatly rounded 
off instead of projecting irregularly as in Gypagus. These 
points will be better understood after an inspection of the accom- 
panying drawings (text-figs. 46-48). It will be seen that my 
reading of the facts differs slightly from that of Mr. Pycraft. 
Articular perfection appears to me to have been arrived at both 
in Cathartes and Sarcorhamphus from a lower stage such as 
persists in Gypagus. 

On the ventral surface of the skull a very conspicuous difference 
defines Clathartes on the one hand from Sarcorhamphus and 
Gypagus on the other. In the two latter birds the basitemporal 
region is deeply excavated, and the sides are prolonged into very 
marked exoccipital processes. In Cathartes, on the contrary, 
this region of the skull is much flatter and there are no such 
conspicuous lateral processes. The inner lamina of the palatines 
is hooked and overhanging posteriorly in Sarcorhamphus and 
Gypagus; in Cathartes this region of the bone does not over- 
hang posteriorly and is merely triangular in form. Moreover, in 
Cathartes the anterior half of the palatine has its broad surface in 
a plane horizontal, in the other genera it is rotated upwards and 
is at an angle to the horizontal plane. <A final point to which I 
desire to direct attention is illustrated in the accompanying 
drawings (text-figs. 46-48). These sketches represent the man- 
dibles of the three genera under consideration viewed from behind. 
The relatively, as well as actually, much greater thickness of the 
internal angle in Sarcorhamphus and Gypagus will be apparent. 

The foregoing account does not at all pretend to be a detailed 
review of the structure of the skull in the American Vultures. I 
have economised space by simply dealing with those facts which 
appear to me to throw light upon the neutral relationships of the 
genera Sarcorhamphus, Carthartes,and Gypagus. Other facts which, 
in my opinion, do not bear upon this subject have been ignored. 
The result is, I think, to show that the Condor and the King 
Vulture are comparatively slight variations of the same type, 
while Cathartes stands equally apart from both, a conclusion which 
is quite in accord with current ornithological opinion. J think it 
is going rather too far, as has been done*, to include both the 
former Vultures in one genus; but it is, in my opinion, manifestly 
absurd to combine Cathartes and Gypagus, and to write of 
Cathartes papa, as has also been done *. 


* “Standard Natural History,’ Boston, 1885, p. 268. 
+ Taschenberg, ‘ Bibliotheca Zoologica,’ v. 1899, p. 3966. 


Acanthoderes 

lateralis, 248. 
Acanthodoris, 358. 
Accia, 256. 
Acheeta, 204. 

bohemica, 222. 

eisenit, 222. 

maorica, 205, 206, 221, 

232. 

(Anacheta) cameranoi, 
Achryson 

surinamum, 248, 
Acomys 

cahirinus, 347. 

dimidiatus, 347. 

nesiotes, 198, 261, 347. 
Acreea 

phlegetonia, 258. 
A&geon 

fasciatus, 29. 

(Crangon) fasciatus, 

24. 


(—) trispinosus, 24. 
7®olosema, 214. 
/Equorea 

albida, 182. 

violacea, 182. 
Agbalus 

sp., 249. 

Aglantha, 177. | 
Aglauropsis, 187. | 
Agouti 

paca, 241. 
Aleces, 273. 

machlis, 262. 
Alestes 

affinis, 329. 

macrolepidotus, 329. 
Alurnus 

nigripes, 249. 

thoracicus, 249. 
Ameiva 

surinamensis, 70. 


Proc. Zoou. Soc.—1903, Vou. II. No. XX VI. 


INDEX. 


Amphicodon 

fritillaria, 168. 
Amphignathodon 

guentheri, 116. 
Anacanthus 

costatus, 248, 
Angiostomum 

brachylaimus, 154, 155. 
Anisodoris, 360. 
Anisomys, gen. noyv., 

imitator, 200. 
Anomala 

sp., 246. 
Anthropopithecus 

troglodytes, 198, 315. 
Aotus 

azare, 234. 
Aphrodite 

aculeata, 148. 
Apostolepis 

assimilis, 70. 
Apterodema 

acuticollis, 256. 

paraguayensis, 255, 256. | 
Aquila, 157, 163. 

chrysaétus, 386. 
Archibuteo, 157, 163. 
Archidoris, 356, 357, 360, 

367 


africana, 354, 356, 361, | 
362, 363. 

australis, 356. 

incerta, 356. 

kerguelensis, 356. 

minor, 394, 356, 361. 

nyctea, 356. 

rubescens, 306. 

tuberculata, 357. 
Artachza, 363. 
Artibeus 

planirostris, 234. 


| 


Asbolia 
villosipes, 259. 


Ascaris 
dipsadomorphi, 158, 
155. 


infundibulicola, 158, 
155. 
’ solitaria, 158, 155. 
Astacus 
gammarus, 195. 
Astana, 246. 


Asteronotus, 3857, 359, 
360, 361, 363, 369, 
379, 3880. 


bertrana, 380. 

cespitosus, 385. 

hemprichi, 355, 384, 

385. 

mabilla, 385. 
Asturina, 157, 163. 
Atherinichthys 

brasiliensis, 60. 

sallei, 60. 
Atollia 

bairdii, 190. 
Aurelia 


flavidula, 169. 


Babirusa, 262. 
Balbiana 
siamensis, 156. 
(Sarcocystis) stamensis, 
155. 


Barbus 
duchesnit, 382. 
erlangert, 331, 334, 
paludinosus, 332. 
Barilius 
loati, 332. 
niloticus, 332. 
Bathycodon, 187. 
Batonota 
ensifer, 249. 
gregaria, 249. 
Belopzeus 
carmelitus, 253. 


26 


394 


Belopzeus 

niger, 247, 252, 258. 
Boa 

constrictor, 336. 
Bos 

bubalis, 155, 156. 

taurus, 263. 
Bostrychopsis 

uncinata, 247. 
Bothriocephalus, 145. 
Bothrioneuron 

americanum, 219. 
Bothynus 

sp., 246. 

ascanius, 246. 
Bougainvillia 

macloviana, 172. 

mertensii, 171, 172. 

superciliaris, 172. 
Brachyenathus 

sp., 245. 

imperialis, 245, 
Branchiura 

coccinea, 210, 211, 

212. 
sowerbyi, 211, 212, 216, 


Bufo 
melanostictus, 150, 151, 
154, 155. 
penangensis, 150, 151, 
155 


typhonius, 69. 
Bulimulus 
corumbaensis, ‘71. 
montivagus, 70. 
Buteo, 157. 
Butherium 
corvinum, 248. 


Cadlina, 855, 359. 
Callichroma 
suturale, 248, 
Calligrapha 
annulata, 54. 
Callithrix 
melanura, 234. 
Callula 
pulchra, 152. 
Calymmus 
cuculatus, 247. 
Camaria 
levipennis, 247. 
Camelopardus 
gira ft, 263. 
Camelus 
hactrianus, 1. 
dromedarius, 262. 
Oancer 
pagurus, 195. 


INDEX. 


Canis 
sladeni, 235, 244. 
vetulus, 235. 
Canthidium 
decoratum, 250. 
sladeni, 245, 249, 250, 
258. 


sp., 245. 

chalybeus, 245. 

edentulus, 245. 

histrio, 245. 
Capra 

hircus, 263. 
Carduelis 

carduelis, 3. 
Cariacus, 273. 

mexicanus, 262. 

rufus, 262. 
Carmarina, 182. 
Carminodoris, 359. 
Carpophaga 

whartoni, 316. 
Caryoborus 

sp., 248. 
Casella, 355, 357. 
Castnia 

dedalus, 260. 
Catablema, 171, 173. 


Cathartes, 386, 388, 389, | 


390, 391, 392. 
atratus, 387, 390. 
papa, 392. 
Cebus 

azare, 234, 

elegans, 234. 

libidinosus, 234. 
Centrinus 

sp:, 247. 
Centropristis 

brasiliensis, 61. 

dispilurus, 61, 62. 

subligarius, 61. 
Centropus 

sinensis, 1538, 154, 156. 
Cephaiophus 

abyssinicus, 194. 

griummt, 268. 
Ceraspis 

cornuta, 246. 
Ceratohyla 

bubalus, 115, 116. 
Ceratophrys 

cristiceps, 69. 
Ceratosoma, 355, 357, 
Cervulus 

muntjac, 2, 262. 
Cervus, 273. 

axis, 262. 

duvauceli, 8. 

manchuricus, 262. 


Cestoplana 
jiliformis, 110. 
rubrocineta, 110. 
unicolor, 3. 

Cheetogaster, 213. 

Chalcolepidius 
limbatus, 247. 

Chalcophana 
viridipennis, 249. 

Chaleophaps 
natalis, 316. 

Cheilospirura 
ophthalmica, 154, 156, 
siamensis, 154, 156. 

Chelethiops 
bibie, 832. 

Chelonarium 
ornatum, 245. 

Cheraphilus 
nanus, 29, 
trispinosus, 29. 

Chersydrus, 327. 

Chiloglanis 
brevibarbis, 333. 
deckenit, 333. 
modjensis, 332, 3883, 

3d4. 
niloticus, 333. 

Chirostoma, 60, 

Chlamys 
cistella, 248. 
hieroglyphica, 248. 
smaragdina, 248. 

Chordodes 
montoni, 152, 155. 
puncticutatus, 153. 
siamensis, 158. 

Chorinemus, 66. 

Chromodoris, 355, 356, 
357, 359. 

Circaétus, 157. 

Cireus, 157, 163. 


Clarias 
robecchit, 332. 
Clavipalpus 
tenuis, 246, 251, 
258. 
Clitellio 
arenarius, 212, 219. 
Codonium 
apiculum, 164, 165, 
166, 167, 168, 
192: 


codonophorum, 168. 
conicum, 168. 
gemmiferum, 168. 
princeps, 167, 168. 
pulchellum, 167, 168. 
Ceelosternus 
exornatus, 252. 
sturio, 247, 252, 258. 


Coendou 
boliviensis, 240. 
brandti, 240, 241. 
centralis, 240, 241. 
prehensilis, 240, 
9. 
Colaspis 
sp., 249. 
Jlavipes, 249. 
14-costata, 249. 
Colastus 
latus, 245. 
tonsus, 245. 
Copturus 
sp., 247. 
Corine 
rosario, 167. 
Cosmogramma 
Julvocineta, 54, 
patricia, 53. 
peruana, 53. 
unicincta, 53. 
Crangon 
allmanni, 28. 
echinulatus, 27, 28. 


fasciatus, 24, 25, 26, | 


27, 28, 29, 30. 
nanus, 24, 25, 27, 
30. 
norvegicus, 28. 
sculptus, 26. 
spinosus, 24, 28. 
trispinosus, 24, 25, 26, 
27, 28, 29, 30. 
Cua, 24, 25, 26, 
30. 


to 


8, 


Crimissa 
piceicollis, 249. 
Crocidura 
russula cypria, 344. 
whitakeri, 344. 
Cryptobranchus, 301. 
japonicus, 306, 307. 
maximus, 306. 
Cryptorhynchus 
sp., 247. 
Oubaia 
aphrodite, 187. 
Cyanea 
bougainvillii, 172. 
Cybium 
immaculatum, 67. 
Cyclocephala 
sp., 246. 
dimidiata, 257. 
laminata, 257. 
melanocephala, 257. 
ovulum, 246. 
araquayensis, 257. 
Cee 246, 
vincenti@, 257. 


‘INDEX. 


Dalaca 
metullus, 260. 
sladeni, 260. 
Damaliscus 
albifrons, 1. 
Dasyprocta 
acouchy, 241. 
azare, 241. 
Dasypus 
encoubert, 242. 
gilvipes, 242, 243. 
sexcinetus, 242, 243, 
Dasyurus 
albopunctatus, 198. 
Deinodrilus, 208. 
Deltochilum 
cupricolle, 246, 
fuscocupreum, 245. 
Dermestes 
vulpinus, 245, 
Desis 
kenyone, 316. 
Desmogramma 
decempustulata, 52. 
Fenestrata, 52. 
inclusa, 52. 
partita, d2. 
peruand, 51, 
stali, 51. 
Deuterocampta 
12-maculata, 249. 
flavidu, 51. 
luteola, 51. 
musicalis, 249, 255. 
nigrimana, 51, 
obscurimana, 51. 
puncticollis, 50. 
sceutellata, 50. 
sladene, 249, 
238. 
Diagramma 
cavifrons, 62. 
goeldii, 62. 
Diaulula, 360. 
Dicotyles, 262. 
Dictyodoris, 361. 
Didelphis 
paraguayensis, 243. 
pecilotis, 243. 
Dinematella 
cavosa, 169. 
Diopatra 
chilensis, 132. 
dorsalis, 131. 
Sragosa, 131. 
glutinatrix, 131. 
madeirensis, 132. 
magna, 131. 
neapolitana, 131, 182, 
144. 
pourtalesii, 131. 


250, 


395 


Diphyes 
bipartita, 189. 
chamissonis, 189. 
Diphyopsis, 189. 
Diporochzxta, 203. 
aquatica, 203, 204, 
205, 206, 226, 232. 
chathamensis, 227. 
intermedia, 203, 204, 
227, 
moroea, 2277. 
pellucida, 227. 
scolecotdea, 227. 
Dipsadomorphus 
dendrophilus, 146, 150, 
158, 155. 
Dipurena 
dolichogaster, 164, 169. 
Discodoris, 357, 359, 360. 
boholiensis, 354, 364. 
cerulescens, var. varie- 
gata, 304, 365. 
Discognathus 
blanfordii, 330, 351, 
334. 
chiarinii, 330, 333. 
dembeensis, 331. 
johnstonii, 331. 


makiensis, 330, 3el, 
33 


quadrimaculatus, 330, 
o3l. 


vinciquerre, 331. 
Disparoplana, gen. nov., 


103 


dubia, 103, 104, 113. 
Dolichotoma 

strigata, 249, 
Dorcacerus 


barbatus, 248. 
Dorcatherium, 262, 281. 
Dorcopsis 

macleayt, 196. 
Doris 

apiculata, 361. 

areolata, 3870. 

carinata, 361. 

maculata, 857. 

osseosa, 861. 

rusticana, 363. 

sordida, 380. 

spongtosa, 370. 

trastis, 3861. 

(Thordisa) villosa, 367. 
Doryphora 

eneofasciata, 89, 46. 

estuans, 42, 

amicta, 34. 

bahiaensis, 40, 41. 

bifasciata, 46, 47.. 

bistriquttata, 39. 


396 


Doryphora 
brevifasciata, 31. 
chalybeofasciata, xxii. 
chevrolati, 46. 
cyaneofasciata, 45. 
dejeant, 45. 
diversipes, 34. 
ecuadoriensis, 37. 
Ffasciatipennis, 47. 
fasciato-maculata, 44. 
flavimana, 35. 
flavoannulata, 41. 
flavofasciata, 46. 
flavozonata, 249. 
Sruhstor fert, 38. 
Sulgora, 36. 
gerstaeckeri, 40. 
histrionica, 38. 
honduraensis, 39. 
imitans, 41. 
instabilis, 37. 
Jucunda, 31. 
lateralis, 44. 
maculata, 47. 
nigrovarians, 32. 
nigroviolacea, 45. 
paraguayensis, 33. 
piceo-maculata, 45. 
prasind, 3). 
sanguinipennis, 42. 
seminigra, 35. 
semiviridis, 32. 
sexmaculata, 45. 
sexplagiata, 45. 
sexspilota, 36. 
signiceps, 32. 
spherica, 31. 
staudingert, 42. 
sternalis, 43. 
submetallica, 36. 
terminata, 40. 
vigintiplagiata, 44. 
vinula, 33. 
viridifasciata, 39. 
viridiornata, 38. 

Dromezus 
nove-hollandie, 327. 

Drymeus 
pecilus, 71. 

Dryotriorchis, 157, 161, | 
162, 163. 
spectabilis, 161. 

Dules 
auriga, 6). 
flaviventiris, 61. 


Echidna 

hystrix, 193. 
Echinodoris, 357, 
Echinorhynchus 


sp., 152, 155, 


INDEX. 


Echinorbynchus 
bufonis, 150, 155, 156. 
cinctus, 149. 
dipsadis, 149. 
heruca, 150. 
heterorhynchus, 149. 
megacephalus, 149. 
oligacanthoides, 149. 
oligacanthus, 149. 
patani, 149, 150, 155, 

156. 
tigrine, 151, 152, 155. 
xenopeltidis, 151, 155, 
156. 

Elasmopalpus 
pyrrhochrellus, 260. 

Elephas 
africanus, 264. 
indicus, 264. 

Elodea, 335. 

Elythrosphera 
cupreata, 59. 

Emarginea 
gammophora, 259, 

Enchytreeus, 204. 
adriaticus, 221. 
albidus, 221. 
hyalinus, 221. 
mobi, 221. 
simulans, 205, 206, 

2119) 232) 

Enhydrina 

valakadien, 147, 148, 
155, 156. 

Enhydrus 
tibialis, 245, 

Epicauda 
sp., 247. 
atomaria, 247. 
strigata, 247. 

Epilachna 
cacica, 245. 

Epitragus 
sp., 247 

Equula, 63. 

Equus 
asinus, 263. 
caballus, 263. 
grevyi, 193. 
quagga burchelli, 196. 

Hrebus 
agarista, 259. 

Erinaceus 
auritus, 348, 344. 
europeus, 344. 

Erythropus, 157, 159. 
vespertinus, 159. 

Eudoxia 
eschscholtzti, 189, 


| Eumolpus 


elavipalpus, 249. 


t 


Eumolpus 
surinamensis, 249. 
Eunice 
teretiuscula, 136. 
Eupemphix 
natterert, 69. 
Huphoria 
lurida, 246. 
Huryscopa 
proxima, 248. 
Kustylochus, 107. 


FHaleo, 157, 18; 19} 
160, 162. 
esalon, 157. 
biarmicus, 157, 158. 
candicans, 157. 
Seldeggi, 157, 158. 
lanarius, 157. 
peregrinus, 157. 
sacer, 157, 158. 
subbuteo, 157. 
Felis 
wiedit, 235. 
Filaria 
corynodes, 154, 156. 
longicirrata, 154, 156. 
sciuri, 154, 156. 
Fornarina, gen. nov., 336. 
phillipsi, 336. 
Fracassa, 357, 359, 361. 
Fridericia 
antarctica, 204. 
galba, 204. 


Galeopithecus 

volans, 154, 156. 
Galera 

barbara, 236. 
Galerucella 

sp., 249. 
Galictis 

barbara, 315. 
Gargamella, 360. 
Gazella 

grant, 119, 120. 

— brightii, 120. 

— robertsi, 119, 120, 

121. 

— smithii, 120. 

muscatensis, 317. 
Gazza, 63. 
Genetta 

genetta, 122, 1238, 124. 

plesictoides, 121, 122, 

128, 124. 

senegalensis, 123. 
Genyatremus 

cavifrons, 62. 

luteus, 62. 


Genypterus 
brasiliensis, 68. 
Geranoaétus, 157. 
Gerres, 63. 
axillaris, 64, 65. 
brasilianus, 64, 65. 
brevimanus, 64. 
embryx, 64, 69. 
lineatus, 64, 65. 
mecxicanus, 64. 
patao, 63, 64, 65. 
plumieri, 64. 
Geryonia, 182. 
Glossina 
morsitans, 316. 
palpalis, 316, 317. 
Glympis 
sp., 209. 
habitalis, 259. 
Giyphodes 
jegia, 260. 
Gobio 
hirticeps, 330. 


quadrimaculatus, 330. 


Gonionema, 186. 

Gonionemoides, 187. 
geophila, 169, 187. 

Gonionemus, 191. 
agassizti, 165, 1885, 

188, 192. 

murbachit, 187, 188. 
suavensis, 137, 188. 


vertens, 165, 183, 185, 


186, 187, 188. 


(Cubaia) aphrodite, 187. 


Gonynema, 186. 
Gordius 

parone, 152, 153. 
Gossea, 187. 
Griburius 

pretectatus, 248. 
Gromphas 

inermis, 246. 
Gymnetis 

sp., 246. 

liturata, 246. 
Gypaétus, 157, 163. 

barbatus, 386. 


Gypagus, 157, 386, 388, — 


389, 390, 391, 392. 
papa, 387, 388, 391. 


Gypohierax, 157, 163, 


386. 
angolensis, 162. 
Gyps, 157, 160. 
rueppelli, 160. 


Halgerda, 357, 360, 361. 


formosa, 358, 373. 
wasinensis, 3909, 


873, 385. 


BOT, | 


INDEX. 


Halgerda 
DB) 


willeyi, 355, 872, 373, 


385. 
Haliaétus, 157. 
Halla, 357, 358, 359. 
Hamadryas 
ophiophagus, 322, 
Hammaticberus 
lacordairei, 248. 
Haplochilus 
antinori, 333. 


Haploplana, gen. noy., 
109 


elioti, 109, 113. 
Haplotaxis, 204. 
gordioides, 223, 225. 


heterogyne, 205, 206, 


223, 224. 
smithi, 204, 223. 
Heliopetes 
sp., 209. 
arsalte, 259. 


| Helotarsus, 157. 


Hemiderma 

perspicilatum, 254. 
Henlea 

ventriculosa, 204. 
Herpestes, 124. 
Herpetotheres, 157, 

158. 

cachinnans, 158. 
Hesperodrilus, 203. 

albus, 208. 
Heterakis 

circularis, 153, 156. 

rimula, 153, 156. 
Heterocephalus 

ansorget, 336. 

glaber, 336, 337. 

phallipsi, 336. 


Hexabranchus, 379, 384. 


Hieracidea, 157, 158. 
berigora, 158, 162. 
Hierofalco 
candicans, 157. 
Hilipus 
sp., 247. 
nevulus, 246. 
validus, 247. 


Hinulia, 125, 126, 128. 


Hippocrene 
bougainvillii, 172. 
mertensiz, 165, 171, 

172. 

Hippopotamus 
amphibius, 262. 
minutus, 122. 

Histiotus 
velatus, 234. 

Homalodes 
brasilianus, 245. 


397 


Homalonotus 

validus, 246. 
Homoiodoris, 360, 363. 
Homolepida 

englishi, 128. 
Hoplocercus 

spinosus, 70. 
Hoplodoris, 357, 359. 
Hoplopactus 

lateralis, 

258. 

Hoploptera 

sp., 247. 
Hyalinvecia, 131. 
Hydrocyon 

Jorskalii, 329. 
Hydrophis, $27. 
Hydrus, 326. 


246, 251, 


| Hyla 


goeldit, 116. 
nasica, 69. 
senicula, 69. 
venulosa, 69. 
Hylodes 
gollmeri, 69. 
Hyomys, gen. noy., 198. 
meeki, 198, 199, 
Hypsioma 
sp, 248. 


Jasciata, 248. 


Ilyodrilus 


coccineus, 212. 


| Iphimeis 


Sulvipes, 249. 
Tsonychus 

griseus, 246. 
Kentrodoris, 389, 360, 

361, 
rubescens, 360, 374, 389. 


Labeo 
cylindricus, 330. 
gregorit, 329. 
neumannt, 829, 334. 
Lachesis 
wagleri, 194. 
Lachnosterna 
Sulvipennis, 246. 
Lamellidoris, 388. 
Lamprosoma 
chameleon, 248. 
viride, 248. 
Lamprospherus 
seintillaris, 249. 
thoracicus, 249. 
Laodice, 177, 184, 190. 
Laodicea 
cellularia, 172. 


| Lepophidium, 67. 


398 


Leptinotarsa 

paraguayensis, 58. 
Leptodactylus 

typhonius, 69. 
Leptomys 

elegans, 198. 
Leptoplana, 100, 108. 
Lepus 

ereticus, B47. 

cyprius, 347. 

timidus, 348. 
Leucopheeus 

scoresbit, 315. 
Leucopternis, 157, 163. 
Leucothyreus 

sp., 246. 

pruinosus, 246. 
Limnodrilus 

sp., 205, 217, 218. 

claparedianus, 215, 217, 


dugesi, 217. 
gotot, 219. 
lucasi, 205, 206, 207, 
216, 232. 
novezelandi@, 218. 
vejdovskyanus, 205, 
207, 218, 216, 217, 
231. 
Liocephalus 
caducus, 70. 
Liogenys 
sp., 246. 
Liognathus, 63. 
Liolepis 
bellit, 154, 156. 
Liolepisma, 127. 
Liophis 
almadensis, 70. 
Lirope, 182. 
Lissonema 
rotundata, 154, 156. 
Lizzia 
blondina, 168. 
Lonchoglossa 
eaudifera, 234. 
Lophius 
piscatorius, 68. 
Lophonocerus 
barbicornis, 248. 
Lucidota 
sp., 247. 
Lutianus 
luteus, 62. 
Lycena 
cassius, 258. 
Lycalopex 
chiloensis, 236. 
Ffulvicaudus, 236. 
Lygosoma 
annectens, 127. 


INDEX. 


Lygosoma 
concinnatum, 126. 
granulatum, 126, 129. 
lorie, 128. 
milnense, 125, 126, 129. 
muelleri, 128. 
pratti, 128, 129. 
pulchellum, 127. 
pulchrum, 127, 129. 
simum, 126. 
Lysidice 
collaris, 143. 
Lystronychus 
ceruleus, 247. 
latipennis, 247. 


Mabuia 
aurata, 70. 


frenata, 70. 


Macduftia 
bonhardi, 136. 
Maeraspis 


brasiliensis, 256, 257. 
cribratus, 257. 
dichrous, 257. 
pantochloris, 246. 
Macrosoma, 256. 
Mallodon 
spinibarbe, 248. 
Mallomys 
rothschildi, 199. 
Maoridrilus, 203. 
Margellium 
octopunctatum, 168. 
Marmosa 
cinerea, 243, 244. 
constantieg, 248, 244. 
regina, 244. 
Marphysa 
acicularum, 136, 137. 
adenensis, 136. 
belli, 180, 136. 
bonhardi, \36. 
corallina, 136, 137. 
Jallax, 136, 137. 
Jurcellata, 136,141,142, 
148, 144. 
goodsiri, 136. 
Januarit, 187, 148. 
macintoshi, 136, 137, 
138, 140, 141, 144. 
mossambica, 130, 136, 
139, 144. 
nove-hollandie, 
139. 
parishi, 137, 148. 
regalis, 136, 1387. 
saxicola, 186, 137. 
simplex, 1386, 140, 141, 
142, 144, 
stragulum, 136. 


136, 


Marphysa 
striata, 136, 137. 
Maypa, 256. 
Mazama 
rufa, 242. 
Medusa 
campanulata, 177. 
digitale, \71. 
Megaderus 
stigma, 248. 
Megalobatrachus, 
309, 313, 315. 
japonicus, 298. 
Megalophrys 
montana, 154, 155. 
Megalostomis 
obesa, 248. 
Megascolex 
antarcticus, 204. 
Melicertum, 186. 
penicillatum, 176, 177. 
Menidia, 60. 
Menobranchus, 305, 307, 
309. 
lateralis, 304. 
Menopoma, 301, 303. 
Merocausta 
vinosa, 259. 
Mesonema 
victoria, 165, 180, 181, 
182, 192. 
(Zygodactyla) cerules- 
cens, 182 
Mesonemella 
bairdit, 183. 
eyaneum, 182, 188. 
eurystoma, 185. 
Metallactus 
sp., 248. 
kollart, 248. 
Miamira, 355. 
Micralestes 
acutidens, 329. 
Micronycteris 
megalotis, 234. 
Milvago, 157, 158, 160. 
chimachima, 158. 
chimango, 158, 162. 
Milvus, 157, 161, 163. 
ictinus, 161. 
Miniopterus 
schreibersi, 343. 
Molge 
blasit, 337. 
marmorata X cristata, 


307, 


337. 
Molpadia, 147, 155. 
Monocrepidius 
sp., 247. 
Monophyes 
primordialis, 182, 


Mormyrus 
kannume, 328. 
Morphuus, 157, 163. 
Morphoides 
sp., 245. 
Moschus 
moschiferus, 262. 
Muggivea 
kochit, 165, 189. 
pyramidalis, 190. 
Murena 
helena, 60. 
insularum, 60. 
Mus 
alexandrinus, 346. 
armandvillez, 199. 
decumanus, 78, 81, 336, 
346. 
musculus, 72, 78, ‘7d, 
85, 96, 346. 
— gentilis, 346. 
nudo-plicatus, 336, 
ratius, 345, 346. 
sylvaticus, 72, 73, 7A, 
75. 
tectorum, 845. 


Mylacrodon, gen. nov., | 
62. 
goeldii, 63, 68. 
Myotis 
myotis, 341, 342. 
Myrmecophaga 


tridactyla, 242. 


Naia, 326. 

tripudians, 319. 
Nais 

elinguis, 213. 
Nasua 

nasua, 236. 
Naupactus 

leucogaster, 258. 

nodicollis, 258. 

perpastus, 246, 258. 

tuberculatus, 257. 
Neacomys 

spinosus, 239, 240. 

— amenus, 239. 
Nectomys 

garleppi, 238. 

squamipes, 238, 259. 

— mattensis, 238. 
Neleus 

punctiger, 245. 
Neobola 

bottegt, 332. 
Neodrilus, 203. 
Nisaétus, 157, 159, 160, 

161, 168. 
fasciatus, 159, 386. 


| CXdopeza 
Okapia 


INDEX. 


Norops 
auratus, 69. 
sladenie, 69. 
Notiodrilus, 203. 
Notoscolex, 204. 
(Hypogzon) orthostt- 
chon, 203. 
Nototrema 
cornutum, 116. 
oviferum, 116. 
testudineum, 116. 
Nyetipithecus 
azare, 234. 
Nyctobates 
gigas, 247. 
variolosa, 247. 


Obelia 
polystyla, 190. 
Oceania, 182. 
gregaria, 179. 
languida, 179, 180. 
Octochetus, 203. 
Octoplon 
flavopictum, 248. 
Odontochila 
discrepans, 245, 
CGidionychis 
12-notata, 249. 
nigropunctata, 249. 
optima, 249. 


pogonocheroides, 248. 


erichsoni, 338, 340. 
Oligoplites, 66. 
Ommatoplana, gen. nov., | 

111, 115. | 

tuberculata, 111, 118. 
Oncideres 

amputator, 254. 

Ffasciata, 254. 

maculosa, 254. 

sladent, 248, 254, 258. 
Ontherus 

sp., 249. 

sulcator, 245. . 
Onuphis, 131. 

holobranchiata, 185 

144. 
Opiiophagus 
bungarus, 319, 320, 
321, 328, 326, 327. 
Orchistoma, 181. 
Orodoris, 355. 
Orvx 
beisa, 336. 
Oryzomys 
laticeps, 239, 


Otosaurus, 126, 


399 


Ovibos 
moschatus, 262. 


| Ovis 


aries, 263. 
musimon, 263. 
ophion, 348. 
steatopyga, 263. 
Oxydia 
sp., 259. 
Oxymerus 
basalis, 248. 
Oxysoma 
tuberculatum, 154, 155. 
Oxyurus 
coronata, 154, 156. 
siamensis, 154, 156. 


Paludicola 

signifer, 69, 
Pandalina 

brevirostris, 24. 
Pandion, 157. 
Pantomorus 

sp., 246. 
Papilio 

arsalte, 259. 

cassius, 298. 
Paraplanocera 

aurora, 102, 108, 118. 

langt, 103. 
Paroaria 

albigularis, 350. 

capitata, &50, 351. 

cervicalis, 350, 391. 

cucullata, 350, 351. 

dentata, 350. 

gularis, 351. 
Paronuphis, 131. 
Pavoncella 

pugnaxr, 3. 
Pectantis, 187. 
Pelidnota 

eruginosd, 246. 

swmptuosa, 246, 
Pelodrilus 

ignatovi, 225. 

violaceus, 225, 
Peltodoris, 360. 

angulata, 354, 365. 

aurea, 354, 366. 
Pentaprion, 63. 
Pentastoma 

montliforme, 154. 
Perameles 

longicauda, 201, 202, 

ornata, 201. 
Pericopis 

sacrifica, 259. 
Peristedion 

altipinnis, 65, 68, 


400 


Petelodoris, 357, 367. 
Phalanger 
carmelite, 198. 
Phanzus 
kirbyt, 246. 
mimas, 246. 
paleno, 246. 
palemo, 246. 
Phasianus 
hagenbeckt, 2. 
Phelsuma 
madagascariense, i, 
Pheretima, 204. 
Phalidium 
buskianuwin, 180. 
cymhaloideum, 180. 
flavidulum, 180. 
gregarium, 165, 179, 
180, 192. 
languidum, 165, 179, 
180. 
temporarium, 180. 
variabile, 180. 
Phialodoris, 369. 
Phreodrilus, 203, 207, 
208. 
lacustris, 204, 205, 207. 
mauianus, 204, 205, 
206, 208. 
subterraneus, 209. 
Phylloplana, gen. nov., 
100, 107. 
lactea, 107, 118. 
Physalia, 170. 
Pinotus 
carbonarius, 246. 
eremita, 250. 
fissus, 246. 
nisus, 246. 
roberti, 246, 250. 
subeneus, 246. 
Pipistrellus 
kuhlii, 342. 
Plagiocheta, 203, 204. 
Planocera 
armata, 100, 101. 
crosslandi, 100, 101, 
elliptica, 107. 
folium, 102. 
graffi, 102. 
nebulosa, 102. 
pelagica, 102. 
pellucida, 101, 102. 
reticulata, 102. 
simrothi, 101, 102. 
Platydoris, 357, 360, 361, 
364, 380, 381, 382, 
383, 384, 
argo, 358. 
arrogans, 376. 
cruenta, 376. 


Platydoris 


Platyomus 


Platyptera, 125. 
Platyurus, 327. 
Plesictis 


Plutellus, 203. 


Polyboroides, 157, 160. 
Polyborus, 157, 158, 159, 
160 


Polycanna 
Polychrus 


Polyorchis, 184. 


Pontodrilus, 203. 
Pontophilus 


Poria 
Porocephalus 
Poropleura 
Prepusa 
Prionotus 
Pristipoma 


Proboscidactyla 


Procavia 


INDEX. 


ellioti, 355, 377. | 

eurychlamys, 355, 374, | 
SD BM To 

formosa, 355, 376; 877. 

incerta, 355, 378. 

papillata, 355, 379, 
382. 

pulchra, 355, 359, 377. 

scabra, 355, 375, 377. 

variegata, 359, 361. 

vicina, 399, 361, 379. 


argyreus, 246. 
transversesiynatus, 246, 


croizeti, 122, 123, 124. 


lacustris, 208, 204, 
205, 206, 228, 232. 


typicus, 160. 


brasiliensis, 159. 
FSungina, 183. 
acutirostris, 70. 


campanulatus, 177. 

minuta, 165, 174, 176, 
192. 

penicillata, 169, 
176, 177. 


175, 


bermudensis, 228. 
insularis, 228. 


norvegicus, 29, 
sptnosus, 29. 


sp., 245. 
moniliformis, 154, 155. 


cuprea, 248. 


distigma, 245. 
beanit, 65. 
cavifrons, 62. 


brevicirrata, 165, 178. 
flavicirrata, 169, 178. 


sp., 264. 


capensis, 264. 


Procayvia 
dorsalis, 264. 
Proechimys 
bolivianus, 240. 
longicaudatus, 240. 
Prosicela 
flavipennis, 5d. 
inornata, 5d. 
maculata, 5d. 
simplicipennis, 55. 
Prostenus 
cyanipes, 247. 
sladent, 247, 
258. 
Pseudochirus 
corinne, 198. 
Pyrophorus 
sp., 247. 
janus, 247. 
noctilucus, 
Python 
reticulatus, 
155. 


243 


Ale 
153, 154, 


Raia 

cyclophora, 60. 
Rana 

tigrina,. 151, 162, 155. 
Rhachidion 

obesum, 248. 
Rhamphobrachium, 131. 
Rhegmatodes 

tenuis, 182. 
Rhiacichthys 

aspro, 125. 

nove-guinee, 124, 125. 

sinensis, 125. 
Rhigus 

latruncularius, 246. 
Rhinoceros 

sp., 204. 

bicornis, 195. 

simus, 194, 195, 316. 

sumatrensis, 264. 
Rhinochenus 

sp., 247. 

reichet, 247. 

stigma, 247. 
Rhinolophus 

blasii, 341, 342. 

ferrum-equinum, 342. 

hipposideros, 342. 
Rhipidomys 

benevoleus, 237. 

macrurus-latimanus, 

237. 

pheotis, 237. 

roberti, 237, 238. 
Rhbizodrilus 

lacteus, 219. 


Rhizodrilus 
(Vermiculus) pilosus, 
219 


Rhynchobothrius, 147. 
Rhynchophorus 
palmarum, 247. 
Rostanga, 357. 
Roussettus 
egyptiacus, 341, 342. 


Sabinea, 28. 
Saecopleura 
sp., 260. 
Sarcorhamphus, 157, 386, 
388, 389, 390, 391, 
392. 
equatorialis, 390. 
Sarsia, 169. 
brachygaster, 166. 
eximia, 190. 
prolifera, 168. 
pulchella, 167. 
rosario, 190. 
(Codonium) pulchella, 
166. 


Sciurus 
caniceps, 154, 156. 
langsdor-ffi, 237, 240. 
Sclerodoris, gen. nov., 
355, 360, 361, 380, 
384. 
coriacea, 355, 388, 385. 
minor, 350, 382. 
osseosa, 355, 380. 
rubra, 355, 382. 
tuberculata, 350, 381, 
382. 
Seolopax 
rusticula, 352. 
Scombroides 
altus, 66, 67. 
mundus, 67. 
palometa, 66. 
saliens, 66, 67. 
saurus, 66. 
Scombromorus 
cavalla, 67. 
immaculatus, 67. 
maculatus, 67. 
regalis, 67. 
Scotinus 
sp., 247. 
Semnopithecus 
albocinereus, 154, 156. 
Sepia 
officinalis, 148. 
Sericoides, 256. 
Serpentarius, 389. 
Serranus 
auriga, 61, 62. 


Proc. Zoou. Soc.—1903, Vou. IT, No. XX VII. 


INDEX. 


| Serranus 


dispilurus, OL. 

jlaviventris, 61. 

subligarius, 61. 
Solea 

Jonsecencis, 68. 
Speothos 

venaticus, 236. 
Spherion 

sladeni, 248, 253, 258. 


Spherodoris, 395, 357, 


358. 
levis, 370. 
Spilornis, 157. 
Spizaétus, 157, 160, 163. 
ceylonensis, 160. 
orientalis, 160. 
Stalachtis 
phlegetonia, 258. 
Staphylinus 
sp., 240. 
Staurodoris, 
360, 874. 
bicolor, 358. 
calua, 354, 356, 364. 
depressa, 354, 363. 
Januarti, 362. 
pecten, 36. 


pseudoverrucosa, 306, 
357, 364. 

pustulosa, 364. + 

verrucosa, 306, 


308. 
Staurophora, 175. 
mertenstt, 174, 180. 

Staurostoma, 173. 
Steirastoma 
stellio, 248. 
Stenoplesictis, 122, 
Stilodes 
ecuadoriensis, 49. 
flavilabrum, 48. 
flavofasciata, 47. 
Sruhstorferi, 48. 
geometre, 49. 
transversofasciata, 48. 
Strongylium 
sp., 247. 
azureum, 247. 
Struthio 
camelus, var. Massaicus, 
198. 
Stylaria 
lacustris, 213. 
Stylochoplana, 107. 
Stylochus 
argus, 107. 
conglomeratus, 107. 
crassus, 107. 
Frontalis, 107. 
limosus, 107. 


307, 


356, 357, | 


401 


Stylochus 
littoralis, 107. 
neapolitanus, 106, 107. 
pilidium, 107. 
plessisiz, 107. 
sargassicola, 107. 
suesensis, 106, 107. 
zanzibaricus, 105, 106, 
107. 
zebra, 107. 
Sus 
porcus, 262. 
serofa, 262. 
Susica 
quadrata, 259. 
Sylvilagus 
brasiliensis, 241. 
minensis, 241, 242. 
— chapade, 241, 242. 
paraguensis, 241, 242. 
Syncoryne 
occidentalis, 167. 
rosario, 167. 
Syndictyon 
angulatum, 164, 168, 
169. 
reticulatum, 169. 


Tamandua 
tetradactyla, 242. 

Tapirus 
americanus, 263. 
éndicus, 263. 


| Tatu 


megalolepis, 243. 
novemcinctus, 243. 
septemcinctus, 243. 


Taupodrilus, gen. noyv., 


203, 205, 209. 
simplex, 205, 206, 207, 
209, 231. 
Tauroma 
sp., 249. 
Tayassu 
albirostris, 242. 
tajacu, 242. 
Tellena 
varians, 248. 
Testudo 
elephantina, 193. 
mydas, 145, 148. 


Tetraonyx 
sexguitatus, 247. 
Tetrarhynchus 
Soy Wass, ide, 11595), 
156. 


erinaceus, 148. 
holothurie, 146, 
158, 156. 
lingualis, 147. 
27 


147; 


402 
Thaumantias | 
cellularia, 165, 172, 
192. 
pilosella, 173, 174. 
Thordisa, 356, 357, 360, | 
369. 
crosslandi, 854, 359, 


360, 368, 385, 
maculigera, 3867. 
stellata, 354, 368. 
villosa, 354, 365, 365, | 

367, 368, 385. 

Thorunna, 357. 
Thrasaétus, 157. 
Thylacinus, 116. 
Tiara 

pileata, 170, 171. 
Tilapia 

nilotica, 339. 
Tinnunculus, 157, 159. 

alaudarius, 159, 162. 
Toxicocalamus 

longissimus, 129. 

stanleyanus, 128, 129. 
Tragelaphus 

scriptus, 263. 
Tragulus 

javanicus, 262. 

kanchil, 262. 

stanleyanus, 262. 
Trichophorus 

electus, 248, 254, 258. 

interrogationts, 254. 

7-signatus, 254. 
Trippa, 357, 358, 559, 

360, 361, 381. 
areolata, 355, 369, 381, 

382, 3838. 


monsoni, 355, 365, 371, 
385. 


INDEX. 


Prippa 
(Phlegmodoris) mephi- 
tica, 370. 


| Tropidurus 


spinulosus, 70. 
Trypanosoma 
brucet, 317. 
castellanii, 316. 
Tubifex, 204. 
sp., 205, 206, 207, 219. 
blanchardi, 219. 
rivulorum, 209, 219. 
(Heterocheeta)  costa- 
tus, 219. 
(Psammoryctes) plica- 
tus, 219. 
(—) velutinus, 219. 
(Spirosperma) ferox, 
DANS). 
Turnix 
taigoor, 154, 156. 
Turris 
breviconis, 165, 
192. 
digitalis, 171. 
neglecta, 171. 
Tyrinna, 350. 


170, 


Uloma 

sp., 247. 

retusa, 247. 
Ungalia, 327. 
Urodera 

Jamiliaris, 248. 
Uromys 

papuanus, 200. 

validus, 200. 
Ursus 

maritimus, 348. 
Urubitinga, 157, 163. 


THE END. 


Vampyrops 
lineatus, 235. 
Vespertilio 
murinus, d42. 
Viverra 
karnuliensis, 124. 
Vulpes 
eyyptiaca, 345, 
melanogaster, 340. 
persica, 345. 
vulpes, 345. 
Vultur, 157, 160, 163, 
386, 387, 388. 
auricularis, 160, 386, 
387. 
calvus, 160, 886, 387. 


Xenopeltis 
unicolor, 151, 155. 
Xestia 
polita, 248. 
suturalis, 248. 


Zophobas 
opacus, 247. 
Zostera, 3867, 382. 
Zygodactyla 
cyanea, 182, 183. 
grenlandica, 182. 
Zy gogramma 
argentinensis, 56. 
brasiliensis, 57. 
burmeisteri, 57. 
ceruleo-vittata, 58. 
curvato-lineata, 56. 
jflavolorata, 57. 
interstitialis, 55. 
myops, 57. 
5-virgata, 58. 


THE ZOOLOGICAL SOCIETY OF LONDON. 


Tu1s Society was founded in 1826 by Sir Sramrorp Rarvtes, 


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Pror. Gzorert B. Howzs, D.Sc., 
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Lr.-Cot. L. Howarp Irsy. 

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2 


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3 Hanover Square, London, W., 
October, 1908. 
MEETINGS 
OF THE 
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FOR 


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1903. 
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« 


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LIST OF THE PUBLICATIONS 


OF THE 


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| October, 1908. ] 


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relies Salle M O29) ui Phe ee LSGE—6O) Epa ome Olas 15 0 O 
ny AU si TO Nero Goon (URES); oon 1D) dO, 13 12 0 

oy WOOL, af iy don bie ieaaga 1G) sl Gc 12: iO 
wp | LDS af CO Khoo Cle) coco IA I o. 1G 20 
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SS Opie ean ciercron een a cicheneaneate 9s. LS es 2 36s. sooo Ghee 
dete W fies stats puter Ue Hae 9s. MD Sy hte eee 36s. nooo “hs 
Wststs Goo oc Aieveradeteletonsualictoersisliite 9s. UD SNe 36s, coco akSs 
SSO) ree Hele vrs nraraine ona are iaae te 9s. spate LS s ee bret, 36s. sooo “kek 
SIO: tecture ea orca eae 9s. UE 2 Gua Pa tie ares OS) pooo abst, 
Index, 1881-1890 ........ caun 465 Oban | kh 
TSO tev Aston suapewetayalene cea reuoraetoieue Cpe Mevcelave: site anton aicna ebenoesdevenene 36s. saan (Gbek, 
OS rc ee reer tener eRe TENCE CPRE RCH OPAH CHET Sasol hinlin al cucatranaenevel aiitevecenette oG6s. OS: 
NSO SM rca a reretrnal stein miei er ere eRe TENG oe. Ula e ave uanatavavmusiste eine 36s. sboo) Ges 
SO ies siecerar eta ale rene AR Re eR Mee ekat snoueliallg) adel etharep over aeneets 36s, noag Gkekh 
US By FRO O Oe OO COO AG-CICiO OLEAN EMER RIP an Ee 36s. sooo, 2s 
MS OGM eae eie ctencraiar crave hen uctavenee tentials. oc gce-are, atbleseconas eusuatetane 36s. das AOS: 
A SS Zaeaietcien creche Mererere er ata Sea ee eT nee Nick. 6 ei cre aict ale Mabe latere 5G Clos b6o0  ahekh 
SOS re Cr sccrevaerareveversen’s She reanWetensteeveual dy si x: seus cum Sueueherateveltever's 36s. sooo Che 
SOO eee trem everstcere Tonsil ceareroe RONEN crit aera auc latobelesananeioten exe 36s. sooo GES 
DUC TOLO ies TMs An en Gid in ea ric aiet  eml a by AHL BR i OOSe ASS: 
index wi SOI NS00 Mwai ae. (th sooo 8S 


PROCEEDINGS or tos GENERAL MEETINGS ror SCIENTIFIC 
BUSINESS or tuz ZOOLOGICAL SOCIETY OF LONDON. 
8vo. 5 vols. 


Price to Price to the 
Fellows. Public. 
113)0) a () One Gennes cnc UBS. caateares 24s, 
25 EO eR PCE CaM Sh Us TOMB E GA ccc Tt ee 24s. 
C028 Ai Be POR MERE HHO RRA Arid ao Tesyen een 24s, 
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Re Th aptadinlt cco, Deke se oad nce ueee sue aaa ee On 12s 


LISTS OF THE ANIMALS IN THE SOCIETY’S GARDENS. 


List of the Vertebrated Animals now or lately Living in the Gardens 
of the Zoological Society of London, (Highth Kdition.) 8vo. 
1883. Price 3s. 6d. 

List of the Vertebrated Animals now or lately Living in the Gardens 
of the Zoological Society of London. (Ninth Edition.) 8vo. 
1896. Price 4s. 6d. 


Catalogue of the Library of the Zoological Society of London. 
(Fifth Edition.) Svo. 1902. Price 6s. 


These publications may be obtained at the Socrery’s OrricF 
(3 Hanover Square, W.), at Messrs. Loncmans (Paternoster low, 
EL.C.), or through any bookseller. 


Mth ZOOLOCIO RE RECORD: 


070400 — 


‘HE object of the Zoonogrcat Recorp is to give, by means of an 
annual Volume, complete lists of the Works and Publications 
relating to Zoology in all its branches that have appeared during 
the year preceding the issue of the Volume, together with full 
information as to the points they deal with, arranged in such a 
manner as to serve as an Index to the literature of Zoology in all 
parts of the globe, and thus vo form a repertory that will retain its 
value for the Student in future years. 


The ‘ Zoological Record’ is published by the Zoological Society 
of London at the price of 30s. per volume. But all Members of the 
Society have the privilege of receiving it, including the cost of 
delivery (within the United Kingdom), at a subscription price 
of 20s. per annum. This Subscription is due on the Ist of July in 
every year, and the privilege of Subscription is forfeited unless the 
amount be paid before the 1st of December following. 


The Zoological Society, having purchased the entire stock of 
the ‘Zoological Record,’ is able to supply complete scts. The 
thirty-seven Volumes to the end of the nineteenth century, and the 
Index-Volume (1880-1900) in addition, will be supplied for £15 
net (or without the Index-Volume, for £14 10s. net). Volumes of 
any single year (exclusive of the last five volumes and Vol. 6) can 
likewise be supplied at 10s. per volume net. 


The price of the Index Zoologicus (Index-Volume 1880-1900) 
is 20s., to Fellows 18s. 


Members of the Society wishing to subscribe to the ‘ Record’ 
are requested to apply at this office for a Form, to be returned 
when filled up and signed by the subscriber. In order to facilitate 
the payment of the subscription, a Banker’s Order Form is also 
furnished to those who prefer that mode of payment. This order, 
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Learned Societies and Institutions and members of the former 
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‘Record’ on the same conditions as are accorded to Members of 
the Zoological Society. 

Beginning with Vol. xxxix., the divisions of the ‘ Zoological 
Record’ may be obtained separately, as shown in the subjoined 


List. 
b 


SEPARATE DIVISIONS OF THE ZOOLOGICAL RECORD. 


At present each Volume of the ZoorocicaL Record consists of 
20 separately paged Divisions. Commencing with Volt xoccen tor 
1902, these Divisions may be obtained separately, in paper covers, 
stitched and lettered. 


The following are the Divisions and their net prices, viz. : 


M7) 


NO en NO N22 Ll NO NO PY 2D NOD QD 


. List of abbreviations of journals, ete. .. 
. General Subjects .. 

. Mammalia 

. Aves ROT Wee 
. Reptilia and Batrachia.. 
Pisces 

. Lunicata 

. Moilusea 

. Brachiopoda .. 

. Bryozoa 

. Crustacea 

2, Arachnida 


. Myriopoda 


OMIA ME Wd 


| 
Ww hore © 


po 


) Insecta <. 

. Echinoderma 

. Vermes .. 

. Coelenterata .. 

18. Spongise 

19. Protozoa wie, softs. ke Sy ay Shera seca 
20. Index of new names of genera and subgenera 


ft pS eH 
“Io OV 


bo bo FE ww Oo 
SOSORSASCASASSSOSCaAZSCaaca!l 


bo 


On receipt of the price any Division will be forwarded as soon 
as ready. 


These separate Divisions can be obtained from the Zoological 
Society, 3 Hanover Square, London, and also from the following 
Agents: Friedliinder & Sohn, 11 Carlstrasse, Berlin ; and Librairie 
A. Hermann, 6 rue Sorbonne, Paris. Cheques and Post-Office 
Orders should be made payable to “The Zoological Society,” and 
crossed ‘* Drummond’s,” 


P. CHALMERS MITCHELL, M.A., D.Sc., 


Secretary. 
October, 1903. 
ZooLocican Society or Lonpon, 
3 Hanover Square, W. 


LIST OF VOLUMES or rue ‘ZOOLOGICAL RECORD.’ 


The Record of Zoological Literature, 1864-1868. Vols. 1.-v. 
Edited by Atsert C. L. G. Giwrmer, M.A., M.D., Ph.D., F.Z.8., &c. 
Price 10s. each Volume net. 


The Record of Zoological Literature, 1869. Volume Sixth. 
Edited by Atsert C. L. G. Gtnrner, M.A., M.D., Ph.D., F.R.S., 
F.Z.8S., &c. London, 1870. Price 30s. 


The Zoological Record for 1870-1872, Vols. vir... Edited 
by Atrrep Newron, M.A., F.R.S., F.L.S., V.P.Z.5., &c. Price 10s. 
each Volume net. 


The Zoological Record for 1873-1883, Vols. x.-xx. Edited by 
Epwarp Carnpwe tt Rye, F.Z.8., M.E.S. Price 10s. each Volume net. 


The Zoological Record for 1884, 1885, Vols. xxt., xx1r. Edited 
by F. Jerrrey Bett, M.A. Price 10s. each Volume net. 


The Zoological Record for 1886-1890, Vols. xx11.—xxvui. 
Edited by Franx E. Bepparp, M.A., F.Z.S. Price 10s. each 
Volume net. 


The Zoological Record for 1891-1896, Vols. xxvit1—xxxi11. 
Edited by D. Suarp, M.A., F.R.S., F.Z.8., &. Price 10s. each 
Volume net. 


The Zoological Record, Volume the Thirty-fourth; being 
Records of Zoological Literature relating chiefly to the year 1897. 
By J. A. Thomson, R. Lydekker, R. Bowdler Sharpe, G. A. Boulenger, 
W. A. Herdman, E. R. Sykes, E. A. Smith, G. C. Crick, A. W. 
Brown, D. Sharp, F. A. Bather, Florence Buchanan, and R. von 
Lendenfeld. Edited (for the Zoological Society of London) by Davin 
Suarp, M.A., F.R.S., F.Z.S., &e. London, 1898. Price 30s. 


The Zoological Record, Volume the Thirty-fifth ; being Records 
of Zoological Literature relating chiefly to the year 1898. By J. A. 
Thomson, R. Lydekker, R. Bowdler Sharpe, G. A. Roulenger, W. A. 
Herdman, E. R. Sykes, E. A. Smith, G. C. Crick, A. W. Brown, 
D. Sharp, F. A. Bather, Florence Buchanan, Rk. T. Gunther, and 
R. von Lendenfeld. Edited (for the Zoological Society of London) by 
Davip Suarp, M.A., F.R.S., F.Z.8., dc. London, 1899. Price 30s, 


The Zoological Record, Volume the Thirth-sixth ; being Records 
of Zoological Literature relating chiefly to the year 1899. By 
J. A. Thorson, R. Lydekker, R. Bowdler Sharpe, G. A. Boulenger, 
W. A. Herdman, E. R. Sykes, E. A. Smith, G. C. Crick, A. W. 
Brown, D. Sharp, F. A. Bather, A. Willey, and R. von Lendenfeld. 
Edited (for the Zoclogical Society of London) by Davip Suarp, M.A., 
F.RS., F.Z.8., &e. London, 1900. Price 30s. 


The Zoological Record, Volume the Thirty-seventh; being 
Records of Zoological Literature relating chiefly to the year 1900. 
By J. A. Thomson, R. Lydekker, R. Bowdler Sharpe, G. A. Boulenger, 
W. A. Herdman, E. R. Sykes, E. A. Smith, G. C. Crick, A. W. 
Brown, D. Sharp, F. A. Bather, A. Willey, and E. A. Minchin. 
Edited (for the Zoological Society of London) by Davin Suarp, 
M.A., F.R.S., F.Z.8., &c. London, 1901. Price 30s. 


The Zoological Record, Volume the Thirty-eighth; being 
Records of Zoological Literature relating chiefly to the year 1901. 
By J. A. Thomson, R. Lydekker, R. Bowdler Sharpe, G. A. Boulenger, 
Alice L. Embleton, E. R. Sykes, E. A. Smith, 8. Pace, Albert 
Brown, D. Sharp, F. A. Bather, and E. A. Minchin. Edited (for 
the Zoological Society of London) by Davin Suarp, M.A., F.R.S., 
F.Z.8., &e. London, 1902. Price 30s. 


Index Zoologicus. An alphabetical list of names of genera 
and subgenera proposed for use in Zoology, as recorded in the 
Zoological Record, 1880-1900; together with other names not 
meluded in the ‘ Nomenclator zoologicus’ of 8. H. Scudder. Com- 
piled (for the Zoological Society of London) by Caartzs OwEn 
Warernouse and edited by Davin Suarp, Editor of the Zoological 


Record. London, 1902. Price to Fellows, 18s. net; price to the 
public, 20s. net. ; 


The Zoological Record for the years 1864-1900 and ‘Index 
Zoologicus’ (38 yols.). Price £15 net; or without ‘Index Zoo- 
logicus’ (37 vols.), Price £14 10s. net. 


These publications may be obtained at the Socrery’s OFFICE 


(3 Hanover Square, W.). 


LIST OF INSTITUTIONS 


TO WHICH 


COPIES OF THE SOCIETY'S PUBLICATIONS ARE PRESENTED. 


AFRICA. 
The South-African Museum, Cape Town. 
The South-African Philosophical Society, Cape Town. 
The Museum, Durban, Natal. 


AMERICA, SOUTH. 
The National Museum, Buenos Ayres. 
The Museum of Natural History, Santiago, Chili. 
The Museum of La Plata, La Plata, Buenos Ayres. 


AUSTRALASIA. 
The Royal Society of Tasmania, Hobart. 
The Royal Society of Victoria, Melbourne. 
The Zoological and Acclimatization Society of Victoria, Melbourne. 
The Linnean Society of New South Wales, Sydney. 
The Royal Society of New South Wales, Sydney. 
The New-Zealand Institute, Wellington. 


AUSTRIA. 
The Hungarian National Museum, Budapest. 
The Imperial Academy of Sciences, Vienna. 
The Zoological and Botanical Society, Vienna. 


BELGIUM. 
The Belgian Society of Geology, Paleontology and Hydrology, 
Brussels. 
The Congo Free State Museum, Tervueren, Brussels. 
The Entomological Society of Belgium, Brussels. 
The Malacological Society of Belgium, Brussels. 
The Royal Academy of Sciences, Brussels. 
‘The Royal Museum of Natural History, Brussels. 


BRITISH INDIA. 
The Asiatic Society of Bengal, Calcutta. 
The Geological Survey of India, Calcutta. 
The Indian Museum, Calcutta. 


CANADA (DOMINION OF). 
The McGill College, Montreal. 


The Geological Survey of Canada, Ottawa. 
The University of Toronto, Toronto. 


2 


CHINA. 
The China Branch of the Royal Asiatic Society, Shanghai. 


EAST INDIES. 
The Royal Society of the Dutch East Indies, Batavia. 


FRANCE. 


The Linnean Society of Normandy, Caen. 

The Agricultural Society, Lyons. 

The Entomological Society of France, Paris. 
The Museum of Natural History, Paris. 

The National Society of Acclimatization, Paris. 
The Zoological Society of France, Paris. 


GERMANY. 


The Royal Prussian Academy of Sciences, Berlin. 

The Society of Friends of Natural History, Berlin. 

The Natural-History Union for Rhineland and Westphalia, Bonn. 

The Senckenbergian Society, Frankfort-on-Main. 

The New Zoological Society, Frankfort-on-Main. 

The Natural History Society, Freiburg-in-Breisgau. 

The Royal Society of Sciences, Gottingen. 

The Imperial Leopoldino-Carolinian Academy of Naturalists, 
Halle. 

The Natural-History Society, Halle. 

The Natural-History Union, Hamburg. 

The Royal Biological Station, Heligoland. 

The Medical and Natural-History Society, Jena. 

The Royal Bavarian Academy of Sciences, Munich. 

The Union for Natural History of Wirtemberg, Stuttgardt. 


GREAT BRITAIN AND IRELAND. 
The Belfast Natural History and Philosophical Society, Belfast. 
The Philosophical Society, Cambridge. 
The Royal Dublin Society, Dublin. 
The Royal Irish Academy, Dublin. 
The Royal Physical Society, Edinburgh. 
The Royal Society, Edinburgh. 
The Free Public Library and Museum, Liverpool. 
The Atheneum Club, London. 
The British Museum of Natural History, LonJon. 
The Entomological Society, London. 
The Geological Society, London. 
The King’s College Library, London. 
The Linnean Society, London. 
The London Institution. 


3 


The Royal College of Physicians, London. 

The Royal College of Surgeons, London. 

The Royal Geographical Society, London 

The Royal Institution, London. 

The Royal Society, London. 

The University College, London. 

The Literary and Philosophical Society, Manchester. 
The Owens College, Manchester. 

The Natural History Society, Newcastle-on-Tyne. 


The Plymouth Institution and Devon and Cornwall Natural-History 
Society, Plymouth. 


The Marine Biological Laboratory, Plymouth. 
The Yorkshire Philosophical Society, York. 


HOLLAND. 


The Royal Academy of Sciences, Amsterdam. 
The Royal Zoological Society, Amsterdam. 

The Dutch Society of Sciences, Haarlem. 

The Dutch Entomological Union, The Hague. 
The Royal Museum of the Netherlands, Leyden. 


ITALY. 
The Royal Institute of Superior Studies, Florence. 
The Civil Museum of Natural History, Genoa. 
The Italian Society of Natural Sciences, Milan. 
The Zoological Station, Naples. 
The Royal Academy of the Lincei, Rome. 
The Royal Academy of Sciences, Turin. 


JAPAN. 
The Science College of the Imperial University, Tokyo. 


RUSSIA. 


The Society of Naturalists, Jurjeff (Dorpat). 

The Society of Sciences of Finland, Helsingfors. 

The Imperial Society of Naturalists, Moscow. 

The Entomological Society of Russia, St. Petersburg. 
The Imperial Academy of Sciences, St. Petersburg. 


SCANDINAVIA. 


The Bergen Museum, Bergen. 

The Society of Sciences of Christiania, Christiania. 
The Royal Danish Society of Sciences, Copenhagen. 
The University Zoological Museum, Copenhagen. 
The Royal Swedish Academy of Sciences, Stockholm. 
The Royal Academy of Sciences, Upsala. 


4 


SPAIN. 
The Royal Academy of Sciences, Madrid. 


SWITZERLAND. 
The Philosophical and Natural-History Society, Geneva. 
The Vaud Society of Natural Sciences, Lausanne. 
The Society of Natural Sciences, Neuchatel. 
The Natural-History Society, Zurich. 


UNITED STATES OF AMERICA. 
The Boston Society of Natural History, Boston. 
The Museum of Comparative Zoology, Cambridge, Mass. 
The Field Columbian Museum, Chicago. 
The Illinois State Laboratory of Natural History, Illinois. 
The American Journal of Science, Newhaven. 
The American Museum of Natural History, New York. 
The New-York Academy of Sciences, New York. 
The Academy of Natural Sciences, Philadelphia. 
The American Philosophical Society, Philadelphia. 
The Entomological Society, Philadelphia. 
The Essex Institute, Salem, Mass. 
The Smithsonian Institution, Washington, D.C. 
The United States Fish Commission, Washington, D.C. 
The United-States Geological Survey, Washington, D.C. 
The United-States National Museum, Washington, D.C. 


WEST INDIES. 
The Institute of Jamaica, Kingston. 


LEI DPI 


The Publications (except in special cases) are sent out direct 
as soon as they are issued. It is requested that they may be 
acknowledged, in order that any mis-delivery may be brought to 
notice. 

Publications sent in exchange to this Society should be addressed 
to the Librarian at this Office. It is requested that they may be 
sent direct by post, as much delay is caused by their transmission 
through booksellers and in other ways. 


By order of the Council, 


P. CHALMERS MITCHELL, M.A., D.Sc., 


Secretary. 
3 Hanover Square, Lonpon, W., 
October, 1903. 


PROCEEDINGS 


OF THE 


GENERAL MEETINGS FOR SCIENTIFIC BUSINESS 


ZOOLOGICAL SOCIETY 
OF LONDON. 
1903, vol. II. 


PART T. 
CONTAINING PAPERS READ IN aaa 
an 


MAY anv J UN 


SMe tia . Mon 
OCTOBER 1903. ee 


PRINTED FOR THE SOCIETY, 
SOLD AT THEIR HOUSE’ IN HANOVER SQUARE. 
LONDON : 


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


Ores [Price Twelve Shillings.] 


visr of oonat 
i 1903-—Vou. 1: 


eek Part I. 


" May 12, 1903. 


1. A Contribution to the Study of Double Monstesitiess in Bishios 
M.A., M.D., EUs in Bebyyela Glasgow epee (Piste 


ren oe trispinosus (Hailstone). By Rosner Gum 


3. Descriptions of New Species of Siuth eae » Coeoptam of of th 
By Marrin Jacopy, FLEAS. 2.00... 2. cess eect cece ee ee ees, 


4, On a Collection of Fishes made : Dr. Gosldi at Rio Janeiro. ‘ 
(Plates VLG VICE) cos vie ee a ee 


ae 26, 1903. 


Grosso, =) presented by Mrs. Perey Sladen to the British J 
Expedition to Central Brazil.) By G. A. Bouuunenr, F.B.S., 


3. The Present State of Knowledge of Colour-heredity in Mice and E 
M.A., F.RS., F.Z.8., Fellow of St. John’s College, Sole 


4. On the Marine Fauna of Farts and Beh Hast Atiiea, from : 


 Acotylea. By F. F, Laipuaw, B. A, Cen! ‘Assston Date on: ay 
ee in the Owens College. (Plate TX) Cee ee 


_ Contents (continued). 


June 16, 1903. 


The Secretary. Report on the Additions to the Society's Menagerie in May 1903... 


1 


sees 


Mr. Frank Finn, F.Z.8. Exhibition oy and remarks ape a es hen-feathered Bantam _ 


aes Pratt a Revarey social Siadeswls oo tinisjetal a'hic'« 


Mr. G. A. Boulenger, F.R.8. Exhibition of, and remarks ie a specimen of the ee 
Ceratohyla bubalus carrying eggs on its back ..........+- . 


Mr. F. HE. Beddard, F.R.S. Exhibition of a bust of the late President of the Society, 
Ins Ebeniower MK O.Bs. jis ec ceases ee x 


Mr, F. B. Beddard, F.R.S. Exhibition of, and remarks upon, sections of the ovary of the 
Thylacine 


eC Ce eC a cy 


. Mr. R. E. Holding. Exhibition of some skulls and horns of, and remarks upon the horn- 
growth in, the St. Kilda four-horned Sheep 


ee a ey 


Dr. A. 8. Woodward, F.R.S. Exhibition of photographs illustrating the exhumation of 
a receutly-discovered Mammoth in Siberia.........ceece scenes c cece cece cence ats 


Mr. Oldfield Thomas, F.R.S. Exhibition of, and remarks upon, the horns of a new form 
of Grant’s Gazelle, proposed to be named Gazella granti robertsi . 


1. On an Extinct Species of Genet (Genetta plesictoides, sp. n.) from the Pleistocene of 
Gyorusr by Dororay Mi. Ay Bann. “CRlatey XG )ils 2) oie. «e's tunics oie disieinels c)/elsieietale eis.» 


oe Description of a new Fish of the Gobiid Genus Rhiacichthys from British New Guinea. 
Byte. DOULMNGHR DE as Vebefies re (elate XD). ec. s jas cele sacs o¢ seisinies welne 


3. Descriptions of new Reptiles from British New Guinea. By G. A. Boutuncer, F.R.S., 
V.P.Z.S. (Plates XII. & XIII.) 


Cr ee i i i i a ee cary 


4. On the Marine Fauna of Zanzibar and British East Africa, from Collections made by 
Cyril Crossland in the Years 1901 and 1902, —Polychstta. Part Il. By Cyrm 
Crosstann, B.A., B.Sc. (Plates XIV. & XV.).. 


5. On the Ento-Parasites collected by the « Skeat Expedition ” to Lower Siam and the 
Malay Peninsula in the Years 1899-1900. By Arruur E. Sureuey, M.A., F.Z.S., Fellow 


and Tutor of Christ’s College, Cambridge, and University Lecturer in the Advanced 
Morphology of the Invertebrata. (Plate XVI.).. 


6. On the Modifications of Structure in the Syrinx of the Accipitres, with Remarks upon 


other Points in the Anatomy of that Group. By Frank E. Beppaxp, M.A., F.B.S., 
EXGsentor) to thei SOclebyy je's! seis. meus Cram fue oaiaicinie: Nelsie eile wists ticle sey oe terenerers Sande 


7. On Medusx from the Coast of British Columbia and Alaska. By Lours Mursacu and 
OressweLt Susarer, F.Z.S. (Plates XVIL-XXIL) .......  eesdatiediee 


See ee ee sees 


CC a Ci rrr 


114 


116 


119 


119 


129 


145 


157 


I It. -} Anatomy of Monstrous Fish Embryos — 
hed r ee TaN 


ooh Caine: lavve ae wee 


oe ue Mylacrodon goeldii. an 
fi AVAL ali Peristedion altipinnis. 2. Genypterus brasiliensis 


seteece CANIS SE CS fe eeee 


10.80 -Planarians from Zabrina RO oa 


Fein oon 1. Genetta genetta (L.); 2-6. G. plesietoides, sp. nov. ; 
Si eye, - sictis croizeti Filhol Paco trle Rnters Wag ite Cuctesemette 


: ji mae mepade (Oe aa 
Se ee ae ive A dels aie bD, ae IELTS 3. L. eds umn 


santnabig TRA Oa pase a 
XIV. Fig. 1. Diopatra neapolitana. 2. Onuphis holobranchiat . 
nhs 3-6. Marphysa macintoshi, DSP. seeeseceteesee eens 
Xv. Hise. 7-10. Marphysa mossambica. 11, 12. M. simplex, n. 
18, 14. M. furcellata, n. sp. 6.02. .00 cece cece ce eee 
XVI _ Ento-Parasites from Lower ela ete eee ee tee ee eter eee 


; RVI Be 
Beige uae Ive th. ©: ORNL, 
xXx + Meduse from British Columbia and Alaska .....,......++ 
a | 
XXII. 7 ‘ 
. NOTICE. /u" ena 
_ The ‘ Proceedings’ for the year are issued i in Sour parts, pears two volumes ae 
me as follows:— 
% | “vou. in 
Part I. containing papers oe in January and February, i in June. 
oy ah May! ag »  Margh and April, in ene 
VOL. I. yea Rtays 4) ats 
Tart I. containing papers read in May and June, in October. . net, 
JU ., ‘ _ November and BG ee ots in April, 


vt Proceedings, 1903, Vol. I. Part 11. was published on chistes 6th, 1908, 


- 


PROCEEDINGS. 


OF THE 


GENERAL MEETINGS FOR SCIENTIFIC BUSINESS 


' OF THE 


" 1O0LOGIOAL SOCIETY 
OF LONDON. 


1903, vol. IT. 


PART IT. 
CONTAINING PAPERS READ IN 


NOVEMBER anv DECEMBER. 


APRIL’ 1904%- 


PRINTED FOR THE SOCIETY, 
SOLD AT THEIR HOUSE IN HANOVER SQUARE. 
LONDON : 


MESSRS. LONGMANS, GREEN, AND CO., 
PATERNOSTHR-ROW. * 


[Price Twelve Shillings}  §& 


LIST OF CONTENTS. 
1903.—Vor. I. 


Part IL. ee 


November 3, 1903. 


The Secretary. Report on the Additions to the Society's Menagerie in June, July, Agony . 


and September, 1903 ....---.--.-.. ee e-e es been eee eect ee ce eee Sialeys la orarabatel - 
Dr. P. L. Sclater, F.R.S. Exhibition of, and remarks upon, the horn of a Rhinoceros 
from the White Nile: ¢..0...2 22s oe eee ee ee a sferenciot sitet 194 By 
The Secretary. Exhibition of, on bebalf of Mr. H. Slade, a series of Se a thes 7% 
Indian Elephant in the act of conress’:. ....4...).%.swalse- ahs be ne eee LOD yA Ae 
i Ks eg 
Mr. Henry Scherren, F.Z.S. Exhibition of, and remarks upon, abnormal claws of ie eee ai: 
Oxrsib and VO DStEr’ cceystae sc oe) ele te etslev tense selene se peeet Beleneta etna aisle, 2) ciie/ahielel ats Node heienehauane 195, asian 
Mr. R. I. Pocock, F.Z.S. Exhibition of, and remarks upon, photographs of a mounted = ese 


specimen of Burchell’s Zebra ’..-. sc suds s0.9n (01a e ede lek ge 196 


Mr. BR. I. Pocock, F.Z.8. Exhibition of a large Geophilomorphous Centipede from 
\{GINGTA0(S Eason Mis erieren as nice tate omc ie waa lease co Rae AAG age eee Tete ah atta ene 


Mr. Oldfield Thomas, F.R.S. Exhibition of specimens and descriptions of new species. of 
Mammals from New Guinea. (Plate XXULI) 3 oo eve ctetelstcpetevele e aeraians cies han 


1. On some new Species of Aquatic Oligochata from New Zealand. By W. B. Benwam, 4: i 
D.8c., M.A., F.Z.8., Hon. M.R.S. Tasm.; Professor of Biology in the University of 
Otago. (Plates XXIV. MVD.) 5 se wed 20 stern wpe in ei etal lace we pe eo eee 20 


. On the Mammals collected by Mr. A. Robert at Chapada, Matto Grosso. (Percy Sladen 
Hxpedition to Central Brazil.) By Ouprimtp Tuomas, F.R.S. (Plate XX VIL.) 


bo 


3. List of the Coleoptera collected by Mr. A. Robert at Chapada, Matto Grosso. aa 


Sladen Expedition to Central Brazil.) By ©. J. Ganan, M.A., and G. J. Arrow. 


(UB Taibe XX ViIETS ee as AS ce ale otay ohn yee cia are ia cae pe Dia RaS A ay fie Aico) sin » 


4, On the Lepidoptera collected at Chapada, Matto Grosso, by Mr. A. Baber . (Perey 


Sladen Expedition to Central Brazil.) By F. A. Heron and Sir Guorcz F. Hanson, 
Bart., F.Z.8. 


. Note on the Cypriote Spiny Mouse. By P. Cnatmuzrs Mircurir, M.A., D.Se., 
Secretary to the Society 


ex 


ee eC CC Cr CC 


6. On the Muscles of the Ungulata, By Brrrram C. A. Winpin, D.Sc., M.D., M.A, 
E.R.S., Professor of Anatomy in the Univer sity of Birmingham, and F. G. Parsons, 
E.R.C. ioe H.Z.8., F.L.S., Lecturer on Human and Comparative Anatomy at St. 
Thomas's Hospital, late Hunterian Professor in the Royal Gees of See 
ae —FPart Il. Musele> of the Hind-limb and Trunk ..... Acdoneoe 


7. On some Points in ths Seantohay: chiefly of the Heart and Vascular See) of the 
Japanese Salamander, Megalobatrachus japonicus. By Frank BE. Bupparp, M.A, 
F.RS., F.B.S.E., Prosector to the Society: 2.5 See ik eerie Bie aioe ares eat 108, TL ee 


November 17, 1903. Be eh ae 
The Secretary. Report on the Additions to the Society’s Menagerie in October 1903 me 3815 ‘ 


Mr, Henry Scherren, F.Z.9. Exhibition of, and remarks upon, a horn of Rhinoceros simus. — 316 


Mr, R. I. Pocock, F.Z.8. Exhibition of. a piece of basalt containing a sae of the Marine sen 


Spider Desis hen YOM ii Heleralnracunieieetueils eleh sini ojanrtal starr tera yards earns Soesa dic Heaeuise Ine eae 31 
Mr. R. T. Pocock, F.Z.8. A new suggestion as to the use of the white rump-patches in the . i 
Bay Ch On ane OMNIS GAM ARMM AAMME CMS SMA) A aagrees Se 31 


Mr. E. E. Austen. Exhibition of, and remarks upon, specimens of the Teetee- which | 
acts as transmitter of “sleeping-sickness” in Uganda ........+2..ceee+cees oleaie ec 316 


Contents continued on page 3 of Wr app: 
art i (ititi—‘—Ss—sOO—:~:~Ci‘S 


ConTENTS (continued). 
“ 
November 17, 1903 (continued). 


- Mr, Oldfield Thomas, F.R.S. Exhibition of, on behalf of Mr. W. H. de Winton, F.Z. ce a 
drawing of a female Gazelle bearing a hair-whorl on the withers .....+6.06.00s.0 0. 317 


Page 


Mr. C. Tate Regan. Notice of a Memoir entitled “Revision of the Fishes of the Family 
MNT COU a aan Meakaferinr sti sfotutstatefeceters lc (ake alontte « 


1. On Early Sanskrit References to the Tiger. By. V.V. Ramanan, M.A, PZS......... 318 


2. On the Trachea, Lungs, and other Points in the Anatomy of the Hamadryad Snake | 
(Ophiophagus bungarus). By Frank E. Brpparp, M.A., F.R.8., Prosector to the 
SHIGDAZGE ES Fb oGihS SUOU ADE Ec DOBSON GOSS ROUT CIUBIND OCIS CEBMIES ARCnet ono ~ Speman tie 319 


3. Report on the Fishes collected by Mr. Oscar Neumann and Baron Carlo von Erlanger 
in Gallaland and Southern Ethiopia. By G. A. Boutuyeer, F.R.S., V.P.Z.S. (Plates 
Oo OO Ga GSS BA ale a hy Sdiprs Beiniciens MOrin GPE MEMNOP np Manca. shin Saint 328 
a December 1, 1903. 
Mr. F. Martin Duncan. Demonstration of the Zoological Bioscope 


Mr, F. E. Beddard, F.R.S. Exhibition of, and remarks upon, a portion of the large 
intestine and the cecum of a Boa 


Mr. F. EB. Beddard, F.R.S. Exhibition of, on behalf of Mr. G. A. Doubleday, a hairless 
Peer PONEEN Coie Hey CO OTIAIVONE Euaitirwic: sta" chev e\oinisi aie: a 0 c\s)e'sy sio.etsiie's' viet Sic sic ciej ae vise Mimteien eee 336 
Dr. Walter Kidd, F.Z.S. Exhibition of a drawing of a Beisa Antelope showing a Teves 
SMSO AMbON. UM OpDA Care tate stasateta olay cietsta rs «Pein (otal clecciee eysletela’ e\/oie (are ie « aiesersie reves 336 


Mr. Oldfield Thomas, F.R.S. Exhibition of, and remarks upon, a specimen of the Rodent 
Fornarina (gen. nov.) phillips: and description of Heterocephalus ansorget (sp. nov.).. 336 


Mr. G. A. Boulenger, F.R.S. Exhibition of a hybrid Newt between Molge marmorata S 


and M. cristata fe) dS, CO er dgacge tne CO ODO Co hociahe OA BeLC ONE Op.ch eo Me Uomo abe es 337 
Prof. E, Ray Lankester, F.R.S. Exhibition of, and remarks upon, two. Cae re- 
presenting the arrangement of the hair on the faces of two specimens of Okapi...... 337 
Prof, H. Ray Lankester, F.R.S. Exhibition of specimens of Medusz from the Victoria 
MME de Nae satire Mela venti ah yee recite ales yah ava watts Bimthialale Sc Miplereer ery uaia els tayale obs 340 
1. The Mammals of Cyprus. By Dororny ME AN SEPAIRE cccb arabs Wea ee he) tein afr ist eo raiat fey nee 341 
2. On the Cause of Death of a Polar Bear recently living in the Society’s Gardens. By 
Repeumre N. Sauaman, M.A., M-B. Cantab., F.Z.S., Acting Director of the Patho- 
logical Institute, London Hospital .. es : 348 


3. On the Development of the Adult Colouring in the Yellow-billed Cardinal (Paroaria 
capitata) from 8. America, By Artuur G. Buruzr, Ph.D., F.LS., F.Z.8., &e. ...... 850 


4, On the Occasional Transformation of Meckel’s Diverticulum in Birds into a Gland. 
By P. Caaumers Mircurnz, M.A., D.Se., Secretary to the Society.................. 352 


5. On some Nudibranchs from East Africa and Zanzibar, Part III. By Sir C. Exroz, 
K.C.M.G., H.M. Commissioner for the Hast Africa Protectorate, F.LS8.—Doridide 
Cryptobranchiate, Evi) (Plates XXX NRT Vi) ales a ateln wa cla’ inte, « Peace eres, O04 


6. A Note upon the Tongue and Windpipe of the American Vultures, with Remarks on the 
Interrelations of the Genera Sarcorhamphus, Gypagus, and Cathartes. . By Franx E.- 


BRPDARD.- MLAs. BRS), Prosector tothe Society? sre ae. «/-/-/s 0) «s+» ac) ae aicisetemtoreiets 386 

TIVE LES eet ig RE RI DE ae oS RRS wt OL SE aa Ae eA MMR MS DA a a! MEE: Sea Le 
Titlepage -....... Siaicisyalal as stelalatayer tevapa tiara yerare i 
List of Council and ‘Officers IE epee neal MCRAE EPR TSI eee y ig. cabs cod after eee ee es ii 
iraiaGinconboniai. 1. Woamer cis a aarenneuee eee seks cisiters és iii 
Alphabetical List of Gees Ba EH net ARTs ENO a ee RT Pa a A em RAL Ue Kila WY ix 
TURE oud TEE asp Sean Ae eee on ab LY Ue ey Pa ESTA ie oe a Be ba a ge eRe xvii 
Petes astra PUTER =... ...' oie Si angle) iaie suaiplele’ anelaidm Rabie eu, Fe 5 Aa Ra Coe aa SAN yeh xix 
See OITA N EDT CH LOLS) oie'eebara sis << cvele tialetu/alaiele lorine wttunbalaue shane Vala tiater shave hu shau dN S| Anan XXI 


Errata 


LIST OF PLAT 


1903.—VOL. II. 


PART II. 
: pebadeene 
Plate Selene S cau aye 
XXIII. 1. Anisomys imitator. 2. Hyomys mecki.....-.. 
XXIV. 2 


XXV. } Aquatic Oligochstes from New Zealand ........+2-- 
MERVAG Pe etree EF AEE D's 
XXVEl. Canis sladent oo... cbiens cae ceteeene nave edee = 
_ XXVIII. Coleoptera from Central oe spiiiela| ale eiaieiwte's ainje'ate 
XXIX. © Labeo newmanii oo.e.sccrecececcecseccercccscece 
KXX, Barbus erlangeri .osceeccrccecseccess beet er enes 
XXXI. 1. Discognathus makiensis, 2. Discognatius 
3. Chiloglanis modjensis..c-+..000.ercccetesere. 
XXXII. 1,2. Thordisa villosa. 3. T, crosslandi. eS Trippa n monso: 
5. Halgerda willeiiicc aes ols wat ice se =e 
XXXII. 1-3. Thordisa villosa. 4-8. T. crosslandi ...... 
XXXIV. 1,2. Halgerda wasinensis. 3. Sclerodoris coriacea. 
ae Copepod. 5, 6. Asteronotus eal 7 


NO TICE. 


Part. I. 


II. 


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way 


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