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PROCEEDINGS
OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
OF THE
AVOLOGICAL SOCIETY
OF LONDON.
1903, vol. 1.
(JANUARY—APRIL.)
PRINTED FOR THE SOCIETY,
AND SOLD AT THEIR HOUSE IN HANOVER-SQUARE,
LONDON:
MESSRS. LONGMANS, GREEN, AND CO,,
; PATERNOSTER ROW,
Ih JES 1b
OF THE
COUNCIL AND OFFICERS
OF THE
ZOOLOGICAL
SOCIETY OF LONDON.
1903.
COUNCIL.
(Elected April 29th, 1903.)
His Grace Tur DuKs or Beprorp, K.G., President.
Witiram T. Buanrorp, Hsq.,
LL.D., F.R.S., Vice-President.
Grorce A. BouLencer, Hsq.,
E.R.S., Vice-President.
Wiiuiam E. pre Winton, Ksq.,
Acting-Superintendent of the
Gardens.
Hersert Druce, Esa., F.LS.
CHARLES DrumMonpD, Ksq.,
Treasurer.
Cures H. Garry, Esq., LL.D.
FREDERICK GILLett, Esq.
F. DuCange Gopman, Esq.,
D.C.L.,F.R.S., Vice-President.
Apert Gtnruer, Esq., M.D.,
Pu.D., F.R.S., Vice-President.
Carr. THe Marquis or Hamit-
ton, M.P.
Pror. Grorce B. Howss, D.Sc.,
LL.D., F.R.S., Vice-President.
Lr.-Cou. L. HowArp Irpsy.
Srr Epmunp G. Lover, Br.
K. G. B. Meapr-WaA po, Esa.
P. CHaumers Mircuet, Ksq.,
M.A., D.Sc., Secretary.
Str THomas PAINE.
E. Lort Pures, Esq.
Davip Suarp, Ksgq.,
OLDFIELD Tuomas, Hsq., F.R.S.
Henry Woopwarb, Hsq., LL.D.,
FE.R.S., Vice-President.
M.D.,
PRINCIPAL OFFICERS.
P. Cuatmers Mircuey, Esq., M.A., D.Sc., Secretary.
W. E. pe Winton, Esq., Acting-Superintendent of the
Gardens.
Frank E. Bepparp, HEsq., M.A., F.R.S., Prosector.
Mr. F. H. Warernouss, Librarian.
Mr. JoHn Barrow, Accountant.
Mr. W. H. Couz, Chief Clerk.
Mr. Grorcr Artuur DouBiepay, Clerk of Publications.
Mr. Artur Tomson, Assistant Superintendent of the
Gardens.
LIST OF CONTENTS.
January 20, 1903.
Page
The Secretary. Report on the Additions to the Society’s
Menarerioiin December 1902 wi. iae. ts sekceeesewesemesneehs i
Mr. P. L. Sclater. On the Zebra-and-Pony Hybrid living
im ther Society siMienarenie | 62. 4ieeswsenewoeese side beaten omens 1
Mr. J. S. Budgett, M.A., F.Z.S. Account of his recent
Journey to Uganda in search of the Okapi and
ELONA PD LCIIUS EN ARAN NOM IE Se GhrM as gah Sete eed vn tne t ins cad seer owes
bo
1. Note on the Spiracles of Polypterus. By J. 8S. Bupcert,
DIE ABE Se suit cee som euens instcat iene ars ates te soso enone 10
2. On the Brains of WNasalis larvatus and of some other
Old World Primates. By Frank E. Bepparp, F.R.S.
(SOS: Soa donads COCEOO COCO ES BOE ROOOH SC OE EOR COR ATEnE HSE GHOC AGREE eon 12
3. On the Fishes collected by Mr. G. L. Bates in Southern
Cameroon. By G. A. Boutencrr, F.R.S., V.P.Z.8.
(ERI DIVES) rth tenance. doecbeen ea tb acartocancis ses aceeeeeebede 21
4. On the Anatomy of the Gephyrean Phascolosoma teres,
n.sp. By W. K. Hurron, M.A., M.B., Senior Demon-
strator of Anatomy in the University of Glasgow.
(elena WAR VAIINY) 8 tomes oz cdoumabnastd scans cece cae ced anaes cure 29
Or
. On Potamon (Potamonautes) latidactylum, a new Fresh-
water Crab from Upper Guinea. By Dr. J.G. pe Man,
of Terseke;: Holland: “(Plate TX.) 12 b.ccccccecseowedececeess 4]
iV
Page
6. On a new “ Bird’s-dung” Snider from Ceylon. By R. IL.
Pocock, F.Z.8., and the Hon. N. C. RorHscuiLpD, B.A,
E.ESS., EZS. ’ (Plate X.) BPE. .bscoa cou Renee perenne Saj00> 48
7. On the Crustaceans of the Genera Petalidiwm and Ser-
gestes from the ‘Challenger, with an Account of
Luminous Organs in Serg gestes challengeri, n. sp. By
Dye dels di Hansen (Copenhagen). (Plates XI. & XII.) 52
February 3, 1903.
The Secretary. Report on the Additions to the Sage s
Menagerie in January 1903 ....... eee cece eee ee ences ee en ens 79
1. Notes on the Hair-Slope of four Typical Mammals. By
WW Areas, [GIDDY IBAgsb sonoscoeacodstocosoosssnscopanoonoopsoeand 79
2. A Prodromus of the Snakes hitherto recorded from China,
Japan, and the Loo Choo Islands; with some Notes.
By Captain F. Watt, Indian Medical Service .........-.- 84
3. Note on the Wild Sheep of the Kopet-Dagh. By R.
IDPADINIEGDG, aaqoanesqsouoséoposbonnoocoeodosoooooosasccedoscbonceonns 102
4. On new Parasitic Copepoda from Zanzibar and Kast
Africa, collected by Mr. Cyril Crossland, B.A., B.Sc.
By Staff-Surgeon P. W. Basserr-Surru, NG eh VA Se coo LOA
5. On the Original Home of the Tiger. By Col. C. EH.
RSimmohyeew, OBS! OMINIKET COLD, — Shscsaosodobnoncedoooccoco0009 109
6. On the Mode of Copulation of the Indian Elephant. By
H. Suave, Conservator of Forests, Maymyo, Burma ... 111
7. On the Coelenterata collected by Mr. C. Crossland in
Zanzibar.—l. Ceratella minima, n.sp. By Sypney J.
Hickson, M.A., F.R.S., F.Z.S., Beyer Professor of
Zoology in the Owens College, Manchester. (Plate
DrCIUTIED)) obodbdobbbndetonus assdsodondasadoqhoudiadsqobadakdesuecosoos 113
8. Contributions to our Knowledge of the Plankton of the
Faeroe Channel.—No. VIII. By G. Herserr Fowter,
TRNgs IEloial Dy ID S/ATSB noncassasesoadescesenonbonaogubdesdodococescd 117
9. On the Present Condition and Habits of the Klk in
Norway. By MH. J. Huwes, FR.S. ...6... 2. sce eeenes 133
Vv
February 17, 1903.
Mr. R. KE. Holding. Exhibition of, and remarks upon, some
Skulls of Mammals showing abnormal dentition .........
1. On some new Species of Spiders belonging to the Families
Pisauwride and Senoculide ; with Ghar acters of a new
Genus. By Freperick Pickarp-CamsBrives, B.A.,
Begs (Plates Rein OXVe)! ieasiigdier. cect ldecenenMteeies oble
bo
On the Marine Fauna of Zanzibar and British East Africa,
from Collections made by Cyril Crossland in the Years
1901 and 1902.—Polycheta. Part I. By Cyrm Cross-
TAND; Bet; Bsc, ii (elatese Navel. do Xe Wels) is, wrerele ate 9
3. On the Axis, Atlas, and Proatlas in the Higher Therio-
donts. By R. Broom, M.D., B.Se., C.M.Z.S. (Plate
DS WAIHI H Ip Vans donecneRepbce eo cbt neabocs contsc spr onceeeor scockos ocean
4. A Revision of the Fishes of the Genus 7’riacanthus. By
CRRT Arnis EGRGUAING Bee Ales ig tutes eaticle seiocciclas tin eae ele Rislalgtetarerstente
5. On the Geographical Variations of the Sand-Viper, Vipera
ammodytes. By G. A. Boutencer, F.RS., V.P.Z.8. ...
6. Notes on the Habits of the Hoolock. By Gro. CaAnDLER,
Wicib (OPnMIIe OF etanghbonbbbcoconataonbs oadbocasocoeosoadeadeeanocccas
March 8, 1903.
The Secretary. Report on the Additions to the Society’s
Menasere mel ebriuaity 190 Si ecctans..) 2 cae ashe nsercet
The Secretary. Extract from a letter from Major C. Delmé
Radcliffe concerning skins of a Monkey and an Otter
PMO TOG OSAMA). cere nacseeieli stance stone shia ne ets'a.s/e palelnwisi'a <6
Mr. F. EK. Beddard, F.R.S. Exhibition of a mounted skin
Olthne Greater Bird Ol PATACISE! sce cmcseie<ce cee sescsine as cies
Mr. J. L. Bonhote, F.Z.8. Exhibition of, and remarks wpon,
a photograph a BKlephants showing considerable growth
of hair, and note upon the Sansky ie name of the Tiger..
Prof. F. Jeffrey Bell, F.Z.5. © Exhibition of, and remarks
upon, a Holothurian of the Genus Actinopyga ............
1. On a new Species of Pigmy Antelope of the Genus eo-
tragus from the Cameroons District, W. Africa. By
WB. De WINTON, BLAS. (Plate XTX.) oid. ccssts.
Page
169
Wel
180
185
187
191
191
192
192
yi
i)
. On the Land Operculate Mollusca collected during the
““Skeat Expedition” to the Malay Peninsula in 1899-
1900. By E.R. Syxus, F.Z.S. (Plate XX.)
eee ceecesese
(su)
. The Significance of the Callosities on the Limbs of the
Equide. By R. lypexnur, PAS. ....-0..1..-.e eens
4. Note on some Remains of Struthio karatheodoris Maj.
of the Island of Samos. By Ruporr Martin, of Basel
University
ee oeeccnnese oer eseooeoeresaseeseassesoreeosessesoseserenaee
Cy
. On a new Genus and two new Species of Harthworms
of the Family Hudrilide, with some Notes upon other
African Oligocheta. By Frank HE. Bepparp, M.A.,
F.R.S., F.Z.8.
ecaeeaee cee oeoes reese soeoooceveearneeooeooesasereoues
March 17, 1903.
Prof. Newton, F.R.S. Exhibition of, and remarks upon,
photographs of the White Rhinoceros, taken by Mr.
OoRe saunders. © oViaG 2 im Zrululamcleee ee eee en eee ane
Mr. Oldfield Thomas, F.R.S. Exhibition of a skin and
description of a new species of Monkey, Rhznopithecus
erie Bors ( LEAN Key O- [a WinaenennoeAbsonadoanondoosdodsssoghasons
Mr. Oldfield Thomas, F.R.S. Exhibition of specimens and
description of a new species of Duiker, Cephalophus
COON [cA RB ROP POR AraC ANNA An innichAnCcuObAnUNT ARdons sh oodeAsKoeooo
1. Observations and Experiments on Japanese Long-tailed
Fowls. By J.T. Cunninenam, M.A. F.ZS. ............
2. On some Nudibranchs from Hast Africa and Zanzibar.—
Part II. By Sir C. Exniotr, K.C.M.G., H.M. Com-
missioner for the Hast Africa Protectorate, F.Z.S. ...
3. Contributions to the Osteology of Birds.—Part VI.
Cuculiformes. By W. P. Pycrart, F.Z8., ALS,
(Plate XXIT.)
eens esoaencecseeec ees er osereceseseseer seo eso ees es eee
April 21, 1903.
The Secretary. Report on the Additions to the Society’s
Mienacerieamy March: LOO Sie. -rs-csensee cece eeeecee ees
1. Linneus and Hunter on Feather-Tracts. By Henry
SCHERREN, BEZiS J 8 oo. A hud. slehia cers aetemnentevlah Miata elem sree ree
Page
194
199
203
210
258
292,
292.
vil
2. On some Mammals collected by Capt. H. N. Dunn,
R.A.M.C., in the Soudan. By Oxprietp Tuomas,
US EvsS eau ne eee pret ole «d\artoctaqise sataste soso coin eaena au San escebines
3. On a Collection of Turbellaria Polycladida from the Straits
of Malacca. (Skeat Expedition, 1899-1900.) By F. F.
Laipiaw, B.A.Cantab., Assistant Lecturer and Demon-
strator in the Owens College. (Plate XXIII.)............
4. On the Phylogenetic Cause of the Transposition of the
Testes in Mammalia: with Remarks on the Evolution
of the Diaphragm and the Metanephrie Kidney. By
W. Wooptanp, University College, London
5. On the Geographical Distribution of Spiders of the Order
Mygalomorphe. By R. I. Pococn, F.Z.8. ............... é
Page
301
319
ERRATUM.
Page 110, line 29, for Archipelago, which read Archipelago which.
APA BH TEC A tal sf
oF THE
CONT hy aU Onis;
With References to the several Articles contributed by each.
BassEtt-Smitu, Staff-Surgeon P. W., R.N., F.Z.8.
On new Parasitic Copepoda from Zanzibar and East
Africa, collected by Mr. Cyril Crossland, B.A., B.Sce.......
Bepparp, Frank H., M.A., F.R.S., Prosector to the Society.
On the Brains of WVasalis larvatus and of some other
OIdaWrorldaerimayess sec. ceacslseuictenis se parece oemoleecen @ake cocme
Exhibition of a mounted skin of the Greater Bird of
{EE ania ISU: Ra eM AER Ee Se, ERT ep aah SO A Re
On a new Genus and two new Species of Karthworms
of the Family Hudrilidw, with some Notes upon other
Atricom OM GGG hear tej. ccenacesielasieisisecisecapeiuccsosisgssinesed ese
Beuy, Prof. F. Jerrrey, M.A., F.Z.S8.
Exhibition of, and remarks upon, a Holothurian of the
GemusrAeriiany gay wre den Worse sees ceeehuicssste seve cstvvanes
Page
104
210
x
Bonuore, J. Lewis, M.A., F.Z.S.
Exhibition of, and remarks upon, a photograph of
Hlephants showing considerable growth of hair, and note
upon the Sanskrit name of the Tiger .................-s..««
Bounencer, Grorce ABert, F.R.S., V.P.Z.S.
On the Fishes collected by Mr. G. L. Bates in Southern
Cameroon a (@elatess]: Ve) nur eeeeeeepenr ere ae nee
On the Geographical Variations of the Sand-Viper,
Vipera animodifles p.8. nigh yekee Saeki
Broom, Rosert, M.D., B.Sc., C.M.Z.S.
On the Axis, Atlas, and Proatlas in the Higher Therio-
Gomis": (Plate DLV WLI NEA Ne er Mente eminent Nadel | Aap rRmeme
Bupeert, J. 8., M.A., F.Z.S.
Account of his recent Journey to Uganda in search of
teins) CUEF OM BuMNOl JZQUNTTICPOIS. sencebonasocoasobocoeoacosotnsoedeseuss
Note on the Spiracles of Polypterus ......................+.
CAMBRIDGE, F. Pickarp. See Pickarp-Campripes, F.
CANDLER, GrorGcE, M.B. Cantab.
Notes on the Habits of the Hoolock ...................0.
CROSSLAND, CyriL, 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 I. (Plates
DOV Dds DVB oo rceeaes dcr (ct cen ee ee eee
CunnineuaM, J. T., M.A., F.Z.S8.
Observations and Experiments on Japanese Long-tailed
Fowls
whesheheieseielsialslaceehejleleiersieleile)eela/elslels)elsleleleleisisl=lallainielalalsiistclsicietsialaletereletotetaterets
Dr May, Dr. J. G., Ierseke, Holland.
On Potamon (Potamonautes) latidactylum, a new Fresh-
water Crab from Upper Guinea. (Plate IX.) ............
Page
192
185
177
bo
10
187
169
227
x1
De Winton, WituiAM E., Acting-Superintendent of the
Society’s Gardens.
On a new Species of Pigmy Antelope of the Genus
Neotragus from the Cameroons District, W. Africa.
(Plate XIX.)
i rr er a
Kuror, Sir Cuartes, K.C.M.G., H.M. Commissioner for
the East Africa Protectorate, F.Z.S.
On some Nudibranchs from East Africa and Zanzibar.—
Part II.
ee
Kuwes, H. J., F.R.S., F.Z.S.
On the Present Condition and Habits of the Elk in
Norway
ee ee i i
Fowuer, G. Hersert, B.A., Ph.D., F.Z.S.
Contributions to our Knowledge of the Plankton of the
Faeroe Channel.—No. VIII
i i i |
Hansen, Dr. H. J., Copenhagen.
On the Crustaceans of the Genera Petalidiwm and
Sergestes from the ‘Challenger,’ with an Account of
Luminous Organs in Sergestes challengeri, n. sp. (Plates
SICTGE oS) Heebpa so vpnebeton doen seeonap cous uaboduanuacicbonocnocceC
Hickson, Sypney J., M.A., F.R.S., F.Z.8., Beyer Professor
of Zoology in the Owens College, Manchester.
On the Coelenterata collected by Mr. C. Crossland in
Zanzibar.—l. Ceraiella minima, n. sp. (Plate XIII.) ...
Ho.prine, R. EK.
Exhibition of, and remarks upon, some Skulls of
Mammals showing abnormal dentition
Hurron, W. K., M.A., M.B., Senior Demonstrator of
Anatomy in the University of Glasgow.
On the Anatomy of the Gephyrean Phascolosoma teres,
Tig eed eR CR AVA ARLIM Aue MEI umeaniiey). coe dcce steele’ neues’
Page
192
250
133
Litre
or
to
Xi
Kipp, Wautrr, M.D., F.Z.S8.
Notes on the Hair-Slope of four Typical Mammals
Laiptaw, F. F., B.A.Cantab., Assistant Lecturer and
Demonstrator in the Owens College.
On a Collection of Turbellaria Polycladida from the
Straits of Malacca. (Skeat Expedition, 1899-1900.)
(GEIB) NO-CINUE)) Goenscoodhaabpoconavoauondooesosndco20D0cen doo cana
LyDEKKER, RicHarD, B.A., F.R.S., F.Z.8.
Note on the Wild Sheep of the Kopet-Dagh ............
The Significance of the Callosities on the Limbs of the
JHE Tes a cocies badeannancneperodonnabon nos oosboSoducquodoMeandcccHscés
Martin, Rupour, of Basel University.
Note on some Remains of Struthio karatheodoris Maj.
obithe Wslamd "Of Samos ii cack on ccncece see eemecee ere sces
Newton, Prof. ALFRED, M.A., F.R.S., F.Z.8.
Exhibition of, and remarks upon, photographs of the
White Rhinoceros, taken by Mr. C. R. Saunders, C.M.G.,
AAW ALOU UNE ANG Leen Son OM Weer NM Rr eninge Eur oa see e
PickARD-CAMBRIDGE, FREDERICK, B.A., F.Z.8.
On some new Species of Spiders belonging to the
Families Pisauride and Senoculide ; with Characters of
apnews Genus: (Plates XGmVards PXOVe) ieeereearercce cess ae
Pococg, R. I., F.Z.8.
On the Geographical Distribution of Spiders of the
Order) My calomorphee: 42.05) eceeeer eget eeee sea as enieee
Pococr, R. I., F.Z.8., and Roruscurip, Hon. N. C., B.A.,
F.E.S., F.Z.8.
On anew “ Bird’s-dung” Spider from Ceylon. (Plate X.)
Page’
79
301
203
tol
48
xili
Pycrarr, WILLIAM PrAne, F.Z.S., A.L.S.
Contributions to the Osteology of Birds.—Part VI.
Ciscoe an (ENC ROR) Nite wosietecunseonceaenasientvineens
Reaan, C. Tarr, B.A., British Museum, Natural History.
A Revision of the Fishes of the Genus Zriacanthus .
Rotruscuitp, Hon. N.C., B.A., F.E.S., F.Z.8., and Pococr,
Rt, B:Z8:
On anew “ Bird’s-dung” Spider from Ceylon. (Plate X.)
ScHERREN, Henry, F.Z.8.
Linneus and Hunter on Feather-Tracts ...............0-
Scuater, Pump Luriey, M.A., D.Sc, Ph.D., F.RS.
(Secretary to the Society till January 20th, 1903).
Report on the Additions to the Society’s Menagerie in
TOyeceerill eee USO: eoncnbonddncebosgsoncdb aococoboundasonostaboodgond
On the Zebra-and-Pony hybrid living in the Socicty’s
IWIGSTOKENETS) seo carnesanfp cone oc. qa00boc Toonp nos scebnomuscee” snoteoc
Scrarer, Wittiam Lutury, M.A. (Secretary to the Society
from January 21st to April 29th, 1903),
Report on the Additions to the Society’s Menagerie in
dicintienyy ISS $cc cnscr ano cenconnpsodoon UB occoousouSOsNaacpUseeOrouD
Report on the Additions to the Society’s Menagerie in
Pebriary OOS yc. s cs ccnecuseescmceemaseenieserocesiocnsoanavensse
Extract from a letter from Major C. Delmé Radcliffe
concerning skins of a Monkey and an Otter from Uganda.
Report on the Additions to the Society’s Menagerie in
Metre hieel’ OO ieae meme vive wieisniete aviecio'e sla siaciels\a slate svielen.e visio vnlaieisielno.vlns
SiapveE, H., Conservator of Forests, Maymyo, Burma.
On the Mode of Copwation of the Indian Elephant ...
Page
180
48
1A
XIV
Stewart, Col. C. E., C.B., C.M.G., C.LE.
On the Original Home of the WMiger (i) ).2ss2-eeee
Sykes, Ernest Rutruven, F.Z.8.
On the Land Operculate Mollusca collected during the
‘Skeat Expedition’ to the Malay Peninsula in 1899-1900.
(JENS 2O3G))sacocota- Hee SNM RG eB Ate Eos Bee ea nas s f+
THOMAS, OLDFIELD, F.R.S., F.Z.8.
Exhibition of a skin and description of a new species
of Monkey, Rhinopithecus brelichi. (Plate XXI_).........
Exhibition of specimens and description of a new species
of Duiker, Cephatophes tgniyer owes. sesdastse-sniaccee eee eos
On some Mammals collected by Capt. H. N. Dunn,
RVAGME CE inet }SOu Catt oa he ree eee eRe Rene eon eee
Watt, Captain F., Indian Medical Service.
A Prodromus of the Snakes hitherto recorded from
China, Japan, and the Loo Choo Islands; with some Notes.
Wooptanp, W., University College, London.
On the Phylogenetic Cause of the Transposition of the
Testes in Mammalia: with Remarks on the Evolution, of
the Diaphragm and the Metanephric Kidney
eee eee er score coes
194
84
hist OF uae s:
1903.—Vot. I.
Plate Page
I. 1. Alestes intermedius. 2. Alestes opisthotenia. 3. Am- |
Phalius LONGWOSULIS: 2. cleo eevee eanteesrnse series:
II. 1. Labeo annectens. 2. Barbus teniurus
ARERR 5
Ill. 1. Barbus progenys. 2. Barbus batesti ee
IV suuitlecrosynomontis Dabestt! \iueniaie = 0 dle eialeels ctorake ieleic einiedils |
V. 1. Anabas pleurostigma. 2. Mastacembelus sclatert.... )
VI.
VII.} Anatomy and Histology of Phascolosoma teres ........ 29
VAD Oe
IX. Figs 1-6. Potamon (Potamonautes) latidactylum. Figs.
7-9. Potamon (Potamonautes) africanum. ...... 6044. 4]
RON SEA y/NATACHNE TOURS CRUG: 6\.15)5 ores hotels vielelnve Vea ott stator le 48
XI. Fig. 1. Petalidium foliaceum. 2. Petalidium jun. 3. Ser-
gestes profundus. 4. S. prehensilis, 5. S. kroyert. oa
Goa Sicunelisl cerns GimiG Nw Sie Ni Uin Glebamanomtardlg. SMW OaWO et areMs st 4
XII. Fig. 1. Sergestes arcticus. 2. S. challengert ..........
PRCLT Le Ger chelate walters Sale eines, 61 Hadden sale ee one gine 113
XIV. Spiders of the Families Pisaurida and Senoculide ....( 45)
XV; | Spiders of the; Pamilly Pisanridies. f2i0 os... ee eles \
XVI. Figs. 1, 2, 8, 4, 7, 8. Phyllochetopterus elioti. Figs. 5,
GS PE TCURG LEM Neve Rt ste at OG oft eRe ols 2.6 FSO Gees 169
PROVE RYU MOCHELOPLEHUBIELIOLE ac Ris: o's ix) che hones) on eral bieies stat ntatel
XVIII. Axis, Atlas, and Proatlas of Gomphognathus and Tri-
HUAN SEDANS LEO ee eee Sao 177
Men PIN CO Umno US JU RLERD: staat, whanau Scthaie ebie em aieie kai ANA ys aaeho Res 192
XX. Mollusca from the Malay Peninsula ......... aie SiARN 194
BONST oy EUIO TAD PEL RECUS UF ELUCNT HM. SUB rian, SRE Pid cain eee oat ate 224
XXII. Osteology of the Cuculiformes ............00.02 0000s 258
XXIII. Polyclads from the Straits of Malacca............000% 301
LIST OF TEXT-FIGURES.
1903.—Vot. I.
Page
1. Hybrid between Equus burchel, 3, and H. caballus, Q ...... 2
2. Upper surface of cerebral hemispheres of Nasalts .........+.. 13
SAnothen, brainvof tu heysame)speciessteraymive oes sirerteiees etalon 13
4, Longitudinal median section of brain of Nasalis.............5 15
5. Upper surface of cerebral hemispheres of Colobus guereza...... U7
6. Longitudinal median section of brain of Colobus guereza ..... ey la
7, Lateral view of brain of Cynopithecus niger...........+s0008- 19
8. Longitudinal median section of brain of Cynopithecus niger .... 19
9. Tentacular wreath of (a) Phascolosoma and (b) Phymosoma.... 88
10. Front view of skull and horns of adult Ram of the Kopet-Dagh
Uital). tu ee hee eee aloo eprom teem aya 103
11. Chondrocarpus aoalson and Chondrocarpus sp. ............ 105
WD, WVAUOGHE LO GOSUMIED, Sean nen nnobcnnonSsunooanoooDOOGONOS 107
13.
14.
15. $Telsons of larve attributed to Thysanoessa longicaudata.:.... 131
16,
Wo
18. Horns of fully adult Elk from Trondhjem, Norway .......... 134
19. Horns of fully adult Elk killed in Bjorndal.................. 184
20. Horns of a full-grown but not adult Elk, killed in Upper
INamidallens, Vols agate aims cr tee RIE eto tick rene 135
91. Horns of young Elk, supposed to be 3 years old.............. 135
22) Tracing of shed horn picked up ati Mo. nee. eee oe 142
93. Horns of a large Elk from Lithuania, in Branicki Museum .... 145
24, Horns of a very old Elk showing degeneration .............. 147
25. Tracing of a cast horn found at Solem in Bangdal by Capt.
TE SARTO as oa oe Seam eben OM 6 arb Si nio HOMO DReOISS 4 5/0.0'd.0 6 148
96) One-homed sh livin ips wach Miurseumayaeiamersin i ibe iereiions 149
97. Side views of heads of (a) Vipera ammodytes f. typica and (6)
VAD DILENECVOTQUES sir ic alle oleic sya seetnM ae Neeenee eves ohailere halter 186
98. Front view of end of snout, showing the lepidosis ............ 186
Or Cr Or
y=! (eS)
Sr Or Or Sr Cr
So ot He Co to
on on
ea
PaO SAE MLE PUOUIESLY cm crnrrrd dale o 1 ARS ace ole a g ele
XVI
Right femur of Struthio haratheodoris, Caudal view ........
Right femur of Strathio karatheodoris. TRostral view ...... oi
. Pelvis of Struthio karatheodoris. Ventral view............ 5
Pelvis of Struthio karatheodoris. Dorsal view ............-.
. Lateral view of the pelvis of Struthto karatheodoris .......44.
. Dissection of Stuhlmannia michaelsent ..... 00. e eves eee ecces
Ventral surface of Bettonia lagariensts ...........05 Aenea a¥e
Termination of the male efferent apparatus of Lettonia lgune a=
ENSIGN, crass Vaulted neiaioneeco ube ter tee FE AMO IAI RISO RCL. ay averers
mopermatophoreiot whanewareus Spiers '\ce oalss ae vir as ais 4s «ale .
Recently-killed RAmoceros simus, sahil Gy onron Se DOr Semon naee
Recently-killed Rhinoceros simus, adult S ....... cece cee
Japanese Long-tailed Fowl. Cock A, photographed April 1905.
Japanese Long-tailed Fowl. Cock B, photographed April 1905.
Left side view of sternum and eon e of Cuculus
CANOPUS RINNE RT ee aves oath Lae Pe utenis Dinu esce oD
Left side view of Senin and shoulder-girdle of Cowa heiraae
Dorsal aspect of the pelvis of Cacomantis merulinus .......4+-
Dorsal aspect of the pelvis of Geococeyx mericanus... 1.6.6.5
IM ORVLC Wy OL SANG ee scccie aerate opelicva wise nce) aysusia'cna/oiaveere| clade eave poyadetavs
Wentral aspectiot same tier. arctan relator o ckaielsiarsy strep vaavaleieys
Topographical diagram Thonn, feather-tracts and c we spaces
in cehematio bird. ahs dvniaetabted Saree teles eaifedel sate ons fotetekevetatands 6
Hye-spots of Riotiyulenes EVANS ..... rd aoeaeereC Sagoo oe Gare
Male organs of Notoplana evansti ...... LY ah cosemeedt ats fe) Speeraeteg? oe
Genital apparatus of Leptoplana malayand ..... 0... cece
. “ Brain-eyes”’ of Semonia penangensis ............2.0sesuve
Genital apparatus of Bergendalia anomala ..........-. ardicrets
BLY CoS POUs) Ol) LeOCESLES ANGUS! 21). erste) sijelo leary as avieta elaine
Kye-spots of Prosthiostomum pallidum ...... siectda ysl Sto ckeyaneaueeaeate
Diagram illustrating Testis Descent . ths Saea clu istt tonat ice SIR
Map to illustrate fie Geographical Misimhunien of the Macon
tteleam cenera Ol MU iiur7a ce rac. aiey~ solsicierstei eg bie ie sles
Map to sinneennte the Geographical Distribution of aN aie
Brachybothriide, and Mecicobothrtid@ oo... cece eee
. Map to illustrate the Geographical Distribution of the Gene
HMO MAGde = so. co olen SRIne ciotciD Ibn Domino a eNeC UIC ees
Map to illustrate the Geographical Mice bution a the iiae
LOTR cleat «s BO OLOD DOU Oem ODI OOh ARCO CCT eee:
Proc. Zoo, Soc.—1903, Voz. I. b
304
306
309
S11
»
jlv
DQ 1-7
517
Oo
ees
LIST OF NEW
GENERIC TERMS
PROPOSED IN THE PRESENT VOLUME (1903, vou. T.).
Page
Acolischnus (Arachn.) ............ 362
Asthenoceros (Vermes) ............ 315
Bergendalia (Vermes) ..........-- 310
Ceratophyllidia (Moll) .........063 250
Chondrocarpus (Copep.) ......... 104
Page
Microsynodontis (Pisces) ......... 26
Notoplana (Vermes) ............+6- 302
Paradossenus (Arachn.)...... 153, 155
Pleurophyllidiella (Moll.)......... 251
Ventriculina (Copep.) ...........+ 106
‘
AD : i
PROCEEDINGS
OF TIIE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
OF THD
ZOOLOGICAL SOCIETY OF LONDON
1908, Vol. I. (January to April).
January 20, 1903.
Prot. G. B. Howes, D.Sc., LL.D., F.R.S., Vice-President,
in the Chair.
The Secretary read the following report on the additions to the
Society’s Menagerie during the month of December 1902 :—
The registered additions to the Society’s Menagerie during the
month of December 1902 were 30 in number. Of these 15 were
acquired by presentation, 1 was born in the Gardens, and 10 were
received on deposit and 4 in exchange. The total number of
departures during the same period, by death and removals,
was 141.
Amongst the additions worthy of notice are two very fine speci-
mens of the One-wattled Cassowary (Casuarius uniappendiculatus),
from New Guinea, deposited by the Hon. Walter Rothschild,
M.P., F.Z.S., on Dec. 30th.
Mr. P. L. Sclater read an extract from a letter from Messrs.
Stagmann, Ksselen & Roos, of Pretoria, dated Dee. 8th, 1902, and
addressed to Major W. H. Birkbeck of the Remount Department,
Johannesberg, from which it appeared that the hybrid Zebra,
placed under the Society’s care by the King on July 19th, 1902
Proc. Zoou. Soc.—1903, Vou. I. No. I. 1
2 MR. J. S. BUDGETT ON [ Jan. 20,
(see P. Z. 8. 1902, vol. ii. p. 225), was the progeny of a male
Zebra and a pony mare. Major Birkbeck added the following
Text-fig. 1.
Hybrid between Equus burchelli, 8, and E. caballus, 9.
particulars:—The hybrid was rising 12 months in July 1900
when he came in. He went home with a pony of which he was
very fond, and if he gets troublesome the company of a quiet
pony will pacify him at once. We gelded him, thinking Her
Majesty might use him, and a stallion hybrid is always a terror.
The late owner Erasmus has tried to claim compensation, but as
the beast was a fair capture he has no case at all. At the
surrender the Boers kept any horses of ours they had and we, of
course, keep theirs.
Mr. J. 8. Budgett, M.A., F.Z.S., gave an account of his recent
journey to Uganda and return by the Nile, which was illustrated
by a large series of photographs taken by him as he went along.
Mr. Budgett made the following remarks :—
The special object of my journey to Uganda was to continue
investigations on the life-history of African fishes and especially
‘
1903. ] HIS JOURNEY TO UGANDA. 3
of Polypterus. “Having made previous attempts to solve this
problem under certain conditions in a confined area, it was
thought that by observing it under varied conditions of latitude
and altitude, new light might be brought to bear upon it.
From Uganda it was possible either to work down the Congo
from its source or to return northwards down the Nile.
On my way to Uganda, and in Uganda, I gathered what infor-
mation I could about the two routes. The southern end of the
Nile Valley really lies at the foot of Ruwenzori, while a short
journey westwards from this point would bring one into the head-
waters of the Congo.
A special inducement to take the Congo route was that infor-
mation might in this journey be obtained about the new animal
Okapia johnstoni, and other interesting forms, believed to exist in
the Semliki Forest.
From information gathered in Uganda, it was clear that it was
useless to hope to meet with Okapi in British territory, and,
moreover, I here learned that the Belgians had found the Okapi
in large numbers in the Welle country.
I found also, what had been very difficult to learn before
leaving home, that the season of the rains and the breeding of
Polypter us were considerably earlier at the source of the Nile than
they were further northwards; that the Semliki Valley was a
most inconvenient place at which to make a permanent camp
by the river-banks, owing chiefly to the scarcity of food; and
that only one species of Polypterus was to be found there, while
at least three species were to be found in the Nile farther to the
north.
The difficulty of taking delicate apparatus through the Congo
Forest to the upper waters of the Congo was incomparably
ereater than to the upper Nile.
And, lastly, it was to be borne in mind that the time of year at
which one might hope to be successful in the main object was
that at which it was well nigh impossible to do much in the way
of collecting the higher Ver tebrata which might be supposed to be
of interest in the Semliki Valley ; for at this time the grass is so
high that moving away from beaten tracks is almost impossible,
while anything smaller than giraffe or elephant is seldom seen.
Bearing these facts in mind, I had little hesitation in deciding to
work down the Nile, striking it at a point farther northwards
than the Semliki River, in or der to take advantage of the lateness
of the season in that region. Accordingly, having fitted out my
safari or caravan at Entebbe, I started for Butyaba, on the west
coast of Lake Albert, on July 10th. I had the advantage of
starting thus along a good road which had just been clear ed for
the greater part of the way, and along which the rest-houses had
been repaired for the convenience “of the Commissioner and
Consul-General of British Kast Africa, Sir Charles Eliot, who
has recently made sucha remarkably rapid journey from Entebbe
to Gondokoro, the frontier station of Uganda on the Upper Nile.
1*
4 MR. J. S. BUDGETT ON [Jan. 20,
At Kampala I diverged, however, for a few days along the old
road to Massindi to the eastward.
T had with me at the start 50 men and boys and my bicycle.
So many books have been written on Uganda, that there is
little need to describe the scenery of these tropical highlands,
especially as Sir Harry Johnston’s wonderfully complete book is
now in everybody’s hands.
Shortly, one may say that, on going northwards from Lake
Victoria, forest is hardly seen after leaving Kampala. We passed
day after day through almost endless elephant-grass, with palm-
groves and papyrus-swamps in the lower parts. The hills are
clothed with clumps and patches of acacia and euphorbias,
while their summits are very frequently covered with huge
granite boulders. There were thunderstorms and rain every
afternoon, and for the first few days I saw little in the way of
animal life: occasionally a Civet cat would cross the path, while
overhead Hornbills and Plantain-eaters of various species were
common. In the valley of the Maangia for a time we were rid -
of the everlasting elephant-grass, and here Cobws thomasi and
Zebra were plentiful. The Cobus thomasi of this region is some-
what different to that met with in the valley of the Nile, the
horns having a wider curve and being stouter and of a lighter
colour, while the animal itself is of a larger build and has more
brilliant markings.
The Maangia River flows northward through wide undulating
plains, covered at this time with hay-like grasses upwards of five
feet in height, and dotted over with very fine acacias, of a cedar-
of-Lebanon appearance and of richest deep-green colour. The
grasses and bushes of the roadside teem with bird-life: as we
rode along, the little Vidua principalis, with his dozen sombre
wives, was a constant companion, flitting just ahead of us for
a mile or more along the road. Likewise the Common Shrike
of these parts (Lanius excubitorius) has exactly the same habit
of driving along in flocks with a caravan as the Corvinella
corvina of the West Coast. In the marshy parts, Scopus wmbrettia
was often seen.
Then, leaving the plain, we struck up over the Bukamva hills,
and at the crest dived into the dense grass at the side of the road,
to travel for four weary hours over a wretched and little-used
track, often obliterated by the tramp of elephants, and where it was
quite impossible to make any progress with the bicycle. Then,
descending rapidly by swampy valleys and thick jungle, we came
suddenly into the new road to Hoima and Butyaba. From here to
Hoima the road crossed the steep hills which form the boundary
between Uganda and Unyoro, passing by abundant plantations of
bananas and through many a lovely valley, at the bottom of which
a stream ran through the richest vegetation, the banks carpeted
with Cannas, winter cherries, and hemlock, while overhead were
many Pterocarpus-trees with blossoms like the Alamanda flower.
As I did not care to get too far from my safari, I would often
1903.] HIS JOURNEY TO UGANDA. 5
bicycle on for an hour, and then sit down in one of these shady
spots, and watch the Mouse-birds hanging like acrobats in attitudes
most quaint, and Sun-birds darting in ‘and out of the great red
blossoms of the Spathodea, while often °a noisy flock of “Pri ionops
plumatus passed hurriedly along. In the more open parts I often
saw several pairs of Ground Hornbills, and each time one had a blue
throat, the other a red one. I can but think that this is a sexual
character, though which is the female and which is the male
I was unable to determine. Other birds seen were Pawocephalus
meyert, a species of Macronyx, Irrisor erythrorhynchus, Hirundo
rustica, and the handsome Snipe Rhynchwa capensis.
A very amusing bird that I watched was Hrythropygia ruficauda,
which is most assiduous in its courting of the female, spreading
its tail before her like a fan.
In the more shady parts one might often see the butterflies, as
I remember seeing them in the forests of Paraguay, covering the
ground with large patches of colour, in flocks according to their
species.
Shortly before reaching Hoima the river Kafu is crossed ;
here a network of papyrus-swamps with good causeways over
them abound in duck, geese, and kingfishers of various kinds.
Then, winding upwards, a high point is “reached from which the
village of Hoima i is seen, and in the very far distance one can
make out the Blue Mountains on the other side of Lake Albert.
At this high point I saw a charming little Widow-bird (Viduwa
hypocherina). Lions round this part are plentiful and somewhat
dangerous, as they usually are in countries where game is not
abundant. At Hoima I heard that several natives had recently
been carried off by the Lions.
From here, two days’ march through the so-called Budonga
Forest brings one to the shores of Lake Albert. This Budonga
Forest is nothing more than rather heavily-wooded scrub. It is
true that in the ravines and gorges there are strips of real forest,
but it is not in any way comparable with the real forest of the
tropics, where the sky can scarcely be seen.
This Budonga woodland teems with herds of Elephant—I
myself caleulated that there were over 200 in one herd which
we came across. Some of the males had enormous tusks, and
these big fellows seem to keep slightly aloof from the rest of the
herd. I knew that we were quite close to this herd, as there were
great roadways through the jungle with quite fresh, smoking
dung; and here J first noticed what struck me many times
subsequently, that when elephant-dung falls on a pathway or
clearing, there within half an hour you will constantly find,
heaped up all round the dung, the earth-workings of a shrew or
mole. What is it the shrew seeks in the dung? Is it the fly-
larve that have been blown upon the dung, or is it the dung
itself? Frequently elephants in these parts appear of a bright
red colour, having covered their bodies with the dust of crushed-
up termite hills.
6 MR. J. S. BUDGETT ON ~- [Jan. 20,
On July 29th I looked from a high point on the road on to
Lake Albert, a vast sheet of glistening water, 1000 feet below,
bordered on this side with level plains of park-land, broken here
and there by lagoons and*swamps, where I was to try first for the
Polypterus.
Of the results of my Polypterws work during this journey I
shall say nothing here; suffice it that I stayed down by the
lake-side from J uly 30th to August 15th, trapping, netting, and
shooting. During this time the ‘fishes most abundantly sae ‘with
were Hydr ocyon for skalii, Alestes baremose, Distichodus miloticus,
Labeo hosei, Bagrus bayad, Hutropius niloticus, Synodontis 1 ignites
Tilapia nilotica, also very large specimens of a Citharinus.
Lates niloticus is frequently caught by the natives here 5 and
6 feet long, usually with the spear. Protopterus and Polypterus
were both obtained here. The River-Tortoise (7'rionyx triunguis),
28 inches in length, and very large specimens of Rana occigtalis
were also common here.
The common Antelopes were Cobus thomasi, Cobus defassa,
Tragelaphus scriptus, Ovibi, and Cephalophus equatorialis. Down
on these lake-side flats the avifauna differs in a marked manner
from that in the highlands. Wyphantornis cucullatus was now
building in hundreds in the water-side bushes ; Laniarius barbarus
and Telephonus senegalus in the low bushes, with Merops albicollis,
Lamprocolius purpureus, Pyromelana flammiceps, Terpsiphone
perspicillata, and Dicrurus assimilis, were the birds most
frequently met with. These birds were seldom seen in the
highlands.
Lanius excubitorius seems to have a curious habit of giving a
peculiar chattering call whenever a wounded animal is near. We
often made use of this indication when tracking wounded beasts,
and I have no doubt of the truth of this fact.
From here I struck due east through the Budonga Forest again
to the Victoria Nile. The journey through this woodland country
was at this time of year most arduous, all the paths being densely
overgrown with rank grass, while in the ravines the creepers and
hanging lianas were a great hindrance to the porters. During the
four days I was in this rank jungle I saw very little in the way of
animal-life except Elephants, a few Baboons, and an occasional
Puff-Adder, one of them 4 ft. 5 in. in length, and a few interesting
insects—Phasmidee and Mantidee.
Plant-life was much more interesting, and almost overwhelming
with its abundance of variety and its beauty.
The handsome Nightjar (Cosmetornis vexillarius) was often seen
at sunset in these forest-camps.
At length we struck the old road from Masindi to Wadelai, and
the bicycle came into use again. At my first camp along this road
there were large numbers of a golden-eyed black Weaver-bird
(Ploceus migerrimus), which I saw nowhere else. In its size and
shape, courting- and nesting-habits, it resembles very closely the
gregarious Hyphantornis cucublatus.
1903. | HIS JOURNEY TO UGANDA. 7
The Masindi road now made straight for a high conical hill,
from the shoulder of which we had this part of Africa laid out
as a map before us. To the south, the Budonga Forest; to the
west, the north end of Lake Albert, with the valley of the White
Nile extending northwards; and immediately below, from east to
west, the valley of the Victoria Nile. Descending from terrace to
terrace, we at length arrived at the village of Fajao, just below
the Mur chison Falls, on the 22nd of August.
This wonderful gorge has been described by Baker, Vandeleur,
and others, and their descriptions are no exaggeration. One looks
down on the swirling, surging water, that, leaving the base of the
falls, sweeps round the hill on which the old fort used to stand,
with a feeling of utter amazement at the vast numbers of leaping
fishes, crocodiles, and hippopotami that have found their way
into this cul-de-sac of the Nile system.
Here I continued my work with more success than on Lake
Albert, the commonest fishes being Alestes baremose, A. macro-
lepidotus, Lates niloticus, Clarias lazera, Tilapia xiblir.
The natives here use enormous wattle-traps, which they set in
certain fixed spots, usually out of the main force of the current,
and often catch very large fish in them.
On August 29th I started again, as this is a most unhealthy
place ; many of my men were on the sick-list, and food was getting
scarce.
Once out of the gorge of the Victoria Nile, we came into open
rolling savannah country of grass and Borassus-palms, baobabs,
and scrubby acacias. Then crossing several rivers with difficulty,
we arrived at Wadelai on September Ist.
During this stage of the journey I noticed several birds not seen
in this part of Africa before : there was Corvinella affinis, Parus
leucopterus, and several species of Capitonidie, all reminding me,
as did the landscape, of the Gambia on the West Coast. Here
also were Melittophagus bullockoides, Macrony croceus, Urolestes
equatorialis, Telephonus and Crateropus.
From Wadelai I sent my porters on to Nimule, about 100 miles
distant, taking my loads and servants down the river by boat.
After a few days’ work at Wadelai, we started down the river
on the 8th of September. The scenery on this part of the Nile is
very charming, the hills in many places coming right down to the
waters edge. Here one sees the process of the er rowth of the sud
in every stage. Beginning with the floating separate plants of
Pistia Bae. the seeds of, first a small floating rush, then of
the “oom soof” grass, settle on and gradually bind together this
carpet of separate plants into a floating island of grass. So
abundant are these floating islands that often we appeared to be
stationary, even when moving at 5 or 6 miles an hour, for all the
visible banks were moving too. Once, however, the mass lodges
against the stationary papyrus, it quickly becomes overgrown by
this, and is converted into permanent swd.
Fishing villages are numerous on this part of the Nile. The
8 MR. J. S. BUDGETT ON [Jan. 20,
natives make very good traps of papyrus-grass, and also hunt the
hippopotamus with long spears with a rope and float of ambatich-
wood attached. Amongst other fishes caught here were Mor-
myrops, Mormyrus, Hyperopisus, and Malapterurus.
Much of this way the hills retreat, and there is nothing seen
but grass floating and grass stationary, not even bird-life to
relieve the monotony. The last 20 miles, however, before reaching
the garrison town of Nimule it is very different. The Nile flows
straight towards the mountains above Nimule, and here widens
into beautiful lagoons covered with water-lilies, in the foreground
sheets of Pistia of the most vivid green, in the background bold
wooded hills. Here and there are rocky islands with schools of
hippopotami basking in the sun; Bee-eaters (Melittophagus
pusillus), the Jacana (Parra africana), and the gorgeous little
Kingfishers (Corythornis cyanostygma) abound. And then the
Nile plunges into the great Nimule gorge, to tumble down
cataract after cataraet, breaking up and pulverizing the floating
vegetation, and issuing again at Fort Berkley free from sud.
I was now getting anxious about catching the Sudan Govern-
ment steamer, which comes up once a month to Gondokoro, and
determined to leave the Nile and go straight overland for
Gondokoro. The actual distance was little over 100 miles, but at
this time of year the difficulties of travelling and crossing over
rivers in flood were such that one could not tell at all how long
the journey would take. After passing through the Nimule
gorge, we came to the affluence of the Assua with the Nile. The
Assua was now in flood, the only way of crossing being by means
of small rafts of ambatch-wood equal to taking one load at a time.
None of my porters were able to swim, and all had likewise to be
crossed on the rafts. After very nearly losing two men down
the rapids, the crossing was completed after eleven hours’ hard
work. Here, again, the bird-life was different. I saw many birds
while on the march that I was unable to identify. There were
great numbers of a Weaver-bird of brownish colour with a white
crown, building innumerable star-like nests made of straight wiry
grasses woven in at a tangent to the nest. There were also seen
in these parts, for the first time, Scoptelus notatus and Crypto-
rhina afra, though amongst these were not seen specimens with
red beaks as was the case on the Gambia.
Just below its confluence with the Assua River the Nile flows on
two sides of a high hill; a fact which strikes one as remarkable,
for the two branches were mountain-torrents of very little depth
of water.
There we left the river, and passed through country with many
villages and a good deal of cultivation, especially ground-nuts and
millet. The aspect of this country of the Madis struck me as
remarkably similar to that of the Gambia: the soil was rich and
sandy, and the nuts produced were of great size. In some of the
valleys we saw quantities of very fine bamboo, while many of the
trees were almost smothered by the beautiful creeping lily
Gloriosa superba.
1903. | HIS JOURNEY TO UGANDA, 9
In one of these villages quite 10 per cent. of the natives had
marked elephantiasis. They were very friendly, and provided me
with whatever I wanted. After three days’ wandering by winding
paths from village to village, we came back to the main path by
the side of the Nile, which for over sixty miles runs along close
under a range of mountains on its western bank. From this point
northwards for some time the beautiful little Parrot Palwornis
docilis was common. The only Antelopes seen in this part of the
journey were Cobus leucotis and a species of Damaliscus. The
grass seemed to get longer and longer, and marching in the early
morning, when the heavy dew was hanging from every blade of
grass in great drops, was most disagreeable.
On September 19th we reached the flourishing village of a well-
known chief named Adimadi. This village was situated in a
hollow on the top of a high hill, with natural rocky fortifications
surrounding it, and overlooking a fertile valley to the east. On
the heights above the village I saw considerable numbers of what
appeared to be a large red Colobus-monkey, a specimen of which
I failed to secure.
Long-horned cattle were plentiful here, and are probably the
same race as the long-horned cattle of Ankoli.
On the hill-sides were numbers of very fine African mahogany-
trees (Aa@ia) and springs of good water. In these trees were many
kinds of Plantain-eaters and Rollers (Coracias caudatus). This
was the first place during the whole journey that I met with any
Rollers. The hitherto daily rainstorms were getting less frequent,
and the dry-season was setting in.
From here we marched through undulating park-like country
with small trees, to a similar isolated group of hills, with the
village of Leju nestling beneath. Here, again, were fine spreading
trees, in which were numbers of beautiful glossy Starlings (Spreo
superbus), and also the King of the Sparrows (Dinemelia dinemelia).
Passing down from the Leju hills again, we marched through
country of a rather barren nature, of rank grass and small
acacias. The whole way the elephant-tracks were very numerous,
and we came suddenly on a herd of twelve with two old tuskers
among them.
The country now became more and more barren, and on
September 22nd we reached the hills again, opposite the Belgian
station of Redjaf. Here I saw several birds I had not seen before,
including Merops nubicus, Vinago waalia, Laniarius erythrogaster,
and, I think, Lanius collurio, though it may have been a different
species.
At Gondokoro I sent back all my porters and Uganda servants,
and after a few days’ work fishing &e., I started northwards, on
the Sudan Government steamer ‘ Abuklea,’ for Khartoum, on
September 27th.
The first few days the steamer passes through firm banks, on
which, notwithstanding the grass, we saw several water-buck
and some buffalo. Many small villages line the banks, while
several old Dervish forts are passed.
10 MR. J. S. BUDGETT ON THE [ Jan. 20,
At Canissa, about 100 miles north of Gondokoro, I changed
into the ‘ Kaibar,’ the post-boat to Khartoum; then, passing
through the sud region in three days, we came to the mouth of
the Sobat and the land of the Shelluks.
On my arival at Khartoum, I set to work to get Arab fisher-
men and servants, fishing-tackle, provisions, &c., and returned in
a few days to Fashoda.
Here I made my final attack on the Polypterus problem. I had
three species of Polypterus to work with, while material was
fairly abundant. However, after several weeks’ work, I finally
packed up my things, and disconsolately returned to England ;
having got a good deal of side-light on the life and habits of
Polypterus, having seen something of the Fauna and Flora of
the most wonderful river in the world, but having again failed in
my principal object—namely, to obtain the early stages in the
development of Polypterus.
In conclusion, I should like to say that, throughout the journey,
I received at the hands of the Uganda and Sudan officials the
most courteous and liberal assistance on all occasions.
The following papers were read :—
1. Note on the Spiracles of Polypterus.
By J. 8. Bupezrr, M.A., F.Z.8.
[Received January 19, 1903. ]
On seeing a letter in ‘The Field’ for November 8th, 1902, by
Mr. Boulenger, in which he says that, after observing Polypterus
in captivity for more than a year, he had not been able to
learn anything concerning the use of the spiracles to Polypterus,
I determined to go over my former observations concerning these
structures and see whether I had by chance been mistaken as to
their use.
I have in captivity a pair of Polypterus senegalus kept in an
aquarium at a temperature of 75° to 80° F. They are quite
tame and regularly take food from a fork.
On December 2nd I watched them for one hour after feeding.
While eating, the spiracles were repeatedly rapidly opened and
closed, though not widely. The movement was apparently inde-
pendent of other masticatory movements. Within the hour each
of the pair came to the surface three times at irregular intervals.
1. Specimen A came to the surface and gulped air with the
mouth ; immediately after leaving the surface, two large
bubbles of air were discharged from under the opercula.
During the descent to the bottom the two spiracles slightly
opened and from each a minute bubble of air issued.
2. Specimen B performed the same movement, but no air was
seen to issue from the spiracles.
1903. ] SPIRACLES OF POLYPTERUS. 11
3. Specimen B repeated the movement, and small bubbles of
air did issue from the spiracles.
4, Specimen A repeated the movement, but no air was seen to
issue from the spiracles.
5. Specimen B repeated the movement violently after some
excitement, and as it met the surface widely opened the
spiracles, forming a triangular aperture, one side being
the side of the head and the two other sides being the two
plates of bone which form the spiracular flap; whether air
passed in or out of the spiracles was impossible to see as
the top of the head was out of the water. No bubbles
passed from the spiracle during descent.
Specimen A repeated the movement, the spiracles did not
open, and no air was seen to issue from them during
descent.
On December 8th I watched them again foran hour. Polypterus
A and B came to the surface for air 8 times, and 4 times the
spiracles were widely opened above the surface of the water, and a
sound produced as of the sucking in of air.
T have often found it convenient to kill Polypterus by piercing
the cranial roof and destroying the brain. During the operation
it is quite easy to stimulate the brain-centres in such a way that
the spiracles are widely opened as described above. It is possible
to stimulate continuously so that the spiracles are retained in the
widely opened condition. I believe, then, that the spiracles are
used to take in and to give out air from the swim-bladder. At
certain times the fish rises quite slowly to the surface in the
horizontal position, when it would be easier for it to exchange
the air in the swim-bladder from the surface of the head than to
turn its head upwards in order to take air by the mouth. By
closing the mouth and opercula, distending the body-wall and
opening the spiracles, I believe the fish is able to inhale air, and |
should suppose that it expires previously during the same move-
ment, as does Protopterus. I think it possible also that in the
very shallow water which this fish frequents at certain times
of year, it may be of use to the fish to change the air in its
swim-bladder in this way. I have often noticed, in changing the
water in a tank in which numbers of these fish are confined, that
when the water is exhausted the spiracles are frequently opened.
The position of the spiracles almost immediately over the long
slit-like glottis is in favour of the view that they are connected
in their functions with the latter. They seem also to be used, as
I at first believed, to let out the excess of air from the pharynx
after the fish has taken air into the swim-bladder, either with the
mouth or with the spiracles.
Observation upon these points is very difficult owing to the
rapidity with which the movement takes place; but the fish 1
have been watching have become very tame, after three years of
captivity, and these movements are now more slow and much
more easily watched.
12 MR. F. E. BEDDARD ON THE [Jan. 20,
2. On the Brains of Nasalis larvatus and of some other Old
World Primates. By Frank E. Bepparp, F.R.S. &c.
[Received January 17, 1903.]
(Text-figures 2-8.)
Among the types of Old World Primates not studied by Messrs.
Kiikenthal and Ziehen *‘ in their otherwise fairly exhaustive survey
of the cerebral convolutions in the Apes, are Vasalis (if it be a
distinct genus), Colobus, and Cynopithecus. I take the oppor-
tunity afforded me by the possession of brains of these three
genera to compare their characters with those of other Old World
genera. The memoir of Kiikenthal and Ziehen has aided me
greatly in this attempt on account of the clearness and accuracy
of their figures, many of which I have been able to verify by an
inspection of brains of the same or allied species. I do not,
however, always find myself able to agree with the selection of
characters which they use to define the various genera of Apes
examined by them’. I cannot distinguish by any tangible
differences the arrangement of the furrows in the genera Macacus,
Cercopithecus, Cercocebus, and perhaps Papio. It appears to me,
in fact, that among the Cercopithecide there are only two plans
of cerebral conformation, one confined to the Cercopithecine and
the other to the Semnopithecine. The facts which I discuss in
the present communication are confirmatory of that view. But
the classificatory results to which they appear to lead are not a
little surprising, and may possibly be regarded as tending to
throw doubt upon the use of the cerebral convolutions as an
index of anything save physiological resemblances.
§. The Brain of NASALIS LARVATUS.
I have been able to compare the brain of the young male
Nasalis larvatus® with those of two other examples, which I owe
to the kindness of Dr. Charles Hose of Borneo, and I find that
there are, as might be expected, some slight differences of detail
in the arrangement of the furrows. The accompanying drawing
(text-fig. 2) illustrates the superior aspect of the cerebral hemi-
spheres of the specimen of Wasalis which forms the subject
1 “Untersuchungen tiber die Grosshirnfurchen der Primaten,” Jen. Zeitschr. f.
Naturw. xxix. (n. s.) 1895, pp. 1-122.
2 For instance, I find that the backward prolongation of the sulcus precentralis,
whose absence is stated as a characteristic of Cercopithecus, is present in C. stairsi.
In Cynocephalus mormon the parallel fissure joins the Sylvian above, so that the
statement “‘a [the parallel fissure] nahert sich dem hinteren Ende von S [the
Sylvian] um S dann im Bogen zu umkreisen”’ is not universal in its application.
In the same species the furrow hitherto lettered 6 (= inferior occipital) is not
“entirely confined to the lateral convexity ”; it also extends ventrally. The fissures
u and a form a complete Y-shaped furrow as in other Old World forms. Messrs.
Kikenthal and Ziehen write :—“ x verschmilzt gar nicht oder nur scheinbar mit w.”
3 Gratiolet (Mém. sur les Plis cérébraux, &c. pl. iv. figs. 1, 2) has figured a brain
of this animal.
1903. BRAINS OF MONKEYS.
Upper surface of cerebral hemispheres of Nasalis.
a, inferior occipital fissure ; c, lateral occipital fissure ; d, intraparietal ;
e, postcentralis ; R, fissure of Rolando; S, Sylvian.
Text-fig. 3.
'
Another brain of the same species.
Letters as in text-fig. 2.
]
3
14 MR. F. E. BEDDARD ON THE [Jan. 20,
of the present communication, while the internal view of one
hemisphere is shown in another drawing (text-fig. 4, p. 15). The
variations that I have noted in the three brains are the
following :—
There is some variability in the exposure on the upper surface
of the brain of the parieto-occipital fissure. This, as will be seen
from the drawing exhibited (text-fig. 2, p. 13), is better marked in
a large female brain, given to me by Dr. Hose, than in the smaller
male brain (text-fig. 3, p. 13). In the third brain there are only
just traces to be observed of this fissure on the superficial view of
the brain. A furrow which I identify with the postcentralis
superior is present here in rudiment (as it is in the Macaques and
other genera) occasionally. It is best developed, but practically
on one side only (the right), in the small male which lived in the
Society's Gardens. In the large female brain there are traces
of the fissure on one side only (the right); the fissure is absent
in the third brain.
The fissure of Rolando curves back and joins the Sylvian
fissure In the large female brain; it does not do so in the two
remaining brains. On the left side of one brain the Sylvian and
the parallel fissures joined superiorly, a state of affairs which is
characteristic of the brains of many monkeys.
On the right hemisphere of one brain only the lateral occipital
fissure was Incomplete, and consisted of the lower arm only, there
being but a faint indication of the upper arm of this Y-shaped
fissure.
These appear to me to be the principal differences in the sulci
of the three brains.
I have compared them carefully with the brains of six species
of Semnopithecus ', of which I owe two to the kindness of Dr. Hose,
while four were extracted from the skulls of specimens which
have lived in the Society’s Gardens.
The species are as follows:—S. maurus, S. femoralis, S. hypo-
leucus, S. rubicundus, S. priamus, and S. entellus. The differences
between these brains and those described by Kiikenthal and
Ziehen ° are slight.
The fissure lettered H by the above-named authors, which is
the presylvian of other writers, is less regular in its occurrence
than it appears to be in Vasahs, but the difference is not sufficiently
marked to permit of any stress being laid upon the fact. In onl
one instance (S. femoralis), and on one side only (the right), did
the Sylvian fissure join the parallel fissure above. The rarity
of this arrangement is exactly as in Vasalis. In one case only
(S. maurus), and also on one side only (the left), were there
indications of the fissure of Rolando joing the Sylvian fissure.
The rarity of this arrangement is again paralleled in Vasalis.
1 Kiikenthal and Zichen (Jen. Zeitschr. 1895, p. 1) refer to the literature, but they
have not included a paper by Lankester (Quart. Journ. Sci. ii. 1865).
2 “ Untersuchungen tiber die Grosshirnfurchen der Primaten,” Jen. Zeitschr. 1895,
p. 1.
1903. ] BRAINS OF MONKEYS. 15
The relations of the parieto-occipital and the Simian fissures
appear to me to be exactly the same in Semnopithecus as they are
in Nasalis. As a rule, they are perfectly distinct from each
other; but in two brains the operculum was more fully developed,
as it is in the Macaques, and thus the two fissures appeared to
coincide,
The postcentralis I found to be always present, and on both
sides, though often asymmetrical.
In four Semnopithecus brains I find a small furrow running
between the intraparietal and the Sylvian, which may represent
the anterior fork of the parallel fissure figured by Kiikenthal
and Ziehen in several Apes (and lettered a’ in their figures), and
stated to occasionally occur in Semnopithecus. The existence of
this fissure, so far as my material allows me to state, differentiates
the brain of Semnopitheeus from that of Vasalis.
Dr. Elliot Smith ' has mentioned in one brain of Semnopithecus
entellus a bifureation of the calearine fissure, such as that which
is constant, or nearly so, in Macacus and Cercopithecus. I have
found this well developed on one side (the left) of a brain of
S. rubicundus, and less marked on the same side of a brain of
S. priamus, and on the right side of a brain of S. femoralis. It
may be observed, however, that in Semnopitheeus the T-shaped
ealearine fissure, when it is T-shaped, is not visible on a dorsal
view of the undivided brain as it is in Macacus. In the largest of
the three brains of Vasalis which I have examined, there were
indications of the same bifurcation on the right side.
Text-fig. 4.
Longitudinal median section of brain of Nasalis.
e, calloso-marginal fissure; Oa, calcarine ; i.p, internal parieto-occipital,
The inferior occipital sulcus is always less in the Langurs than
in the Macaques. In one Semnopithecus brain (the left side of
S. priamus) this furrow showed an unusual character, in that. it
joined the parallel fissure. The same occurred on the right
1 Cat. Physiol. Ser. R. C. S. ed. 2, vol. ii. p. 426.
16 MR. F. E. BEDDARD ON THE [Jan. 20,
hemisphere of a brain of S. hypoleucus. 'This I have not seen in
Nasalis.
This same brain (of S. priamus) and a brain of S. entellus
show on both sides an unusual condition of the inferior temporal
fissure, which is well developed and runs parallel with the Sylvian
and parallel fissures. The more usual condition in this genus isa
much shorter fissure which is more transverse in direction, and
that is the condition which obtains in Wasalis. The fissure is
not figured at all by Kiikenthal and Ziehen, but is mentioned
as being very feebly developed. Another furrow which I find to
vary in the Semnopithecus brains at my disposal is the calloso-
marginal sulcus; it sometimes bends up and cuts the surface of
the brain. In Wasalis it always does.
§. The Brain of CoLoBuS VELLEROSUS.
Broadly speaking the brain of this monkey is very like that
of a Macaque. ‘There are, however, a few small points of
difference, of which one at any rate may be of some little
significance. The resemblance is so close in most particulars that
it is really unnecessary for me to describe the brain in detail.
The drawing exhibited herewith (text-fig. 5, p. 17) will adequately
prove my statement. I may, however, remark that the fissure of
Rolando quite cuts the inter-hemispheral suleus: that the post-
centralis is well marked on both sides: that the precentralis
superior is recognizable and has a direction parallel with the long
axis of the brain. On one side (the right) the Sylvian and
parallel fissures join above, as is so common on both sides with
the Macaques. The occipital lobe is very smooth, as is often the
case with the Macaques, and the lateral occipital sulcus is hardly
marked at all. The cross-piece of the characteristically Macacine
calearine fissure is visible when the brain is inspected from above.
A small fissure, which I have not observed in other Old World
Monkeys, is to be seen on either side, behind the Simian fissure,
and running parallel with the longitudinal axis of the brain.
Kiikenthal and Ziehen figure, but give no name to an apparently
similar fissure (lettered 5) in the brain of Lagothrix humboldti.
I am able to confirm their demonstration of fact. They do not
figure the same fissure in any of the Cercopithecide.
Tam unwilling to seem to emphasize too strongly this point of
likeness between the African Colobus and the New World
Lagothrix ; but I may remind zoologists that likenesses between
Colobus and the New World Monkeys have been pointed out.
More important than this, however, is the light which the
brain-structure of Colobus appears to throw upon its relationship
to other Old World genera. On account of the structure of its
stomach, and for some other reasons also, Colobus has been
associated with Semnopithecus into a subfamily Semnopithecine,
contrasted with the Cercopithecine which embraces the remaining
Cercopithecidee. It should be plain from the statements made in
1903. ] BRAINS OF MONKEYS, 17
the present paper, and from the illustrations which support them,
that so far as its brain is concerned Colobus cannot be placed
in close proximity to Semnopithecus. From this point of view,
Text-fig. 5.
Upper surface of cerebral hemispheres of Colobus guereza.
g
Ca, calearine fissure. Other letters as in text-fig. 2.
Text-fig. 6.
Longitudinal median section of brain of Colobus guereza,
Letters as in text-fig, 4.
Colobus is most emphatically to be placed among the Cercopithecine.
In fact, it is probably safer to follow those zoologists who do not
subdivide the family Cercopithecide at all.
Proc, Zoot. Soc.—1903, Vou. I. No, IL, 2
18 MR. F, E. BEDDARD ON THE [Jan. 20,
§. The Brain of CYNOPITHECUS NIGER.
IT have been able to examine a single brain of the Celebesian Ape,
of which drawings are exhibited herewith (text-figs. 7, 8, p. 19).
The Simian fissure runs completely across and contributes to
a perfect operculum, as in the Macaques. But the brain of Cyno-
pithecus differs from the brains of the latter and agrees with that of
Semnopithecus in the fact that the infraparietal fissure of each side
bends sharply outwards before joining the Simian fissure ; this sug-
gests an exposure of the parieto-occipital, and a very faint groove
(better marked upon the left side) is probably to be looked upon
as a representative of this. The lateral occipital sulcus is a single
fissure, that is to say, it has not the Y-shape exhibited in so many
monkeys. There is, however, a faint depression suggesting the
upper arm of this ¥. The inferior occipital sulcus is in some respects
rather peculiar. Dr. Elliot Smith has justly pointed out} that in
Semnopithecus “the inferior occipital sulcus has dwindled to most
insignificant proportions, and unless the student examines a large
series of brains he will hardly recognize in the little arc around
the lower end of the Simian sulcus the representative of the deep
operculated infra-occipital suleus in the Macaques.” Cynopithecus
niger has an even smaller semicircular representative of this
furrow, which just arches round the lateral termination of the
Simian fissure, and is nowhere near to reaching the posterior
extremity of the occipital lobe. Further than this, a straight
furrow, either connected (left side) or nearly connected (right
side) with this, runs down the temporal lobe for some distance,
exactly parallel with and between the parallel and collateral sulci.
This I take to be the inferior temporal sulcus of brain anatomists.
In the Macaques and Cercopitheci this sulcus is (so far as my
own experience goes) quite constantly represented by a short
furrow at the lower end of the temporal lobe, and this furrow is
a marked character of those brains, and of Cercocebus and Cyno-
cephalus. This more ventrally placed furrow is not to be seen in
my specimen of Cynopithecus. The existence of an upper portion of
the inferior temporal sulcus is not, however, absolutely distinctive
of Cynopithecus. It is indicated in a brain of Cynocephalus
mormon which I have at my disposal, and in Vasalis and Semno-
pithecus. Ina brain of Cynocephalus porcarius this furrow is as
well developed as in Cynopithecus, but on one side only (the
right). But, with this exception, the furrow is nowhere so fully
developed as in Cynopithecus. So far, therefore, it appears to be
characteristic of this genus. It is, however, apparently impossible
to lay much stress upon the reduced inferior occipital sulcus of
Cynopithecus as a point of resemblance to Semnopithecus in the
absence of a large series of brains of the former genus. Messrs.
Kiikenthal and Ziehen figure a brain of Cynocephalus sphinx in
which the fissure is fully as reduced as it is in Cynopithecus or
Semnopithecus. I have already pointed out (supra, p. 12 foot-
1 Cat. Phys. Series Mus. Roy. Coll. Surg. vol. ii. p. 426.
1903.] BRAINS OF MONKEYS, 19
note) that this state of affairs is not universal in Cynocephalus ;
but it has still got to be proved that it is universal in Cyno-
pithecus,
Text-fig. 7,
Lateral view of brain of Cynopithecus niger.
&, inferior temporal fissure; co, collateral; P, parallel.
Other letters as in text-figs. 2, 3.
Text-fig. 8,
Longitudinal median section of brain of Cynopithecus niger,
Letters as in text-fig. 4.
The collateral swlews, as compared with that of the Macaques
and Cercopitheci, is particularly well developed. It has, so to
speak, taken advantage of the feeble development of the inferior
occipital to thrust itself forward upon the occipital lobe, where it
is plainly visible when the brain is viewed from behind; it is also
visible where it curves upwards upon the lateral aspect. It is
the rule among Monkeys for this fissure to be largely concealed
by the cerebellum. Its exposure for its whole length in Cyno-
pithecus appears to me to be a characteristic feature of the brain
of that monkey,
Oo
~
20 ON THE BRAINS OF MONKEYS. [Jan. 20,
The parallel sulcus extends considerably beyond the Sylvian
fissure, and does not—as is so commonly the case with the
Macagues—join that fissure dorsally. An interesting fact about
this fissure is that it bends forward at its dorsal extremity, instead
of being continued on in a straight line. In this bending I see
a point of likeness to the Semnopithect (cluding Vasalis). No
absolute distinction between Cynopithecus and Macacus can be,
however, drawn on account of this furrow, simce Kiikenthal and
Zichen ‘figure a brain of Macacus inwus in which there is this
same hending forwards, and, moreover, a bifurcation of the furrow
superiorly, such as I note in my example of Cynopithecus niger.
The fissure of Rolando presents no noteworthy characters ; it does
not nearly cut the inter-cerebral groove. The postcentralis 1s
better developed on the right side of the brain than on the left,
and is twansverse in both cases. The precentralis superior is, on
the other hand, better developed upon the left side than upon the
right. It is transverse in position. Among Monkeys this fissure
is more commonly parallel with the long axis of the brain.
The median parieto-occipital sulcus, visible when the brain is
bisected longitudinally, presents what I regard as rather an
interesting and suggestive character. This fissure, in the Macaques,
&e., has a distinctly forward inclination, often making an angle
of quite 45° with the vertical. On the oihes hand, in the Raine
pithect this furrow is nearly vertical (a brain of Vasalis) or with
a distinctly backward inclination. In Cynopithecus niger this
same furrow is nearly vertical, but with a slightly backward
inclination, thus resembling the Semnopithecidee more than the
Cercopithecidee.
The calcarine sulcus of Cynopithecus is not at all like that
furrow in the brains of Cynocephalus, Macacus, Cercopithecus,
and Cercocebus. In the four last-named genera it is a T-shaped
sulcus, the cross of the T appearing almost, sometimes in fact
quite, upon the upper surface of the occipital lobe. This furrow is
very character istic of those genera. in Cynopithecus the furrow
is simple and oblique in direction, asit is,as a general sue, among
the Semnopithecide.
Tt may be convenient to tabulate the likenesses shown in the
brain of Cynopithecus to that of Semnopithecus.
The brain resembles that of the Semnopithecit in :—
(1) The form of the intraparietal fissure.
(2) The backward direction of the internal parieto-occipital.
(3) The absence of any junction between the Sylvian and
parallel fissures.
(4) The simple form of the calearine fissure.
(5) The shortness of the inferior occipital fissure.
(6) The presence of a well-marked superior portion, and the
absence of an inferior portion of the inferior temporal
fissure.
The first two characters are absolutely distinctive of Cyno-
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1903. ] ON FISHES FROM SOUTHERN CAMEROON, at
pithecus and Semnopithecus ; the third and fourth very nearly so,
the last two characters are found also in Cynocephalus, but are
more characteristic of Cynopithecus and Semnopithecus.
It will be seen that if one were permitted to base a classification
upon cerebral charactersyCynopithecus would have to be removed
from its position among the Baboons and placed nearer to the
Langurs; this is not too extreme an interpretation of the brain-
characters considered on purely morphological grounds. We may
possibly regard Cynopithecus as occupying a somewhat basal
position with regard to Cynocephalus on the one hand, and
Semnopithecus on the other. For, in fact, its characters occur
in both, though the Baboon-like characters are on the whole less
marked,
3. On the Fishes collected by Mr. G. L. Bates in Southern
Cameroon. By G. A. Boutencur, F.R.S., V.P.Z.5.
[Received November 28, 1902. ]
(Plates I.-V.)
The freshwater fish-fauna of Cameroon is still very imperfectly
known. A small list published by Peters in 1876" and another
by Linnberg in 1895* are the only contributions that have
hitherto appeared on this subject. The collection made by Buchholz
and reported upon by Peters was important as yielding the first
specimen of the curious Pantodon buchholzi, since rediscovered in
the Niger Delta and in the Upper Congo and Ubangi. It has now
been ascertained that this little fish flies or darts through the air,
and is, in fact, a freshwater flying-fish. . Dr. Pellegrin, of the Paris
Museum, has kindly informed me that, according to the notes of
M. J. de Brazza, the specimen obtained in the Congo by this
explorer was caught by means of a butterfly-net whilst moving
like a dragonfly above the surface of the water.
Mr. G. L. Bates, whose previous collections included some very
remarkable Batrachians described in these Proceedings, has now
made, at my request, a rather extensive collection of freshwater
fishes in Cameroon, of which I here give a list, together with
descriptions of nine new species, one of which deserves to be made
the type of a new genus.
The specimens were obtained mostly in the Kribi River, some
15 miles from the sea; others are from a small tributary of the
Campo River, near Efulen, Bulu Country, 1500-2000 feet ; whilst
others again are from the Mvile River, a small stream flowing
southwards into the Campo, at about the same altitude as the
preceding. .
1 Mon. Berl. Acad. 1876, pp. 195 & 244.
© (Efvers. Vetensk,-Ak. Forh. Stockholm, 1895, p. 179.
22 MR, G. A. BOULENGER ON [Jan. 20,
MorMyrip&.
1. PETROCEPHALUS SIMUS Sauv.
bo
. IsSICHTHYS HENRYI Gill. y,
. MARCUSENIUS SPHECODES Sauv.
m oO
. MARCUSENIUS BRACHYHISTIUS Gill.
CHARACINID&®.
5, BRYCON/®THIOPS MICROSTOMA Gthr.
6. ALESTES LONGIPINNIS Gthr.
7. ALESTES INTERMEDIUS, sp.n. (Plate I. fig. 1.)
Depth of body 3 times in total length, length of head 4 times.
Head as long as deep, twice as long as broad ; snout shorter than
diameter of eye; latter 22 to 2 times in length of head ; adipose
eyelid indistinct ; interorbital width 3 length of head; width of
mouth equal to diameter of eye; maxillary not extending quite
to below anterior border of eye; 14 teeth (£) in the upper jaw,
8 in the outer row of the lower jaw; length of lower border
of second suborbital less than diameter of eye. Gill-rakers
moderately elongate, 12 or 13 on lower part of anterior arch.
Dorsal II 8, above ventrals, originating a little nearer end of
snout than caudal, middle branched rays much produced (13 to 1
as long as head) in the males. Adipose fin small, 13 to twice as
distant from the rayed dorsal as from the caudal. Anal IIT 19-21,
the outline very convex in the males. Pectoral shorter than head,
not reaching ventral; latter produced into a long filament in the
males. Caudal forked. Caudal peduncle a little longer than deep.
Scales 33-34 a, 2 between lateral line and ventral. Silvery; a
large black spot on the caudal peduncle and on the median rays
of the caudal; latter yellow, blackish at the end; dorsal black
and red.
Total length 85 millim.
Three specimens from the Kribi River.
Intermediate between A. longipinnis Gthr. and A. tholloni
Pellegr. Differs from the former in the smaller scales, from the
latter in the smaller number of anal rays.
8. ALESTES OPISTHOTHNIA, sp.n. (Plate I. fig. 2.)
Depth of body 34 or 33 times in total length, length of head 4
or 41 times. Head a little longer than deep, twice as long as
broad; snout a little shorter than diameter of eye; latter 22 or
3 times in length of head; adipose eyelid very feebly developed ;
interorbital width 2 or nearly 3 length of head; width of mouth
equal to diameter of eye or a little less; maxillary not extending
quite to below anterior border of eye; 16 teeth (}) in the upper
1903.] FISHES FROM SOUTHERN CAMEROON, 23
jaw, 8 in the outer row of the lower jaw; length of lower border
of second suborbital equal to diameter of eye. Gill-rakers
moderately elongate, closely set, 20 to 23 on lower part of anterior
arch. Dorsal IT 8, entirely behind base of ventrals, originating a
little nearer CEN than end of snout, twice as deep as long.
Adipose fin small, 24 or 25 times as distant from the rayed dorsal
as from the caudal. Anal ITT 13-15. Pectoral as long as head,
reaching ventral or nearly so far, Caudal forked. Caudal peduncle
as long as deep. Scales 25 a, 2 between lateral line and ventral.
Silvery, brownish on the back; a black humeral spot and a black
lateral band, commencing under the dorsal and extending on the
median rays ‘of the caudal : dorsal, caudal, and ventral fins lemon-
yellow.
Total length 130 millim.
Four specimens; Kribi River and Mvile River.
Very closely allied to A. fuchsii Blgr. and A. kingsleye Gthr.
Distinguished from the former by the less massive form and the
longer pectoral fin; from the latter by the more posterior position
of the dorsal fin and the larger eye; from both by the more
numerous gill-rakers,
9, ALESTES MACROLEPIDOTUS Cuy.
10. DisticHopus NovrospiLus Gthr.
11. XenocHaArAx sprtuRuS Gthr,
CyPRINID&.
12. LABEO ANNECTENS, sp. n. (Plate II. fig. 1.)
Body compressed, its depth nearly equal to length of head, 43
to 5 times in total length. Head 1} as long as broad ; snout
rounded, strongly projecting beyond the mouth, with numerous
nuptial tubercles ; eye supero- -lateral, in the second half of the
head, its diameter 5 to 7 times in length of head, 24 to 34 times
in interocular width ; width of mouth, with folded lips, 2 to 2
length of head ; rostral flap and posterior border of lip feebly
denticulated; inner surface of lip with numerous feeble, transverse
plice ; a small barbel hidden in the folds at the sides of the
mouth. Dorsal III 9-10, with strongly notched border; the
longest ray equals the lenoth of the head and twice that of
the last; the fin equally distant from end of snout and from root
of caudal. Anal IL 5; longest ray } length of head. Pectoral
rounded, a little shorter than head, not reaching ventral; latter
reaching or nearly reaching vent, ie first ray falling angen the
ninth (sixth branched) ray “of the dorsal. Caudal deeply forked,
with pointed lobes. Caudal peduncle 13 as long as deep. Scales
36-394, 3 between lateral line and ventral, 12 round caudal
peduncle. Olive above, whitish beneath, with a rather indistinct
darker lateral band ; fins greyish,
Total length 210 muillim.
24 MR. G, A. BOULENGER ON [Jan. 20,
Two specimens from near Efulen.
This new species, allied to LZ. parvus Blgr., is interesting as
completely connecting Labeo with Tylognathus, and showing that
the latter genus can no longer be distinguished.
13, BARBUS PROGENYS, sp. n. (Plate III. fig. 1.)
Depth of body 33 times in total length, length of head
32 times. Snout rounded, depressed, 33 times in length of
head ; eye supero-lateral, its diameter equal to the inter-
orbital width and 5 times in length of head; mouth large, its
width 32 times in length of head, lower jaw projecting beyond
the upper; lips well-developed, lower interrupted on the chin ;
barbels two on each side, anterior # diameter of eye, posterior as
long as eye, the distance between them $ diameter of eye. Dorsal
TIT 10, last simple ray feeble, not stronger than those following,
2 length of head; free edge of the fin emarginate; its distance
from the occiput less than its distance from the caudal. Anal IT 5,
longest ray 4 length of head. Pectoral 3 length of head, not
reaching ventral; latter slightly posterior to origin of dorsal.
Caudal peduncle 1% as long as deep. Scales 33 2, 23 between
lateral line and ventral, 12 round caudal peduncle. Silvery,
brownish on the back, dorsal and lateral scales brown at the base ;
dorsal and anal edged with blackish, the former with an ill-defined
dark band across the middle; other fins whitish.
Total length 180 millim.
A single specimen from the Kribi River.
Allied to B. bowkeri Blgr., from Natal. Distinguished by the
more anterior dorsal fin with 10 branched rays, the shorter anal
fin, the narrower interorbital region.
14, BARBUS TENIURUS, sp. n. (Plate IT. fig. 2.)
Depth of body 33 to 34 times in total length, length of head
23 to 4 times. Snout rounded, 34 to 4 times in length of
head; diameter of eye 4 times in length of head, interorbital
width 21 to 24 times; mouth inferior, its width 3 to 33 times
in length of head; lips moderately developed, interrupted on
the chin; barbels two on each side, anterior 14, posterior 14
to 2 diameters of eye, the distance between them equal to diameter
of eye. Dorsal IIT 8, last simple ray ossified and moderately
strong, but much thicker than the first branched ray, a little
shorter than head; free edge of the fin emarginate; its distance
from the occiput much less than its distance from the caudal.
Anal III 5, longest ray 2 to # length of head. Pectoral # to 4
length of head, not reaching ventral; latter a little posterior to
origin of dorsal. Caudal peduncle 13 as long as deep. Scales
24-27 aoe 23 between lateral line and ventral, 12 round caudal
peduncle. Olive-brown above, yellow on the sides and below; a
series of black dots on the lateral line, and a black band on each
side of the caudal peduncle; fins white.
bo
i |
1903. | FISHES FROM SOUTHERN CAMEROON,
Total length 120 millim.
Several specimens from the Kribi River and from Efulen.
Allied to B. camptacanthus Blkr. and B. potamogalis Cope.
Differs from both in the stronger third simple dorsal ray, the
more numerous scales in the lateral line (24-27 instead of 21-24),
and the coloration. From the first in the longer posterior barbel,
from the second in the longer barbels, the smaller eye, and the
broader interorbital region.
15. BarBus BATESI, sp. n. (Plate III. fig. 2.)
Depth of body equal to length of head, 33 times in total length.
Snout rounded, 3 times in length of head; diameter of eye 5
times in length of head, interorbital width 3 times ; mouth
inferior, its width 4 times in length of head ; lips well-developed,
lower continuous; barbels two on each side, subequal, nearly
twice as long as eye, the distance between them half their length.
Dorsal IV 8, last simple ray strong, bony, not serrated, slightly
curved, 2 length of head; free edge of the fin emarginate ; its
distance from the occiput less than its distance from the caudal.
Anal IIL 5, longest ray 3 length of head. Pectoral # length of head,
not reaching ventral ; latter below anterior rays of dorsal. Caudal
peduncle 14 as long as deep. Scales 30 a 3 between lateral line
and ventral, 12 round caudal peduncle, - Brownish above, the
scales darker at the base, white beneath ; fins greyish.
Total length 235 millim.
A single specimen from the Kribi River.
This is the first discovered West African representative of the
B. bynni group. Its nearest ally is B. tanensis Gthr., from the
Tana River, East Africa.
16. BarBus KESSLERI Stdr.
17. Barus GurrRALI Thomin,
18. BARILIus UBANGENSIS Pellegr.
19. BARILIUS KINGSLEY Bler.
SILURID!.
20. CLARIAS LIBERIENSIS Stdr.
21. CHRYSOBAGRUS LONGIPINNIS Bler.
22. AUCHENOGLANIS BALLAYI Sauv.
23. AMPHILIUS LONGIROSTRIS Blgr. (Plate I. fig. 3.)
Anoplopterus longirostris Bouleng. Ann. & Mag. N. H. (7) viu.
1901, p. 447.
Depth of body 9 times in total length, length of head 4 times.
Head longer than broad; eyes small, in the second half of the
head, two diameters apart; interocular width 2 length of snout,
26 MR. G. A. BOULENGER ON [Jan. 20,.
which is rounded and projects a little beyond lower jaw ; posterior
nostril midway between eye and end of snout; premaxillary teeth
forming a very short band, measuring about 1 width of mouth ;
maxillary barbel # length of head, reaching root of pectoral; mandi-
bular barbel 4 length of head. Dorsal I 6, nearer end of snout than
root of caudal, first ray $ length of head. Adipose short, as long
as dorsal. Anal I 5, midway between root of ventral and root of
caudal. Pectoral a little longer than ventral, $ length of head.
Caudal forked. Caudal peduncle 14 as long as deep. Dark olive-
brown above, mottled with black, white beneath ; dorsal, pectorals,
and ventrals light, with two transverse series of blackish spots ;
caudal whitish, with some black spots, black at the base, with a
large black blotch on each lobe.
Total length 77 millim.
A single specimen from hills in the Bulu country, near Efulen.
24, SYNODONTIS OBESUS Bler,
MIcROSYNODONTIS, gen. nov.
Closely allied to Synodontis, differing only in the absence of a
free orbital border and of suborbital bones, in the more elongate
form and the rounded caudal fin, and in the curious modification
of the transverse processes of the fourth vertebra.
25. MIcROSYNODONTIS BATESII, sp. n. (Plate IV.)
Depth of body 5 to 6 times in total length, length of head 43
to 5 times. Body subcylindrical or feebly compressed in the
precaudal region, strongly compressed behind; vent in the middle
of the total length. Head broader than deep, 14 to 14 as long
as broad, without ridges or keels; skin on vertex, occiput, and
nuchal shield adherent to the finely rugose bones; a small frontal
fontanelle ; eye directed upwards, in middle of head, its diameter
6 or 7 times in length of head, twice in interorbital width. Lips
moderately developed ; maxillary barbel simple, nearly as long as
head; mandibular barbels with long, slender branches; outer
mandibular barbels + length of head, inner 2. Premaxillary
teeth small, forming a villiform band; mandibular teeth much
shorter than the eye, 20 to 30 in number. Gill-cleft restricted to
the sides. Occipito-nuchal shield a little longer than broad,
ending in two rounded processes. Humeral process narrow,
sharply pointed, rugose. Skin smooth. Dorsal I 6; spine strong,
straight, striated but not serrated, about } length of head. Adi-
pose dorsal low, elongate, 2 to 3 times as long as its distance from
the rayed dorsal. Anal III-IV 8-9. Pectoral spine § to 3 length
of head, strong and striated, with feebly serrate outer edge, with
12 to 17 strong retrorse teeth on the inner side. Ventral not
reaching anal. Caudal rounded. Dark brown or blackish, with
5 yellowish cross-bars above, the first on the occiput, the second
at the base of the dorsal; the body lower down with round
yellowish spots ; throat and belly greyish, spotted or marbled wit’.
1903.] FISHES FROM SOUTHERN CAMEROON, 27
dark brown; fins spotted with dark brown, caudal yellowish at
the base.
Total length 100 millim.
Several specimens from the Mvile River.
The vertebral column consists of 38 vertebre, 13 pracaudals
and 25 caudals. ‘The first 7 are completely united, and the
transverse processes of the 4th, which form the spring-mechanism
in Synodontis, have a very extraordinary form. This process bears
a spheroidal expansion in front, whilst behind, where it presses
against the bladder, it is slightly excavated or cup-shaped. It
may be described as similar to its homologue in Synodontis, but
with a large, bell-shaped, bony knob attached to its anterior surface.
The air-bladder is large, as in Synodontis. The male genital gland
is very peculiar, being lacerated into numerous digitiform lobes,
CYPRINODONTID&,
26. HAPLOCHILUS SEXFASCIATUS Gill.
Epiplatys seafasciatus Gill, Proc. Acad. Philad. 1862, p. 136
(Gaboon R.).
Pacilia sexfasciata Peters, Mon. Berl. Ac. 1864, p. 396
(Liberia).
Haplochilus infrafasciatus, part., Giinth, Cat. Fish. vi. (1866),
pp. 313 & 357 (Old Calabar).
Haplochilus sexfasciatus Giinth. 1. ¢.
Lycocyprinus sexfasciatus Peters, Mon. Berl. Ac. 1868, p. 146
(Gaboon).
Epiplatys infrafasciatus Cope, Proc. Amer, Philos. Soc. xi.
1871, p. 457.
Haplochilus infrafasciatus Steind. Notes Leyd. Mus. xvi.
1894, p. 76 (Liberia); Lonnberg, Gifv. Vet.-Ak. Foérh. Stockh.
1895, p. 188 (Cameroon).
27, HAPLOcHILUS ELEGANS Bley,
OPHIOCEPHALID,
28. OPHIOCEPHALUS oBscURUS Gthr.
ANABANTIDA.
29, ANABAS MACULATUS Thomin.
30. ANABAS PLEUROSTIGMA, sp. n. (Plate V. fig. 1.)
Closely related to A. kingsleye Gthr., but snout longer, as long
‘as the eye in the adult, at least two-thirds the interorbital
width. Dorsal XIV—X VI 10-11; anal VITI-IX 10-11. Scales
27-29 =; lateral line 14-17/10-12. A large round blackish spot
on the middle of the side, above the extremity of the pectoral fin ;
no dark spot at the base of the caudal fin,
Total length 170 millim.
Several specimens from the Kribi River.
28 ON FISHES FROM SOUTHERN CAMEROON, [Jan. 20, .
CICHLIDA.
31. PELMATOCHROMIS BATESII Bler.
Recently deseribed from the Benito River.
32. PELMATOCHROMIS SUBOCELLATUS Gthr.
33. TILAPrA LATA Gthr.
MASTACEMBELIDE.
34, MASTACEMBELUS LOENNBERGII Bler.
28 dorsal spines. Length of head 33 times in its distance from
vent,
35. MASTACEMBELUS SCLATERI, sp. n. (Plate V. fig. 2.)
Depth of body 12 to 13 times in total length, length of head
(without rostral appendage) 6 to 7 times; vent equally or nearly
equally distant from end of snout and base of caudal; length of
head 24 to 2? times in its distance from vent, and 4 to ? im its
distance from first dorsal spine. Snout 3 times as long as eye,
ending in a trifid dermal appendage which is longer than eye;
cleft of mouth extending hardly to below nostril; a strong pre-
orbital and two strong preopercular spines. Vertical fins united
with rounded caudal. Dorsal XXVI-XXVII 85-90; spines
very short, Anal If 80-90. Pectoral not quite 3 length of head.
Seales very small, 19 or 20 between origin of soft dorsal and
lateral line. Olive-brown, whitish on the belly; a dark band on
each side of the head, passing through the eye, sometimes
continued on the anterior part of the body; a more or less
distinct series of large, dark, light-edged ocelli along the base of
the dorsal.
Total length 225 millim.
This new species, named after our retiring Secretary, Mr. P. L.
Sclater, in recognition of many favours received from him during
his tenure of office, is based on four specimens from the Mvile
River. J/. sclateri differs from I. marchii Sauv. in the more
numerous dorsal spines, from JZ. eryptacanthus Gthr., liberiensis
Blgr., and loennbergi Blgr. in the larger head, and from JM. con-
gicus Blgr. in the presence of only two anal spines and the still
smaller scales.
EXPLANATION OF THE PLATES.
Puate I.
Fig. 1. Alestes intermedius, p. 22, natural size.
2. Alestes opisthotenia, p. 22, 3.
3. Amphilius longirostris, p. 25, natural size.
3a. # Z Upper view of head and pectoral fin, x 13.
35. ss 33 Lower view of head, X 1}.
Prats II.
Fig. 1. Labeo annectens, p. 23, +.
2. Barbus teniurus, p. 24, natural size.
VW K.H. del
IPAS, ISOS vO. LP, Vu.
JXNUNVOMNESAINID! ISUIL'S WO ILO” Ole
PRAS COLO'S OMA MERGES .
P ZS, 19038, vol LL Vil.
Bale & Danielsson. Lt hath.
W.K.EL. dell.
JRNUNIPOIMNS FNM) Tels WOW, '© Cx Or
PHASCOLOSOMA TERES.
PA. So LGOS VO), f JP Wa.
Bale & Danielsson Lt? lith
ZUINUAIE OIE ZNINUD) SUIS EOI GX OF
PIELAS COO SOMA We IRIS -
1903.] ON THE ANATOMY OF A NEW GEPHYREAN WORM, 29
Prate III.
Fig. 1. Barbus progenys, p p. 2A, 7 5.
2. Barbus batesii, p. 25, 3 B
Prats IV.
Microsynodontis batesii, p. 26.
a. The whole fish, natural size.
6. Mouth, x 2.
ce. Skeleton, natural size.
d. Upper view of skull, shoulder-girdle, and nuchal shield, x 1}.
e. Side view showing air-bladder and anterior part of vertebral column, X 2.
f. Left transverse processes of coalescent anterior vertebrae, X 3.
PLATE V.
Mev 1. Anabas pleurostigma, p. 27, &.
Mastacembelus sclateri, p. 28, ‘natural size.
On the Anatomy of the Gephyrean Phascolosoma teres,
n.sp. By W.K. Hurroy, M.A., M.B., Senior Demon-
strator of Anatomy in the University of Glasgow.
[Received November 6, 1902. ]
(Plates VI-VIII.’ & Text-figure 9.)
Some time ago, while engaged in dredging-operations in the
Firth of Clyde, Dr. J. F. Gemmill obtained two specimens of the
Gephyrean worm figured in Plate VI. fig. 1. These he handed
over to me, and at his suggestion I undertook the task of
identifying them. My examination having led me to the con-
clusion that I was dealing with an animal hitherto undescribed,
it seemed advisable to give some account of its anatomy; and
this I have essayed to do in the following pages.
Unfortunately the presence of sand in the alimentary tract,
no less than the leathery nature of the animal’s skin, formed an
almost insuperable obstacle in the way of obtaining continuous
series of sections: as a result, my account is in some respects
incomplete. The worm was dredged from a depth of 60 fathoms,
nine miles to the south-west of the Corsewall Light. The bottom
was fine mud.
External Characters and Integument.
The worm is shaped like a very long-necked Florence oil-flask
and measures 40 mm. in length. Since, however, in both indi-
viduals examined the introy ee was almost completely i invaginated,
in order to estimate the true length probably 8 mm. must as
added to this figure. The body of the worm is seen (Plate VI.
fig. 2) to consist of three portions which differ externally, and are
found upon dissection to have definite relations to the contained
viscera.
First, occupying the anterior third or more of the animal’s
1 For explanation of the Plates, see p. 40.
30 MR. W. K. HUTTON ON THE [Jan. 20,
length, and containing within it the more important organs, comes
fa airly stout and muscular portion 2 mm. in diameter (Plate VI.
fig. 1, A). This gradually passes posteriorly into a much slighter,
less muscular, and longer part, which extends for more than the
middle third of the length and measures 1 mm. in thickness ;
while lastly, there is a short, thick, ellipsoidal piece, measuring
5 mm. by 2°5 mm., having very thin walls which are transparent
enough to allow the coils of gut within to be dimly seen. This
terminal portion is quite sharply demarcated from the rest of the
body, and forms a sack in which the coiled portion of the intestine
(Plate VI. fig. 2, G) is contained.
The colour of the spirit-specimens is a pale yellow over the
anterior three-fourths of the animal’s length; on the thick
terminal part the tint is a dirty bluish white, the contents of the
intestine appearing through the body-wall as brownish markings.
The skin has a slightly iridescent sheen; it is entirely devoid
of papille and is of a leathery toughness. It is studded with the
openings of innumerable cuticular glands (Plate VII. fig. 6, C.G.).
These glands (Plate VI. fig. 3) are spheroidal, and measure ‘05 mm.
in diameter. Each is lined with a single layer of rather flattened
cells (LL), and the majority are completely filled with small yellow,
highly refractive granules (I). The glands on the introvert are
very much smaller, and granules are not so plentiful in them as
elsewhere.
Though definite papille are not found, tegumentary appendages
are represented on the introvert by chitinous hooks. These are
arranged in a series of ten transverse rows (Plate VI. fig. 4).
The individual hooks vary in length (Plate VIT. fig. 5) from -06
to:08 mm.; they are brown in colour, almost black at the tip, are
but slightly curved, and the isolated hooks shown in the figure
are grooved on their concave edge for about two-thirds of their
length (Plate VII. fig. 5, F). Each hook has a small multicellular
epidermal core. The first part of the introvert, that nearest the
“head” of the animal, is perfectly smooth, and, when invaginated,
abuts against the outer surface of the tentacular crown through-
out two-thirds of its length; for the remaining distance, however,
the tentacles touch the hook-bearing part. Laceration of the
tentacles during evagination and invagination is averted in some
Sipunculids by the presence of a “collar,” which intervenes
between these delicate structures and the rough introvert. Here
there is no such Jprovision; the end is gained by a transverse
furrowing of the skin on the hook-bearing introvert during
invagination ; and by this means the hooks, projecting from the
base of the groove, barely come into contact with the tentacles.
An indication of this condition is represented in fig. 6, H.I.
(Plate VII.).
General Arrangement of Organs.
Figure 2 (Plate VI.) is a drawing of a dissection of the worm.
It has been opened along the ventral aspect, and the two retractor
1903. | ANATOMY OF A NEW GEPHYREAN WORM. 31
muscles (D.R., V.R.) of the left side have been cut through. The
anterior part of the alimentary canal has been twisted aside and
turned so that it presents to view its dorsal surface. The intro-
vert is almost completely invaginated, the real “head” of the
worm being about the position of the two eye-spots (E).
The internal aspect of the body-wall has a glancing appearance
like mother-of-pearl ; the longitudinal muscles form a continuous
sheet with no division into strands. There are fowr retractor
muscles (D.R., V.R.), two dorsal and two ventral. The latter are
more than twice the length of the former, stretching from the
termination of the anterior muscular part of the body (between
A & B, fig. 1) to the anterior part of the cesophagus, where they
fuse with the dorsal muscles to form a sheath for that tube. The
arise, one on either side of the nerve-cord. The dorsal muscles
are attached to the body-wall on either side of the hind-gut, a
little way behind the anus. In fig. 2 the only part of the
nervous system seen is the eye- spots, which are visible in a
dissection as a couple of black specks shining through the walls
of the introvert.
The alimentary canal (G.F.), with the exception of the anterior
part of the cesophagus, is shown in full. It is, like the worm
itself, divisible into three parts, an anterior and a posterior
portion, comparatively straight, and a middle piece (G) exceed-
ingly coiled.
In the drawing, for the sake of clearness, that part of the gut
(F) contained in the thin portion of the worm is shown as though
absolutely straight; it is really shghtly crumpled. Nowhere is
there any intestinal spiral; there is no spindle-muscle, and the
intestine is entirely free except for about 2 mm. behind the anus,
where the rectum is attached to the body-wall by thin radially-
disposed strands of muscular fibre, which in sections (Plate VII.
fig. 6, M) appear to divide the colon into dorsal and ventral
portions. The nephridia are noticeable objects in a dissection
(fig. 2, B.T.) on either side of the anus. All that is visible of
the blood-vascular system is the tortuous dorsal vessel (H). The
generative organs are visible only in microscopic preparations.
Musculature.
The dermo-muscular tube has the arrangement common to the
Phascolosomidee, Underneath the integument there lies a layer
of circularly-disposed fibres (Plate VIT. fig. 6, R.M.), and beneath
this again one of fibres having a longitudinal direction (L.M.).
As already mentioned, this latter layer 1s a continuous one, with no
division into strands such as are seen, for example, in the genus
Sipunculus. The musculature is thickest and strongest in the
anterior third of the animal’s body ; becoming thinner as the
body narrows, it reaches the extreme of tenuity over the ellipsoidal
posterior part, the thinning of the tube appearing to take place at
the expense of the circular layer,
32 MR. W. K. HUTTON ON THE [ Jan. 20,
Avound the anus and the external nephridial apertures, both
layers furnish special sphincter-fibres ; while along either side of
the nerve-cord, as is afterwards noted, there runs a slender longi-
tudinal strand of muscle-fibre (N.C., fig. 6).
The general arrangement of the four special retractor muscles
of the introvert and of the radial rectal muscular attachments
have already been described ; there remains only, in regard to the
former, to speak of the special appearance presented by the
muscular fibres in this worm. On comparing the ventral
retractors with the general longitudinal musculature, one is
struck by the great similarity of texture presented by both. A
glancing, bright suggestion of firmness, reminding one of the
surface of fresh tendon among the higher forms, is noticeable.
The dorsal retractors, on the other hand, have a dull, almost a
translucent appearance. Microscopical examination of transverse
sections reveals little difference between the two sets of muscles,
save only that in a given area fewer muscular fibres are present
in the dorsal muscles, and the amount of Intermuscular connective
tissue appears to be greater than in similar sections of the ventral
set (Plate VII. fig. 7, A & B). On the whole, both from the
relatively marked shortness of the dorsal muscles and from the
appearances just described, I incline to suspect that they cannot
be functionally very active. They contain a sufficient amount of
connective-tissue between the proper muscular bundles to make
the suggestion that they are in a state of metamorphosis and
regression at least probable (wide J, Bland Sutton’s ‘ Ligaments,’
&e., 2nd ed., 1897, p. 3 et seq.).
Alimentary Canal.
Surrounding the mouth, which hes at the bottom of the depres-
sion formed by the union of their bases, are several simple
leaf-shaped tentacles. So far as could be ascertained by the
examination of a series of sections, 15 tentacles are present, and
of these 11 are longer than the others. Hach tentacle contains a
branch of the blood-sinus and a slender nerve (Plate VII. fig. 8).
This figure, which represents a typical transverse section of a
tentacle, shows that on the mesial aspect the cells covering its
surface are narrow, long, arranged in palisade form, and are
ciliated, while those of the opposite side form a simple columnar
epithelium. For the greater part of their extent the tentacles
are free, but round the mouth their bases are fused together, and
form a series of gutters which lead to the oral aperture.
The mouth itself lies on the top of a papilla, whose sides are
grooved by the above-mentioned furrows. The cells lining the
grooves are cubical and ciliated, and are continuous with the
ciliated epithelium on the mesial aspect of the tentacles. Succeed-
ing the mouth is a short cesophagus with very muscular walls; in
section (Plate VII. fig. 10) it appears to be star-shaped, the lumen
of the tube being encroached upon by the inward projection of a
1903.] ANATOMY OF A NEW GEPHYREAN WORM. 33
series of papillee whose epithelial covering is apparently devoid of
cilia.
For the reason mentioned at the beginning of this paper, it was
no easy matter to gain a certain knowledge of the histology of the
alimentary tract. Figure 9 A (Plate VII. ) shows the character of
the epithelium of the mid- -gut generally : lying upon the thin mus-
cular wall are narrow cells of varying length, with a basal nucleus
and granular cytoplasm; the outlines between the several cells
were in many places most indistinct. The epithelium of the rectum
(fig. 9 B) is composed of cubical cells, with a distinct margin on
the surface next the lumen of the gut. They possess cilia, and
were the only cells in the alimentary tract in which the presence
of a ciliated border could with certainty be determined,
Circulatory System.
This consists of a dorsal vessel, a sinus surrounding the base of
the tentacles, and tentacular vessels.
The dorsal vessel is single, and though convoluted it is devoid
of diverticula: coursing along the dor Aol wall of the fore gut, it
surrounds the base of the fentacles with a cireular sinus into
which the brain dips (Plate VIII. figs. 11, 15, 16); from this
circular vessel tentacular sinuses arise, they form noticeable
objects in sections of the tentacles (Plate VII. fig. 8, B.8.).
Nervous System.
This is of the usual type, and consists of a brain, a ventral
nerve-chain, and a couple of sense-organs.
The brain is a small, somewhat cordiform body, measuring in its
long axis, which corresponds with that of the animal, three-tenths
of a millimetre; it les on the dorsal wall of the pharynx at the
base of the tentacles; it is surrounded by a fine capsule of con-
nective-tissue, and the anterior surface in addition is covered by
an epithelium (Plate VIII. fig. 13, E.), which is continuous with
that of the outer non-ciliated aspect of the tentacles; laterally, the
muscular tube formed by the union of the retractors abuts upon it
(fig. 15, R.M.); below the brain is the blood-sinus (B.S., figs. 11,
13, 15) from which the tentacular vessels arise.
The majority of the cerebral cells are small, with a clear cyto-
plasm and relatively large nuclei; they are massed anteriorly,
dorsally, and posteriorly: the central and greater part of the
ventral aspect of the brain are composed of fibres, but behind
the point of origin of the circumcesophageal commissures there
are to be seen in the lower cortical part several very large
ganglion-cells (G.C., fig. 11): these giant cells measure ‘03 mm.
in diameter; some are pyriform, others irregular in shape, and
these appear to be multipolar, unlike those of Phymosoma
(Shipley, No. 7%"), but on this point I cannot be absolutely
definite.
? This number has reference to the list of authors quoted on p. 40.
Proc. Zoou. Soc.—1903, Vou. I. No. III. 3
34 MR. W. K. HUTTON ON THE [Jan. 20,
From the brain two pairs of nerves arise. ‘The first pair
(Plate VIII. fig. 16, B), which are very slender, take their origin
from the ventral aspect of the ganglionic mass just anterior to the
eye-spots; from them twigs pass to supply the epineural canal and
apparently also the two dorsal tentacles. The remaining tentacles
receive their nerves from the two stout cir cumesophageal con-
nectives (fig. 16, A), which constitute the second pair of nerves,
and arise ventro-laterally behind the eye-spots, coursing round
the base of the tentacular crown, between it and the muscular
tube formed by the fusion of the retractor muscles. Ventrally,
the circumecesophageal commissures unite to form the nerve-cord.
This shoavs no trace of segmentation. Dorsally it is composed of
fibres, while ventrally nerve-cells are distributed evenly along its
whole length. It is accompanied laterally by two fine muscles, as
is usual in the Sipunculide.
Sense-Organs.
In tracing back series of sections, there becomes apparent on
the dorsal wall of the pharynx a patch of long-celled columnar pig-
mented epithelium. The cells of this layer are found (Plate VIII.
figs. 12, 13, 14, 15, 8.E.) to be the dorsal wall of a slit-like canal,
whose ventral wall is formed by cells which, on the one hand, cover
the brain in front, and on the other are continuous with the non-
ciliated epithelium which clothes the outer aspect of the two
dorsal tentacles (8.V.), which are seen in fig. 12 to have united
along their mesial margins.
Posteriorly the canal thus formed (which may be called the
epineural canal) becomes very narrow centrally, and ends blindly
by dividing into two little cals-de-sac (8.P., fig. 15), which
project downwards and outwards into the cerebral cortex, and are
lined by a columnar epithelium the cells of which are deeply laden
with granules of a reddish-black pigment.
The cup-like organs thus formed are usually called ‘“ eye-spots,”
and are present in at least four other species of Phascolosoma, but,
so far as ] am aware, no account of their structure has hitherto
been published. Their existence In an animal which lives at a
depth so great as that of 60 fathoms is noteworthy, and appears
to me to argue for the organs in question some function other than
even merely photometric ; but asmall fraction of diffused daylight
can permeate so far. ‘They are always to be found in the genus
Phymosoma, and from the above-given description it will be seen
that the sense-organs of Ph. teres resemble in the main those of
Phymosoma varians (Shipley, 7). They differ, however, from
those of Phymosomea in their relations to the tentacles.
In the genus Phymosomea the cells entering into the formation
of the eye-spots are continuous, on the one hand, with theepithelium
covering the anterior aspect of the brain, and, on the other, with
those which constitute the lming of the mesial surface of the
tentacles. In Phascolosoma, on the contrary, the sensory epi-
1903. | ANATOMY OF A NEW GEPHYREAN WORM. 35
thelium is furnished by a tentacular surface which is apparently
external,
sap seeming discrepancy is readily explained by a reference to
the following diagrams (text-fig. 9), modified from Selenka, in
which A represents the position of the mouth, B that of the
cerebral ganglion, while thick-dotted lines indicate the relations
of the tentacular wreath.
Tn text-fig. 9 @ this wreath is seen surrounding the mouth with
a simple circlet, as in Phascolosoma teres. In such a case the
infundibuliform approach to the cerebral organ must be lined by
cells, continuous not only with cells of the introvert epidermis
but with those covering the outer aspect of the tentacular
chaplet.
Text-fig. 9,
ger ae ad Ne.
is XN io in \ ‘
/ a Fah eet uae
| 5 \ ] ‘
1 A as ea
\ | : ~. =? ;
\ / A A
& A ie of
N Pa ee Bos ;
ae Ebel a Senn ee Sees
a b
Tentacular wreath of (a) Phascolosoma and (6) Phymosoma.
In text-fig. 95 the crown of tentacles is seen to have been in-
dented in its upper (neural) part, and the edges of the indentation
have been bent up in such a manner as to form a horseshoe-
shaped lophophore which, partially encircling the brain, overhangs
the mouth instead of surrounding it. The finely dotted line in the
diagram represents the position of the “lower lip” in Phymosoma ;
this may be looked upon as corresponding to the fused bases of
aborted tentacles. Here, then, the brain, lying at the base of the
lophophore, can only be approached by a canal whose walls are
coated by an epithelium continuous with that covering the mesial
surface of the tentacles.
The arrangement seen in Phascolosoma is the more primitive.
Briefly stated, tentacular surfaces which are inner and outer in
the genus Phascolosoma, become outer and inner respectively in
Phymosoma, owing to the secondary indentation of the crown of
tentacles.
According to Shipley (Joc. cit.), the space included within the
concavity of the lophophore of Phymosoma becomes the repre-
sentative of the ‘“ preeoral lobe ” of Phoronis.
Qe
34
36 MR. W. K. HUTTON ON THE § ~ [ Jan. 20,
Although nowadays, in virtue of our knowledge of the totally
distinct modes of development of the tentacles in Phoronis and
the Sipunculids, it does not seem possible to institute any
homology between the lophophoral crowns of Phoronis and
Phymosoma, yet it may be of interest to note that in the species
of Sipunculid under consideration the homologue of the pigmented
epithelium, which in Phymosoma covers Shipley’s ‘ preeoral lobe,”
is to be found in that layer of pigmented cells (Plate VIII.
fig. 12, S.E.) which forms the dorsal wall of the epineural canal
as well as in that layer continuous with it, bounding the anterior
surface of the brain (fig. 13, E.).
Nephridia.
These are small (Plate VI. fig. 2, B.T.), measuring less than the
twentieth part of the length of the worm; the right nephridium
is the longer and more convoluted of the two. Hach organ 1s
retort-shaped (Plate VIII. fig. 17), having a bulbous portion (C)
attached to the body-wall, and a tubular part (D) which, arising
from the posterior part of the bulb, bends round it mesially and
projects freely forwards into the ccelom. The wall of the bulb is
thin, covered externally by the cells of the peritoneum (A, fig. 18),
and internally by a single layer of granular-looking columnar
epithelium (C, fig. 18); it is composed of slender intercrossing
muscular bands. The cavity of the bulb is packed with fine
vesicles (H, fig. 18), which, it will be seen, are derived from the
secreting epithelium of the tubular portion of the nephridium.
The bulb communicates with the exterior by a fine canal (A, fig. 17),
lined with a small-celled epithelium, which pierces the body-wall
immediately in front of the anus, and its cavity is, further,
placed in connection with the ceelomic space through a slit-like
opening (B, fig. 17) in front of the excretory canal. The lips of
this opening, lined with ciliated epithelium, le between the body-
wall and the bulbous part of the nephridium, and are concealed
by its overhanging anterior extremity. The tubular portion of
the organ has a wall which is formed by a peritoneal lameiia,
heneath which is a layer of muscles (B, fig. 18). This muscular
layer consists of a meshwork of strands, between interstices of which
the lining-epithelium bulges outwards, forming little crypts lned
with a single layer of secretory cells (C). In cross section there
are seen projecting into the lumen of the tube feathery columns
of cells (D, fig. 18), which have an appearance comparable to that
of goblet-cells. The nucleus is small and irregular in shape,
placed basally; while the cell-contents may be seen to become
clearer near the free margin, and are evidently extruded ultimately
as the thin-walled vesicles which are found filling the lumen of
the organ.
Shipley aptly compares the process, which takes place also,in
Phymosoma, to the formation of the granules excreted by the
cells of the mammary gland.
os
~~]
1903. ] ANATOMY OF A NEW GEPIHYREAN WORM.
The Generative Organs.
Stretching across the ventral aspect of the celom, at the point
where the ventral retractors are attached to the body-wall, is a
band of cells with relatively large nuclei. These cells constitute
the genital band and are derived from the peritoneal epithelium.
In the single worm examined, neither free ova nor spermatozoa,
were to be seen in the body-cavity.
Systematic,
The presence of hooks and the possession of four retractor
muscles are characters which at once place Phascolosoma teres in
the first of the five “ groups” into which Selenka (5) divides the
genus. Atthe time of publication of his ‘ Die Sipunculiden’ three
species alone (PA. vulgare, Ph. elongatum, and Ph. cylindratum)
composed the group. To these must now be added, as well as
Ph. teres, Ph. sanderi Collin (1) and Ph. lobostomum Grube
(Fischer, 2). The position of Phascolosoma abnormis Sluiter (6)
must in the existing state of classification remain doubtful, as the
whole anterior end of the animal was wanting.
The nearest ally of Ph. teres in the group is Ph. elongatwm
(see Keferstein, 3). Both have the same arrangement of tentacles
and the same number of rows of hooks, while the hooks themselves
are very similar. Lh. teres, however, differs widely in the pro-
portionate length of its introvert, and, as regards its imternal
anatomy, in the small size and unequal bulk of the nephridia, in
the absence of any definite spiral coiling of the intestine, in the
great length of the hind-gut, and in the absence of a spindle-
muscle; and, as well as in these particulars, in its characteristic
habit of body. From every other member of the group except
Ph. cylindratum it differs in the complete absence of papillie.
These peculiarities warrant the systematic zoologist in assigning
to the animal under discussion a distinct place in the genus, and
I propose for it the name of Phascolosoma teres.
That the genus Phascolosoma is an eminently variable one is
evident to any investigator who makes a systematic study of the
Sipunculide ; and that this tendency to variation may result in
the inclusion in one small sub-group of animals which differ in an
astonishing extent in external habit, is abundantly exemplified
by the fact that Phascolosoma teres and Ph. sanderi (Collin) two
worms so dissimilar that a casual observer would be pardoned for
placing them in different genera—must yet, in the present rather
unsatisfactory state of classification, be united in the group called
by Selenka “ Group I.” of the Plisiscolosomidas : yet, while the
latter was considered by its describer Collin to present points of
resemblance to that curious Sipunculid Goljingia macintoshii (8),
and seems to have points in common with <Aspidosiphon, the
former appears to me to present in its structural peculiarities
definite affinities to the sedentary Phascolion. NRoule (4), who
38 MR. W. K. HUTTON ON THE [Jan. 20,
investigated the Phascolosomide obtained by the ‘Travailleur’
and ‘Talisman,’ in speaking of classification, goes the length of
saying, “‘ L’espéce bien affirmée n’existe pas dans le genre Phasco-
losome”; and by this somewhat strong assertion he would
indicate that the genus is in such a plastic condition—in a state
of fluctuating variability as one may term it—that it is capable
of being influenced by what Semper called ‘“ the conditions of
existence” in such a way, that within the genus we may have
numbers of types, differing on the whole but little from one
another, the links of their genetic affinity easily supplied, which
yet point the way toward the probable origin of the other
Sipunculid genera. As Roule expressed it: ‘“‘ Ce genre est encore
en voie de transformation, il se morcelle en un grand nombre de
types peu dissemblables, et ne se divise point en espéces précises,
dont les variations s’enserrent dans des limites peu éloignées.”
Roule’s investigations led him to the conclusion that the Phasco-
losomidee examined by him formed a series of links connecting
the parent Phascolosoma stock with Aspidosiphon. In the phylo-
genetic table drawn up provisionally by Selenka in his beautiful
monograph ‘ Die Sipunculiden’ (page 6), a polyphyletic origin is
suggested for both Phascolion and Aspidosiphon. The former
may have arisen directly from the parent Phascolosomid, or from
a common stem (similarly derived) which branches into Onchne-
soma, Tylosoma, and Phascolion; while Aspidosiphon may have
been derived from the directly produced Phascolion, or froma stem
common to itself, Phymosoma, and Cloeosiphon.
In Phascolosoma teres we have, as it appears to me, not perhaps
an actually intermediate type, but at the least a species which
gives us hints as to the probable path taken in its phylogenetic
course by Phascolion ; while the species examined by Roule play
the same role as.regards the genus Aspidosiphon. Phascolosoma
sanderi, though considered by Collin to approach Golfingia, has a
distinct but superficial resemblance to <Aspidosiphon, and was
rightly, as I think, placed by its describer in the genus Phascolo-
soma.
In the worm which forms the subject of this paper the follow-
ing characters suggest Phascolion-like aftinities:—The differing
bulk of the two nephridia, the relatively feeble development of
the dorsal retractors (a character upon which Roule placed much
reliance in his investigations), the lack of spindle-muscle, and the
coiled, but not spirally-wound gut,—all these are peculiarities
which place the worm in closer relationship with Phascolion than
‘with any other of the groups derived from the original Phasco-
losomid stock.
Information as to the habits of Phascolosoma teres is wanting ;
but in its mode of life it is probably sluggish, lying buried in soft
ooze with its anterior end alone projecting. The thin-walled
bag-like posterior portion, endowed with but feeble muscularity,
so sharply demarcated from the rest of the body, so relatively
large, would undoubtedly act as a hindranee to active locomotion
1903. ] ANATOMY OF A NEW GEPHYREAN WORM. 39
through sand or mud. Most species of Phascolion live with the
posterior part of the body either embedded in mud or protected
by some cast-off mollusk-shell; and if we assume, as the external
habit of Phascolosoma teres seems to me to warrant, this similarity
of mode of life between the two types, certain peculiarities of
structure in Ph. teres become of easy explanation.
Selenka, long ago, pointed out that those Sipunculids which
live in shells usually have only one nephridium. Jn other words,
in animals of sluggish habit, the needs of metabolisin are served
by one excretory tube.
Now in Ph. teres, while there are two nephridia, they are hoth
small and are of different sizes; the left has retrograded to a
greater extent than the right. It is to be noted that when only
one nephridium persists—as in Phascolion—it is that upon the
right side.
Enfeeblement of the muscular system is correlated with inactive
habits; and in worms lying half buried in mud a strongly
developed system of retractors ceases to be a clamant need. The
extreme case is reached in Phascolion, where often the ventral
retractors alone remain, while an indication of regression of the
dorsal retractor system is seen (vide supra) in Phascolosoma teres.
Lastly, in this worm, the coils of intestine, completely filling
up the sharply-delimited posterior part of the body, could scarcely
be efficiently acted upon by any spindle-muscle, which, we see, is
absent.
In these structural characters, then, we seem to obtain in
Ph. teres an indication of the path taken by Phascolion in its
derivation from the more primitive Phascolosoma.
Summary.— With the characters of the Group.
PHASCOLOSOMA TERES, n, sp.
Body 40 mm. long, introvert ca. 8 mm. Posterior portion
stout, sharply demarcated, ellipsoidal, containing the coiled
portion of the gut.
Colour yellowish, merging to dirty bluish white over the
posterior part.
Skin smooth, iridescent, devoid of papille, translucent posteriorly,
beset with numberless glands. A simple wreath of 15 tentacles
surrounds the mouth. Forward on the introvert 10 rows of
backwardly directed, slightly curved, brownish hooks, ‘(06-08 min.
long.
Two eye-spots. Alimentary canal free, without spindle-muscle
or mesenterial attachment save close to the anus. Spiral winding
absent, but the tube much coiled in its middle portion, Hind-
gut very long. Twonephridia, the right the larger, one-twentieth
of the body-length. Simple contractile vessel.
Hab, Firth of Clyde. Dredged from 60 fathoms.
40 ON THE ANATOMY OF A NEW GEPHYREAN worm. [Jan. 20,
WORKS REFERRED TO.
(1) A. Cottry.—“ Gephyreen gesammelt von Herrn Stabsarzt
Dr. Sander,” in Archiv fiir Naturgeschichte, lvili. p. 197,
1892.
(2) Fiscuer.—‘‘ Die Gephyreen des Naturhistorischen Museums
zu Hamburg,” in Abhandlungen aus dem Gebiete der
Naturwissenschaften &c., Hamburg, Bd. xii. p. 1, 1895.
(3) Kurersteiy.— Beitrage zur Kenntniss der Gattung Phascolo-
soma F. 8. Leuck.,” in Zeitschrift fiir wiss. Zoologie,
Bd. xii. p. 35, 1862.
(8a) Kererstern.— Beitrige zur anatomischen und _syste-
matischen Kenntniss der Sipunculiden,” in Zeitschrift fiir
wiss. Zoologie, Bd. xv. p. 404, 1865.
(4) Rovts.—“ Sur le Géphyriens des grands fonds de la mer
recueillis par le ‘Travailleur’ et le ‘Talisman,’” in Comptes
Rendus, exxvii. p. 197.
(5) SrnenKka.— Die Sipunculiden,” in Semper’s ‘ Reisen.’ Wies-
baden, 1883.
(6) C. Pu. Sturrer.— Beitrage zu der Kenntnis der Gephyreén
aus dem Malayischen Archipel,” in Natuurkundig Tijdschrift
voor Nederlandsch-Indié, Deel xlv. p. 472, 1886.
(7) Surptey.—‘ On Phymosoma varians,” in Quart. Journ. Micr.
Seis volypcxxie mess pale O90}
(8) Lanxesrer —“ Golfingia macintoshu, &e.” Trans. Linn. Soe.
Lond. 2nd ser., Zool. vol. ii. p. 469, 1885.
EXPLANATION OF THE PLATES.
PuateE VI.
Fig. 1. Theentire animal, magnified about 6; times. A, the anterior muscular part.
B, the middle thinner portion. C, the bag-like posterior piece. AN., the
anus. pp. 29, 30..
Fig.2. Semi-diagrammatic figure of a dissection, magnified about 13 diam. I, the
introvert. E, the eye-spots. B.T., the nephridia. H, dorsal vessel.
D.R., dorsal retractor muscles. V.R., ventral retractors. AN., position of
anus. G, coiled part of gut. F, straight part. pp. 30, 31.
Fig.3. Cuticular glands in vertical section. x 290. R.M., radial muscles.
C, cuticle. L, secretory cells. I, granular contents. p. 30.
Fig. 4. Hooks from introvert. Surface view. X 154. p. 30.
Prats VII.
Fig. 5. Isolated hooks. X 290. F, furrow. p. 30.
Fig. 6. Transverse section 1 mm. behind the anus. X 53. C.G., cuticular glands.
R.M., radial musculature. IL.M., longitudinal do. M, muscles attached
to rectum. G, rectum. 8B.T., nephridia. TT, tentacle cut across.
H, hooks. J,imtrovert. N.C., ventral nerve-cord and muscles. p. 31.
Fig. 7. Histology of the retractor muscles (Powell & Lealand 7';-imm., eyepiece A).
A. Transverse section of dorsal retractor, showing amount of interstitial
connective between the bundles.
B. Ditto of ventral retractor, showing almost no intermuscular connective.
p. 32.
Fig. 8. Transverse section of tentacle. x 290. B.S., blood-sinus. N, nerve.
A, ciliated mesial cells. 3B, columnar outer cells. p. 32.
Vig.9. A. Epithelium of mid-gut. > 290. B. Epithelium of hind-gut. Xx 290.
p. 33.
Fig. 10. Transverse section of esophagus. X 290. Part only of the stout muscular
wall is shown. M, muscular wall. EH, the epithelium thrown into papille.
p. 32.
1903. | ON A NEW FRESHWATER CRAB FROM UPPER GUINEA, 41
Puate VIII.
Fig. 11. Transverse section of brain behind origin of cesophageal commissures, to
show giant ganglion-cells. 290. R.M., retractors. .S., blood-sinus.
G.C., giant cells. p. 33.
Figs. 12, 13, 14, 15. Four transverse sections through the brain and base of the
tentacular crown, to show the epineural canal and formation of the eye-
spots. X 53. B.S., blood-sinus. V.C.,nerve-cord. R.M., retractor muscles.
S.E., sensory epithelium of the dorsal, and §.V., that of the ventral wall of
the epineural canal. ‘T, tentacles. S.P., eye-spots. E, epithelium of
anterior cerebral surface. p. 34.
Fig. 16. Transverse section through brain and bases of the tentacles, showing the
origin of the circumcesophageal commissures (X 53 and reduced). p. 34.
A. Circumcesophageal nerves.
B. Nerves to the epineural canal and dorsal pair of tentacles.
C. Two nerve-strands, in cross section, which seem to be connected with
the sense-organs.
D. Fused retractor muscles.
i}. Blood-sinus.
Fig. 17. Diagram of a nephridium. _ A, excretory canal. B, calomic pore.
C, vesicle. D, tubular part. E, body-wall. p. 36.
Fig.18. To show histology of tubular portion. x 290. A, peritoneal cells.
B, muscular wall. _C, secreting cells which at D are scen forming
feathery columns. E, vesicles. p. 36.
5. On Potamon (Potamonautes) latidactylum, a new Fresh-
water Crab from Upper Guinea. By Dr. J.G. pu May,
of Ierseke, Holland."
[Received November 15, 1902.]
(Plate 1X2)
In the year 1881, Potamon africanum A. M.-E. was known only
by the short diagnosis and the figures in the ‘ Nouvelles Archives
du Muséum,’* made from a quite young individual from the
Gaboon. The anterior legs had neither been described nor figured,
and it is therefore not surprising that some older specimens of a
Potamon from Liberia were referred by me erroneously to this
species *.
Some time since, three adult specimens of a Potamon were sent
me for examination by Prof. Jeffrey Bell ; they had been collected
in the River Prah, in the south of Ashanti, West Africa, These
Crabs not only proved to belong to the same species as that
described by me in 1881, when compared with a female of medium
size and a very young male from Liberia in the Leyden Museum,
but they proved also to be new, as a typical specimen of P. afri-
canum, a middle-sized female from “ Ogoué” (evidently the
River Ogowé, just below the Equator), was kindly sent me by
Prof. Bouvier, and as a more complete description of P. africanum
was published in 1887, in which, however, the legs have not been
1 Communicated by F. Jerrrey Bert, F.Z.8.
For explanation of the Plate, see p. 47.
3 Vol. v. p. 186, pl. xi. fig. 2 (1869).
Telphusa africana de Man, Notes Leyd. Mus. iii. p. 121 (1881).
bo
nS
42 DR. J. G. DE MAN ON A NEW [Jan. 20,
figured (A. Milne-Edwards, ‘ Observations sur les Crabes des
eaux douces de l'Afrique,’ Paris, 1887, p. 4, pl. 2. fig. 8). I
therefore propose the name of Potamon latidactylum for this new
species that inhabits Liberia and Ashanti, on account of the
characteristic shape of the hands, and as this feature was not alluded
to in my somewhat incomplete description of 1881, another will,
I think, be welcome.
The rivers of West Africa and of the Soudan are inhabited by
several species of Potamon which differ from the other species
of this subgenus in the existence of éwo epibranchial teeth behind
the external orbital angle, instead of one as is usual. I was
at first inclined to create for these species a new subgenus, but
the differences from Parathelphusa are, indeed, of too little
importance. These species are the following :—
Potamon aubryt H. M.-K. Gaboon.
» pelit Herklots. Gold Coast.
BP africanum A. M.-K. Gaboon ; French Congo.
a decazet A. M.-K. French Congo.
43 emarginatum Kingsley. West Africa.
» floweride M. Soudan.
35 latidactylum, n. sp. Upper Guinea.
It must, however, be observed that P. emarginatwm is con-
sidered by Miss Rathbun (Proc. U.S. National Museum, xxii.
1900, p. 285) to be the same as P. aubryi. P. latidactylum
may at first sight be distinguished from P. africanum by the
different shape of the eaxtraorbital and epibranchial teeth and by
the different form of the hands, especially of the fingers, but there
are still other differences.
The cephalothorax is a little more enlarged than in the species
from the ‘“Ogoué,” as is readily shown by the measurements.
The upper surface is somewhat less depressed, the branchial regions
are somewhat swollen, especially the anterior ones, and the gastric
and cardiac areas are, in aged individuals, also slightly convex and
not so much depressed as in P. africanum. The oblique furrows
or depressions limiting off the protogastric areas from the anterior
branchial lobes are scarcely distinguishable in P. africanwm ; in
the other species, however, they are quite distinct, and, in adult
individuals, rather deep. The urogastric lobes and the cardiac
area are a little broader in proportion to the breadth of the
carapace than in P. africanum. In the latter the gastric region
is faintly granular or rugose just behind the postfrontal ridge,
but in the new species it appears everywhere smooth, though
finely and sparsely punctate. The anterior branchial area is
slightly rugose in P. africanum, but not marked with oblique
wrinkles ; these are quite distinct in the species from the River
Prah and Liberia ; on the posterior branchial lobe these rugosities
occur in both species, they are, however, thinner and finer in
P. latidactylum. The posterior part of the upper surface
igo 1905, vol, LPlDe
BESS
Z SN
| NEG: de Man del.
io
“TAMON (POTAMONAUTES) LATIDACTYLUM.
NNT NA KT TLARAA
1903. ] FRESHWATER CRAB FROM UPPER GUINEA. 43
appears, on either side of the cardiac and intestinal regions,
somewhat granular and rugose in the Congo species, but quite
smooth, though finely punctate, in P. latidactylum.
In the large aged female from the River Prah the oblique rug
near the lateral boundaries of the carapace show a tendency to
disappear.
In both species the postfrontal ridge is rather prominent,
smooth, and nowhere granulated. In the type specimen of
P. africanum it extends in a nearly straight line towards the
second epibranchial tooth, uniting with that of the right side,
but ending about 1 mm. short of that on the left. The postfrontal
ridge of P. latidactylum usually curves slightly forwards mesially
and at each end; it never unites with the epibranchial teeth,
ending about 13 or 2 mm. short of the middle of the first epi-
branchial tooth. It is situated a little more distant from the
orbits than in P. africanum, so that the furrow behind the orbits
is somewhat narrower in the latter species. The front has the
same form and breadth in both, and shows in the middle a broad,
though shallow bay; both the front and the furrow behind the
orbits are a little granular in P. africanum, but quite smooth,
though finely and sparsely punctate, in P. latidactylum.
The extraorbital and the two epibranchial teeth have a quite
different form (Pl. IX. figs. 1-3 & 7, 8). In P. africanum
(Pl. IX. figs. 7, 8) they are more prominent and separated from
one another by much deeper incisions. The first epibranchial
tooth is, in this species, a little smaller than the extraorbital, but
has about the same form; its straight or slightly arcuate outer
margin makes a right angle with the anterior margin of this
tooth. In P. latidactylwm, however, the extraorbital tooth and
the anterior epibranchial one are much less prominent and much
lower, the anterior margin of the first epibranchial tooth being
considerably shorter than its outer margin, measuring only one-fifth
of it. In the aged female from the River Prah the extraorbital
tooth appears somewhat longer than the epibranchial one, and in
the two males (Pl. TX. fig. 1) they have about the same length ;
in the much younger female from Liberia (Pl. IX. fig. 3) the
epibranchial tooth, however, is almost twice as long as the extra-
orbital, but in the quite young male from the same locality
(Pl. TX. fig. 2) the extraorbital tooth appears again once and a
half as long as the epibranchial. As regards the relative length
of these teeth, P. latidactylwm presents therefore considerable
individual differences. In both species the second epibranchial
tooth passes backward into an arcuate crest, bounding the cephalo-
thorax laterally; this crest, which in young individuals of P. lati-
dactylum (Pl. TX. fig. 3) appears distinctly denticulate, the den-
ticulations or crenulations disappearing in more aged individuals,
reaches in P. africanum to the posterior boundary of the uro-
gastric lobes and does not curve inward on to the surface of the
branchial region: in P. latidactylum, however, it is much shorter,
44 DR. J. G. DE MAN ON A NEW [ Jan. 20,
scarcely reaching to the level of the mesial crescentic portion of
the cervical suture, and i distinctly curves inwards on to the
upper surface of the carapace. The orbits have the same form
in both species, but in P. africanwm the incision that separates
the extraorbital tooth from their lower margin is deep, much
deeper than in the new species, so that in the former the outer
angle of the orbits strongly projects beyond this margin.
In P. africanum the transverse furrow, limiting off the sub-
hepatic region from the branchial floor, is bordered by a row of
small granules, the subhepatic area is covered with prominent
rugosities and | granules ; in our new species the latter appears
almost smooth, and the row of granules is also less distinct. The
rugosities on the inflected portion of the cephalothorax and on
the outer part of the pterygostomian regions are also much less
distinct in the species from Upper Guinea than in P. africanum.
The outer foot-jaws have the same form and characters in both
species, and the furrow on the ischium-joint runs in both a little
closer to the internal than to the outer margin of this joint.
The type of P. africanwm is a female, the abdomen of which
has not yet obtained its full development and size. The sternum
of P. latidactylwm is smooth, punctate, and in both species a
transverse furrow unites the postero-external angles of the buccal
frame. The male abdomen (PI. IX. fig. 4) somewhat resembles
that of P. consobrinum de M. (Notes Leyden Museum, 1899,
pl. 10. fig. 10,2). The lateral margins are slightly concave. The
terminal joint has about the same length as the penultimate; it
is triangular, rather pointed at the tip, not rounded like that of
P. consobrinum; the lateral margins are very slightly arcuate
posteriorly, nearly straight towards the tip. The posterior margin
of the penultimate joint is a little broader than the anterior ; the
lateral margins, slightly convex anteriorly, are a little concave
posteriorly. The length of the antepenultimate joint measures
about two-thirds that of the penultimate, its posterior margin is
a little concave, so that the lateral margins are somewhat shorter
than the length of this joint in the middle line. The lateral
margins of this and of the following fourth joint are straight, but
those of the fifth are rounded; in P. consobrinuwm, however, this
latter is also straight. As regards the coarse punctation, P. lati-
dactylum agrees with P. consobrinum.
In the two males from the River Prah the right chelipede is a
little larger than the left; in the very young one from Liberia
the left isa little larger. The upper margin of the arm of the
larger male from the River Prah bears some transverse rugosities,
and one observes a small tubercle on the concave upper surface at
the proximal end, somewhat nearer to the upper than to the
anterior margin; the latter is also somewhat tubercular on its
proximal half. The anterior surface of the arm bears, near the
articulation of the wrist, a conical tubercle not far from the
anterior margin, and a lower, broader tubercle between it anc
1903.] FRESHWATER CRAB FROM UPPER GUINEA, 45
the under margin; the latter is also slightly tubercular along
its distal half. The three faces of the arm are quite smooth.
The carpus is also nearly smooth above, though sparsely and
rather finely punctate; the inner margin is armed with a rather
small pointed spine in the middle and beneath it with a small
acute tubercle. The larger chela (Pl. IX. fig. 5), measured
horizontally, is a little longer than the length of the cephalothorax
and the fingers are a little longer than the palm; the latter is
just as long as high near the articulation of the fingers. The
convex outer surface of the palm is quite smooth, though finely
punctate, similar to the upper margin, but the lower edge of the
palm is obsoletely tubercular. The fingers are somewhat compressed,
especially the immobile one. The immobile finger is rather high
at the base; its flattened outer surface appears minutely and
densely granular under an ordinary Jens, and one observes on it
three longitudinal rows of impressed points running to the tip of
the finger, the middle one of which is situated a little closer to
the lower margin of the finger than to the upper, and looks
like a very shallow furrow. The 15 or 16 teeth are small, two
or three are somewhat larger than the others, and one in the
middle of the finger is the largest of all. The fingers leave a
narrow interspace between them when closed, the pointed tips
being only in contact and crossing one another. The dactylus is
somewhat arenate and tapers rather slowly to the tip; both on
the outer and inner surfaces and on its upper edge this finger
appears minutely granular under a lens, and one observes on it
several rows of small shallow puncta. The teeth agree in size and
in number with those of the index, three or four being somewhat
larger than the others. The little convex inner surface of the
palm is quite smooth,
The left chelipede is somewhat smaller, the hand being 41 mm.
long; it fully agrees with the right one, but the fingers are in
contact and the rows of puncta are a little more distinct,
In the young male from Liberia the fingers are in contact in
both chele, the upper margin of the palm appears distinctly
granulate under a lens, and the longitudinal furrows on the
fingers are already visible to the naked eye.
In the young female from Liberia the right chelipede is a little
larger than the left, the fingers are in contact, and the furrows
on the index, as also the rows of puncta on the mobile finger,
ave distinctly visible to the naked eye. The upper margin of the
palm of the left hand appears finely granular.
In the aged female from the River Prah the left chelipede is
much larger than the right; the hand, indeed, is 45 mm, long,
just as long as the cephalothorax, and 203 mm. high, but the
right chela (Pl. IX. fig. 6) is only 31 mm. long and 13 mm. high,
In both the fingers are in contact throughout their whole length
and the pointed tips cross one another; the strongly compressed
and much flattened immobile finger is not furrowed, though the
46 DR. J. G. DE MAN ON A NEW [Jan. 26,
rows of puncta are still visible. The fingers are but little longer
than the palm, and the latter is on its outer surface and on its
upper margin smooth, sparsely punctate. The fingers of the
smaller chela, however, are once and a half as long as the palm
(Pl. TX. fig: 6).
The female type-specimen of P. africanwm compared with the
female from Liberia, which is about the same size, shows the
following differences :—The anterior legs (Pl. IX. fig. 9) are equal,
as regards their shape and size. The transverse rugosities on
the upper margin of the arm are more distinctly granular, and
the anterior and lower margins are also more distinctly tuber-
culated ; the conical spine on the anterior surface, near the articu-
lation of the wrist, is more pointed, and several small tubercles
are seen between it and the anterior margin of the arm, that are
wanting in P. latidactylwm. The upper surface of the carpus is
slightly granular at the base of the spine on the inner margin,
and one observes, moreover, everywhere fine impressed lines and
points; the spine is somewhat larger than in P. latidactylum,
and instead of a small tubercle, there is beneath it a smaller,
pointed spine. The hands (Pl. IX. fig. 9) have a quite different
form. The fingers, almost in contact, are distinctly somewhat longer
than the palm, and the latter is distinctly less high than long. 'The
upper margin of the palm is covered with depressed granules,
and the puncta on the convex outer surface are partly arranged
in longitudinal rows. The fingers are not compressed, slender,
nor curved, except at the pointed tips which cross one another.
The immobile finger is much lower at its base than in the
other species, and its convex outer surface is distinctly furrowed ;
of the three furrows visible to the naked eye, the middle one is
deeper than the others. This finger is armed with 25 or 26 small
teeth, some of them on the proximal half are somewhat larger
than the others. Finally, the dactylus tapers regularly to the tip
and is also longitudinally furrowed by rows of impressed points ;
the teeth agree with those of the index. The fingers are almost
smooth.
The ambulatory legs, the measurements of which are given
on p. 47, apparently agree with those of P. latidactylum, but the
meropodites are more granular along and near their upper edges.
Potamon aubryt AH. M.-E., from the Gaboon, P. pelii Herklots,
from the Gold Coast, and P. flowert de M., from the Soudan, are
different species (vide de Man, Proc. Zool. Soc. Lond. 1901, vol. i.
p- 94), being at first sight distinguished by the upper surface of
their cephalothorax being very convex from behind forwards,
smooth and shining, by the different form of the fingers, of the
abdomen, &e. The young male of Potamon pelii described J. e.
p. 99, I have re-examined for the purposes of this memoir,
1903.] FRESHWATER CRAB FROM UPPER GUINEA, AT
Measurements in nrillimetres.
Perea ei al gl @
aban ios Galrd. |S
|
Breadth of the cephalothorax ...............0..sc0cee eee .-| 67 |58 |51 | 483) 34) 53
Length of the cephalothorax without the abdomen...) 45} | 39 |385 |33 | 24+) 39
Distance between the external orbital angles ......... 40% | 354 | 31 | 303/23 | 32
* posterior epibranchial teeth ...)574|50 | 434 | 423 | 314] 433
Breadth of the anterior margin of the front............ }18 |16 | 14 |i4 | 103) 14
Distance, in the median line, between the anterior :
margin of the front and the postfrontal ridge ...... 6 | 5 | 45) 4 | 83] 43
Distance between the outer angle of the orbits and ;
UieMpOstivOn bale Ges ee clececeaaecceeeees- aickaes: 5y| 43] 46) 4 | 382) 38h
Distance between the outer angle of the orbits and
the posterior epibranchial tooth ......................55 10 3s | OF) 7e| 54) 6S
Breadth of the urogastric ares) 2.0... ..cc.cccsesssesace ses 11 8k] 7 6s} 44] 7
a 99 CATING ANGI Goi iivcccceeecetnereecnsaeene-| a) [12° || 10%) 10° || 751 9
55 NO OMAIIES recdascs csedPennebe seareGeosciacee ag a bLAR PREY ailamtcy all mtsteal Wats) alpiarsy
Height of the orbits, near the lower internal angles .| 6 | 5% 5 + | 32/1 6
Length of the terminal segment of the abdomen...... 7) be 4A,
» penultimate segment .............0..0005 63 6 44
Breadth of the anterior margin of this segment ...... 81 | 72 5 |
33 » posterior margin of it soriceal| pan |) MOP El eos |) st
Length of the larger chela ................c.csseeceeee-2-- 45 146 | 39 31} | 223 | 34
- of) TEENA ” oon nan eresaqucd nec boucdeele GOMER Cea RB IG Ce, Wien si tal eo) al (gl ced bal OYE wl bn IC)
Height of the palm, near the articulation of the
GLYCHAVJNEIS: sco shoo neo nde nstined doeebee a Mbet sce COI A CERES ae AB Mane CAO a Racy al Letlccy alfa ayo al amp 3
Length of the legs of the penultimate pair ............ 86 |76 |71 |66 |50 | 80
ss » meropodites of the penultimate pair...)27 | 25 | 235/21 | 16} | 27
Breadth _,, 35 ‘3 43 AS call tek || Shall Nels 5] 8h
Length of the propodites ss S » ---|16 |13 |124)12 | 94) 154
Breadth _,, » ” » oo >| 6 5z| 53) 4 67
Length of the dactylopodites ,, Bs 5) ---| 208 | 17 | 164) 154) 112118
5 By ees) On Gheplastipaiceaeancrserereetres| Cou Olio ole (99) 1160
= » meropodites of the last pair ............|214]193/18 |164/13 | 20
Breadth ,, : PP an sip Seo ouGeal ha tesenl |e Clos Ih a Aat ae Véos IN (tats
Length of the propodites __,, a eeeceeens es | (La LO RN OS.) ORM NR lO
Breadth ,, ae ef a Sosessancesal i | Ger INGE | ERNE
Length of the dactylopodites ,, 53 sonccnosocee| I SMA Wale aloes |) fs} haat
Nos. 1-3, River Prah; Nos. 4, 5, Liberia; No. 6, type-specimen of Thelphusa
africana A. M.-E., Ogoué, from the Paris Natural History Museum.
EXPLANATION OF PLATE IX.
Figs. 1-6. Potamon (Potamonautes) latidactylum, n. sp—Fig. 1. Dorsal view of
the anterior part of the carapace of the largest male specimen from the
River Prah, X 13. Fig. 2. Left antero-external angle of the carapace of
the young male from Liberia, xX 8. Fig. 3. The same of the young female
from Liberia, x 3. Fig. 4. Abdomen of the largest male from the River
Prah, X 1}. Fig. 5. Larger chela of the same male from the River Prah,
x 14. Fig. 6. Smaller chela and carpus of the adult female from the
River Prah, x 1}.
Figs. 7-9. Potamon (Potamonautes) africanum A, M.-E., female type-specimen of
the Paris Museum, from Ogoué.—Fig. 7. Dorsal view of the anterior part
of the cephalothorax, x 1}. Fig. 8. Left antero-external angle of the
carapace, X 3. Fig. 9. Right chelipede, x 1}.
48 MR. R. I. POCOCK AND HON. N.C. ROTHSCHILD oN [Jan. 20,
6. On a new “ Bird’s-dung” Spider from Ceylon. By R.I.
Pocock, F.Z.S.,and the Hon. N. C. Roruscuitp, B.A.,
Ie Sio5 Leis
[Received November 18, 1902. ]
(Plate X<5)
During a recent visit to Ceylon, the Hon. N. C. Rothschild, in
company with Mr. EK. H. Green, discovered a specimen of a
Spider (Phrynarachne) imitating, for purposes of allurement and
concealment, a patch of bird’s-dung. A photograph of the spider
resting upon the leaf tn sitw was obtained, and the leaf with its
patch of web was brought home with the spider (Pl. X. figs. 1, 2).
The film of white web upon the leaf, with its irregular outline
and marginal, often apically thickened projections, closely assimi-
lates the splash of the fluid components of the excrement; while
the spider itself, with its mottling of black and white, resting
upon the centre of the film, with its legs tucked in, represents
the semi-solid central core. So close, indeed, was the imitation
that both observers were for a time deceived, until closer scrutiny
revealed the true nature of the deceptive patch.
It is commonly assumed that the purpose of this imitation is
purely alluring or pseudepisematic. No doubt, however, it is an
equally important procryptic factor, serving to protect the spider
from various enemies, especially the wasps of the family Pompilide.
The spider, which appears to represent an undescribed species,
may be named after one of its discoverers *.
PHRYNARACHNE ROTHSCHILDI, Sp. n.
® — Colour. Carapace with a broad central black patch, with
irregular lateral margin, extending from the eyes to the posterior
slope, and forming an angular expansion on each side imme-
diately behind the head; and with its posterior border passing
into a median longitudinal stripe, continuous with a trans-
versely arcuate black stripe which extends from the middle of
the posterior slope of the carapace on to its lateral portion,
breaking up into fainter patches separated by radiating pale
bands; clypeus, sides of head, and area between central patch and
lateral patches yellowish white. Mandibles yellow in basal half,
with a large black patch, not reaching the fang, in the distal half
in front, blackish brown beneath. Sternum mostly yellowish white,
with a large subtriangular black patch, divided by a narrow pale
line in its posterior half, and some black at the bases of the legs.
Labium blackish, maxille blackish internally, whitish externally.
Palp with its femur white below, black above in its basal half;
patella white, tibia and tarsus black, variegated with yellow
patches. First and second legs with the coxe black, variegated
1 For explanation of the Plate, see p. 51.
2 Mr. R. I. Pocock takes the sole responsibility for the naming and description of
the species.
Pin, 13> vol, 1 PL xX
5
F.Pickard-Cambridge del et lith West, Newman imp
PHRYNARACHNE ROTHSCHILDI.
1903.] A NEW SPIDER FROM CEYLON. 49
with dirty yellow below, whitish with a black spot above ; femora
jet-black variegated with paler patches above, in front, and
behind, with a narrow whitish line between the tubercles below,
the distal end yellowish white; patelle yellowish white, with
two palely fuscous patches above and some behind; tibie and
protarsi mostly jet-black, with a pale median inferior longitudinal
stripe between the spines; tarsi pale above, black below; third leg
with coxa and trochanter blackish below, the femur amber-yellow
in its basal two-thirds, yellowish white with a black patch, much
broader behind than in front, in its distal third; tibia black and
whitish, the black predominating posteriorly; protarsus and tarsus
mostly yellowish; fourth leg, with exception of its coxa, mostly
yellowish white, lightly spotted above. Almost the whole of the
upperside of the abdomen jet-black from its anterior margin
backwards to the spinners, with the postero-lateral tubercles and
the sides yellow; the under surface (i.e. the area between the
lung-sacs on the epigastric region and behind the genital fold
nearly back to the spinners) black, this black field showing a trans-
versely oblong pale patch behind the genital aperture (Plate X.
figs. 3 & 4).
Carapace not strongly tubercular, flattish longitudinally along
the middle line, with a shallow transverse depression behind the
ocular area of the head; area between the ocular tubercles
depressed ; eyes of posterior line (fig. 5) subequally spaced, the
medians about 5 diameters apart, lying about their own diameter
in front of a tangent joming the anterior borders of the laterals,
which are distinctly larger than the medians; eyes of anterior line
(fig. 6) strongly recurved, the inferior edges of the laterals as high
as the superior edges of the medians, the latter considerably smaller
than the laterals, about 24 diameters from each other and about
2 diameters from the laterals'; distance between the medians
about equal to the height of either above the edge of the clypeus,
which is weakly tubercular and furnished with prominent angles,
Mandibles prominent basally, scarcely tubercular ; margins of
fang-groove thickly fringed, the posterior armed with one fang, the
anterior with teeth, remote from base of fang; the posterior fringe
continuous with a scanty band of hair passing up the inner side
of the posterior surface of the mandible.
Legs. Femur of Ist leg thickly covered in front with small
tubercles, with a few large tubercles above and two rows of about
five tubercles each below, the posterior surface smooth; tibia
bowed, armed beneath with two rows of about 4-5 spines, the
anterior and dorsal surface also spined ; protarsus a little longer
than tibia, armed beneath on each side of the middle line with a
band of longer and shorter irregularly arranged spines, those on
1 Owing to a want of definition of the border of the corneal lens, the precise size
of the eyes of the anterior line, and consequently of the width of the interspace
between them, is hard to ascertain, varying apparently in different lights and with
lenses of different power. In the above-given description the eyes are described as
seen under a 4-inch objective and a platyscopie lens.
Proc. Zoou, Soc.—1903, Vou. I. No. IV. t
50 MESSRS. R. I, POCOCK AND N.C. ROTHSCHILD oN [Jan. 20,
the posterior side being much less numerous than on the anterior,
the anterior and dorsal surfaces of the segment also spiny ;
tarsus covered with rows of spinules; 2nd leg like the Ist, but
with the femur almost quite smooth ; 3rd and 4th legs with tarsi
bristly beneath, tibia of 3rd spined below, of 4th scarcely spmed.
Abdomen as wide behind as it is long, and twice as wide as it 1s
in front, the anterior border transverse and armed with four
tubercles, the lateral border with four small tubercles, the
postero-lateral angle with two large superior and two small
inferior pale tubercles ; between the former are four pairs of
shining black tubercles, two pairs of which are much larger than
the Others: in addition to these the dorsal surface is furnished
with three pairs of small scattered tubercles; lateral surface
pitted ; inferior surface with two rows of muscular pits. Vulva
undeveloped.
Measurements in mm.—Total length 9, carapace 4; posterior
width of abdomen 5°5, anterior width nearly 3; length of Ist leg
14, its femur 4, patella + tibia about 5, 3rd leg 6, tarsus + pro-
tarsus about 5
Loc. Kandy.
Tn size and shape this species does not appear to differ appreci-
ably from P. ceylonica O. P. Cambridge (P. Z.S. 1884, p. 201,
pl. xv. fig. 3, sub Ornithoscatoides), which was recorded from
Ceylon. The colouring of the two, however, is very different.
In P. ceylonica the anterior legs are heavily blackened only upon
the protarsi and distal half of the tibie, the mandibles have no
lack patch, and the black-and-yellow pattern of the carapace
and abdomen, so conspicuous in P. rothschildi, is but little differ-
entiated ; also the cephalic tubercles between the two rows of
eyes are much higher in P. ceylonica (see also Simon, Hist. Nat.
Araign. 1. p. 1043, fig. 1087, 1895).
Another Ceylonese species, O. nigra O. P. Cambr. (loc. cit.
p. 202, pl. xv. fig. 4), was based upon the male sex, and cannot,
therefore, be compared with those based upon females. <A third
species, P. fatalis ©. P. Cambr. (P.Z.8. 1899, p. 525, pl. xxx.
fig. 7), from Ceylon, differs so markedly in form, coloration,
tuberculation, &e., as to need no comparison with this new species.
Also P. peeliana Stol. (J. A. S. Bengal, xxxviii. 1869, p. 229,
pl. xx. fig. 4), from Sibsagar, Assam; P. tuberosa Blekw. (Ann,
Mag. N. ‘Hist. (3) xiv. p. 38, 1864, & PLAS. 1884, pl. xv. fig. 2),
from the Hast Indies; and the Burmese form P. papulata Thorell,
are all different from this new form.
To Dr. H. O. Forbes, F.Z.S.’, belongs the credit of the discovery
that the coloration of the species of Phrynarachne belongs to the
pseudepisematic category. The pattern of yellow and black which
decorates the dorsal and ventral sides simulates that of the sem1-
solid central portion of a patch of bird’s-dung splashed upon a leaf,
the paler more fluid portion being represented by a thin irregular-
shaped carpet of white silk, in the centre of which the spider
takes its stand. The spider discovered by Messrs. Rothschild
1 P.Z.S. 1883, p. 586, pl. li.
1903.] A NEW SPIDER FROM CEYLON, 51
and Green was resting upon the leaf in the normal position, that
is to say back uppermost (figs. 1 & 2).
Forbes, however, expressly states that P. decipiens, the species
he discovered, lies back downwards on the web, holding itself
in place by means of the spines with which the anterior upper
surfaces of the legs are furnished, and he adds that the under-
side of “its rather irregularly-shaped and prominent abdomen
is almost all white, of a pure chalky white” (p. 587), and that
“its pure white abdomen represents the central mass of the bird’s
excreta, the black legs the dark portion of the slime.” Refer-
ence, however, to his figure, which was apparently drawn partly
from memory, the leaf "and web having “ gone astray” during
the transport home, shows that the under side of the abdomen, so
far from being almost all white, is furnished with a large black
sub-oblong patch, which extends from its anterior border con-
siderably past the middle. It is noticeable, too, that from the
ventral aspect the abdomen is not irregularly shaped, but evenly
oval in outline and devoid of tubercular excrescences. The dorsal
surface, on the aes as represented in the figure given by
Mr. Cambridge (BZ. 1884, pl. xv. fig. 1), might very well
be described as almost “il white and rather irregularly shaped.
The white colour largely predominates, and the tubercles project
prominently from the expanded posterior portion. Mention is
made of these discrepancies between the figure and description,
to justify the suspicion that Dr. Forbes may have mistaken the
dorsal for the ventral surface of the spiders he saw. Moreover,
one cannot but wonder how the spider maintains a secure hold
back downwards, especially when the powerful prehensorial legs of
the first and second pairs are released, as released they must
surely be, to seize an alighting butterfly. One would think that
the flapping of the insect’s wing would pull the spider, now
insecurely anchored, from its hold and bring both to the ground
together. On the other hand, if the spiders of this kind always
rest in the normal position seen and photographed in the case of
P. rothschildi, the simulation of the bird’s-dung is equally perfect,
and the spider, while seizing a butterfly with its fore legs, can
maintain itself securely in place by grasping the web with the
claws of the remaining pairs and by gluing its spinning-mamillee
to the subjacent silken threads.
These, then, are the points for further investigation which we
would ask residents in the tropics to take note of :—Do the species
of Phrynarachne assume indifferently the dorsal or the ventral
attitude? Is it a peculiarity of one species to lie back uppermost,
and of another belly uppermost? If they lie, as Forbes asserts,
back downwards, how is a secure hold maintained when an insect
has to be seized ?
EXPLANATION OF PLATE X.
Phrynarachne rothschildi, sp. n., p. 48.
Figs. 1, 2. The ue resting on Jeaf. Fig. 3. Upper surface.
Fig. 4. Under surface. Figs. 5, 6. Byes.
4%
52 DR. H. J. HANSEN ON CRUSTACEANS [Jan. 20,
7. On the Crustaceans of the Genera Petalidium and Sergestes
from the ‘Challenger,’ with an Account of Luminous
Organs in Sergestes challengeri, n. sp. By Dr. H. J.
Hansen (Copenhagen).
[Received November 29, 1902. ]
(Plates XI. & XII.")
During a stayin London in July and August, 1902, I examined
various groups of Crustacea in the British Museum (Natural
History). I beg the Director, Professor E. Ray lLankester,
and Mr. F. Jeffrey Bell to accept my sincere thanks for the free
use of the collection and for their kind help.
In the paper “On the Development and the Species of the
Crustaceans of the Genus Sergestes” (Proc. Zool. Soc. Lond.
1896, pp. 936-70) I have given a revision of this extensive
genus. I had studied a very rich material of pelagic forms
belonging to the Copenhagen Museum, among which are all the
types of Kroyer; besides I had examined types of 5 species
established by Chun, Metzger, and Ortmann.
Among other things, I proved that ‘of the 59 (or 60) hitherto
described species only about 20, or one-third of the total number,
have been established on adult animals, such as have almost or
entirely arrived at sexual maturity ; and that almost all the other
species are true larve, and even of these a considerable number are
larval stages of species already established on adult specimens, .. .”
Of earlier authors, C. Spence Bate has produced a very large con-
tribution on the genus Sergestes, extending to eighty-eight quarto
pages and seventeen plates, in his ‘“ Report on the ‘Challenger’
Macrura.” He established the genus Petalidiwm on a new species,
described 24 new species of Sergestes &c. In 1896 I wrote
(p. 939): ‘This large contribution is of course of great
importance, but unfortunately neither the descriptions nor the
figures are so good as could be wished, and in numerous instances
...a re-examination of the type specimens is absolutely necessary
—the greater part of the new species are but larve.” I have now
studied all types which are preserved in the British Museum, and
the present paper contains the results of my examination.
Bate describes 31 species of Sergestes as examined by himself :
of these, 24 are established as new to science, 6 are considered
to be Kroéyerian species, and one is referred to S. atlanticus
H.M.-Edw. The types of 9 of the species established by Bate do
not exist in the British Museum; some specimens mentioned in
his work and belonging to other species are also absent; but
several specimens belonging to various species and omitted in the
Report were found in the collection. I am therefore only able
to give more or less incomplete notes, based on the study of the
1 For explanation of the Plates, see p. 78.
as 4 .
PA, t903 voll Plex
Zor 2l 36r 3br'
Zs eoS\ :
Ae
H.J.Hansen del. Edwin Wilson , Cambridge
FIG.1.PETALIDIUM FOLIACEUM. 2.PETALIDIUM gun. 3.SERGESTES
- PROFUNDUS. 4.S.PREHENSILIS. 5.S.KROYERI. 6.S.SIMILIS.
P.Z.5.1903,vol.I. Pl. XI.
Edwin Wilson, Cambridge
FIG.1.SERGESTES ARCTICUS. 2.S.GHALLENGERI.
4i.J.Hansen del.
1903.] IN THE ‘ CHALLENGER’ COLLECTION. 53
specimens, on 22 of the species in question, 15 of which were
established by Bate himself. Furthermore, he refers Petalidiwm
Bate, Sciacaris Bate, Acetes H. M.-Edw., and Lucifer Vaugh.
Thomps. to his family Sergestide ; of Lucifer he describes two
species, but the genus must be more thoroughly studied than has
hitherto been done before the examination of the ‘Challenger’
specimens ; Sciacaris Bate has one species, which is only a larva of
a Sergestes, and the type seemed to be wanting in the Museum ;
of Acetes, Bate had no specimens; and Petalidium is mentioned
below.
Of Bate’s 31 species of Sergestes only 6 are really mature forms,
25 being larvee. Special attention has been paid to the adult speci-
mens preserved in the Museum and enumerated by Bate; on two
of these specimens I have established two new species, and
besides I add notes and some drawings to the representations of
Bate. Unfortunately nearly all the specimens of rare and
especially interesting species were very much mutilated.
Our present knowledge of the adult species of Sergestes of the
Atlantic fauna is far from complete, but yet we are acquainted
with so many species that it was possible for me in my earlier
paper either to refer the J/astigopus-forms examined to the
mature species, or to describe the older larval stages and some-
times the black-eyed but still immature forms, so that they can be
recognized with certainty and referred to the mature forms when
these are discovered in the future. All the Atlantic larval forms
from the ‘Challenger’ seen by me have now been referred in a
similar way. But many larve established by Bate as valid species
of adult or sub-adult animals have been secured in the Pacific.
Our knowledge of the mature stage of the species living in that
vast ocean is still rudimentary; and I have therefore not been
able to refer the larvee of three of Bate’s species to any species
established on adult specimens. Bate’s types of his species
established on larval forms are often either defective or very
young, wherefore I thought it of little use to describe and figure
them again; but I have generally added some notes on their
affinities, and sometimes also a few corrections to his descrip-
tions. When the Pacific has been moderately well explored
by further expeditions, many adult forms and their larval stages
will be discovered ; and a future student of the group will then be
able to refer at least some of the larvee, which I cannot interpret,
to their adult forms. To the young larve described by Bate as
species of JJastigopus I pay no attention at all: the types seem
to be lost.
I think it convenient first to deal with all the ‘ Challenger’
species in the same consecutive order in which they are described
in Bate’s Report ; then to put together some results of the investi-
gation; finally, to mention more fully the luminous organs in
Sergestes challengeri, n. sp
In order to abridge the descriptions, in the following pages—
as in my earlier paper~-I make use of some abbreviations :—
54. DR. H. J. HANSEN ON CRUSTACEANS [ Jan. 20,
antenn. ped. = peduncle of the antennule; mxp.’-mxp.*’=the
second and third pair of maxillipeds; trl.\-trl.’=the first to the
fifth pair of trunk-legs; br.=the first, br." =the second branchia
above the same legs; ext. br. of urp.=external branch of the
uropods.
I. Notes on the Species of Petalidium and Sergestes.
a. PETALIDIuM Bate.
To this genus Bate has referred only one species. It will be
convenient to deal with the characters of the genus together with
those of the species.
PET. FOLIACEUM Bate, pp. 348-50, pl. lx. (Plate XI. figs. la—1g.)
Bate mentions five specimens from two stations: Stat. 146,
lat. 46° 46’ S., long. 45° 31’ B., 1375 fathoms; and Stat. 159, lat.
47° 25' S., long. 130° 22’ E., 2150 fathoms. All these specimens
are at present in the Museum, but even Bate’s text and his figure
of the entire animal show that most of the appendages presenting
specific characters are wanting or have been mutilated. Bate
writes (p. 849): “The great distinction between this genus and
Sergestes exists in the form, character, and arrangement of the
branchial plumes, which consist of a series of plates and eylin-
drical filaments, situated side by side in a series of rows at right
angles to the stalk. There is but one plume to each of the five
anterior somites of the pereion, the posterior two somites having
none; between some of the somites is a large foliaceous plate.”
But this description is difficult to understand: his figure 3,
representing the branchie, is defective, and his tabular view
(p. 349) is wrong. He was of opinion that the foliaceous plates—
of which he had seen only three—were pleurobranchie, while the
“branchial plumes” were arthrobranchiz; but this is incorrect:
they are decidedly pleurobranchie asin Sergestes. Petal. foliacewm
Bate differs from all species of Sergestes in one quite unimportant
feature, that no trace of branchie is found above trl.*, and in one
important character, viz., the structure of the pleurobranchial
plumes. This structure is very interesting (fig. le). The most
developed branchie are, as usual, those above trl.* and trl.*; each
of these consists of an anterior and a posterior half, and each half
of five (to six) transverse rows of branchial plates, generally five
or six in each row, and these plates (some of the lowest excepted)
are directed upwards. ‘The pleurobranchie above mxp.’ and
especially trl.* are less developed, with a lower number of trans-
verse rows, and partially with a lower number of plates in the
rows. The pleurobranchial plumes in VPetalidiwm look very
different from those in Sergestes; the real differences are: a
much lower number of rows, a much lower number of plates in
the rows, and that the plates are much larger, curved upwards, and.
look much more independent. Behind the upper part of each of
these four pleurobranchie originates a pleurobranchial lamella (/.),
1903.] IN THE ‘CHALLENGER COLLECTION. 55
which is a reduced branchia; these lamelle are very long above
trl.? and trl.*, while the two anterior lamelle above mxp.’ and
trl.‘ are much less prominent. On mxp.” is found an epipod (ep.),
with a branchia consisting of a few plates, and above it a rudi-
mentary pleurobranchia consisting of one very small lamella.
Fig. la and fig. le show the rostrum, consisting of a carina
with a short or ve1 ‘y short terminal spiniform process. There are
no supraocular or hepatic spines; the gastro-hepatic groove is
well developed. A comparison of fig. 1 ¢ with fig. 1 d shows that
the eye-stalks are considerably depressed; they are from two and
a half to nearly three times longer than the eyes at the inner
terminal angle, with a small rounded knot turning inwards and
a little wpwards (fig. 1 6; fig. 1 d), and besides (always ?) with an
exceedingly small tubercle somewhat in front of the inner angle
and more downwards on the inner side (fig. 1d). In the antenn.
ped. the basal joint is very broad (fig. 1 6), decidedly shorter than
the outer margin of the two other joints together ; the third joint
is scarcely three times longer than broad, considerably longer
than the outer margin and a little shorter than the inner margin
of the second joint, which is stout, with its Inner margin only a
little more than twice as long as its breadth. (lee specimen
from which figs. 1¢ and le were drawn measures 51 mm. in
length, and was captured on Stat. 146; figs. 1 a-16 were drawn
from the specimen secured on Stat. 159).
But besides these five specimens of Petalidiwm, I found still
two smaller specimens of the same genus among the ‘ Challenger’
animals. One of these had been determined as Sergestes japonicus
Bate, but is not referred to that species in his text. It was
captured at Stat. 158, 7/111., 1874, 1800 fathoms, thus near one
of the above-named stations. It measures only 21°5 mm. in
length. Figs. 1fand 1g show that its rostrum has the terminal
process somewhat longer than in the large specimens; the eyes
are a little longer as compared with the length of their stalks, and
these are pro portionately somewhat broader, without a distinct
knot at the end of the inner margin. There is no supra-ocular
spine, but the hepatic spine is moderately developed ; the gastro-
hepatic groove is very distinct. No branchiew above trl.” This
small specimen belongs certainly to P. foliaceum Bate.
Bate established (pp. 428-31) his Sergestes profundus on two
specimens, both badly mutilated. He describes each Bpeinen
separately, beginning with one captured at Stat. 137, lat. 35° 59'S.,
long. 1° 34' E., depth 2550 fathoms. But, unfortunately, ne
specimen does not belong to Sergestes but to Petalidiwm; the
type is besides so mutilated that I should have preferred to
omit it, if it had not been described by Bate. It measures
17-5 mm. in length. The rostrum is shown in fig. 2@; it is
described by Bate: ‘“ It consists of a short fine point projecting
horizontally for about one-fourth the length of the ophthalmopod,
and is dorsally furnished on ihe crest with a small tooth.” The
eye-stalks are aS in the small specimen from Stat, 158 just
56 DR. H, J. HANSEN ON CRUSTACEANS [Jan. 20,
described ; the posterior branchie to a large degree are destroyed,
and as to the other features, I refer to Bate’s description (p. 429).
I cannot say with certainty whether the animal is a very young
specimen of P. foliacewm Bate, or belongs toan unknown form; the
rostrum deviates considerably from the type of P. foliacewm, but
its shape presents a stage between those in the adult and im a larva,
nearly agreeing with that in a larval form mentioned below.
In my earlier paper on Sergestes I described S. obesus Kr.
(=S. sanguineus Chun). I stated that it was a larva, a Masti-
gopus, and added (p. 968): ‘It is easily seen that this species
cannot remain in the genus Sergestes, but whether it should be
referred to Petalidium Bate, or a new genus should be established
for its reception, is impossible to decide with certainty. The
branchial plates recall the plates found in Petalidiwm, and there-
fore I provisionally transfer it to that genus....” I had not
seen any adult Petalidiwm, but suggested (p. 967) that the
branchial plumes interpreted by Bate as arthrobranchie are in
reality pleurobranchie. I can now decide that S. obesus Kr. is
the Mastigopus of Petalidiwm: the branchie prove it; besides,
trl.2 in the Mastigopus is exceedingly elongate, with the basal
part very thick, and in the adult form the basal joint of the same
pair (fig. Le) is exceedingly thick, very much thicker than those
of trl.) and trl.?. Finally, the shape of the rostrum of S. obesus
Kr. agrees nearly with that of the young Petalidiwm from
Stat. 137 just mentioned. I described the branchiz of S. obesus
Kr., stating that in the largest specimen a rudimentary branchia
was found above trl.*; in the largest but one of the specimens I
have now looked for this branchia, and foundit. Finally, I stated
that P. obesum (Kr.) is “decidedly distinct from P. foliaceum Bate,”
and I maintain this opinion, founding it on the fact that P. foli-
aceum does not possess any branchia above trl.*; and it is very
improbable that such a branchia begins to develop itself in the
Mastigopus and disappears again in the adult.
6. Sereestes H. M.-Edw.
SERG. INTERMEDIUS Bate, p. 383 (no figure).
Bate mentions one specimen, 5 mm. in length, from the
“China Sea, off Luzon.” The specimen has been preserved ;
according to the rostrum, the eye-stalks, and the ext. br. of urp.,
it belongs to “ Serg. brachyorrhos Kr.,” which is the youngest
Mastigopus of S. edwardsii Ky. (P. Z.8. 1896, pp. 963-64).
SERG. PREHENSILIS Bate, p. 385, pl. xxi. (Plate XI. figs. 4a,
4b.)
Bate has examined one specimen from Stat. 236, lat. 34° 58’ N.,
Jong. 139° 29' E., off Japan, 775 fathoms. The type, an adult
male, has been preserved; it differs from all other species known
tome. It will be useful to redescribe its most essential characters,
and two new figures are given (Pl. XI. figs. 4a, 46). The
1903. | IN THE ‘ CHALLENGER’ COLLECTION, 57
rostrum is rather long, directed forwards and _ considerably
upwards; its terminal portion is produced into a spiniform
process, and the upper margin of the rostrum has at the base of
that process a sharp angle as a rudiment of a spine; the lower
margin of the rostrum between its base and the apical spine is
strongly convex, the upper margin nearly straight. Supra-ocular
and hepatic spines are wanting, the gastro-hepatic groove slightly
developed. The eyes are large, considerably depressed ; seen from
the side (fig. 4.@), they are somewhat longer than the whole stalk ;
seen from above, their basal margin is very oblique (fig. 4 6), so
that the interior margin of the distal joint of the stalk is as long
as the outer margin of the eyes. The antenn. ped. with the outer
margin of the first joint is a little longer than that of the two other
joints together, and only a little shorter than their inner margin ;
the second joint with the inner margin is three times longer than
the breadth, and somewhat longer than the third joint, which is
about two and a half times longer than broad. The antennal
squama reaches nearly to the end of the antenn. ped., with its
distal portion broad and the outer spine well developed. Mxp.° is
a little shorter than trl.* The branchiz present a transition-form
between those of S. arcticus Kr. (Pl. XII. fig. 1 c) and S. robustus
Smith; the pleurobranchial lamella above mxp.* is very small.
Of the branchie above trl.’ , br.’ is scarcely two-thirds as long as
br.; of those belonging to “Viel ay OTS iS slightly longer than 67."
and br. a little shorter than br.’ above trl.’ The ext. br. of urp.
is almost five times longer than broad, and the hairy portion of its
outer margin is a little more than one-fourth of the total length.—
Length 36°5 mm.
This species occupies an intermediate position between S. arcticus
Kr. and S. robustus Smith. Bate’s fig. 4, showing an antennule
and an eye, is misleading, the antennular peduncle being too
slender, with the basal joint too short, the third joint too long.
SERG. JAPONICUS Bate, p. 387, pl. lxx. figs. 1, 2.
Bate enumerates three specimens from two localities: Stat. 232,
lat. 35° 11' N., long. 139° 28' E., 345 fathoms; and Stat. 207,
lat. 12° 21' N., long. 122° 15’ E., 700 fathoms. All have been
preserved, and belong to one species. In 1896 I wrote that
S. japonicus Bate must be identical with S. mollis Smith (taken
in the Atlantic, off the United States), and gave reasons for
my view. On comparing Bate’s specimens with Smith’s elaborate
description (Rep. U.S. Comm. Fish & Fisher. for 1882, p. 419,
1884) and his figures (Rep. U.S. Comm. Fish &c. for 1885, pl. xx.
figs. 3-5), I arrived at the same result. It should be especially
mentioned that an examination of the branchiz showed the most
complete agreement with Smith’s description and drawing. For
full information on S. japonicus Bate, I refer, therefore, zoologists
to the papers of Smith.
As already mentioned, I found in the bottle with S. japonicus
from Stat. 232 a tube containing a smaller animal determined
58 DR. H. J. HANSEN ON CRUSTAOEANS [Jan. 20,
to be 8. japonicus and labelled Stat. 158. It is not mentioned:
in Bate’s work. On a closer examination it turned out to be a
specimen of Petalidiwm, and it has been described above.
SERG. KROYERI Bate, p. 388, pl. Ixx. figs. 3, 4. (Plate XI.
figs. 5a, 56.)
Bate established this species on one large specimen, from
Stat. 170, lat. 29° 55'S., long. 178° 14’ W., 520 fathoms. The
specimen is very mutilated; a new description with two figures
(Pl. XI. figs. 5a, 5 6) is here given.
The rostrum (fig. 5 a) is rather low, rounded above, with the
upper front angle blunt and slightly projecting and the anterior
margin concave; but it could not be settled whether the upper
margin of the rostrum had been damaged or presented its natural
shape. Supra-ocular and hepatic spines are wanting, the gastro-
hepatic groove is strongly developed. The eyes (fig. 56) are
large, somewhat depressed; seen from above nearly as long as
broad, slightly longer than the outer, and decidedly, but not
much, longer than the inner mar ein of ‘the stalk, On the upper
side of the stalk, close behind the eye and near the inner margin,
is seen a rather small, subeylindrical, distally rounded process,
directed obliquely forwards, inwards, and upwards; it seems to
terminate in an organ. In the antenn. ped. the basal joint is
broad, with the outer margin a little shorter than that of the
two following joints together, but only two-thirds as long as the
inner margin of the same joints; the second joint is stout, with
the inner margin scarcely three times longer than the breadth;
the third joint is stout, but yet considerably more slender than
the second, scarcely three times longer than broad, somewhat
longer than the outer and somewhat shorter than the inner
margin of the second joint. The squama of the antennz seems
to be nearly as broad at the distal end as in S. japonicus Bate,
but it could not be seen whether the outer spine is developed.
Mxp.’ and all tr unk- legs are wanting. The pleurobranchial
lamella above mxp.” is small. The branchiz above trl.’ have been
broken off on both sides; of the branchiz above trl.*, 6r. is nearly
three-fourths as long as 67. above trl.*, while br." is slightly more
than half as long as the same. The ext. br. of urp. with the
apical part is wanting ; the branch seems to have been at most four
times longer than broad, perhaps without marginal spine, and with
the haired part of the margin unusually short.—Length 60 mm.
The species is allied to S. prehensilis Bate and S. robustus
Smith, but it is easily distinguished from all species hitherto
discovered by the process on the eye-stalks,
SerG. artanticus H. Milne-Edw., Bate, p. 389, pls. Ixvili. &
Ixix,
In my earlier paper I wrote (p. 947) that “‘ I am not convinced
that all the specimens from the localities enumerated (p. 390)
belong to S. atlanticus,” and I produced some grounds for that
1903. ] IN THE ‘ CHALLENGER’ COLLECTION, 59
opinion. The investigation of the material preserved in the
British Museum proved the correctness of my disbelief, but, it
must be admitted, to a degree not supposed.
Of the specimens enumerated by Bate, the following have not
been preserved in the British Museum: “ Stat. 42, lat. 35°58’ N.,
long. 70° 35' W., 2425 fathoms,” “length 25 mm.”—and “On
May 6-18th, 1876, in lat. 32° 41’ N., long. 36° 6’ W... one
specimen ... at the surface; and on the 7th of the same month,
near the Azores....two other specimens were taken at the
surface.” I can now state with certainty that if the specimen
from Stat. 42 lived near the bottom in that enormous depth, it
did not belong to S. atlanticus; perhaps it was captured near the
surface, but at all events the locality must be omitted as uncertain.
Most probably the specimens captured in May 1876 belonged to
S. atlanticus. The specimens from the other stations enumerated
by Bate belong to four species, and each station must be mentioned
separately,
“North Atlantic .... Stations 62 and 63, on the passage from
Bermuda to the Azores. Three specimens.” In a bottle labelled
“ Between Bermudas and Azores” I found eight partly mutilated
specimens of S. atlanticus.
“Between Teneriffe and St. Thomas.” In a bottle with the
same locality, three specimens of S. atlanticus.
“Station 320,.... lat. 37° 17'S., long. 53° 52’ W., off Monte
Video; depth 600 fathoms.” Bate does not directly state the
number of specimens, but he writes ‘‘ Length 38 mm.,” and the
meaning is probably that he had one single specimen. Ina bottle
labelled “ off Monte Video” I found six small and badly preserved
specimens of S. atlanticus M.-Edw., and one large specimen of
S. arcticus Ky., and it is decidedly the last-named specimen which
was procured from 600 fathoms. As to S. arcticus Ky., the
student is referred to Kréyer’s work, to the description and
drawings in various papers of 8. Smith, to notes in my earlier
paper, and to some remarks below, in the description of S. similis,
n. sp., together with figs. 1 a—c on Pl. XII.
“Station 159, ... south of Australia ; depth 2150 fathoms. ...
Three specimens... Length 43 mm.” In the collection three
large specimens are present, but they belong to S. arcticus Kr.,
which thus has been proved to be distributed through the deeper
Atlantic, from the southern part of Greenland to lat. 38° 8., and
to south of Australia.
Bate enumerates still two deep-water stations, viz. Stat. 252,
off Japan, and Stat. 173, off Matuku, Fiji Islands; he examined
one specimen from each of these localities, but the animals belong
to two new species to be described below.
But before giving these descriptions I will sum up the results
of the examination of the specimens referred by Bate to S. atlan-
ticus H. Milne-Edw. (=S. fristi Kr.). The Copenhagen Museum
possesses some hundreds of adult specimens of S. atlanticus, taken
at a large number of places in the Atlantic, the Indian Ocean,
60 DR. H. J. HANSEN ON CRUSTACEANS [Jan. 20,
and the western part of the Pacific, and all these were taken
near the surface; the ‘Challenger’ specimens taken near the
surface and referred by Bate to S. atlanticus really belong to that
species, but his specimens secured at the deep-sea stations 320,
159, 232, and 173 belong to three other species.—I have found it
unnecessary to attempt to point out what parts of Bate’s long
description (and which of his drawings) can be applied to S. atlan-
ticus; the zoologist is referred to the good description of Kroyer
together with the additional notes in my earlier paper.
SERG. SIMILIS, n. sp. (Plate XI. figs. 6 a6 d.)
The type of this species is the above-mentioned specimen on
which Bate writes (p. 320): “Stat. 232,... lat. 35° 11’ N., long.
139° 28’ K., off Japan; depth 345 fathoms... . Length 50 mm.”
The species is closely related to S. arcticus Kr. The rostrum
(fig. 6 5) is nearly oblong-triangular, a good deal longer than in S.
arcticus (Pl. XII. fig. 1 6) and directed more upwards ; the anterior
margin of the scutum below the rostrum is strongly convex (fig. 6a)
and protruding, while it is nearly vertical in S. arcticus (Pl. XII.
fig. 1a). The supra-ocular and hepatic spines are well developed ;
the gastro-hepatic groove distinct. The eyes are nearly as in
S. arcticus, large, broader than deep, scarcely as long as broad
(fig. 6c), decidedly shorter than the outer margin of the distal
joint of the stalk, and one-half shorter than its inner margin.
The antenn. ped. (fig. 6c) is about as in S. arcticus; the outer
margin of the basal joint is as long as the same margin of the two
distal joints together ; the second joint is rather slender, its inner
margin almost four times longer than its breadth and scarcely
longer than the third joint, which is slender, about six times
longer than broad. The squama of the antenna is moderately broad
at the end, with the outer spine projecting beyond the terminal
margin. Of the long appendages, mxp.’, trl.*, and trl.’ are wanting,
and the remaining thoracic legs are about as in S. arcticus. The
branchie above trl.’ and trl.* (fig. 6d) present excellent differences
from those in S. arcticus (Pl. XII. fig. 1c); br.* above trl.’ is not
quite half as long as br. above trl.*, and in spite of this not incon-
siderable length it is semi-rudimentary, having only some posterior
branches, nearly all short, and no anterior branches. Of the
branchiz above trl.’, br. is long, four-fifths as long as br. above
trl.?; 6r.1 is more than half as long as br., very well developed, at
least as long and more than twice as broad as br." above trl.’
[In S. arcticus (Pl. XII. fig. 1 c)—from a specimen obtained in the
most northern part of the Atlantic—br.’ above trl.* is about three-
fourths as long as br. above trl.*, with well-developed branches
on both sides; of the branchie above trl.*, br. is only a little more
than half as long as hr. above trl.*, while 6r.! is small, considerably
narrower and shorter than br.’ above trl.°] The ext. br. of urp.
has the apical portion broken off, but it is narrow, and seems to
have been a little more than five times longer than broad, thus
slightly narrower than in S. arcticus, but otherwise of the same
aig SE Se es ST SD a PS
1903. ] IN THE ‘ CHALLENGER’ COLLECTION. 61
shape and with the marginal spine well developed.—Length
o4 mm.
The species is easily distinguished from S. arcticus Kr. by the
shape of the rostrum, together with the anterior margin of the
scutum, and especially by the branchiz above trl.* and trl.’; in
all other features these two species are closely allied.
SERG. CHALLENGERI, n, sp. (Plate XII. figs. 2 a—2 n.)
The type of this species is the specimen on which Bate writes
(p. 390): “Station 173, July 24th, 1874; lat. 19° 9' 35” S.,
long. 179° 41’ 50” E.; off Matuku, Fiji Islands; depth 315
fathoms; bottom, coral mud. One specimen, male. Dredged.
Length 24 mm.” He referred it to S. atlanticus. I have the
pleasure of appending the name of the renowned ship to this
Sergestes, which is one of the most interesting species of Crustacea
secured by the expedition. Unfortunately the single specimen is
very mutilated.
The rostrum (fig. 26) is rather low, short, obliquely triangular,
turning somewhat upwards; its apex is acute and very slightly
produced. The supra-ocular spine is wanting; the hepatic spine
is rudimentary; the gastro-hepatic groove (fig. 2 a) is deep, and
the cervical groove very distinct. The eyes have been broken off,
only the basal part of the stalks being left. In the antenn. ped.
(fig. 2c) the outer margin of the basal joint is a little longer than
that of the two other joints together; the second joint is
moderately robust, its inner margin a little more than three
times longer than the breadth; the third is slightly more than
two and a half times longer than broad, a little shorter than the
outer margin of the second joint, and only three-fifths as long as
the inner margin of the last-named joint. The squama of the
antenna is distally very broad (fig. 2c), with the outer spine
scarcely projecting beyond the terminal margin. Mxp.’ and trl.*—
trl.’ are wanting; of trl.’ the apical part has been lost, but these
legs seem to have been a little longer than mxp.*, and to be more
slender than in S. arcticus, but otherwise not showing any
difference of importance. Of the branchie (fig. 2d), br. above
trl and trl.’ are long and narrow; br.’ above tri.? is as usual a
lamella; br." above tril.’ is slightly more than one-third as long
as br., especially with its anterior branches well developed; of the
branchie above trl.*, br. is about three-fourths as long as br. above
trl.’, while br." is proportionately large, much longer and broader
than br.' above trl.*, and even more than half as long as br. above
trl” The ext. br. of urp. (fig. 22) has the apical portion
wanting, but the branch seems to have been nearly five times
longer than broad, with the marginal spine well developed and
situated as in S. arcticus.—Length 23 mm.
By the shape of the joints of the antenn. ped., the development
of the branchie above trl.* and trl.*, and the shape of the ext. br.
of urp., this species is related to S. robustus Smith, 8. prehensilis
Bate, and S. kréyert Bate. But it is sharply distinguished from
62 DR. H. J. HANSEN ON CRUSTACEANS [Jan. 20,
all other species hitherto known by possessing an enormous multitude
of luminous organs arranged regularly on the lower surface and
near the lower lateral margins of the cephalothorax, on the six
abdominal segments (figs. 2d, 7, &), on the sides of the shield
(fig. 2a), and on all the appendages preserved (figs. 2c, d,e, f, g, h)
with exception of the maxillule and maxille. The organs are
easily seen; they look almost similar to the eyes in Aranee, and
they differ much in size and direction. I have deemed it advisable
to deal with the special arrangement and the structure of these
organs in a separate section of this paper.
SERG. DORSISPINALIS Bate, p. 394, pl. lxxil. fig. 1.
Bate does not state directly the number of specimens, but he
had probably only one. The length is “9 mm.” and the locality
“south of Australia, March 1874.” An animal with a label of
exactly the same contents was preserved in a microscopical pre-
paration. It isa Mastagopus related to S. arcticus Kr., S. similis,
n. sp., &e., but it could not be referred to any adult form. In
the preparation the abdomen is seen from the side, and the
cephalothorax essentially from below; the spine on the scutum
on which Bate writes “just anterior to the [cervical] suture, in
the median dorsal line, is a small, anteriorly directed tooth,” is
in reality the gastro-hepatic spine, which besides has been drawn
in a position too remote from the front. Bate’s figure is not
correct in some other respects: in the antenn. ped. the second
joint is about as long as the third, and both together somewhat
shorter than the first; the eyes reach to the end of the first joint ;
the squama is longer than in the figure, reaching almost beyond
the second joint of the antenn. ped. and distally narrow; the ext.
br. of urp. is a little longer and conspicuously more narrow than
in the figure, while its marginal tooth is indicated correctly.
SERG. LATERODENTATUS Bate, p. 395 (no figure).
Bate has examined one specimen, measuring 8 mm. in length,
and captured “ south of Australia, March 1874.” The type could
not be found in the Museum. According to the description it is
a Mastigopus belonging to the arcticus-group, probably a younger
stage of the same species as that to which the preceding larva,
S. dorsispinalis Bate, belongs.
SuRG, NASIDENTATUS Bate, p. 398, pl. Ixxn. fig. 2.
Bate does not state directly the number of specimens, and
presumably he had only one; the length was 10 mm. and the
locality the Pacific Ocean, between Valparaiso and Juan Fernandez.
The type does not exist in the Museum. The species is a
Mastigopus ; in wy earlier paper I had already referred it to the
same group as S. arcticus, and nothing further can now be added.
Sura. prapontius Bate, p. 399, pl. lxxil. fig. 3.
Bate does not state the number of specimens, but presumably
1903.] ; IN THE ‘CHALLENGER’ COLLECTION, 63
he had only one. The length is 18 mm. and the locality the
Atlantic Ocean, April 7, 1876. In a preparation bearing the
name of the species, and besides “ Surface, 7 April, 1876,
Atlantic,” one specimen is preserved: it must be Bate’s type,
but it measures only 16°5 mm. in length. In Bate’s figure the
armature on the dorsal line of the abdomen is not correct ; on
the third segment a spine has either been broken off or is rudi-
mentary, the base being distinct ; the spine on the fourth segment
is only half as long as that on the fifth, shorter than in the figure,
and directed obliquely backwards; on the sixth segment a very
short spine is visible. The basal part of the rostrum is somewhat
ascending, the distal spiniform and horizontally porrected. The
ext. br. of urp. has the outer margin hairy in about § of its
length, and the marginal tooth is very small. The distal part of
mxp.° is very incorrectly drawn by Bate in his fig. 37; it is four-
jointed; the third joint is short, much shorter than the fourth,
and both together about as long as the second; the first of these
joints terminates below in two strong setiform spines, both some-
what longer than the second joint, which terminates in two spines
of the same length as the preceding pair; the fourth joint
terminates also in two spines, somewhat, but not much, shorter
than the four just mentioned. (I cannot understand how Bate
drew his misleading figure; it must prevent every student of
his Report from arriving at a correct judgment.) In the British
Museum I have drawn tolerably accurate sketches of the rostrum,
the ext. br. of urp. and the distal part of mxp.*, and I have
compared them with a few specimens in the Copenhagen Museum,
previously determined and shortly described by me as the Wasti-
gopus of S. penerinkti Bate, H. J. H.: I found the agreement to
be so close, that I must consider JS. diapontius Bate and S. pene-
rink Bate (the type of the latter form unfortunately is not present
in the Museum) as two Mastigopus-stages of the same species ;
the type of S. penerinkii Bate measured only 8 mm. in length
and is a young Mastigopus, while S. diapontius Bate is the large
larva.
Bate describes S. diapontius on p. 399, S. penerinkit on p. 418;
and the name S. diapontius must therefore be accepted for the
species. In my earlier paper I described the black-eyed adult
form as S. penerinkii Bate, H. J. H., but I think that jt must
now be necessary to adopt the name JS. diapontius, not only for
the Mastigopus, but also for the mature form, which therefore in
the future must be named S. diapontius Bate, H. J. H.
SERG. ARMATUS Kroyer, Bate, p. 401, pl. Ixxiii. fig. 1,
Bate enumerates three localities: one of them is “September
12, 1875, between Japan and Honolulu, South Pacific Ocean ” ;
the second is “Station 256, July 21, 1875; .... north of the
Sandwich Islands; depth 2950 fathoms.” But in the collection
I found a statement aberrant from both, viz. : “Surface. J apan
to Honolulu, July 1875. Type.” This bottle contained two
64 DR. H. J. HANSEN ON CRUSTACEANS [ Jan. 20,
small specimens, but neither of them can be the type for Bate’s
figure. Both specimens belong to Group II. in my earlier paper,
but neither of them belongs to S. armatus, both having on the
rostrum a well-developed sub-basal dorsal spine, which is absent
in S. armatus Kr. and in Bate’s figure. I have been unable to
refer the specimens, which measure about 8-5 and 11 mm., to any
species known to me, and I thought it useless to describe and
figure them.—One specimen from Port Jackson, Australia, the
third locality in Bate’s report, measures at most 5°2 mm. without
the rostrum, which has been broken at the middle, but possesses
a very fine sub-basal spine. It is so small and so badly preserved
that a reference to any species has been impossible.
Finally I found a specimen from “Sidney,” determined by Bate
as S. armatus, but not mentioned in his work. It is only as long
as the preceding specimen and impossible to determine.
The result is that I have perhaps not seen the specimen figured
by Bate, which may belong to S. armatus Kr., and, according to
the explanation of the plate, measured about 10 mm. in length,
while his specimens of this length examined by me disagree with
his figure by possessing a sub-basal upper spine on the rostrum.
SERG. EDWARDSII Kroyer, Bate, p. 403, pl. Ixxiil. fig. 2.
Bate enumerates three localities. The first 1s ‘“‘ North Atlantic,
April 1873”: in the Museum I found a specimen labelled
“14 April, 73, off Africa, surface,” which most probably is that
indicated in the text, and it belongs to S. edwardsi Kr. The
second locality is “‘ Pacific Ocean, surface, September 1875”: in
the collection a specimen bearing the same inscription is 5. edwardsi
Kr. From the third of Bate’s localities, Cape Verde Islands, I
found no specimen, but a specimen without locality and signed
“type” is an adult specimen of S. edwardsii Kr. (That Bate’s
statement “ Greenland (Av dyer)” is wrong here, and in almost all
other places, has already been pointed out both by Ortmann and
myself.)— Furthermore, I found two small specimens of S. oculatas
Kr., the Mastiyopus of S. edwardsii Kr., which had been deter-
mined by Bate as S. edwardsti and labelled “ Aug. 23, 1873,
lat. 2° 25’ N., long. 20° 1’ W., 100 fathoms,” but these specimens
are not mentioned in Bate’s Report..
Bate’s description of the characters of S. edwards Kr. is in-
complete; the reader is referred to my earlier paper.
Sere. RINKU Kroyer, Bate, p. 404, pl. Ixxii. fig. 3.
Bate mentions two localities: “‘ New Hebrides, August 23,
1874,” and “South Pacific, 1875.” From the first of these
localities the anterior half of a specimen was present. Further-
more, I found two specimens labelled “ Oct. 19, 1875, S. Pacific,
drawn,” and one and a half specimen labelled “Oct. 18, 1875,
surface”: both these localities are in all probability identical with
the second one in Bate’s Report. It may be very possible that all
these specimens belong to S. rink Kr., which is the Mastigopus
1903. ] IN THE ‘ CHALLENGER’ COLLECTION. 65
of S. arcticus Ky., but having in London no material from the
North Atlantic of S. rinkit Ka., for direct comparison of minute
details, and some other species allied to S. areticus being known
from the Indian Ocean and the Pacific, I could not determine the
larvee enumerated with absolute certainty.
One specimen from Cape York, determined as S. rinkii, but
not mentioned in the Report, is S. corniculum Ky., im a stage a
little younger than that figured by Bate as the last-named species,
One specimen, labelled “ 25. 8. 73,” determined as S. rinkit,
but not mentioned in the Report, measures nearly 7 mm.; it
belongs to S. penerinkii Bate, the young larva of S. diapontius
Bate, H. J. H.
Sere. ocuLatus Kroyer, Bate, p. 406, pl. lxxiv. fig. 1.
Bate enumerates six localities. From Stat. 106 three specimens
were obtained, from Stat. 257 one specimen, from Stat. 103 one
specimen, finally from ‘“ September 12, 1875, South Pacific,” one
specimen: all six specimens correctly referred to the Kroyerian
species, which is the Wastigopus of S. edwardsti Kr. (compare my
earlier paper). From the two remaining localities, viz. ‘“‘ North
Pacific, near the Sandwich Islands, August 21, 1875,” and
“ August 27, 1873... off St. Paul’s Rock,” no specimens could
be discovered.
Sere. ovatocuLus Bate, p. 408, pl. xxiv. fig. 2.
Bate gives the locality “The North Atlantic Ocean.” In the
collection I found three specimens with the label ‘ 14 June, 1873,”
which agrees with my quotation from the text. These three
specimens are identical with S. ancylops Ky., which is the
Mastigopus of S. atlanticus H. Milne-Edw.
SERG. PARVIDENS Bate, p. 409, pl. xxiv. fig. 3.
Bate has the. following localities: ‘The tropical part of the
Atlantic; Pacific Ocean, north of the Sandwich Islands; off
Sydney and Wellington, Australia.” Just below he writes:
“Specimens of this species or variety were taken during the
passage from Teneriffe to St. Thomas”; and in the collection one
specimen from the last-named locality is present: it belongs to
S. vigilax Stimpson, the Mastigopus of S. vigilax Stimps., H. J. H.;
it agrees with Bate’s description and figure, and it had already
been pointed out by Ortmann and myself that S. parvidens Bate
belonged to S. vigilaw Stimps. Examples from the other localities
mentioned by Bate could not be found.—From “ Cape York” two
specimens determined as S. parvidens were present: one of them
is a young S. oculatus Ky. (the Mastigopus ot S. edwardsii Kv.);
the other is allied to S. incertus H. J. H., but is so badly preserved
that a determination was impossible. In a preparation a small
specimen from the “ China Sea,” determined as S. parvidens, was
preserved ; it seems to be S. ocwlatws Ky., but it is in a very bad
condition.
Proc. Zoou. Soc.—1903, Vou, I. No. V. 5
66 DR. H. J. HANSEN ON CRUSTACEANS [Jan. 20,
Sere. cornnicuLum Kroyer, Bate, p. 410, pl. lxxv. fig. 1.
Bate writes: ‘Cape York; north of New Guinea; North-west
Pacific.” In the collection I found one specimen from Cape York
labelled “type,” furthermore, one and a half specimen from the
same locality, all correctly referred to the Kréyerian species. In
a preparation I found a specimen determined as S. corniculum
from “N.W. Pacific”; it measures 10°4 mm. in length, but
according to the shape of the ext. br. of urp., the eyes, &e., it is
not that species but S. ancylops Kr., the Mastigopus of S. atlanticus.
From the third locality, ‘north of New Guinea,” no specimen
could be found.
SrrG. ancyLors Kroyer, Bate, p. 413, pl. Ixxv. fig. 2.
Bate has two localities: “New Hebrides; Pacific, July 20,
1875.” I found one specimen from “ New Hebrides” correctly
referred to the Kréyerian form.—In three other tubes, specimens
not mentioned in Bate’s text were present; they belonged to
three other species, but are mutilated and could not be determined.
SERG. LONGIRosTRIS Bate, p. 415, pl. Ixxv. fig. 3.
Bate writes: ‘Mid Atlantic, April 1876,” and according to
the following line he had one specimen. But in the collection I
found three tubes, each containing one specimen, all determined
as S. longirostris, and two of them from ‘ N. Atlantic,” while the
third had no locality ; all three specimens are the young MZastigopus
of S. corniculum Kr., H. J. H. The specimen without locality
seems to be the type for Bate’s figure of S. longirostris, but it
measures scarcely 3mm. in length, the rostrum not included, and
in Bate’s figure the eye-stalks are too long, the eyes too small, and
the third joint of the antenn. ped. too short (it is in reality as
long as the second).
SERG. JUNCEUS Bate, p. 416, pl. lxxvi. fig. 1.
According to the text Bate has seen one small specimen from
the “South Pacific Ocean.” It could not be detected in the
collection. But in my earlier paper I pointed out that it must
be the young Mastigopus of S. tenwiremis Ky., H. J. H.
SERG. LONGISPINUS Bate, p. 417, pl. lxxvi. fig. 2.
Bate has two localities: ‘Station 106... Mid Atlantic”...
and “Station 354.... North Atlantic Ocean.” The collection
contained two specimens referred to this species and labelled
“93 Aug. 1873, 70 meters ”; in the text we find “Station 106,
August 25, 1873; lat. 1° AT’ N., long. 24° 26’ W.; Mid Atlantic
Ocean”; but in spite of the small difference as to the date
(probably originating from a misscript) I am sure that one of
the two specimens was captured at Stat. 106. The other
specimen is probably from Stat. 354. Bate writes: “the specimen
from the tropical part of the Atlantic” and “that from the North
Atlantic,” which indicates that he had only one specimen from
1903. ] IN THE ‘CHALLENGER’ COLLECTION. 67
each of the two stations mentioned ; and I suppose, therefore, that
both specimens have later on been put together in the same tube.
Both specimens, which are in a very bad condition, belong eprrectly
to S. longispinus Bate (the direction of some of the dorsal spines
on the abdomen is very characteristic), and this form is in reality
the Mastigopus of S. cornutus Ky. (compare my earlier paper,
p- 953).
SERG. PENERINKII Bate, p. 418, pl. lxxvi. fig. 3.
Bate records the length to be 8 mm. and the locality ‘* North
Atlantic Ocean.” No specimen referred to this species could he
found in the collection. But above I have mentioned a specimen
of this species referred by Bate to S. rinkii, and it may perhaps
be that described by him as S. penerinkii. He writes (p. 419)
that the last-named “species bears a strong resemblance to
Sergestes rinkit Kroyer, but differs . . .”.. The specimen in question,
referred by him to S. rinkii, measures nearly 7mm. in ype and
was captured ‘“ 25. 8.73,” that is to say, Mid Atl
earlier paper I have described the JJastigopus, and ae the
mature form as S. penerinkit Bate, H. J. H.; but above it is
pointed out that this name must be cancelled as a synonym, and
the species be named S. diapontius Bate, H. J. H.
SERG. FERMERINKILE Bate, p. 419, pl. Ixxvi. fig. 4.
Bate has examined one specimen, captured in the ‘ Pacific
Ocean, lat. 24° 8., long. 148° W.,” and measuring 5mm. The
specimen is not to be found in the Museum collection. According
to the figure it is a very young Mastigopus belonging either to
S. diapontius Bate, H.J.H., or toa closely-allied species of the
same group.
SERG. LONGICOLLUS Bate, p. 421, pl. Ixxvii. fig. 1.
Bate enumerates two localities. The first of them is “ South
Atlantic Ocean, October 5, 1873; near Station 131 i lat. 29° 35°
S., long. 28° 9' W.” In a preparation labelled “5.10.73” I
found the large specimen drawn by Bate; in a tube labelled
“surface, at night, 5 Oct. 1873, South Atlantic,” a small specimen
was preserved. Thus both specimens are from the first locality
in the text, and both belong to S. longicollus Bate, which is the
Mastigopus of S. tenuiremis Kroyer, H. J. H. In the large
specimen drawn by Bate the eyes present a thick yellowish layer
around the black central part; the dorsal spine on the sixth
abdominal segment is exceedingly small.
The second locality in Bate’s text is ‘‘ Station 295, November 5,
1875; lat. 38° 7'S., long. 94° 4" W.; South Pacific Ocean ; depth
1500 fathoms; ... Taken at night.” In the collection a specimen
is preserved labelled “5. Nov. 75, night, 8. Pacific, surface.”
The determination is correct; and the specimen, which measures
9-2 mm. in length, is evidently that mentioned by. Bate; but
the label proves that it was taken at the surface, wherefore
~
5*
68 DR. H. J. HANSEN ON CRUSTACEANS [Jan. 20,
the depth recorded in the text is most misleading.—In a third
tube two specimens labelled 18/10 75 were preserved; they had
been correctly referred by Bate to his S. longicollus, but are not
mentioned in his text.
A preparation contained a specimen of the same species,
labelled ‘“ Sergestes tenwiremis, South Atlantic, 6.10.73.” It is
not mentioned by Bate, who even writes (p. 421) that ‘“ No speci-
men in the ‘Challenger’ collection corresponds precisely with the
description and figure given by Kroyer” of S. tenwiremis. On
the other hand, S. tenwiremis Kr. and S. longicollus Bate are, as
already stated, the same species.
SERG. PRECOLLUS Bate, p. 423, pl. Ixxvil. fig. 2.
Bate has examined one specimen, measuring 25 mm., from the
“North Pacific Ocean.” The specimen could not be found.
The shape of the eyes in Bate’s figure shows that it has been a
large Mastigopus. In my earlier paper I write (p. 958): “is at
least rather nearly related to S. corniculwm Kr., from which it seems
to differ by a somewhat different shape of the ext. br. of urp.,
and by having the fifth abdominal segment ‘dorsally produced to
a point.’”
SerG. semrarmtis Bate, p. 423, pl. Ixvii. fig. 1.
Bate has two localities: ‘“‘ West Pacific Ocean” and “ Station
oan May iG UISiO is)... Maids NorchieAtlamties 70) Auber mulme
description he writes: ‘ Observations——A specimen (pl. Ixvii.
fig. 2) very similar to the type was taken in the Atlantic ...”;
and then he describes the specimen. I must suppose that he
considered the specimen from the ‘“ West Pacific Ocean” to be
the type of his S. semiarmis, and that the specimen described
separately (p. 425) is that from “... May 6, 1876....” Neither
of the specimens could be detected in the Museum; both are
larve, but a reference is impossible. In the collection I found
one specimen determined as S. semiarmis and labelled “13 April
1876, Atlantic, off coast of Africa, surface”; it is a MJastigopus
of S. diapontius Bate, H. J. H.
SERG. LAVIVENTRALIS Bate, p. 425, pl. Ixvii. fig. 3.
Bate had probably only one specimen, 7 mm. in length, from
“North of New Guinea”; it could not be found in the Museum.
It isa young Mastigopus belonging to a species closely related to
S. arcticus Kr.
SERG. SPINIVENTRALIS Bate, p. 426, pl. Ixvii. figs. 5 & 6.
Bate has only one locality: “North Pacific Ocean”; the
animal described measured only 3-5 mm. in length, and some of
its parts are shown in figs. 5a, 51, and 5v, but fig. 5 v, repre-
senting the ext. br. of urp., does not correspond at all with the
description in the text (p. 427). The animal is not to be found
in the collection: it is a small Mastigopus related to S. vigilax
Stimps. and allied species. Bute’s fig. 6a represents the head
1903. ] IN THE ‘ CHALLENGER’ COLLECTION. 69:
of “ Sergestes spiniventralis var.” from the “ West Pacific”; it
has been suggested that the animal is allied to—or identical
with—the Mastigopus of S. vigilaw Stimps., H. J. H., but the
specimen could not be found. The collection contained one
specimen determined as S. spiniventralis and labelled ‘Sidney to
Wellington, 17. 6. 74” ; it measures about 7°5 mm. in length,
and the naked part of the outer margin of the ext. br. of urp. is
slightly more than one-fourth of its length. The specimen is in
all probability a young MJastigopus of S. vigilax Stimps., H. J. H.
SreRG. PROFUNDUS Bate, p. 428 (no figure). (Plate XI. figs. 3a,
30.)
Bate has referred two specimens to this species. He describes
each specimen separately: the first of them, from Stat, 137,
belongs to Petalidiwm, perhaps to P. foliacewm, and has been
dealt with above. ‘The other specimen, from “Stat. 300,
December 17, 1875; lat. 33° 42’ 8., long. 78° 18° W.; west of
Valparaiso; depth 1375 fathoms; .... Trawled,” is a real
Sergestes, related to S. inows Faxon, but differing in the shape of
the rostrum. Having removed the first-mentioned specimen
from the genus Sergestes, I should think it justifiable to apply
the name S. profundus Bate to the last-named specimen, instead
of proposing a new name. ‘The animal is quite membranous,
and is crimson everywhere—a fact proving that it lives in the
depth of the sea, and that its colour has been durable to the
highest and most unusual degree. The posterior part of the
abdomen is wanting, and the animal is on the whole so mutilated
and flabby, that I would have omitted it if it had not been
described by Bate; but for that reason I have thought it necessary
to add some notes with two figures (Pl. XI. figs. 3a, 36). It
agrees with S. mows Fax. as to the membranous quality of the
skin and the posterior branchie, but differs from it in the shape
of the rostrum, which is of moderate length, with the upper
margin somewhat, and the lower margin partly, strongly convex,
and distally it is produced in a moderately short spine (Pl. XI.
fig. 36). (Unfortunately I have not seen any specimen of the
gigantic species S. inous Fax., and can therefore not decide whether
the specimen described by Faxon had the rostrum uninjured.)
Supra-ocular and hepatic spines are wanting. The eyes (Pl. XI.
fig. 3a) are black, comparatively small, somewhat shorter than
broad, not half as long as the eye-stalks, and not broader than the
distal end of the stalks. In the antenn. ped. the outer margin
of the first joint is almost as long as that of the two distal joints
together, therefore somewhat shorter than their inner margin ;
the third joint is a little shorter than the inner margin of the
second, and seems to be about three and a half times longer than
deep. The squama does not reach the end of the antenn. ped.,
and the outer distal spine is well developed. According to Bate
the part preserved measures 18 mm., and the probable length of
the entire animal is 24 mm.
70 DR. H. J. HANSEN ON CRUSTACEANS [Jan. 20,
SERG. VENTRIDENTATUS Bate, p. 431 (no figure).
Bate gives the locality “north of the Sandwich Islands” and
the length “7 mm.” <A specimen labelled in accordance with the
text is preserved in balsam; it is a young S. oculatus Kr., the
Mastigopus of S. edwardsti Kr.
SERG. UTRINQUEDENS Bate, p. 433 (no figure).
Bate gives the locality “ North Pacific Ocean,” and the length
“3:5 mm.” Nospecimen could be found in the collection. Bate’s
specimen is a very young J/astigopus ; in my earlier paper I
have placed it near S. corniculum Ky., but a final interpretation
is Impossible to me.
SERG. DISSIMILIS Bate, p. 437 (no figure).
Jn the collection one specimen, from Cape Verde Islands, is
preserved; it is certainly the type described by Bate. In my
earlier paper I had determined it as one of the larval stages of
S. arcticus Kr., but this is not correct. The rostrum is siightly
more than one-third the length of the eye-stalks, its basal part
somewhat ascending, with a very small spine on the upper angle,
its distal part 1s a slender horizontal spine. The hepatic spine 1s
short. The eyes are only a little higher than the distal end of
the eye-stalks, and these increase gradually in thickness from the
base outwards. In the antenn. ped. the distal half of the first
joint has the margins nearly parallel, and the two other joints are
a little thicker than in the corresponding Mastigopus of S. arcticus.
Fourth to sixth abdominal segments each with a very small dorsal
spine directed backwards; the spine on the sixth segment is the
longest. The ext. br. of urp. is almost, but not quite, five times
longer than broad. All these characters agree with those found
in a Mastigopus of S. mediterraneus H. J. H. preserved in the
Copenhagen Museum ; and sketches drawn in London of the shape
of the distal part of the squama and the telson agree also with
the last-named form. I must therefore consider S. dissimilis Bate
as identical with S. mediterraneus H.J.H. The result is that
the last name must be withdrawn as a synonym, and the species,
of which the sub-adult stage has been described in my earlier
paper, must be called S. dissimilis Bate, H. J. H.
II. Some Results of the Investigation.
In my earlier paper on Sergestes I have paid as much attention
as possible to the animals described by Bate; in nearly all cases
I was able to state whether the specimen was an adult form or a
Mastigopus, and several of the species were interpreted. After
the study of the material preserved in the British Museum, I have
now been able to confirm most of my earlier statements, to
interpret an additional number of the specimens mentioned by
Bate, and to correct two faults committed by myself. I had
erroneously referred S. dissimilis Bate to S. arcticus Kyr., instead
of identifying it with S. mediterranews H. J, H. (see above).
1903.] IN THE ‘ CHALLENGER’ COLLECTION. 71
Furthermore, I had divided a number of species, enumerated on
p- 949 as Group I. A. 6. 3, into two sections, according to
difference in the thickness of the distal joints of the antenn. ped.;
but a study of the types of S. prehensilis Bate and S. kréyeri
Bate showed that Bate’s drawings of the antennule of these
species are incorrect and misleading, wherefore my arrangement
of them was wrong.
It may be useful to put together the alterations and additions
which may now be accepted in the Conspectus on pp. 949-51 in my
earlier paper. In Group I. the following particulars must be
added or altered :—To S. atlanticus H. M.-Edw. belongs only a
part of S. atlanticus sens. Bate, besides the form referred by Bate
to S. aneylops Kr. From S. arcticus Kr., 8. dissimilis Bate must
be removed, and the latter species is to be established separately
with S. mediterraneus H. J. H. as a synonym ; furthermore, some
of the specimens referred by Bate to S. atlanticus belong to
S. arcticus. Near S. arcticus Kr., must be inserted S. similis
H. J. H., established on one of Bate’s specimens of S. atlanticus.
S. prehensiis Bate and S. kréyeri Bate must be removed from
their place and inserted above near S. japonicus Bate, together
with S. profundus Bate, in its new restriction, and S. challengeri
H. J. H., established on one of the specimens referred by Bate to
S. atlanticus. SS. longirostris Bate must be withdrawn as being
a Mastigopus to S. corniculum Kr., H. J. H., and S. cornieulum
sens. Bate is the same species. As uncertain remain: S. dorsi-
spinalis Bate, S. laterodentatus Bate, S. nasidentatus Bate, S. levi-
ventralis Bate, S. rinkii Bate, ¢ Ky, all larval forms belonging to
species related to S. areticus Kr., or perhaps partly belonging
to S. arcticus itself; furthermore, the larve S. precollus Bate,
S. utrinquedens Bate—both at least rather closely related to
S. corniculum Kr., H. J. H.,—and S. semiarmis Bate.
In Group IT. there is hardly anything to alter, but some additions
to make. To S. edwardsii Kr. belongs S. edwardsii Ky., Bate,
S. oculatus Kr., Bate, S. intermedius Bate, and S. ventridentatus
Bate. WS. penerinkii Bate must be cancelled as synonymous with
S. diapontius Bate; and the adult form described as S. penerinkii
in my earlier paper must be named S. diapontius Bate, H. J. H.
The rest of Group II. remains unaltered. S. fermerinkii Bate,
S. spiniventralis Bate, and the species referred by Bate to
S. armatus Ky. could not be interpreted.
Besides, the investigation has yielded some results on the
bathymetrical and geographical distribution of some of the species.
It has been proved that the large specimens (exceeding 30 mm.
in length) referred by Bate to S. atlanticus Ky. are deep-sea
forms belonging to other species: S. atlanticus is common near
the surface; according to Ortmann it has been taken in the
intermediate net from 700-500 m., but it does not live in the
greater depths of the sea. S. arcticws Kr. has a very wide
geographical range, through the deeper to very deep tracts of the
Ocean (see above); S. japonicus Bate has been captured in
the northern part of the Atlantic and the northern part of the
72 DR. H. J. HANSEN ON CRUSTACEANS. [Jan. 20,
Pacific. S. atlanticus H. Milne-Edw., S. edwardsti Kx., S. vigilax
Stimps., H. J. H., 8. tenwiremis Kr., H.J. H., and S. corniculum
Kr., H. J. H., have been proved to be distributed through the
tropical and subtropical parts of the Oceans almost around
the globe, viz., from lat. 23° N., lat. 32° N., or even lat. 42° N.
in the Atlantic, throughout the Indian Ocean to Australia, New
Hebrides, and “South Pacific.”
III. Luminous Organs im Sergestes challengeri, n. sp.
(Plate XII.)
The luminous organs briefly mentioned above are generally
easy to discover; each resembles a very convex, vitreous, faintly
yellowish lens, which is circular and sharply defined. They differ
much in size, some of them being very small and many pro-
portionately large. It may be advisable, first, to give a con-
spectus of the organs observed on the single and unfortunately
very mutilated specimen, next to add some remarks on their
distribution, then to describe their structure, finally to compare
them with luminous organs in other Crustacea.
Conspectus of the Organs observed.
Organs.
On each side of the scutum a row with four organs. 8
Omitherclypeus one organi rece tee leper ee eee eeeEer
On the lower side of the head one unpaired organ
and one near the lateral margin.................. 3
On the lower surface of the thorax and on its infero-
lateral amano Sip.sec ie cicinde uae eeeeteck eee eeny eee 26
On the lower side of the third joint of the antenn.
[QOL OLS) WIAA SH coh adadoos4un oHsosodosobosoGCoHADE +:
On the lower side of each squama four organs ......
On the mandible and its palpus two organs .........
On the first maxilliped two organs.....................
On the second maxilliped five organs .................. Ih
On the first trunk-leg three organs ..................
On the lower surface and on the lateral wall of the
firstiabdominaliseamlentipree.asscen---eeee eee eee
Do., do. of the second abdominal segment ............
Do., do. of the third abdominal segment ............
Do., do. of the fourth abdominal segment............
On the lateral wall of the fifth abdominal segment.
Along the median line of thesixth abdominal segment.
On the outer side of the basal joint of each of the
pleopodsiomejorsamyten cena: cpiieiyieee an eer eee
On the peduncle and on the inner branch of each
VEO} COLO IO) CLAZENAS, "Sodan quads sasecouscsogonoeosococ 4
On the lower side of the outer branch of each uropod
UV OROLESIINS 655 4NG soc an dgd CoGuaRREROOa y yanooEaoCaseod 75 4
AO, FS Or
or ES Sermon)
—
(=)
Total number of organs ... 117
1903. ] IN THE ‘ CHALLENGER’ COLLECTION, (3)
The eye-stalks, the maxillipeds, and the four posterior pairs of
the trunk-legs have been broken off. I am convinced that at
least most, and perhaps all, these appendages possess some organs,
and the lowest number the species possesses must therefore be
about 150!
On the distribution and direction of these organs the following
remarks may be offered. The four organs on each side of the
scutum are arranged rather close in a longitudinal row situated
on the ridge bordering the branchial cavity above (fig. 2a). The
organ on the clypeus is large: one organ is situated on the
segment bearing the antennul, and one on the lower surface of
the head near the lateral margin a little in front of the mandible ;
these four organs look essentially downwards. The arrangement
of the organs on the three posterior thoracic segments can be seen
in figs. 27 and 2d. It is observed that two large organs are
placed at the lower margin of br. above trl.* and of 67." above
trl.“ These organs look outwards and downwards, ard the part
containing the glandular mass &e. behind the lens is somewhat
protr uding, which produces an aspect as if these organs had been
inserted on the end of a kind of rather thick, short stalk. The
remaining organs on the segments mentioned are found on the
lower surface (fig. 27); those placed at the inner angle of the
legs are small or very small, while some in the median line are
large. In fig. 22 fifteen thoracie organs have been drawn; the
remaining eleven thoracic organs are situated on the anterior
segments and arranged in a rather similar way. The four organs
on the lower surface of each squama of the antenna are arranged
for some distance In a row; two of them are seen through the
squama in fig. 2c, The mandible (fig. 2¢) has one organ below
at the antero-inferior margin near the insertion of the palp;
another organ is seen on the lower surface of the first joint of the
palp near its distal end. The first maxilliped (fig. 2,/) has on
the upper side one organ just at the origin of the exopod, and one
on the lower side of the following joint of the endopod; the first-
named organ looks forwards and a little inwards (fig. 2 g), and
the upper margin above it is produced nearly as a lamella, over-
lapping the major part of the lens when seen from above (figs. 2 9
& 2). Of the organs on the outer—in the natural position of
the appendage in reality the lower—surface of mxp.” (fig. 2),
that at the base of the third joint is very small and the others
large; the organ on the first joint looks essentially downwards,
and is “stalked,” as the above-described organs near the lower
margin of the posterior branchie. Of the three organs on trl.’,
two are placed on the inner side of the long fourth joint, one near
the base and the other near the distal end; the third organ is
situated on the anterior side near its end. Hach of the five
anterior abdominal segments has a large organ on the oe eh
of the anterior margin of the Jateral plate (figs. 2d & 27%);
looks forwards and somewhat downwards, besides comietiies a
little outwards. Hach of the two anterior segments has besides
74 DR. H. J. HANSEN ON CRUSTACEANS [Jan. 20,
on the lower surface (fig. 27) four organs, one of which is large
and two of the others very small; on the third and fourth
segments these organs are gradually reduced in number, and none
of them are found on the fifth. The sixth abdominal segment
has a median row of six organs, which are seen in fig. 24 with
the exception of the first, this being hidden by the lateral wings
of the preceding segment. The very short basal jomt of the
peduncle of each of the pleopods has on the outer side a small
organ, looking outwards and at least sometimes a little down-
wards. Hach uropod has one organ on the peduncle, situated on
its inner surface near the lower margin and close to the base,
besides one organ on the interior (lower) surface of the inner
branch near its base; finally, two organs on the interior (lower)
surface of the outer branch, one of them near the middle and the
other on the distal narrowing part.
From the preceding description it is seen that most of the organs
look downwards, a smaller number somewhat outwards or forwards,
rather few almost totally outwards, and none upwards, with
the exception of one on the first maxilliped. With the exception
of the few lateral organs on the scutum, all the others are found
on or near the lower surface of the body and on the appendages.
The structure of the organs is very interesting, and very
different from all hitherto known in any invertebrate animal. I
have examined more closely three organs, viz., that situated on
the third joint of the first maxilliped, one of the “stalked”
organs near br. above trl.*, and one from the antero-inferior
margin of the fourth abdominal segment. The two last-named
organs have been cut off, most of the adhering tissue removed
by dissection, and the organs examined with moderately high
magnifying power. I have found no difference of any importance
between the three organs from such distant parts of the animal’s
body.
The organ taken as type is that from the infero-lateral margin
of the thorax; it has been drawn (fig. 2 m) in optical vertical
section. The skin forms a chitinous, large, and very thick biconvex
lens (a), which is vitreous and a little yellowish; the major
central part is covered by a rather thin limpid layer (a’), but this
layer I could not perceive on the two other organs examined.
The lens is circular in outline; its diameter is about two-thirds
as long as that of the inner portion of the organ at its thickest
part. “The inner side of the biconvex lens is covered by a large
and rather thick coneavo-convex lens (6), which is sormewlant
thinner than the outer lens, but with the diameter a little longer
than that of the same; the lateral margin of the inner lens is
oblique, very broad, touching the external chitin around the
outer lens. This inner lens, which consists of two layers, is
homogeneous, vitreous, and slightly greyish ; but the difference
between the colour of the outer and the inner lens is in the figure
purposely a little more strongly marked than in nature. These
two lenses remind one of optical struments in which the lens is
1903.] IN THE ‘ CHALLENGER’ COLLECTION. 75
composed of crown-glass and flint-glass. Behind the inner lens
is found a thick layer of glandular cells (d), which are light
greyish, very large, and most of them elongate, radiating towards
the centre of the outer lens. The diameter of this layer is some-
what larger than that of the inner lens; and when the luminous
organs are examined in their natural position with a strong
pocket- lens, this layer can often be seen through the skin as a
whitish ring around the lens. Between the laye er of large cells
and the inner lens a thin layer (c) seems to exist, but its quality
could not be made out, and I do not venture to propose any
hypothetical explanation. Behind the glandular layer is seen
another (e), which is yellowish, with numerous transverse fine
stripes, and without trace of cellular structure; it is rather thick
in the middle, but thin around the sides of the glandular layer.
The internal surface of the organ seems to be covered by a thin
layer (f) of connective tissue. That the posterior layer with the
transverse stripes is—as in the Euphausiidee—a reflector can be
taken for granted. But it is impossible to decide whether the
light is produced by the glandular layer or by the inner lens.
Whether the thin layer enveloping the whole organ is pigmented
or not cannot be seen on this old material, which has been
preserved twenty-eight years in spit; that the organs are
immovable scarcely needs to be stated. Future investigation of
living animals and of sections of fresh material must elucidate
whether the organs are especi ially innervated or not, and, besides,
fill up the other gaps in the interpretation of the function and
structure of the layers in these compound structures.
A brief comparison of the luminous organs in Sergestes
challengert with those in other Crustacea is not without interest.
Of animals belonging to that class, luminosity has been observed
in some Copepods, a few Ostracods, Euphausiide, and one
macrurous Decapod. Giesbrecht has published a thorough and
interesting study: ‘ Ueber das Leuchten der pelagischen
Copepoden und das thierische Leuchten im Allgemeinen” (Mitth.
zool. Stat. Neapel, 11 B., 1895, pp. 648-689). He has examined
a small number of pelagic Copepoda—necessarily restricting
himself to all the luminous forms which he could procure in a
living state in the Gulf of Naples—showing that these animals
possess a number of small dermal glands, the secretion of which
produces the luminosity when it, by exhaustion from the glands,
comes in contact with the surrounding water. In a few Ostracoda
a brilliant luminosity is produced in a similar way from glands in
the labrum ; it was already suggested by G. W. Miiller in 1890,
and has since been observed and published by another author.
Furthermore, 1 can mention that during a voyage in the Indian
Ocean, Dr. Th. Mortensen met with a vast number of a pelagic
Ostracod which showed a most brilliant light, and he observed
how this was produced. Finally, the present writer has observed
nearly the same in a number of a large Ostracod which had
been procured in Davis Strait. Late in the evening I observed
76 DR. H. J. HANSEN ON CRUSTACEANS [Jan. 20,
luminous points in recently sieved bottom material from
318 fathoms, and undertook instantly some manipulations with
the animals: the luminous fluid came from the head, probably
from the large labrum, and flowed backwards between the shells,
illuminating brilliantly the space between them, especially on the
ventral side. As to the Entomostraca, we can therefore not speak
of “luminous organs” in the common sense of the word, the light
being always produced in the above-described way.
In almost all Euphausiide real and highly-developed luminous
organs are found, but they differ in structure very much from
those in Serg. challengert. The reader is referred to Sars’s
Report on the ‘ Challenger’ Schizopoda, and to Chun, “‘ Leucht-
organe und Facettenaugen” (Bibliotheca Zoologica, Heft xix.
lief. 4, 1896). The highest number of organs met with in this
family is only ten: one on each eye-stalk, two on each side of the
thorax near its inferior margin, and one in the median line of
each of the four anterior abdominal segments. The light is
produced by the “Streifenkérper ” (Chun)—“a flabelliform bunch
of exceedingly delicate fibres, exhibiting in fresh specimens a most
beautiful iridescent lustre” (Sars, op. cit. p. 71)—situated a little
behind the centre of the organ in a mass of large cells; a biconvex
lens, which is present in the organs with the exception of those on
the eye-stalks, is internal, while the outer chitinous skin is thin ;
a reflector is developed nearly as in Sergestes.
The Danish zoologist Cand. mag. Ad. 8. Jensen has directed
my attention to a book by a French author, and lent me a
German translation: ‘ Die Leuchtenden Tiere und Pflanzen. Von
Henri Gadeau de Kerville. Aus dem Franzosischen tbersetzt
von W. Marshall, 1893.’ In this popular treatise I saw that the
‘Talisman’ had captured a deep-sea shrimp with numerous
luminous organs. I attempted in the ‘ Zoological Record’ and
elsewhere to discover where that animal had been described, but
in vain, and [ will therefore reprint the passage in question from
the German book :—‘‘ Wahrend der wissenschaftlichen Hxpedi-
tionen des Talisman fing man in einer Tiefe von 500 m. einen
langschwinzigen Krebs (Acanthephyra pellucida A. Milne-
Edwards), welcher ein lebhaftes Licht um sich zu verbreiten 1m
stande war und zwar mittels folgender verschiedener Leucht-
organe; erstens befand sich eins am Vorderrand einer Deckschuppe
der Augen, zweitens verlief eine lange leuchtende Linie am
Aussenrand des Tarsus des fiinften Beimpaares, an dessen innern
Basis sowie an der des vorhergehenden Beingliedes sich weiter
leuchtende Flecke befanden, drittens lagen ganz ahnliche Flecke
an der Basis des zweiten Gliedes des dritten und vierten Bein-
paares und ebenso je einer an der Basis des Tarsus derselben
Gliedmassen, viertens sah man einen langen Fleck an der Basis am
Endgliede des hintersten Kieferfusspaares, fiinftens verlief ein
schimmerndes Querband tuber die Hiifte des hintersten Thorakal-
fusspaares, sechstens war eine Doppelreihe glanzender Punkte an
jedem Gliede der Aussengeissel der Brustfiisse, sowie an dem
ausseren Blatt der Bauchfiisse vorhanden, siebentens zeigte sich
1903.] IN THE ‘ CHALLENGER’ COLLECTION, 77
eine doppelte Linie leuchtender Punkte entlang der iiusseren
Geissel der Innenfiihler und achtens verlief eine im hinteren
Teile zusammenhingende, im vorderen in Punkte aufgeloéste Linie
parallel zum Unterrande des Riickenpanzers.” The arrangement
of the organs in Acanthephyra pellucida is rather different from
that in Sergestes challengeri, but it shows yet more resemblance
to it than to that in the Euphausiide. Both in S. challengeri
and in A. pellucida an astonishingly high number of organs exist,
but as to the structure of the organs in the last-named form
unfortunately nothing is known. I suppose that they are real
compound organs, not dermal glands as in the Entomostraca.
If we look for comparison between all luminous animals, it will
be found that only some deep-sea or pelagic fishes and two
Cephalopoda of the genus /istioteuthis (according to Verany and
Joubin) possess a number of real organs which can be compared
with that found in Serg. challengeri and Acanth. pellucida.
I have looked for luminous organs in all adult species of
Sergestes known to me and in Acanthephyr a purpurea A. M.-Kdw.,
but the result was absolutely negative. It is a very curious f: act
that about 150 very compound organs are found in one species of
Sergestes, while they seem to be quite absent in all other species
hitherto known of the genus. It may be added that the luminous
species does not deviate from some of the other forms in any other
character of importance: it belongs, even within the genus, to a
group which contains several deep-sea forms closely allied to each
other. Considered in this light, the existence in one species of
about 150 compound organs seems to me a most astonishing
feature.
Supplementary Note.
In the preceding section it has been mentioned that among the
Cephalopoda two species of MHistioteuthis possess a large number
of compound luminous organs. My friends Prof. G, “B. Howes
and Rey. Ph. R. R. Stebbing have very kindly directed my
attention to two papers, which ‘contain some additional knowledge
of the same topic and may be quoted here. W. HE. Hoyle (Mem.
& Proc. of the Manchester Liter. & Philos. Soc. vol. xlvi. part vi.
1902) points out and describes the structure of twenty-nine
luminous organs in Plerygioteuthis margaritifera, but this number,
though rather high, is yet considerably lower than that in
Ser gestes challengeri. C. Chun (‘ Aus den Tiefen des Weltmeeres,’
Jena, 1900) describes and figures (p. 532) the arrangement and
the colours of twenty- four laemaous organs 1 * Enoplotenthis
diadema Ch., n. sp.” Furthermore, he writes (pp. 5382-383):
“ Abnliche, wenn auch etwas kleinere Organe, besetzen bei
Vertretern der Gattung Calliteuthis die ganze Korperoberfliiche
von den Armen bis zu den Schwanzflossen. Die Bauchseite ist
auch hier wieder reichlicher mit ihnen ausgestattet, als die
Riickenfliiche.” An accompanying figure of a “ Calliteuthis n. sp.,
seen from the ventral side, shows a number of organs considerably
surpassing that in Serg. challengeri.
78 ON CRUSTACEANS IN THE ‘CHALLENGER’ COLLECTION. [Jan. 20,
EXPLANATION OF THE PLATES.
Prate XI.
1. Petalidium foliaceum Bate. p. 54.
a. Rostrum of a specimen from Stat. 159, from the side.
6. Kye and left antennular peduncle of the same specimen, from above.
c. Rostrum of a specimen from Stat. 146, from the side; X 5.
d. Eyes of the same specimen, from above; slightly more than X 5.
e. Basal part of mxp.2—mxp.® and _ trl.4—trl.®, together with the branchial
apparatus of the last-named specimen; X 5. 67., rudimentary pleuro-
branchia to mxp.2; ep., epipod on mxp.? with its branchia; /., four
pleurobranchial lamelle belonging to mxp.? and trl.i—tr]3
1f. Lateral view of the front part of the scutum and the eye of a young speci-
men—measuring 21°5 mm. in length—trom Stat. 158.
lg. Rostrum and eyes of the same specimen, from above.
2. Petalidium sp. p. 55.
Fig. 2a. Front portion of the scutum of a young specimen from Stat. 137, described
by Bate as Sergestes profundus Bate.
3. Sergestes profundus Bate. p. 69.
Fig.3a. Front part of the scutum, eye, and antennular peduncle of the specimen from
Stat. 300; 15/2.
35. Rostrum of the same specimen.
4. Sergestes prehensilis Bate. p. 56.
Fig. 4a. Rostrum, eye, and base of the antennular peduncle, from the side.
4,6. Front end of the scutum, eyes, peduncles of the antennulee and squamz,
from above.
5. Sergestes kroyeri Bate. p. 58.
. Rostrum, from the side.
. Front end cf the scutum, left eye, and the peduncles of the antennulz, from
above; X 4.
6. Sergestes similis, n. sp. p. 69.
Or
oa
Fig.6 a. Front end of the scutum, left eye, and peduncle of the antennula, from the
side; X 14/3.
6 6. Rostrum and supra-ocular spine, from the side; x 12.
6c. Front end of the scutum, eye, peduncles of the antennule and squame,
from above; X 5.
6d. Branchie above trl2—trl4; x 11/2. 2br., first branchia to trl2; 27.,
branchial lamella to tr]2; 36,r., first branchia to trl.3; 3 br.1, second
branchia to trl; &c.
Prats XII.
1. Sergestes arcticus Kr. p. 60.
Fig.1q@. Front end of the scutum, eye, and basal portion of the antennular peduncle,
from the side; x 9/2.
1 6. Rostrum and supra-ocular spine of the same specimen; X 11.
1c. Branchie above trl.2—trl; x 8. The lettering as im fig. 6d on the pre-
ceding Plate.
2. Sergestes challengeri, n.sp. p. 61.
In the following figures o signifies luminous organs.
Fig. 2a. Scutum, with the basal parts of the eye-stalk, antennula and antenna;
x 5.
26. Front end of the scutum, basal part of the eye-stalk, and antennular
peduncle; X 17/2.
2c. Front end of the scutum with the anterior appendages, from above; x 9.
Of the eye-stalks only the basal joint remains; of the left squama the
distal half is omitted; on the right squama the two distal ones of the four
luminous organs are seen through the plate.
1903.] ON THE HAIR-SLOPE IN MAMMALS. 79
Fig. 2d. The basal parts of the three posterior thoracic legs, the branchie above trl2—
trl.4, and the lower part of the two anterior abdominal segments, from the
side; X 11. The lettering of the branchiw as in fig. 6d on the preceding
Plate.
2 e. Left mandible, from below; X 13.
2f. Left first maxilliped, from below; x 13.
2g. Middle part of the first maxilliped, from above; X 44. 01, upper proximal
luminous organ ; 0°, distal luminous organ seen through the endopod.
2h. Left second maxilliped with its epipod and branchia, from the outer side ;
x 13.
2%. The three posterior thoracic segments and the two anterior abdominal
segments, from below, showing 31 luminous organs; xX 9. Of the
pleopods, only the basal joint—with its huninous organ—has been drawn.
2k. Sixth abdominal segment with the basal part of the telson and the uropods,
and the posterior part of the fitth abdominal segment with a part of the
pleopod; X 13/2.
21. Exterior branch of the left uropod, from the outer side; 17/2. The
apical part is wanting.
2m. Luminous organ from the side of the thorax near 67. above trl.3, seen in
optical vertical section; X 180. a, chitinous lens; a!, its thinner
external layer; 6, inner lens; ¢, thin layer between the inner lens and the
thick glandular layer d; e, reflector; f, enveloping thin layer.
2n. Sketch of the luminous organ from the upper side of the first maxilliped ;
x 180. a, lens, partly overlapped above by the protruding chitinous
plate 2.
February 3, 1903.
Howarb Saunpers, Esq., F.L.S., Vice-President,
in the Chair.
The Secretary read the following report on the additions to the
Society's Menagerie during the month of January 1903 :—
The registered additions to the Society’s Menagerie during the
month of January 1903 were 58 in number. Of these 15 were
acquired by presentation and 9 by purchase, and 34 were received
on deposit. The total number of departures during the same
period, by death and removals, was 120.
The following papers were read :—
1. Notes on the Hair-Slope of four Typical Mammals.
By Wauter Kipp, F.Z.S.
[Received November 27, 1902.]
The Otter, Domestic Dog, Ox, and Horse have been selected
for consideration as showing very different arrangements of their
hairy coverings, and as affording by their environments and habits
the probable explanation of the differences found. Two Carnivores
and two Ungulates are thus compared and contrasted.
(1) In the Otter, taken as a type of the long-bodied hairy
mammal with very short limbs, the hair presents an unbroken
slope from the snout to the tip of the tail, On the head and
80 DR. W. KIDD ON THE [ Feb. 3,
trunk it passes from cephalic to caudal, and on the limbs from
proximal to distal extremities in a perfectly uniform manner.
This arrangement obtains in a great number of long-bodied or
primitive forms, such as rodents, smaller carnivores, insectivores,
marsupials, lemurs and lower monkeys. As to the hair-slope in
the Otter, nothing further requires notice, and the type may be
looked upon as that of the primitive hairy mammal.
2) When a Carnivore of a different form, such as a short-bodied
close-haired Domestic Dog, is investigated, a few departures from
the primitive type of hair-slope are found. The Dog is taken as
representing a more highly developed carnivore form, and it
presents, as do all Canidz possessing sufficiently short hair :—
(i) A whorl situated at the extremity of the snout with a
feathered arrangement proceeding from it towards
the orbits.
(ii) Reversed bilateral area of hair on the pectoral region.
(iii) Reversed bilateral area of hair on the ventral surface of
the abdomen in the inguinal region.
(iv) Bilateral symmetrical whorl in the gluteal region lying
exactly over the tubera ischi, and, proceeding towards
this spot, a reversed area of hair on the extensor
aspect of the thigh.
(v) Reversed area of hair on the extensor surface of the
ulna.
These have been fully described, and explanations of their
etiology put forward, in our Proceedings’.
The Domestic Dog thus presents an advance in several directions
upon the primitive arrangement of hair in the Otter.
(3) The Domestic Ox shows certain peculiarities and departures,
both from the primitive and specialized Carnivore type :—
(i) In the frontal region a whorl and a feathering which
proceeds from it towards the level of the external ears,
terminating here in a crest.
(ii) Short longitudinal crest or mane, commencing as a tuft
at the level of the horns.
(iii) This crest or mane is interrupted abruptly in its normal
backward course at about the middle of the neck by a
ridge. At this point there is produced a meeting of
two streams of hair proceeding in opposite directions,
and the forward or “abnormal” stream is found to
commence at a whorl which hes in the median plane
at about the middle of the dorsal region.
(iv) From the whorl situated on the dorsal region the slope
of hair resumes its normal direction and passes to the
tail, on which it is so arranged that a central longi-
tudinal crest is usually found, and it terminates in a
scanty tuft of hair.
1 Proce. Zool Soc. 1902, vol. 1. pp. 148, 149, 150.
1903. ] HAIR-SLOPE IN MAMMALS, 81
The Ox presents no pectoral whorl as do the Dog and Horse, the
dewlap in this region being covered with perfectly smooth and
uniformly arranged hair.
These are the only four constant peculiarities of hair-slope, but
occasionally the flank and post-humeral regions, and extensor
surface of the thigh, show small whorls and featherings connected
apparently with the action of the panniculus carnosus muscle.
(4) The Horse is more specialized in the matter of hair-slope
than any other animal except man.
It shows ¢
(i) Frontal whorl, feathering, and crest.
(ii) Tuft between the external ears.
(iii) Longitudinal crest or mane.
(iv) Strong efficient tail with bushy hair, reaching to the
fetlock.
(v) Inguinal whorl, feathering, and crest.
(vi) Pectoral whorl, feathering, and crest.
Frequently but not constantly :—
(vii) Post-humeral whorl, feathering, and crest.
(vill) Cervical whorls, feather ings, and crests in several
situations,
(ix) Gluteal whorl, feathering, and crest.
(x) Tuft on lateral aspect of the abdomen.
In passing from the simple ty pe found on the Otter to that of
the Horse, a very significant series of changes is thus displayed.
Such facts as are here referred to have little or no intrinsic
interest or importance, but the phenomena of Nature, small and
great, demand explanation in accordance with the methods of
science, and it is impossible to ignore the peculiarities of hair-
slope which have been taken here as typical of a vastly greater
number in other animal forms. Any other interest they may
have is subordinate to their relation to the problems of heredity.
The four groups of facts suggest apparently two explanations as
to their zetiology. The first and most obvious is that some of them
are adaptive modifications of value to the animal; the second,
that others are produced by its habits; and it is not difficult to
distinguish these two classes in the four typical forms chosen for
consideration.
1. In the Otter the uniform trend of hair requires no other
explanation than that this arrangement of hair offers the least
ossible resistance to movements in the water and in burrows.
2. In the case of the Domestic Dog the departures from a primi-
tive type can hardly be ascribed to anything else than to use or
habit: they are adapted by the habits of the animal, not for its
needs.
Proc. Zoot, Soc.—19038, Vou. I. No. VI. 6
82 DR. W. KIDD ON THE [Feb. 3;
3. The Domestic Ox presents certain points of interest intimately
associated with adaptive modifications, but many of the phenomena
are evidently not so associated. Thus, along the median lne of
the dorsal aspect of the Ox is seen a tuft, crest or mane, whorl,
and long efficient tail, the length of the last being such as to
reach beyond the dorsal whorl. The large hairy external ears,
which can be flapped backwards so as nearly to reach the middle
line, may also be included. These may be looked upon as adaptive
modifications, existing for the purpose of defending the animal
against injurious insects; and a few observations on several oxen
and horses have been made as to the importance and frequency of
these, and as to the use of the panniculus carnosus, even in this
temperate climate.
In moderate summer weather and an exposed wind-swept
situation, the number of occasions on which certain oxen and
horses flapped backwards and forwards their ears, corrugated their
skins by the action of the panniculus, and flicked their tails
on to their backs was observed and noted with the following
results :—
Tails flicked by oxen at the rate of 348, 468, 504, 540, 720,
780, 1082 times in an hour.
Ears flapped by oxen at the rate of 684, 816, 840 times in an
hour.
Panniculus acted in oxen at the rate of 984 times in an hour.
Tail flicked by horse at the rate of 1108 times in an hour.
A rough idea may thus be gained as to the importance of certain
of these mechanisms for defence against injurious insects in hot
countries, if in a temperate climate and exposed situation so
frequent a use is made of them.
4, The Horse also presents in the median plane several similar
modifications, as a tuft, mane, and long bushy tail, which, when not
docked, reaches almost exactly to the spot where the mane termi-
nates. The panniculus mm the Horse is much more active in that
part of the flank and in the forequarters where the tail does not
reach than elsewhere.
It remains only to point out the distinction maintained here
between modifications of hair-arrangement which are themselves
adaptive, as in the Otter, and others which are, so to speak, by-
products of habits of the animals exhibiting them, such as whorls,
featherings, and crests. The only meaning properly assigned to
adaptive modifications is that such modifications are adapted for
the needs or comfort of an animal. When they are adapted dy
the habits of an animal, and have no thinkable relation to its
needs, they must be classed strictly as non-adaptive phenomena.
It should be further stated that all the phenomena referred to
are congenital, and would seem to have an intimate connection
with the problems of heredity.
1903.] HAIR-SLOPE IN MAMMALS. 83
List I.
Orrur:).252 No modifications of primitive type of hair-slope.
DOG sees 2 whorls &e.
3 reversed areas of hair.
Oixis... Siasae 2 whorls &e.
Tuft and small mane.
Crest and tuft on tail.
Horse 3 whorls &e.
Tuft.
Constant.
Mane.
Bushy tail.
4 whorls &e. Occasional.
List II.
[aaa te l 7 yer aR
a |
| | Departures | Eton of |
| from Primitive t o vd) .| Envtronments. | Habits
iBadie-slope.. eee » adaptive or |
| OOEN Ge | non-adaptive.
(eee ce ae es |
\Orrer.... None. All adaptive. Homogeneous. | Simple.
|
as peta | 5 | All non-adaptive.) More varied. More varied.
|
}Oistewsas: _4constant, several 1 non-adaptive. | Still more varied. Still more varied.
| occasional. 3 adaptive. |
|
|
| Non-adaptive.
| | |
Horse. | 6 constant, 4 oc-) 3 non-adaptive. | Most varied of | Complex, © com-
| casional. | 3 adaptive. the four types. | bining the habits
of its wild life
{ | Non-adaptive. | with those of
| domestication.
List IIT.
Observations as to use of External Kars, Panniculus
Carnosus, and Tail in Ox and Horse.
OxeEN flapped external ears at the rate of...... 684, 816,840 times
an hour,
In OXEN panniculus acted at the rate of ...... 984 times an hour.
OxeEn flicked tails at the rate of ............... 348, 468, 504, 540,
720, 780, 1082
times an hour.
Horsk flicked tail at the rate of ............ ,.. 1108timesan hour.
84 CAPT. F. WALL ON THE [ Feb. 3,
2. A Prodromus of the Snakes hitherto recorded from China,
Japan, and the Loo Choo Islands; with some Notes.
By Captain F. Watt, Indian Medical Service.t
[Received December 15, 1902. ]
Whilst attached to the China Expeditionary Forces from
1900-1902, I had opportunities of examining the Snakes pre-
served in the three Museums in China, and also others in private
collections. My notes on these, together with those on specimens
obtained myself, form the subject of this paper, which I have
arranged so as to form a complete prodromus of the species
hitherto recorded from the countries above specified.
In spite of the large aggregate of specimens J examined, it is
perhaps worthy of remark that I failed to discover one species
new to science’, and this only serves to show how extensively and
thoroughly this branch of natural history has been worked out.
In the City Hall Museum in Hongkong, out of about one
hundred specimens from the territory above mentioned, I found
many misnamed and others unidentified. I was informed that
during a typhoon some years previously a large case, containing
specimens, was blown over and. the contents wrecked, Out of
the débris labels were recovered as far as possible and replaced,
but some were evidently incorrectly reattached and others were
destroyed. This circumstance may render the accuracy of some
of the records open to question. That this collection is far from
representative is evidenced by the fact that during five and a half
months’ residence in this Port I obtained six species which were
not to be found in the Hongkong Museum.
In the Museum in Shanghai I found about fifty specimens
which were for the most part old, and concerning which there
was practically no information regarding their habitat.
In the Museum belonging to the Jesuit Fathers at Siccawei,
near Shanghai, I examined some two or three hundred specimens,
but these, again, furnished practically no record of their habitat ;
however, the late Pére Heude informed me that they had all been
collected in the Yangtse Valley, and he mentioned Ning-ko-foo
(which I find is on the southern bank of the Yangtse River) as
being the most northern limit. He could not define limits either
to the south or west.
As regards habitat, where I do not cite the authority, so far as
the territory this makes reference to is concerned, it is to be under-
stood that examples exist in the British Museum Collection at
South Kensington.
I have only made reference to points in these specimens which
do not absolutely agree with the descriptions to be found im
1 Communicated by G. A. BouLENGER, F.R.S., V.P.Z.S.
2 The species referred to hereafter as Dipsadomorphus kraepelini, No. 65, which
I had hoped to have the honour of describing for the first time, was recorded by
Stejneger shortly after I began to prepare this paper .
1903.] SNAKES OF CHINA AND JAPAN, 85
Mr. Boulenger’s work ‘Catalogue of Snakes in the British
Museum,’ 1893-1896, and I have also adopted the nomenclature
of that work.
I am indebted to Mr. Boulenger for much valuable information
and assistance in drawing up this synopsis. My thanks are also
due to Mr. F. Bowley, Hongkong, Dr. Barchet, Shanghai, and
the late Pére Heude, Siccawei, for thei courtesy in eranting me
access to the collections in the Museums under their super-
intendence, as well as to Mr, A. Owston, Yokohama, and
Mr. Armstrong, Hongkong, for placing thei specimens at my
disposal.
Part I.—List or CHINESE OPHIDIA,
TYPHLOPIDS.
1. Typuiors Lingeatus, Malay Peninsula and Archipelago,
Hongkong ?
2. TyPHLors BRAMINUS. Southern Asia from Arabia to Malay
Archipelago, South China, Formosa, Hongkong, Hainan, Mau-
ritius, Comoro Islands, Madagascar, Cape of Good Hope.—I
obtained one small specimen three and a half inches long on the
Peak in Hongkong Island,
Boip&.
3. PyTHON RETICULATUS. Burma, Indo-China, Siam, Malay
Peninsula and Archipelago, 8. China, Hongkong (City Halt
Mus.).—I saw three specimens, one in the City Hall Museum,
Hongkong, labelled from that island, one in the Shanghai
Museum, habitat unrecorded; and one in the Siccawei Museum,
consisting of a head only. From the information given by the
late Pere Heude concerning this collection, I must include this
species among the Chinese fauna. In all the specimens there are
three rows of preefrontals, and the lower labials from the 2nd to
4th and from 13th to 18th are pitted. The frontal is divided
longitudinally in two of the specimens. Ventrals 315-319.
Subcaudals 92-100.
4, PyTHoN MoLuRUS. Southern Asia from India to China,
Hainan, Hongkong (City Hall Mus.), Formosa (Swinhoe).—
The Chinese name given me was “ Hiang zo,” or ‘“ aromatic
snake,” perhaps in reference to their cooked flesh, which is highly
esteemed by these people. In Formosa it is called “ Uang’
(Swinhoe).
CoLUBRID”,
5. ACROCHORDUS JAVANICUS. Malay Peninsula, Java, New
Guinea, Hongkong (City Hall Mus. No. 293).
6. POLYODONTOPHIS COLLARIS. Himalayas as far west as Simla,
Assam, Arrakan, Upper Burma, 8. China, Hongkong.—I ob-
tained one specimen on Hongkong Island. There is one anterior
86 CAPT, F. WALL ON THE [ Feb. 3,
temporal on both sides, which comes in contact with the eighth
labial only ; this latter is unusually high and is probably the
result of a confluence with the normal inferior anterior temporal.
The first lower labials do not meet behind the symphysis. Belly
unspotted.
7. EROPIDONOTUS SWINHONIS. Formosa.
8. Tropiponotus NUCHALIS. China.
9. TRopmponoTUS VIBAKARI. Manchuria, Japan, Formosa.
10. Troprponorus piscaror. Southern Asia from India to
China, Hainan (Herz & City Hall Mus.), Hongkong (Hallow. &
City Hall Mus.), Formosa (Stejneger).— Apparently common in
the extreme south. I obtained three specimens from the main-
land opposite Hongkong, and saw one in Mr. Armstrong’s col-
lection which he assured me he procured from Hongkong Island.
J found only two specimens in the large Siccawei Collection. In
one specimen (from Kowloon, opposite Hongkong) there are five
postoculars on one side, in another four on both sides. In one
there are eight upper labials, with the fourth only touching the
eye on both sides.
11. Troprponotus ANNULARIS. China, Formosa.—This must
be a very common snake in the Yangtse Valley, judging from the
large number of specimens in the Siccawei Museum, but in the
extreme south it appears to be rare or absent. There are two
preoculars on both sides in one specimen, five postoculars on
both sides in one specimen, and two postoculars on one side
in one specimen. Ventrals 132-164. Subcaudals 70 in one
specimen.
12. Troprponotus tTIGRINUS. Siam, Cochin China, China,
Hainan, Hongkong ? Manchuria, Corea, Japan.—Called “ Yeh-
chi-po” by the Chin:se, signifying pheasant’s neck, also ‘“‘ Ch’ing-
ch’ang-chung (J/éll.). Apparently as common as JZ’. annularis
in the Yangtse Valley, from the number of specimens at Siccawel.
One specimen in Mr. Armstrong’s collection was, he informed me,
captured on Hongkong Island; however, it was not labelled, and
his collection contained some species he had procured in Japan.
13. Tropiponotus stotatus. Southern Asia from India to
China, Hainan, Hongkong, Formosa, Chusan Archipelago, Philip-
pines.—I obtained one specimen from the mainland opposite
Hongkong. The labials on one side were nine in number, and
the fourth, fifth, and sixth touched the eye. I saw no specimen
in the Shanghai or Siccawei Collections.
14. TRoprponotus suBMINIATUS. Hastern Himalayas, Assam,
Burma, Malay Peninsula and Archipelago, 8. China, Hongkong.
15. TrRoprponorus cuRysarcus. Eastern Himalayas, Assam,
Burma, Malay Peninsula and Archipelago, $8. China, Haman.
1903.] SNAKES OF CHINA AND JAPAN. 87
16. Troprpoxorus BALTEATUS (Cope), Proc. Ac. Philad. 1894,
p- 426. Hainan.
17. TrRoprpoNoruS CRASPEDOGASTER (Blgr.), P. Z.8. Lond. 1899,
p. 163, pl. xvi. fig. 1. China.
18. Troripexorus PERCARINATUS (Blgr.), P.Z.S. Lond. 1899,
p- 163, pl. xvii. fig. 2. China.—I saw six specimens in the
Siccawei Collection which exactly fit the description of Mr. Bou-
lenger’s specimen except in the following details:—Length of
internasals sometimes equals prefrontals. Postoculars and sub-
oculars: the numbers of shields in contact with the back of the
eye, intervening between the supraocular above and the fifth
labial, are four in two specimens on both sides, four in one
specimen on one side, five in two specimens on both sides, five
in two specimens on one side, six in one specimen on one side.
Labials eight with third and fourth touching the eye on one
side, nine with fifth only touching the eye on one side. Ventrals
and subeaudals 140+73, 139+ 2%, 139 +73, 138+ 2, 1388+72 (tip
slightly docked).
19. Taprnopais LAvoucuit (Blgr.), P.Z.S. Lond. 1899, p. 164,
pl. xvii. figs. -le. China.
20. PseupoxENopon mAcrors. Himalayas, Khasi Wills, Hills
in Burma, Yunnan (de Scabra*), and 8.W. China.
21. PSEUDOXENODON DORSALIS. China.
22. OpistHoTROPIS ANDERSONII. Hongkong,—TI obtained five
specimens, all from Hongkong Island. They were all captured in
a swamp near the Sanatorium on the Peak, whilst being drained
during the campaign against malarial mosquitoes. One was dug
up ata depth of about 2 feet below the surface. They accord
with Mr. Boulenger’s description except in the following parti-
culars :—Labials are inconsistent in arrangement. Thereare nine
in one specimen on one side, eight in all the rest. The fourth
and fifth touch the eye in one specimen on one side, the fifth
only touches the eye in two cases on both sides, and in one case
on one side, no labials touch the eye in two specimens on one
side. Preoculars (including the suboculars of Boulenger) are
two in two specimens on both sides, two in two specimens on one
side, one in one specimen on one side, and three in one specimen
on one side. The anterior chin-shields are in contact with five
lower labials in two specimens on both sides, and in one specimen
on one side, with four in one specimen on both sides, and in
one specimen on one side. Ventrals and subcaudals 167+ 60,
149+53, 164+59, 1654+59. Colour uniform dull olive-blhuish
above, lower half of ultimate row and belly yellow. Lower labials
and throat-scales with dull bluish mottling. Sparse mottling
beneath tail.
1 Bull. Mus. H. N. Paris, 1897, iii. p. 2165.
wv
88 CAPT. F. WALL ON THE [ Feb. 3,
93, TRIRHINOPHOLIS STYANI (Blgr.), P. Z. 8S. Lond. 1899, p. 164,
pl. xviii. figs. 2& 2a. China.
24, ACHALINUS RUFESCENS. Hongkong.—TI obtained four
specimens in Hongkong Island, found in the low vegetation on
the slopes of the Peak. In one specimen both anterior temporals
touch the eye on both sides. Scales somewhat irregular, 23-25
in mid-body. Ventrals and subcaudals 150 +56, 158+58, 158+
61, 154+58. Colour uniform olive-brown above, slightly darker
vertebrally, and with iridescence in reflected light. Head same
colour above, merging to chestnut on temporal regions.
25. ACHALINUS BRACCONIERI. SS. China.—I think that this
species will have to be united with dA. spimalis; the differences
between the two shown by Mr. Boulenger' are considerable but
not constant, being shared by individuals of both supposed species.
For instance, in all the four specimens of A. spinalis that I have
examined (one of which I presented to the British Museum,
which has been seen and identified as such by Mr. Boulenger)
the scales are 23, and not 21. The relative length of the sutures
between the internasal and prefrontal has, in my opinion, little
or no weight. The specimen of A. spinalis I sent to the British
Museum has the internasal suture about two-thirds the length of
the prefrontal; and in more than one specimen of bracconiert in
the British Museum a similar condition exists. The specimen
figured by Stejneger* from Japan which he calls spinalis is more
like Mr. Boulenger’s bracconieri, but has 23 scales.
26. ACHALINUS SPINALIS. China, Japan.—I saw three speci-
mens in Mr. Owston’s collection from Mount Fuji, Japan, and
one I found in the Siccawei Collection. In all there is a large
shield on the postero-lateral region of the parietals similar to
those described under A. bracconiert, and, ike them, separated by
one scale in the median line. Ventrals and subcaudals 154+ 58,
165+44, 165448, 1704 44.
27. Lycopon Aauticus. Southern Asia from India to Malay
Archipelago, Philippines, Formosa (City Hall Mus.), Hongkong ?
(Boettger *), 8. China ? viz. from Amoy (Steindachner *).
28. Lycopon Fascianus. W. Yunnan (Anderson), Assam,
Burma.
29. Lycopon suscincrus. Malay Peninsula and Archipelago,
Sumatra, Borneo, Java, Timor, Hongkong, Philippines.—There
are two specimens in the City Hall Museum from Hongkong and
one from Timor. I also examined a fourth, which Mr, Armstrong
1 Cat. Snakes Brit. Mus. vol. 1. pp. 308 & 309,
2 Ann. Zool. Japon. ii. p. 29.
3 The specimen alluded to by Boettger (Mat. herp. Faun. von China, 1888, p. 84)
in the City Hall Museum is no longer m the collection; the only specimen of this
species in that Institution is from Formosa.
4 Reise der Novara, Rept., Wien, 1869, p. 74.
1903.] SNAKES OF CHINA AND JAPAN, 89
informed me he had captured in the filter-beds near Bowen Road
on Hongkong Island. In two specimens the third, fourth, fifth,
and sixth supralabials touch the eye on both sides. Ventrals
192-209 °,
30. Dinopon ruFozonatus. China, Hainan, Formosa, Chusan,
Corea, Tsu Shima, Japan, Loo Choo Islands.—This is a very
common snake about Shanghai, where I encountered it frequently.
I picked up several and allowed them to crawl about my arms
without their exhibiting the least malice or alarm, and they
made little attempt to escape in the first instance. Riding my
bicycle one night I saw one crossing the road; I dismounted,
flashed my lamp on the snake, and, while holding my machine
with my right hand, captured it easily with my left. It made no
attempt at escape, though cover was within a yard or two. It
was full-grown and not desquamating. My servant caught one
in camp one night just outside my hut; it encircled the man’s
leg but did not bite. It contained a large toad (Bufo vulgaris).
I found a large number preserved in the Siccawei Museum.
found one in Mr. Owston’s collection obtained from Japan, and
four others procured from Ishigaki Island in the Loo Choo group.
I noted the following :—Internasals half or less than half the
length of prefrontals. Loreal, in Japanese and Loo Choo speci-
mens, does not touch the eye in all (five); in Chinese does not
touch the eye in five, touches eye in eleven. Postoculars three
on both sides in one specimen, three on one side in one specimen.
Temporals one on one side in one specimen. Labials normal in
all. Anterior chin-shields in contact with four lower labials in
one specimen on both sides, with six lower labials on one side
in two specimens. Ventrals in Japanese and Loo Choo speci-
mens 180-190; in Chinese specimens 192-209. Subcaudals in
Japanese and Loo Choo specimens 76-87 ; in Chinese specimens
64-76. Colour: there appear to be two very distinct varieties.
All the Chinese conform to the following description :—Alternate
bars of jet-black and coral-red (white in old spirit-specimens)
dorsally, breaking into a coarse mottling on the flanks. The black
bars involve two or three scales in the length of the snake, and the
red one scale or slightly more. There are 53-74 black bars on
the body and 18-24 on the tail, the first is broadest and forms a
chevron on the nape. Head black, fading to whitish on labials ;
sutures on crown coral-red. A light temporal streak usually.
Belly whitish, with some lateral mottling. The Japanese and
Loo Choo specimens agree :—Alternate darkish-brown and dirty
whitish (perhaps red when fresh) bars dorsally, breaking up into
a mottling laterally. The brown bars involve four or five scales
(more quite anteriorly), the light one scale. There are 24-33
brown bars on body, 15-18 on tail. Crown of head brown, fading
1 Giinther records also Lycodon (Ophites) albofuscus from Formosa (Ann. Mag.
Nat. Hist. (4) vol. i. 1868, p. 426), but gives no description nor authority for
recording it.
90 CAPT. F. WALL ON THE [ Feb. 3,
to whitish on labials. A light temporal streak. Belly whitish,
with or without sparse lateral mottling, except beneath tail
where this is abundant.
31. DINODON SEPTENTRIONALIS. China, Formosa, Himalayas ?
32. ZAocys DHUMNADES. S. China, Chusan.—A very common
snake about Shanghai and evidently throughout the Yangtse
Valley, for there are many specimens in the Siccawei Museum.
I have encountered it frequently, and consider its generic name
most apt, as it is very active, swift, and clever to elude capture.
I saw as many as four in a day’s ramble in the spring, when
batrachians were clamouving in the water engaged on matrimonial
matters. The snakes were each coiled up on the banks close to
the water, and in low vegetation, awaiting the excursion of some
unwary individual. One I saw coiled up similarly two yards or
so away from two toads (Bufo vulgaris), whose curious behaviour
it was that first attracted my attention. In spite of every care
my presence was always detected by the snake before I was
aware of its presence, and it immediately made off in great haste
for the water, and disappeared among the roots of the aquatic
vegetation. On one occasion in the summer I watched one for
some time in a strip of grass in the open, myself unobserved ;
and it was most interesting to notice the method and care
with which it beat the patch of grass like a harrier, prying into
every recess in the ground or tussock that might harbour some
possible prey. I have seen Chinese jugglers with this snake in
their stock in trade, and I believe that it is this species that
is even now occasionally met with in houses in the heart of
the town of Shanghai. On October 3rd, 1901, I captured a
young one recently hatched in camp which closely resembled the
adult in colouring. Loreal: in one specimen there are two
superposed shields on one side. Temporals: a single anterior
in one specimen on one side. Anterior chin-shields in contact
with four lower labials on both sides in one specimen. Ventrals
190-199. Subcaudals 96-119.
33. ZAMENIS KorROS. Sikkim Himalayas, Assam, Burma,
Siam, Malay Peninsula, Sumatra, Java, W. Yunnan, S. China,
Hainan, Hongkong, Formosa (City Hall Mus.).—I obtained one
specimen on the mainland opposite Hongkong. There are two
specimens in the Siccawei Museum. Loreal single on both sides
in one specimen, three on one side in one specimen. Subcaudals
100-116,
34, ZAMENIS MuUcosUS. Southern Asia from Transcaspia to
Malay Peninsula, Java, 8. China, Hainan (City Hall Mus. & Herz),
Hongkong, Formosa, Chusan (Cantor & Giinther).—I obtained
two examples from Stonecutter’s Island in Hongkong Harbour,
and three from the mainland opposite Hongkong. There are two
specimens in the Siccawei Museum. Labials nine, the fifth and
sixth touching the eye in one specimen on one side.
1903.] SNAKES OF CHINA AND JAPAN. 91
3). ZAMENIS SPINALIS. Siam, Hainan, China, Corea, Mongolia.
This snake is evidently very common in the Yangtse Valley, as
there are ten specimens in the Siccawei Museum. Concerning
one which I obtained from Huangtsun, N. China, my donor wrote
me that he had found it in his verandah, it having dropped, he
believed, from the roof about 7 feet above. He discovered it in the
act of devouring a lizard (Gecko subpalmatus). It had seized the
gecko by the body and was encircling it in its coils. A little later,
when the captive had exhausted its futile struggles, the sn: ake
seized it by the snout and began to swallow it. Frontal three-
fourths to four-fifths parietals. Loreals: two (anterior and
posterior) on both sides in one specimen, Subocular absent in one
specimen on one side. Preeoculars : the upper touches the frontal
in two specimens on both sides. Temporals: two anterior in all
specimens. Labials: eight, with the fourth and fifth touching
the eye in two specimens only, the normal arrangement being
nine, with the fifth and sixth touching the eye. Anterior chin-
shields in contact with four lower labials in two specimens on both
sides. Ventrals 179-207. Subcaudals 84-100.
36. CoLUBER PoRPHYRACEUS. HE. Himalayas, Hills in Assam,
Burma, Malay Peninsula, Sumatra, Yunnan (4 2derson).
37. COLUBER MANDARINUS. Chusan, China (N.W. Fokien,
Bigr.; Prov. Chihli, WGéll.). I saw one specimen in the Siccawei
Collection. It is called by the Chinese “‘’ Hua-tai-tsze,” meaning
variegated girdle (JZdll.).
38. CoLUBER RUFODORSATUS. KE. Siberia, China, Chusan, For-
mosa, Hainan.—EHvidently a very common snake. There are five
in the Shanghai Museum, and a large number at Siccawei.
Frontal often greater than distance to end of snout. Anterior
chin-shields in contact with four infralabials in one specimen out
of fifteen examined.
39. CoLUBER DIONE. 8. Russia, Transcaucasia, Temperate Asia,
China, Hainan, Japan (Digr. _—Evidently a very common sn: ake
in the Yangtse Valley, as the Siccawei Collection contains a large
number of specimens. I obtained two from an officer in Ching-
wang-tao, N. China, who told me the snake was common there.
One specimen contained the brood of some small bird, four in
number, one of which was still partly enveloped in its shell.
Frontal often greater than its distance to end of snout, sometimes
equalling parietals. Preeoculars: three on one side in one speci-
men. Postoculars: one on both sides in one specimen out of
sixteen examined. Labials nine, with fourth and fifth touching
the eye on one side in one Specimen ; nine with fifth and sixth
touching the eye on one side in two specimens only. Anterior
chin-shield often greater than posterior, in contact with six
‘infralabials on both sides in one specimen, and on one side in one
specimen. Scales in mid-body 23 in three specimens, 25 in ten,
92 CAPT. F. WALL ON THE [ Feb. 3,
and 27 in one specimen. ‘This species is called “ Huang-ch’ang-
ching” by the Chinese (J/6l.).
40. CoLUBER THNIURUS. Sikkim, Cochin China, Siam, Malay
Peninsula, Borneo, Sumatra, China, Formosa (Senckenberg Mus.,
Boetig.), Chusan, Manchuria. This snake is common in the
Yangtse Valley. I encountered one near Shanghai, which, however,
1 failed to capture as it took to water and disappeared in the
submerged vegetation. Loreals confluent with preefrontals on both
sides in one specimen. Temporals: single on both sides in one
specimen, and on one side in one specimen. lLabials: nine with
the fourth and fifth touching the eye in one specimen on both sides,
Ventrals 226-242,
41, CoLuBER SCHRENCKII. Amoorland, Corea, N. Japan.
42. CoLUBER PHYLLOPHIS. China.—I saw seven specimens in
the Siccawei Museum, the largest measuring 8 feet linch. There
is also a stuffed specimen in the Shanghai Museum. Scales 21
in mid-body in one specimen. Ventrals 221] in one specimen.
43, CoLUBER DAVIDI. China (Sauvage).
44, CoLUBER MGLLENDORFFII. China (Herz & Broeckelmann).
45, CoLUBER MELANURUS. Burma, Malay Peninsula, Borneo,
Sumatra, Java, 8S. China.— I saw one specimen in the City Hall
Museum from Java. Subcaudals 88.
46. CoLuBER RADIATUS. EH. Himalayas, Bengal, Assam, Burma,
Cochin China, Malay Peninsula, Sumatra, Java, 8. China (J/iill.,
Herz, Moll., Kaufm., Broeckl.), Hongkong (City Hall Mus.).—
There are five specimens in the City Hall Museum from Hong-
kong. I also obtained one specimen from that island wiioht
contained four blind and callow offspring, the brood of some small
mammal, probably a rat’.
47, Denpropuis pictus. S. Asia from India to Malay Archi-
pelago, China (Gunth.), Hongkong, Philippines.—One specimen
of this snake in spirit was given me by a lady, who assured me
she had obtained it on Hongkong Island.
48, SIMOTES PURPURASCENS. Siam, Cochin China, Malay
Peninsula, Sumatra, Borneo, Java, 8. China *.
49, Srmotes cycLurus. Bengal, Assam, Burma, Siam, Cochin
China, Malay Peninsula, Sumatra, 8. China.
50. SIMOTES FORMOSANUS. 8. China, Formosa.
51, SrmorEs viotAceus. Bengal, Assam, Burma, Camboja,
? Coluber climacopharus vel Hlaphis virgatus, described by Boettger (Mat. herp.
Faun. von China, 1888, p. 72) from Corea, is, I think, an error. The specimen
alluded to is evidently Coluber schrenckii.
2 Duméril & Bibron, ‘ Hrpétologie,’ p. 632.
1903. ] SNAKES OF CHINA AND JAPAN. 93
S. China, Hainan, Hongkong.—I saw one specimen in the City
Hall Museum, and obtained two myself on the mainland opposite
Hongkong. One was caught in grass on the hillside and suffered
itself to be handled without retaliation. The other was caught on
a shrub poised over a flower. Frontal: length greater than dis-
tance to end of snout in three specimens. Labials: six with the
third and fourth touching the eye on one side, seven with the
fourth and fifth touching the eye on the other side in one speci-
men. Nasals semidivided on both sides in one specimen.
52. SImMores CHINENSIS. China, Hainan.—Rostral: visible
portion seen from above equals distance to frontal in two
specimens. Labials seven, with the fourth only touching the eye
on both sides in one specimen. Subcaudals 50 in one and 51 in
another'specimen. I saw two specimens in the Siccawei Museum,
and obtained one myself from Kiangyin on the southern bank of
the Yangtse River, to the north- meu of Shanghai.
53, Sumores yartuantr. China (Sauvage).
54, ABLABES MAJOR. China, Hongkong (Giinth., Hallow., City
Hall Mus.), Formosa, Climeen Archipelago. — Evidently fairly
common in the Yangtse Valley. I saw one specimen of this snake,
belonging to Mr. Styan, in Shanghai, and many others in the
Siccaw ei Collection. There are three in the City Hall Museum,
two of which are from Formosa, the other of uncertain resi inet
I saw one specimen in Mr, Armstrong’s collection in Hongkong
obtained from that island, where, as he told me, it is not
uncommon. Jabials: in one specimen seven, with the third
and fourth touching the eye on one side, Subcaudals 89 in one
specimen, and 90 in two others.
ABLABES DOoRI®. Kachin Hills, Assam, China.—I saw one
eT specimen 8 or 9 inches long in the Siccawei Collection.
Ventrals 1602 Subcaudals 852 It agrees in every particular
with Mr. Boulenger’s description. This species has not been
previously recorded from China.
56, CALAMARIA PAVIMENTATA. Burma, Siam, Cochin China,
Malay Peninsula, Java, 8. China (J/dll.), Riu Kiu Archipelago
(Steyneger *).
57, CALAMARIA BEREZOWSKI. §, China (Giinth.’).
58. CALAMARIA SEPTENTRIONALIS. China, Chusan Archipelago,
Hongkong.—I saw two specimens in the Siccawei Collection, both
quite typical.
59, SPANIOPHOLIS SOULIEI Yunnan (de Scabra “)
1 Stejneger (Proc. Biol. Soe. Wash. xiv. p. 191) describes a snake as new to science
under the name of Calamaria pfefferi, which, in my opinion, is a specimen of
C. pavimentata.
= Ann Ac. St. Pétersb. 1896, p. 205, pl. i. fig. a.
3 Bull. Mus. N. H. Paris, ili. 1897, p. 216.
94 CAPT. F, WALL ON THE [Feb. 3,
60. HypstrHINA PLUMBEA. Burma, Indo-China, Malay Penin-
sula and Archipelago, 8. China, Hainan, Hongkong, Formosa.—I
saw two specimens in the City Hall Museum, one labelled China
and one Hongkong, and I obtained four myself on the mainland
opposite Hongkong. All my specimens were caught in or about
a sluggish stream near our camp at Kowloon. Frontal: length
greater than distance to end of snout in six specimens. Anterior
chin-shields in contact with six infralabials in two specimens on
one side. I noticed that in some specimens the lateral scales
about the anal region had minute central tubercles, reminding
one of the condition in Aspidura copti and A. trachyprocta.
61. HyrsrrHINA ENHYDRIS. India, Ceylon, Burma, Siam,
Cochin China, Malay Peninsula, Borneo, Celebes, 8. China,
Hongkong.
62. HypsIrHINA BENNETTI. China, Hainan (Herz).
63. Hyrstrina CHINENsIsS. Siam, China, Hainan, Hongkong
(Steindachner ’).
64, Homatopsis BuccATA. Bengal ?, Burma, Indo-China, Malay
Peninsula, Sumatra, Borneo, Java, Hongkong (Hallow. & City
Hall Mus.),—I\ examined the specimen in the City Hall Museum
which is labelled Hongkong.
65. DrpsADOMORPHUS KRAEPELINI: Boiga kraepelini Stejneger
(Proc. Biol. Soc. Wash. 1901, xv. p. 15). Formosa,—TI found one
old specimen in the City Hall Museum labelled Formosa. This
specimen almost exactly accords with Stejneger’s description, the
only points of difference being :—Temporals are six on one side.
Labials ten on one side. Anterior chin-shields in contact with
five lower labials on both sides: Ventrals 244. Subcaudals 140,
66. DrrpsADOMORPHUS MULTIMACULATUS. Burma, Indo-China,
Malay Peninsula, Sumatra, Java, Celebes, 8. China, Hongkong.—
A common snake in the south of China. I obtained six from
Hongkong Island, one from Stonecutter’s Island in that harbour,
and one from the mainland opposite Hongkong. There are
several specimens in the City Hall Museum, and I saw others in
spirit and in captivity belonging to Mr. Armstrong.
67. PSAMMODYNASTES PULVERULENTUS. EH. Himalayas, Khasi
and Assam Hills, Burma, Indo-China, Malay Penimsula and
Archipelago, Formosa.
68. CHRYSOPELEA ORNATA. Ceylon, Hills of 8. India, Bengal,
Assam, Burma, Indo-China, Malay Peninsula and Archipelago,
S. China, Hongkong?*—I found one specimen in the Siccawei
1 Reise der Novara, Rept., Wien, 1869, p. 68. en
2 The specimen referred to by Boettger (Mat. herp. Fauna von China, p. 142)
is no longer in the City Hall Musewn.
1903. ] SNAKES OF CHINA AND JAPAN. 95
Museum and one in the Shanghai Museum. Subcandals 144 in
one specimen,
69. Arpysurnus ANNULATUS. Loyalty Islands, Loo Choos, Seas
around Formosa (Stejneger) *.
70, Hyprus prarurus. Obok, Red Sea, Indian Ocean, Straits
of Malacca, Tropical and Subtropical Pacific from Loo Choo to
Australia and New Zealand, Malay Archipelago to Central
America,—TI saw two specimens in the City Hall Museum from
Hongkong and Formosa, and one in the Shanghai Museum. Also
two or three specimens in Mr. Owston’s collection in Yokohama,
obtained from the Japanese shores or the Loo Choos. All belong to
variety E of the British Museum Catalogue of Snakes. Frontal:
length less than distance to end of snout in one specimen, less than
parietals in two, Labials nine on one side in two specimens ; none
bordering the eye on one side in two specimens. Anal tetrafid.
71. AcAtypropnis Peroni. Western Tropical Pacific.
72. Hyproruis rascratus. Coasts of India to China and New
Guinea.
73. HypROPHIS GRACILIS, Coasts from Persia to China.
Hainan (//erz).
74, Hypropnis MELANOCEPHALUS. Indian Ocean, Pescadores,
(Stejneger)*, Loo Choos.
75. Hyproruis ogscurus. Bay of Bengal to China. Canton
(Peters) *.
76. Distrra stoKesit. Mekran Coast to Chinese Sea and North
Coast of Australia.
77. DistrrA oRNATA. Coasts of Asia from mouth of Persian
Gulf to Loo Choos, New Guinea, and N. Australia.—This snake
evidently is common in the Loo Choos, as I saw more than twenty
specimens from that region in Mr. Owston’s collection.
78. Distira suscincra, Indian Ocean, Japan, Loo Choos
(Stejneger)*.
1 Stejneger (Journ. Se. Coll. Tokyo, xii. 1898-1900, p. 223) deseribes as a new
species under the name of Hmydocephalus ijime certain specimens, the description of
which I have consulted My. Boulenger about; and he is of opinion that these will be
found to be specimens of Aipysurus annulatus. The enlarged vertebralsand presence
of four prefrontals, found in these specimens, were also present in some specimens
I examined belonging to Mr. Owston in Yokohama, which I regarded at the time as
annulatus. An examination of the specimens in the British Museum shows that a
certain slight enlargement of the vertebrals is present at some spots, but no specimen
has four priefrontals. The weight of such an opinion compels me to modify the views
1 had formed on the subject.
2 J. Coll. Japan, xii. p. 224 (Mierocephalophis melanocephalus).
3 Monatsb. Berlin. Akad. 1872, p. 859 (#1. diadema).
+ Stejneger (Proc. Biol. Soc. Wash. xiv. p. 191, 1901) deseribes certain specimens
obtained from the Riu Kiu Seas as Distira orientalis, which he considers a new
species. From his description I am of opinion that these belong to D. subeineta.
96 CAPT. F, WALL ON THE ~~ [ Feb. 3,
79. DisTIRA BRUGMANSII. Persian Gulf to Chinese Sea.
Hainan (Boettg.)’.
80. DistIRA cyANocINcTA. Persian Gulf to Chinese Seas,
Japan, Papuasia.—I saw one specimen of this snake in the City
Hall Museum labelled Hydrus major from Hongkong, and one in
the Shanghai Museum. One conforms to type A, and the other
to type C of the British Museum Catalogue.
81. DisTrRA VIPERINA. Persian Gulf to Chinese Sea.
82. ENHYDRIS HARDWICKII. Bay of Bengal to Chinese Sea and
Coast of New Guinea.—There are four specimens in the City Hall
Museum: two, labelled Hydrus major, are from Manila, and two
from Bangkok. There is one specimen in the Shanghai Museum
with very markedly spinose tubercles on the median six ventral
rows of scales. These rows are also enlarged. Labials eight in
two specimens, in one on both sides and in the other on one side
only.
83. PLATURUS LATICAUDATUS”*. Bay of Bengal to Chinese Sea,
Loo Choos, New Guinea, and Western South Pacific Ocean.
There are three specimens in the City Hall Museum labelled
Formosa, and I saw several specimens in Mr. Owston’s collection
obtained from the Loo Choo Islands. Ventrals 232-246. Sub-
caudals 32-46.
84. Puarurus coLtuBRINUS. Bay of Bengal to Chinese Sea,
Western South Pacific Ocean.—I examined two specimens, both
in the City Hall Museum, one from Penang and the other of
uncertain habitat.
85. Bunearus FascrAtus. Southern Asia from Bengal to
China, Hongkong (City Hall Mus.).—I obtained one specimen
from the mainland opposite Hongkong, and I saw two others in
the City Hall Museum, one from Hongkong and the other from
the mainland opposite.
86. Bunearus cANDIDUS. Southern Asia from India to China,
Hainan, Hongkong (City Hall Mus.), Formosa.—I saw two speci-
mens in the Shanghai Museum, one in the Siccawei Collection,
and one in the possession of Mr. Styan in Shanghai. There are
four specimens in the City Hall Museum, one from Hainan and
three from Hongkong. All these specimens are of variety B of
the British Museum Catalogue, viz. multicincta. Postoculars:
the normal lower shield confluent with the fourth labial on both
sides in one specimen. Labials six, with the second and third
1 Under the name of Hydrophis cyanocinctus Boettger (Mat. herp. Faun. von
China, 1888, p. 88) describes three specimens of what I consider Distira brugmansti,
collected by Herz in Hainan.
2 The species described by Boulenger (Cat. Snakes Brit. Mus. i. p. 809) as Platurus
muelleri | do not believe to be valid, and I think the specimens on which it is
based will prove to be P. laticaudatus. The only definite point he mentions to
characterize it is the presence of a median ventral keel in the posterior half of the
body, and I have found this peculiarity in at least three specimens of what I consider
undoubted P. laticaudatus.
1903.] * SNAKES OF CHINA AND JAPAN. 97
touching the eye on one side in one specimen, and occasioned by a
confluence between the first and second labials. Anterior chin-
shields in contact with four infralabials on both sides in three out
of four specimens examined.
87. Nara tripupIANs. Southern Asia from Transcaspia to
China, Hainan, Hongkong (Hallow., Steindach., City Hall Mus.),
Formosa, Chusan, Philippines. —In the Siccawei Museum I saw
seven specimens, and in the Shanghai Museum three. There are
fifteen in the City Hall Museum, of which eleven are from Hong-
kong, two from Hainan, and two from China. Of the ten
specimens I examined three have scales in twenty-one rows in
the middle of the body, and three in nineteen. In the remainder
IT have failed to record the number. Nine have more or less
distinct (some very well defined) buff or pale yellowish cross-bands
dorsally. These numbered from 13-21 on the body, and 5-8 on the
tail. They involve one or two rows of scales along the length of
the snake, and are most conspicuous in the posterior third or so
of the body. The intervals involve from ten to twelve scales, and
are sometimes separated from cross-bars by a blackish line. Hood
marked with modified black ocellus. Belly yellow, with one or two
plumbeous bands ventrally, or mottled toa variable extent with
black. In one specimen the whole belly uniform black. One
specimen is evidently a Sputatrix. This I found in the Siceawei
Collection. Scales over hood 21, mid-body 17. Ventrals 177.
Subcaudals 45. No hood-marks. Nearly uniform black dorsally,
with no suspicion of cross-bars. One very broad plumbeous ventral
band involves from the 9th to the 47th shield, and then breaks up
into a mottling and disappears.
88. NAIA BuNGARUS. Southern Asia from India to 8. China,
Hongkong (City Hall Mus.), Philippines.—A newly-hatched ex-
ample is preserved in the City Hall Museum, habitat Hongkong.
I found the head of a large specimen in the Shanghai Museum ;
and whilst I was in Hongkong a gentleman encountered one on
the mainland opposite, which he killed and brought home. I
examined and identified it, and estimated it at betwe een seven and
eight feet in length.
89. CALLOPHIS MACCLELLANDI. Nepal, Sikkim, Assam, Burma,
S. China, Formosa.
AMBLYCEPHALIDSE.
90. AMBLYCEPHALUS MOELLENDORFFII. T'enasserim, Siam, Cochin
China, 8. China, Hongkong, Hainan.—This is a common snake
in the island of Hongkong. I collected ten myself, all of which
were found in low jungle on the slopes of the Peak. I saw three
specimens in the City Hall Museum. Internasals form a suture
with the loreal in all specimens. Postoculars and suboculars :
usually one long semilunar shield extends from the supraoculars
behind, skirting the eye to a point about halfway up its anterior
aspect. Sometimes this scale is divided so as to form a small
Proc. Zoou. Soc.—1903, Vou. I. No. VII. 7
98 CAPT. F, WALL ON THE . [ Feb. 3,
postocular above this shield. Temporals: the antero-superior -
extends back as far as the extreme border of the parietals or
further. Labials: six in two specimens on both sides. Sub-
caudals 36-50. First infralabials meet in four, fail to meet in
one, not recorded in eight.
91, Pseuporarzas yacus. Hongkong (Jan).
VIPERIDG.
92, ANCISTRODON acuTUS. China.—Called by the Chinese
** Qo-woo-shay,” or five-pace snake, owing to the reputed rapidly
fatal effects of its bite. I saw one specimen in the City Hall
Museum labelled A. blomhoffii, habitat Szechuen. There are also
five specimens in the Siccawei Collection. Two are labelled Ou-
Yuen’, and one of these dated 1882 (7. e. six years before Giinther
first described it). One is labelled Chowtong* 1883, and another
Kien-té® 1882. The fifth consists of a dried skin. In talking
to the late Pére Heude about this creature, he told me it is not
uncommon in hilly parts of the Yangtse Valley, and is feared by
the natives more than a leopard. He narrated an experience of
one of the Jesuit Fathers who heard frantic cries for help issuing
from some dense jungle he was riding through. On dismounting,
and proceeding to the spot, he found a Chinaman being pursued
by one of the snakes, which he attacked and killed. Rostral: as
far as I could ascertain there is only one shield beneath the dermal
appendage, which, however, is frequently bent over, causing a
furrow which makes the shield appear divided into two. Labials :
six in one specimen on both sides, and in another on one side.
Subcaudals 56 in one specimen, with Ist-llth entire and rest
divided ; 53 in another, with Ist—9th entire and rest divided; 58
in another, with Ist—28th entire and rest divided; 57 in another,
with Ist—12th entire and rest divided.
93. ANCISTRODON BLOMHOFFII. LE. Siberia, Mongolia, China,
Hainan, Formosa, Japan, Loo Choos, Siam.—Mollendorff says it
is called by the Chinese “ Fei-shang-ts’ao,” snake which flits on
the grass, and ‘ Ch’i-ts’un-tsze,” or seven-inch snake. It is very
common in the Yangtse Valley. There are at least three specimens
in the City Hall Museum from Japan. There are five in the
Shanghai Museum, and a very large number in the Siccawei
Collection. In Mr. Owston’s collection I saw several obtained
from Japan and the Loo Choos. I obtained five myself about
Shanghai in our camp. Two of these, which I found close together
on October 16th, 1901, appeared to be newly hatched. On the
Ath October, 1901, my servant informed me that there were several
snakes lying together dead close to the camp. I investigated the
matter, and found an adult of this species with the remains of
1 Perhaps Tao-yuan in Province Hunan ?
2 Perhaps Chao-tung in North Yunnan ?
3 In Province An-Hui.
1903. ] SNAKES OF CHINA AND JAPAN, 99
twelve young scattered around her. In spite of every endeavour,
I failed to trace the author of this butchery. It is sad luek that
such opportunities as these for observing habits rarely fall to the
lot of those who specially seek them. Residents about Shanghai
occasionally lose dogs under circumstances suggesting a poisonous
bite, and I think these fatalities are frequently due to this little
snake. One specimen I captured had the penis bifid on both sides,
I kept young on two or three occasions, but could not find suitable
food with which to tempt them, and finally resorted to feeding
them with boluses of raw meat, a treatment they displayed their
distaste for by repeatedly disgorging the morsels some time later.
T observed that, when molested and excited, all my specimens
vibrated the tail, but I could never get one actually to strike ;
howeyer all the specimens I had alive were very small. Labials
seven, the third only touching the eye; eight in one specimen on
both sides,
94. Lacnesis gervonii. Khasi Hills, Assam, Thibet; Upper
Yangtse, China.
95, LacnEsis MucrosquamaAtus. Naga Hills, Assam; Formosa.
96. LACHESIS GRAMINEUS. South-eastern Asia, China, Hainan
(Herz & City Hall Mus.), Hongkong, Formosa.—This is a very
common snake in Hongkong, where I procured three specimens.
There are also twelve in the City Hall Museum, obtained locally.
In one of these the skin of the neck had been perforated and
rent by the beak of a bird which it is seen in the act of devouring.
The distension of the mouth and neck is enormous, so that the bird
appears as if it were being thrust through the rent instead of into
the gullet. There is only one specimen in the Shanghai Museum,
and only one in the Siccawei Collection. The residents of Hong-
kong call this the Bamboo-snake, which is a very good name for it
since it is almost always to be found in the foliage of bamboo
vegetation. I am told too that the Anglo-Indians in Assam have
also given this name to the snake.
Part I1.—List or JAPANESE AND Loo CuHoo JstANDS OPHIDIA.
CoLUBRID&.
1. Troprponotus VIBAKARI. Japan, Formosa, Manchuria.—I
found one in the City Hall Museum from Nagasaki, and saw six
others in Mr. Owston’s collection, all from Mount Fuji. Ventrals
152 in one specimen. Subcaudals 80 in one specimen; second to
fifth entire in one specimen.
2. TROPIDONOTUS TIGRINUS. Siam, Cochin China, China, Hainan,
Hongkong ?, Manchuria.—In my opinion this is the commonest
snake in Japan. I frequently met with it, and on one occasion
captured five in a couple of hours. It is called by the Japanese
“yamakagashi,” “ uwabami,” “ orochi,” “ja,” all of which terms, I
7*
i
100 CAPT. F. WALL ON THE [ Feb. 3,
am informed, signify “the largest variety”; also ‘ tora-no-kuchi-
nawa,” or “tiger-snake”; also ‘“miza kuchinawa,” or “ water-
snake”; also “atsuki kuchinawa,” or “thick brown snake.” (See
also No. 12 of Chinese list.)
3. TROPIDONOTUS PRYERI. Loo Choos.—I saw one specimen in
Mr. Owston’s collection from Nawa in the Loo Choos.
4, ACHALINUS SPINALIS. China, Japan. (See No. 26 of Chinese
list.)
5. Dinopon RuUFozONATUS. China, Hainan, Formosa, Chusan,
Corea, Tsu Shima, Japan, Loo Choos. (See No. 30 of Chinese list.)
6. Dryopon sEMICARINATUS. Loo Choos.
7. Dixopon saponicus. Japan.—I found one specimen on the
Southern Island (Kiu Siu), and saw another in Mr. Owston’s
collection. Ventrals 209. Subcaudals 75? and 77.
8. DINODON TESSELLATUS. Japan.
9. COLUBER DIONE. S. Russia, Transcaucasia, Temperate Asia,
China, Hainan, Japan (Bigr.). (See No. 39 of Chinese list.)
10. CoLuBER scHrENcKH. Amoorland, Corea, N. Japan.
11. CoLUBER cCoNSPIcILLATUS. Japan, Corea.—I saw one
specimen in the City Hall Museum labelled Nagasaki and another
in Mr. Armstrong’s collection captured in Japan. Called by
Japanese “jimuguri” (“earth borer”) and “ kawara kuchinawa”
(“‘ dry-river-bed snake ”).
12. CoLUBER CLIMACOPHORUS. Japan.—A very common snake
in Japan, where it is called “aodaisho,” ‘‘nezumi tori” (or “ rat-
catcher”), ‘‘sato meguri” (or “village idler”), and “mugiwara hebi.”
I frequently encountered it. I saw and captured one reclining
on a stone parapet in one of the Nikko temples. I discovered
another in thick jungle in the act of swallowing a half-grown
leveret, truly an enormous meal, since the mammal was more
than twice the weight and girth of the reptile. J thought on
all occasions it was far less wary and active than other nearly
allied snakes of a similar size with which I am familiar, such, for
instance, as Zamenis mucosus and Z. korros, Coluber radiatus,
Zaocys dhumnades, &c. Temporals, in one specimen, three on
both sides.
13. CoLUBER QUADRIVIRGATUS. Japan, Corea.—This species is
almost if not quite as common in Japan as 7’ropidonotus tigrinus.
I scarcely went a day in the country without seeing one, and
often I saw three or four. It is called by the Japanese “shima
hebi” (or “striped variety”), ‘‘karasu hebi” (or “black snake”),
“sukuro hebi” (or “medium black snake”), “‘mugiwara hebi” (or
“‘ wheat straw snake”), and “kuro kuchinawa.” Subocular absent
in one specimen on one side. Labials eight, with third, fourth,
and fifth touching the eye on one side in one specimen.
1903.] SNAKES OF CHINA AND JAPAN, 101
14. CoLuBER scumaAckERI. Loo Choos.
15. ABLABES SEMICARINATUS. Japan, Loo Choos.
16. ABLABES HERMINE. Loo Choos.—I saw two specimens in
Mr. Owston’s collection from the Loo Choos. Ventrals 162-163.
Subcaudals 47—51.
17. CALAMARIA PAVIMENTATA. Burma, Siam, Cochin China,
Malay Peninsula, Java, 8. China (J/6ll.), Riu Kiu Archipelago
(Stejneger). (See No. 56 of Chinese list and footnote.)
18. Hyprus pLaturus. Coasts of Asia from Red Sea to Loo
Choos, Australia, New Zealand, Malay Archipelago to Central
America.—Called ‘“umi hebi” or “sea snake” by Japanese. (See
No. 70 in Chinese list.)
19. HyprorHis MELANOCEPHALUS. Indian Ocean, Pescadores
(Stejneger), Loo Choos.—This is evidently a common snake about
the Loo Choos, since I saw upwards of twenty specimens in
Mr. Owston’s collection from this locality. Postoculars one or two.
Temporals: one anterior usually, sometimes two. Scales two
head-lengths from snout 23-27, mid-body 31-35. Ventrals 3805—
343. Anal tetrafid. (See No. 74 of Chinese list.)
20. DistirA cyANocIncTA. Persian Gulf to Chinese Seas,
Japan, Papuasia. (See No. 80 of Chinese list.)
21. Distira orNATA. Coasts of Asia from mouth of Persian
Gulf to Loo Choos, New Guinea, Northern Australia. (See
No. 77 of Chinese list.)
22. DistirA suBcrncta. Indian Ocean, Japan, Loo Choos
(Stejneger). (See No. 78 of Chinese list and footnote.)
23. ArpysuRUS ANNULATUS. Loyalty Islands, Loo Choos, Seas
around Formosa (Stejneger). (See No. 67 of Chinese list and
footnote.)
24, PLATURUS LATICAUDATUS. Bay of Bengal to Chinese Sea,
Loo Choos, New Guinea, Western South Pacific Ocean. (See
No. 83 of Chinese list.)
25. Puarurus coLtuBrinus. Bay of Bengal to Chinese Sea,
Western South Pacific Ocean. (See No. 84 of Chinese list.)
26. PLATURUS SCHISTORH¥NCHUS. Western Pacific Ocean, Loo
Choos.
27, Hemrpuncarus saponicus, Loo Choos, Japan ?
VIPERIDAE.
28. ANCISTRODON INTERMEDIUS. Japan, Mongolia, Eastern
Siberia, Central Asia.
102 MR, LYDEKKER ON AN ASIATIC WILD SHEEP. [Feb.3,
29. ANCISTRODON BLOMHOFFII. astern Siberia, Mongolia,
China, Hainan, Formosa, Japan, Loo Choos, Siam.—Called by
the Japanese ‘‘mamushi,” “hami,” “ kuchibami,” “hiraguchi.”
All synonymous terms for the “ beautifully marked variety.”
30. LACHESIS OKINAVENSIS. Loo Choos.—Called by the J: apanese
“habu.”
31. LAcHEsSIS FLAVovIRIDIS. Loo Choos.—Also called “ habu”
by the Japanese.
32. LAcHESIS MUCROSQUAMATUS. Naga Hills, Assam; Formosa.
33. LAcHESIS LUTEUS. Loo Choos.
3. Note on the Wild Sheep of the Kopet-Dagh.
By R. Lypexker.
[ Received December 4, 1902. |
(Text-figure 10.)
Through the generosity of Mr. St. George Littledale the collec-
tion of the British Museum has recently been enriched by a very
fine skull, with the horns (text-fig. 10, p. 103), of a ram of the Wild
Sheep of the Kopet-Dagh range, which forms the boundary between
Turkestan and Northern Persia to the eastward of the Caucasus.
Mr. Littledale also brought home the skin of the same animal,
but it unfortunately was so badly injured by vermin that it had
to be destroyed. ‘The skull is the finest of the series obtained
during the trip.
The Kopet-Dagh sheep was named Ovis arkal in 1857 by
Blasius, and is evidently allied to the Urial, with the Punjab
yace of which I have indeed proposed to identify it*. At that
time I had, however, never seen an adult skull; and Mr. Little-
dale’s specimen indicates the right of this sheep to rank as a
distinct race of Urial. It will be remembered that the Punjab
race of the Urial (O. vignet cycloceros), as exemplified at any rate
by specimens from Peshawer and Afghanistan in the British
Museum, differs from the typical O. vignei of Astor and Ladak
by the much greater prominence of the two front angles of the
horns, which are often raised into nodose beads, between which
the front surface of the horn is depressed and carries bold and
widely separated transverse ridges.
In the Kopet-Dagh Urial this prominence of the front angles
of the horns is still more pronounced, though the beading is
somewhat less conspicuous. Moreover, the front surface of the
horn is unusually broad and flattened, with the transverse
1 * Wild Oxen, Sheep, and Goats,’ p. 173.
1903.] MR. LYDEKKER ON AN ASIATIC WILD SHEEP. 103
wrinkles very low and indistinct. The length of the horn is
33 inches along the inner front angle, with a basal circumference
of 11 inches, a basal width of 3 inches, and a basal depth of
4 inches.
The last two dimensions are considerably greater than in a
skull of the Kelat Urial measured by Mr. Hume, in which the
length along the curve is 35} inches.
Text-fig. 10.
Front view of skull and horns of adult ram of the Kopet-Dagh Umial.
+ nat. size.
The Kopet Dagh Urial decidedly appears to be a distinct form,
connected with the typical Ovis vignei by means of the Punjab
race of that species. On these grounds I regard it as a local race
rather than a species; its name will accordingly be O. vignet arkal
(or perhaps arcal). The suggestion of M. Dauvergne’, that this
sheep is identical with the Kelat Urial, is not borne out by a
comparison of the present specimen with a skull of that form in
the British Museum, in which the angles of the horns are much
rounded off. This leads me to think that the Kelat Urial
(O. vignet blanfordi) is, after all, distinct from the Punjab
animal,
1 See p. 131 of my book on ‘Game of Europe, N.W. Asia, and America.’
104 MR, P. W. BASSETT-SMITH ON [ Feb. 3,
4. On new Parasitic Copepoda from Zanzibar and East Africa,
collected by Mr. Cyril Crossland, B.A., B.Sc. By Statt-
Surgeon P. W. Bassert-Smiru, R.N., F.Z.S.
[Received December 4, 1902. |
(Text-figures 11 & 12.)
Mr. Cyril Crossland, in his recent examination of the marine
fauna of Zanzibar and British East Africa, obtained several
specimens of parasitic and semiparasitic Copepods, three of which
he has been kind enough to allow me to examine.
These curiously deformed and often grotesquely-shaped animals
are frequently found attached to the gills, &c., or to the surface of
fish and other marine animals.
A large number from the former which are now described from
a variety of different hosts, and from wide geographical areas,
I enumerated in Proc. Zool. Soc. 1899, p. 438; to this paper
1 have appended a list of addenda, which I have drawn up with
the kind assistance of Mr. E. Bergrotti, of Tammerfers, and
others.
Not only fish but a number of other marine animals are un-
doubtedly infested with these parasites, though at present little
information concerning them has been collected ; the specimens of
Mr. Crossland are therefore particularly interesting.
Gerstacker, in Bronn’s ‘ Klass. und Ordn. des Thier- Reichs,’
1866-79, Crustacea, vol. v. Copepoda, p. 774, mentions five genera
found on Nudibranch Mollusca: Doridicola Lyd., Holidicola Sars,
belonging to the family Ergasilide; Artotrogus Boeck, to the family
Ascomyzontide ; and Splanchnotropus Hance. and Jsmaiha Bergh,
to the family Chrondracanthidz, Also nine genera from various
Vermes, p. 773.
Of the three specimens of Mr. Crossland, two were taken from
the kidneys of species of Pleurobranchids (not determined) and
one from the skin of a Sipunculid (Aspidosiphon).
As they were only single specimens it was impossible to dissect
them, and therefore the descriptions are necessarily mcomplete.
The first two evidently belong to the family Chondracanthide,
but do not fall in with the descriptions of any known genus; they
appear to be most nearly related to the genus Splanchnotropus of
Hancock (Trans. Linn. Soc. vol. xxiv. pp. 51, 55), two species of
which he describes, S. gracilis and S. brevipes, taken from Nudi-
branchs; the present specimens differ from them, however, in the
complete absence of antennze and articulate limbs, and in having
the external ovaries elongated and the eggs arranged in single
series.
I have therefore provisionally placed them in a new genus,
Chondrocarpus, following closely after Splanchnotropus Hane. and
Diocus Fabr.
CHONDROCARPUS, gen. nov.
. Cephalothorax coriaceous, elongated, with four lateral short
1903. ] NEW PARASITIC COPETODS, 105
lobe-like processes on either side; abdomen one-fourth length of
whole, biarticulate, tapering; no distinct antennz or thoracic
limbs; mouth placed on under surface, with minute maxille ;
external ovaries as elongated filiform sacs containing ova in a
single series.
3. Pigmy.
CHONDROCARPUS RETICULOSUS, sp. n. (Text-fig. 11, A-G.)
Hab. Zanzibar: from large Pleurobranchid.
Length 12 mm., breadth 4 mm.
2. Cephalothorax indistinctly segmented, elongated, tapering,
dorsally convex, laterally produced into 4 rounded truncated
processes; the 4th pair being most widely separated and the
smallest; the whole having a peculiar reticulate appearance from
the network of ramifying tubules; anterior extremity rounded;
no visible antennz; mouth as a papilla placed between the Ist
pair of processes ; upper lip triangular; only one pair of maxille
and mandibles (?) could be made out, each terminating in minute
claws. No thoracic limbs; there is a short genital segment, or
ring, from which spring the long filiform ovaries. Abdomen in-
distinctly biarticulate, tapering, without caudal plates or sete.
3. Pigmy. One was seen attached to the last abdominal
segment but was partially hidden, the bifid articulate caudal
extremity only being visible.
Text-fig. 11.
Chondrocarpus reticulosus 9, gen. & sp.n. A. Dorsal surface. B. Lateral view.
C. First segment and mouth-organs. D. One of the lobes showing reticu-
late appearance. E. Second maxilliped. EF. Abdomen showing fixed ¢.
G. Posterior extremity of g. H. Chondrocarpus sp. 2: dorsal surface
(specimen incomplete).
CionprocarPus sp. (Text-fig. 11, H.)
A specimen of a second species of this genus was taken from a
106 MR. P. W. BASSETT-SMITH ON [ Feb. 3,
Pleurobranchid; it was much broken both at the anterior and
posterior extremities. It differs from C. reticulosws in not having
the peculiar reticulate appearance and in having a pair of lateral
lobes on either side of the genital segment.
The third specimen also appears to be new, and belongs to the
family Dichelesthiide, the animal resembling most nearly the
genus Enterocola of Van Beneden (Bull. Acad. Roy. de Belg.
tom. ix. 2nd ser. p. 151), found by him in the respiratory cavity of
“ Aphidium ficus,” than any other form I have been able to find
recorded,
Mr. Crossland’s specimen appears to be much more degenerate
from its parasitic habits; the articulate limbs are excessively
small, difficult to make out, and the ova are carried in long spiral
thread-like processes in a single series as in the Caligidee, and
not in dilated sacs. Unfortunately there was only this single
female specimen for examination. I would provisionally create
for it a new genus “ Ventriculina,” giving the specific name of
“‘ crosslandi” in recognition of the collector.
VENTRICULINA, gen. nov.
Head small, ‘rounded; neck indistinct; 3 thoracic segments,
the first amalgamated with the head; genital segments lobed,
equal in breadth with the thoracic; no dorsal plates; abdomen
short, biarticulate; external ovaries spiral, ova uniserial; first
antenne 4-jointed, simple, second antenn 3-jointed ; maxillipeds
very small.
Three pairs of minute thoracic limbs, first biramose, second and
third uniramose.
VENTRICULINA CROSSLANDI, sp.n. (Text-fig. 12, p. 107.)
Hab. Zanzibar: from a Sipunculid.
Total length 4mm. Colour white.
Head small, rounded in front, broadest behind, from under
which project the anterior antenne. Thoracic segments three,
the first united with the head, the genital segments trilobed; the
whole forming an oblong body without lamellar plates, showing
five distinct rounded lateral lobes, the last pair being slightly
wider and more acute. Dorsal surface convex, marked by five
distinct grooves showing the position of the union of the segments.
Abdomen biarticulate, narrow, one-sixth the total length, the -
last joint terminating in two small caudal plates provided with a
marginal fringe of short bristles. External ovaries long, spiral,
springing from papille at the angle of the genital segments and
abdomen. Ova large, arranged uniserially as in the Caligide.
Anterior antenne 4-jointed, non-setose, the first joint being
the longest and broadest, rising from the underside of the head
just in front of the mouth; second, third, and fourth joints
progressively decreasing in size.
1903. ] NEW PARASITIC COPEPODS. 107
Posterior antenne 3-jointed, rising just in front of the upper
lip; at the distal end and anterior border of the first and second
joints are two short sete, the third terminating in two bristles,
the anterior being very long.
Mouth and appendages placed rather far back; the labrum is
triangular, projecting backwards ; labium simple, rounded. Iwas
able to make out only 2 pairs of maxillipeds; the first very small,
biarticulate, terminating in two short hairs: the second uncinate,
with large globose basal joint, to which was articulated a sharp
curved claw.
Text-fig. 12.
Ventriculina crosslandi 9, gen. & sp.n. A. Ventral surface, X 10. B. Dorsal
surface. C. Ventral surface much enlarged, showing articulate appendages.
D. Posterior antenn.
Only three pairs of thoracie limbs present: the first rising from
the posterior under surface of the cephalic segment, minute,
biramose, each ramus terminating in a single bristle; second and
third pairs uniramose, made up of two articulations, the distal
terminating in two small bristles.
3 not known.
ADDENDA to Systematic Enumeration of Species of Parasitic
‘ Copepoda found on Fish (Proc, Zool. Soc. 1899, p. 438).
ERGASILIDA,
1. Eucanthus marchesettii Valle, Atti Mus. Civ. Trieste, vil.
p. 245 (1885). On Motella tricirrata.
2. Ergasilus centrarchidarum Wright, Proc. Canad. Inst. (2) i.
p. 243 (1883).
3. Lrgasilus biuncinatus Gadd. Meddelanden af Societas pro
Fauna et Flora Fennica, xxvii. pp. 181-182 (1901). On
Gastrosteus aculeatius.
108 ON NEW PARASITIC COPEFODS. [ Feb. 3,
4,
5.
€.
die
8.
9.
10.
23.
29.
: Teaceroe carcharii glaucus Hesse,
Bomolochus onosi T. Scott, 20th Ann. Rep. Fish. Board of
Scotland, p. 289, pl. xiii. figs. 19-22 (1902). . On Onos
mustelus and Onos cimbrius, Firth of Forth.
Bomolochus zeugopteri T. Scott, loc. cit. p. 290, pl. xiii. figs. 23-25.
On Zeugopterus punctatus.
CALIGIDE.
Caligus pacificus Gissler, Amer. Nat. p. 886 (1883). On
Salmo.
Caligus labracis T. Scott, J.c. p. 291, pl. xiii. figs. 26-29. On
Labrus mixtus and il maculatus.
Anchicaligus nautili Stebbing, Willey’s Zool. Res. pt. v. pp. 667—
670, pl. lxxi.
Dinematura musteli levis Hesse, Rev. Sci. Nat. Montpellier,
(2) i. pp. 6, 11 (1880).
Cecrops acimthice vulgaris Hesse, Ann. Sci. Nat. (6) xv. 3,
p. 26 (1883).
Ue. 105 Isis
musteli levis Hesse, l. c. p. 23.
spinacis acanthic Hesse, by Os 10s NO;
9
unicolor Hesse, 1. c. p. 20. On Galeus vulgaris.
”?
DICHELESTHIID®.
. Lernanthropus polynemi Rich. Zool. Anz. iv. 1881, p. 505.
On Polynemus.
. Lernanthropus tetradactylus (probably L. trifoliatus B.-8.
1898).
. Lernanthropus micropterygis Rich. Atti Soc. Tose. Sci. Nat.
iv. p. 82 (1884). On Micropteryx dumerili.
. Lernanthropus tylosuri Rich. l. c. p. 83. On Tylosurus im-
perialis.
. Kréyeria (Lonchidium) galii vulgaris Hesse, Ann. Sci. Nat.
(6) xvi. 3, p. 2 (1883).
. Clavella cluthe T. Scott, U. c. p. 292, pl. xu. figs. 26-31. On
Ctenolabrus rupestris.
. Pagodina (Nemesis) charcharie glauci Hesse, l. c. p. 13.
. Ludactylina carcharie glauci Hesse, l. c. p. 11.
. musteli levis Hesse, l. c. p. 8.
X squatina angeli Hesse, l. c. p. 5.
3) similis T. Scott, l.c. p. 295, pl. xii. figs. 1-19.
On Raia radiata.
. Eudactylina acanthii T. Scott, l.c. p. 296, pl. xiii. figs. 1-9.
On Squalus acanthias.
. Bassettia congri Stebbing, Willey’s Zool. Res. pt. v. pp. 671,
672, pl. xx.
PHILICHTHYIDA.
. Philichthys fialole Rich. Zool. Anz. iii. p. 69 (1880). On
Stromateus fialola.
Philichthys doderleini Rich. Zool. Anz. vi. p. 558 (1883). On
Labrus turdus.
1903.] ON THE ORIGINAL HOME OF THE TIGER. 109
LERNzIDS.
30. Lernea abyssicola Brady, Chall. Rep. viii. p.137. On Ciralias
uranoscopus.
31. Lernea minuta T. Scott, 18th Rep. Fish. Board of Scotland,
p- 161, pl. vii. fig. 13 (1900). On Gobius minutus.
32. Lernea lumpi T. Scott, 19th ditto, p. 128, pl. vii. fig. 12
(1901). On Cyclopterus lumpus.
33. Hemobaphes ambiguus T. Scott, 18th ditto, p. 162, pl. vii.
fig. 15. On Callionymus maculatus.
34, Peraderma petersi Rich. Zool. Anz. iv. 1881, p. 387. On
Gobius buccatus.
35. Peraderma bellottii Rich. Zool. Anz. v. 1882, p. 475. On
Scopelus benotti.
CHONDRACANTHID.
36. Chondracanthus bleekeri Rich. Zool. Anz. iv. p. 387 (1881).
On Chilium chlorurus.
37. Chondracanthus ninnit Rich. Zool. Anz. v. p. 504 (1882). On
Gobius.
38. Chondracanthus ornatus T. Scott, 20th Ann, Rep. Fish Board
of Scotland, p. 298, pl. xin. fig. 34. On Callionymus
maculatus.
LERNEOPODIDS.
39. Achtheres sandre Gadd. Med. af Soc. pro Fauna et Flora
Fennica, xxvii. (1901).
40. Lernceopoda extumescens Gadd. l. ce. On Coregonus.
41. Tracheliastes gigas Rich. Zool. Anz. iv. 1881, p. 504.
42. Charopinus dubius T. Scott, 19th Ann. Rep. Fish Board of
Scotland, p. 130, pl. vii. fig. 15. On Raia circularis.
5. On the Original Home of the Tiger.
By Col. ©. E. Stewarz, C.B., C.M.G., C.LE4
[Received December 6, 1902.]
The ordinary idea of English people that the Tiger was origin-
ally an Indian animal, is, I believe, quite a mistake. After
careful enquiry, I have come to the conclusion that the Tiger is a
comparatively late intruder into India.
Firstly, after enquiry, I can discover no Sanscrit word for the
Tiger. If tigers had existed in India in the days when Sanscrit
was a spoken _ language, there would be a name in Sanscrit for it,
while there is only a modern Hindustani name. There is a
Sanscrit word for Lion, “Singha,” which would point to the
fact that lions were certainly more common than tigers in time
long past. At present lions are not found in India, except a
very few, which are strictly preserved in Googerat, one extreme
1 Communicated by Col. Hinn James, F.Z.S.
110 ON THE ORIGINAL HOME OF THE TIGER. [ Feb. 3,
corner of India, though I will allow that lions were probably
commoner than they are now in the olden time, though probably
never very numerous.
I remember, when I first went to India, nearly 50 years ago, a
lion being killed not very far to the southward of Allahabad, but
this was even then a rare occurrence. I have studied the
question of the habitat of lions and tigers in Persia, where I
resided for a good many years. lions are found only in the
very south of Persia, near the Persian Gulf, and Arabia; while
tigers are only seen in the very north of Persia, near the Russian
border, and especially near the Caspian Sea, on the north of
Persia, and they are more numerous within British territory
than within the Persian boundary, and tigers are more common
in Southern Siberia than they are anywhere in Persia.
Tigers are more numerous in cold countries. They are plen-
tiful in Corea, which has a severe winter climate, and still more
plentiful in the Island of Saghalien, belonging to Russia, and
further north than Corea, and which has almost an arctic climate
in winter. The tiger is mentioned by Marco Polo in his travels, but
nowhere as an Indian animal, and I very much doubt whether
tigers were found in India at the time Marco Polo visited it.
In the Sanscrit works treating of the fighting between Rama
and Rawun, the Demon King of Ceylon, though many animals
are mentioned, such as bears, monkeys, and several others, I
have been unable to find any mention of the tiger ; and the tiger
is not found in the Island of Ceylon, though the leopard is; nor
is the tiger found in the larger island of Borneo, which would
seem to point to its only inhabiting the islands of the Indian
Archipelago, which it could reach by swimming. Thus it would
seem that tigers did not exist in India before the time that Ceylon
was separated from India. Tigers did not exist in the island of
Singapore until about 1809, when apparently they swam over
from the mainland. Tigers are such good swimmers that they can
cross a considerable body of water. J do not think any allusion
to tigers in India can be found in the Greek historians. I should
feel much obliged if anyone could find me such a reference.
In the monuments of the Assyrian Kings, and of the Kings of
Persia, there are constant references to lion-hunts by those kings,
but never allusion to a tiger-hunt. Of course there is an existing
Persian word for tiger, but there is nothing to show that it is at
all ancient.
My own idea is that the tiger was originally a purely northern
animal, which has gradually extended southward. I fancy that
no allusion to a tiger in India can be traced to a period anterior
to the early Mahommedan conquerors of India. I should be
much obliged to anyone who will help me to clear up this
question. We English have so completely assumed the idea
that the tiger is an Indian animal, that we have called him the
Royal Bengal Tiger, though I firmly believe he is as much an
intruder from the north into Bengal as we are ourselves.
1903. ] ON THE COPULATION OF THE INDIAN ELEPHANT, iGl
6. On the Mode of Copulation of the Indian Elephant.
By H. Sxave, Conservator of Forests, Maymyo, Burma.*
{Received December 15, 1902. ]
Some few months ago I was, on several occasions, able to witness
tame Elephants in the act of copulation, This sight has been so
seldom witnessed by Europeans, and is so variously described by
Burmans, that these remarks, supported by a series of photographs
deposited with the Society, may be of interest and value.
As tuskers are usually reputed to be shy of copulating before
eye-witnesses, an account of the manner in which these photo-
graphs were procured may be interesting.
I was in camp at the time with one tusker and four female
Elephants, which were being used regularly for transport purposes.
From the time the tusker was reported to be seeking the company
of the females he was never let loose to graze, but was kept tied
up. I had promised the Burmese mahouts liberal rewards if they
would assist me in procuring some “ snap-shots”; and one day last
February one of them came to tell me the tusker was showing
undoubted excitement ; he was reported to have been tugging at
his chain and looking “nastily” at his keeper. There were,
however, absolutely no signs of ‘‘ must” and no exudation of fluid
from the hole in the temple.
The tusker was said to have shown a preference for one of the
females that had calved about 12 months previously. So I had
her tethered*fore and aft in a small glade, and erected my camera
quite in the open, about 10 yards off. Having focussed her and
got everything ready, the tusker was quietly ridden up behind
the female, and as the mahout slipped off, he slowly advanced
towards her, However, this was evidently the wrong female, for
on the approach of the tusker she showed most unmistakable
signs of fear by persistently screaming and straining at her chain.
The tusker was therefore secured, and this female marched off,
whilst one of the others was caught and tied up in her place.
The tusker was then again brought out, and immediately proceeded
to mount the female.
I had many opportunities of witnessing the operation, and the
following is a general description :—'The female, when in season,
remains perfectly still and quiet, merely signifying her sense of
the tusker’s approach by moving her tail slightly to one side and
gently shifting her hind feet a few inches further apart. The
tusker creeps up behind the female and begins to show signs of
sexual excitement. He then raises his head and lightly places
his tusks on the female’s back, one on either side of her backbone,
with his trunk lying along the dorsal ridge and reaching to her
shoulder. In this position he remains a length of time, varying
with his state of sexual excitement. From my observations I am
convinced that when he has unrestricted access to females, and is
1 Communicated by the SECRETARY.
2 ON THE COPULATION OF THE INDIAN ELEPHANT. [ Feb. 3,
not therefore greatly excited, he may remain a very considerable
time in this position: this has probably given rise to the tales
that are told of the great length of time occupied by the act of
copulation, which I have heard put at two hours.
Eventually, however, the time arrives when the tusker proceeds
to action. First one fore foot, and then the other, is lifted off the
ground, and slowly swung from side to side across his front,
exactly as though he were deliberating from which foot to take
off. Then, levering himself up by his tusks, he places both fore
feet on the female’s flanks, at the same time sinking down on his
hind legs to an almost sitting position. It is at this moment that
he shows the most violent sexual excitement. His organ, which
up till now has been bent back in the form of an arc, with its end
near the ground, and a few inches only in front of his hind feet,
is shot forward, and jerked up and down in a most violent manner.
It is flourished in the air, and often bent into the form of an §
like the thong of a whip, during which semen may or may not be
emitted. At one moment the organ appears outside the tusker’s
shoulder, the next it is seen protruding out behind his hind legs.
It is kept in a state of continual agitation, and not for one single
moment is it still. Hventually it is jerked up straight forward
and hits the female organ from below. Penetration is then
immediately effected without further difficulty. The tusker then
raises himself until his hind legs are quite straight, his organ is
pushed home, and his fore feet are slid along the female’s back
until they rest on her shoulders. Arrived at this position, he
begins to work very much after the fashion of a dog, and I
usually counted from six to eight horizontal motions. Connection
being then complete, the tusker lifts his tusks off the female’s
back and raises his head into the air. In this position he remains
for a few seconds, and then slowly withdraws his organ, letting
himself gently down to the ground in the same way that he
mounted, and quietly moves off. It is then that the female
shows her only signs of excitement: she trumpets softly as with
pleasure, thrusts forward her ears and stiffens her tail, her whole
conduct being indicative of pleasure and pride. The tusker is
throughout perfectly silent.
It is noteworthy that at no time does the tusker use his fore legs
to steady himself or to grip the female as does a dog or a stallion ;
the feet are invariably kept close together on the top of the female’s
back, and I can quite believe the statement of the Burmans that
it makes no difference whether his feet are fettered or not.
I was tinable to distinguish any sign by which I could tell when
a female was in season, but three out of the four females in camp
did come into season during the period of 10 days in February
during which they were under observation.
The operation was accurately timed on one occasion, and was
found to last exactly one minute.
From my own observations, I am convinced it is the female
that comes in season, and that until she does so come the tusker
P.Z.8.1903-vol.1. P). XI.
1D) Ge S.di Jet, clei Bale & Danielsson L™ lith .
CERATELLA MINIMA.
1903.] ON COELENTERATA FROM ZANZIBAR. 113
will take no notice of her: also that he is ready whenever she is.
I can offer no opinion as to whether the companionship of the
tusker tends to bring the female into season or not.
I must draw attention to the way in which both the tusker and
female absolutely ignored our presence. As I have already stated,
the camera was set up within 10 yards of the tethered female,
quite out in the open without any attempt at concealment; and
though there were sometimes six or eight spectators walking about
and talking within this short distance, on no occasion did either
the tusker or the female take the slightest notice of us or even look
ow way. Directly the act of copulation was over, the mahout
called out to the tusker, who, at the word of command, came towards
him, knelt down, and allowed himself to be mounted and ridden off,
As I have said, the female was always tethered fore and aft
with a long chain, but this was solely with the object of prevent-
ing her swinging ‘round and so getting out of focus; but in no
single instance did she attempt to do this, nor was the chain once
needed except, of course, in the case of the first female who was
not In season.
7. On the Coelenterata collected by Mr. C. Crossland in
Zanzibar.—I. Ceratella minima, n. sp. By Sypney J.
Hickson, M.A., F.R.S., F.Z.S8., Beyer Professor of
Zoology in the Owens College, Manchester.
[ Received December 16, 1902. ]
(Plate XIII’)
Our knowledge of the remarkable family of Hydrozoa, the
Ceratelladie, has been ably summarized in the memoir published
by Prof. Spencer in the Transactions of the Royal Society of
Victoria, 1892 (4). The hitherto recorded species are distributed
as follows :—Cer atella fusca: Coogee, Bondi (N.S.W.), Broughton
Island, Flinders Island, Lord Howe Island. C. procumbens « C: ape
of Good Hope, Natal. CO. spinosa: Port Natal. Chitina ericopsis :
New Zealand. Dehitella atrorubens: Delagoa Bay. In brief,
the family has hitherto been known to occur only in Australasian
and §. African waters.
The discovery of a new species of the genus Ceratella in the
tropical waters of the Zanzibar coast is in itself worthy of note,
but especially so in view of the fact that it is associated with
corals, aleyonarians, and other animals characteristic of the
tropical belt of the Kast African coast. I am indebted to
Mr. C. Crossland, of Clare College, Cambridge, who collected the
three specimens “during his recent expedition to Zanzibar, for
permission to examine and describe them.
The African species of the family have not yet been accurately
described, but the accounts of Gray (1) and Carter (2) are
! Wor explanation of the Plate see p. 116.
Proc. Zoot. Soc.—1903, Vou. I. No. VIII: 8
114 PROF. 8. J. HICKSON ON | Feb. 3,
sufficient to show that the Zanzibar specimens should be described
as belonging to a new species.
They appear to be most closely related to the Australian
species Ceratella fusca, but differ from it in one or two characters
which Spencer and his predecessors regarded as of generic
importance.
The genus was defined by Spencer as follows :—Colony
irregularly branching; more or less expanded in one plane;
growing from a creeping base. Main stem flattened, branches
rounded and beset with bracket-like hydrophores.
In C. minima from Zanzibar the main stem is not flattened
but perfectly cylindrical in form, and the hydrophores are so
extremely reduced or rudimentary that they are little more than
ridges on the proximal lips of the hydropores. The branching,
moreover, appears to be strictly in one plane, and the terminal
branches are much more slender and delicate than in the other
species. Before passing on to the specific characters, the size of
the specimens must be considered.
The measurements are as follows :—
Specimen A. Specimen B. Specimen C.
mm. mm. mm.
Height of the colony ......... 29 22 3D
Maximum expanse of the
branchesian eee creer 65 38 50
Diameter of main stem..... : 1:2 0:75 1
The colonies of Ceratella fusca are from 11 to 5 inches in
height; the largest Specuems of C. procumbens described by Carter
were 11 inches long by 5 inches broad, and of C. spinosa 44 inches
long by 2 broad. The height of Chitina ericopsis is 14 inches,
with a trunk (main stem) 1 inch in diameter. The size of
Dehitella is not given by Gray, but from the figures it may be
judged that it is larger than Ceratella fusca. From these figures
it is clear that the Zanzibar specimens are much smaller than the
average size of the adult colonies of the other species. Are they,
therefore, to be regarded as young colonies or as the representa-
tives of a dwarf species? If they are young colonies, it is quite
possible that the main stem or trunk becomes somewhat compressed
in the plane of branching as the colony grows; but the fact that
all the three specimens obtained are of approximately the same
size, suggests that they have reached or nearly reached their
maximum growth. The dwarfing of the tropical species of a genus
that is principally distributed in temperate waters is not without
parallel in the group of Celenterata. The very rudimentary
character of the hydrophores, however, cannot be explained by
the suggestion of immaturity, and must be regarded as of specific
importance. It is true that no gonophores have been discovered
in the specimens, but it is quite probable that, as in other
Ceelenterates of the tropics, their production is rapid and strictly
seasonal, so that no argument can be deduced from this character,
either for or against the theory of juvenility.
The Colony.—The branching is not very profuse, and strictly
1903. ] COBLENTERATA FROM ZANZIBAR. 115
confined to one plane (Pl. XIII. fig. 1). The main stem and the
larger branches seem to have divided dichotomously with the pre-
dominance of the most favoured branch. The terminal branches
are mery, delicate, branches of 6 mm. in length gradually attenuating
from 0:2 mm. to 0°1 mm. in diameter. Each terminal branch ends in
a facultative growing point, and it appears probable that the growth
is continuous. I have compared my specimens from Z anzibar
with a beautifully preserved specimen of Ceratella fusca, for which
the Manchester Museum is indebted to Prof. Spencer ; and I have
noticed that the Australian species is much coarser in appearance,
especially in the region of the terminal branches. I have seen
nothing in the Zanzibar species corresponding to what Spencer
calls “The growing ends of the smaller branches” in Ceratella
fusca, which are flattened in a plane at right angles to that in
which the general growth takes place and are entirely devoid of
zooids. It is possible that the difference may be accounted for
on the supposition that in Ceratella fusca the growth is seasonal
or periodic.
The branches of C. minima are invariably rounded. I have
seen no evidence of a compression or flattening in any region.
The surface is relatively smooth and free from any spines. The
hydrophores are represented by very narrow ridges on the
proximal border of the hydropores. The principal horny fibres,
running longitudinally with a slightly spiral twist, may be clearly
seen through the superficial ectoderm, and in the spirit-specimens
give a ribbed appearance to the surface (Pl. XIII. fig. 2).
The Zooids are numerous on the terminal branches, less
numerous on the thicker branches, and very scarce on the main
branches, as in Ceratella fusca. On the terminal branches they are
arranged slightly to one side of the two lateral lines at intervals
of about one millimetre opposite or alternate to one another. A.
few zooids occur more irregularly distributed.
On comparing such a terminal branch with one of Ceratella
Jusca, it is at once apparent that in the Australian species the
zooids are more numerous and much more irregularly distributed
on all sides of the branch. Hach fully expanded zooid projects
about 0°'7 mm. from the hydropore, and is about 0°14 mm. in
diameter. It bears a variable number, but usually nine capitate
tentacles, each about 0-1 mm. in length.
No gonophores were found on any of the three specimens I
have examined. The skeleton in the terminal branches consists
of one or two main longitudinal horny rods supporting numerous
looping and irregular bands, which maintain the eylindr ical form
of the branch (PI. S00 fig. 3).
In the larger branches the main longitudinal rods are more
numerous, and, being chiefly superficial in position, give a slightly
spiral, longitudinally striated or ribbed appearance to the surface,
in this respect offering a marked contrast to Cerateila fusca.
A series of sections thr ough a small branch shows that there is
present a thin continuous coat of ectoderm covering the whole
branch, as described and figured by Spencer in Ceratella fusca,
8 *
116 ON CGELENTERATA FROM ZANZIBAR. [ Feb. 3,
The arrangement of the canals and the general histology does not
appear to differ materially from the description given by the same
author ; but as the state of preservation of the Zanzibar specimens
was not perfect, a detailed and critical examination of the sections
was not made. I was fortunate enough to find, however, a few
nematocysts that were exploded but remained in situ. They are
(Pl. XIII. fig. 4) very similar in form to the small nematocysts
of Millepora, and exhibit a vesicle and neck 0:01 mm. in length,
armed with four barbs or spines. The thread was invariably
broken, and I have no means of measuring its extreme length.
The endoderm of the tentacles is solid. .
The diagnosis of the species is as follows :—
‘CERATELLA MINIMA, Sp. nov.
Colony probably erect, branching strictly in one plane, irregularly
and not very profusely. The main stems and all the branches
cylindrical in form. Hydrophores very slightly developed.
Skeleton consisting of dark brown longitudinally disposed horny
fibres, united by loops and bands, forming in the larger branches
a dense and firm but flexible skeletal plexus.
Zooids situated slightly to one side of the plane of branching of
the colony, alternately or in pairs, at distances of about 1 mm. on
the terminal branches. Largest colony 29 mm. in height, with a
maximum expanse of 65 mm.
Locality. Ganzibar, shallow water.
LITERATURE.
1. J. HE. Gray.—Notes on the Ceratellade, a Family of Keratose
Sponges. Proc. Zool. Soc. 1868, p. 575.
2. H. J. Carrer.—Transformation of an entire Shell into
Chitinous Structure by the Polype Hydractinia, with short
descriptions of the Polypidoms of five other species. Ann.
& Mag. Nat. Hist. ser. 4, 1873, x1. p. 1.
3. W. M. Bate.—Some new and rare Hydroida in the Australian
Museum. Proc. Linn. Soc. N.S. Wales, 1889, p. 748.
4. W. B. Spencer.—On the Structure of Ceratella fusca. Trans,
Royal Soc. of Victoria, 1892, p. 8.
EXPLANATION OF PLATE XIII.
Fig. 1. Drawing of the whole colony of Ceratella minima (specimen A) twice the
natural size. The base of attachment is covered by an encrusting polyzoon.
Vig. 2. A portion of a large branch and smaller branchiet taken from the region
marked * in fig. 1, more highly magnified, showing the polyps P partially
extended, bearing aa irregular number of knobbed tentacles, ¢.¢. At .h. are
seen the rudimentary hydrophores. The figures show the general arrangement
of the fibres of the horny skeleton. In the large branch they are closely
crowded together. In the smaller branches the principal fibres are separated
by considerable spaces but maintain a parallel arrangement.
Fig. 8. A still smaller branch more highly magnified, in which there is only one
primary longitudinal fibre.
Vig. 4. One of the nematocysts of the tentacles. The diameter of the cyst is about
0°01 mm.
1903.] ON THE PLANKTON OF THE FAEROE CHANNEL, 117
8. Contributions to our Knowledge of the Plankton of the
Faeroe Channel.—No. VIII." By G. Herserr Fow er,
BoA Phe De ok: Zea:
[Received December 20, 1902.]
(Text-figures 13-17.)
The present paper contains notes (in some eases due to the
valued help of friends) on Beroé, Arachnactis, Podon, the Ostracoda,
the Copepoda, the Amphipoda, and the Schizopoda, captured by
H.M.S. ‘ Research’ in 1896 and 1897 in the Faeroe Channel.
CTENOPHORA.
Brro# cucumis Fabricius.
This species, characteristic of cold Arctic currents *, was taken in
the following hauls :—
16 @iu., 300 to 170 fathoms, seven specimens.
13 e, 400 to ¢ fathoms, one fragment.
20 d, 500 to 400 fathoms, one specimen.
Most specimens showed the characteristic brick-red or vose tint,
and though much battered and in some cases inverted, were
referable with a fair amount of certainty to this species.
ANTHOZOA.
ARACTIINACTIS ALBIDA M. Sars.
‘Some information as to the developmental succession of the
mesenteries in this form was given in No. III. of this series.
Since its. publication, IT am glad to say that it has been substanti-
ally corroborated by Prof. van Beneden °.
He agrees with my suggestion to separate the Channel and
North Sea Arachnactis from albida of the Faeroe Channel, and
describes it under the name of loydii, under the idea, which is
probable, but at present unproved, that it will be shown eventu-
ally to be the larva of Cereanthus Uoydii. Till this has been
proved, I venture to think it better to retain my provisional name
of bowrnet for this form.
The occurrences of A. albida are shown in the table (p. 118):
it occurred in over 61 per cent. of epiplankton hauls, never in a
mesoplankton haul, and may fairly be taken to be a purely
epiplankton form. It was present in considerable quantity, as
many as 50 specimens having been taken in one haul.
1 The references to previous papers in the Society’s Proceedings are :—No. I., 1896,
p- 991; No. II., 1897, p. 523; No. IIT., 1897, p. 803; No. LV., 1898, p.540; No. V.,
1898, p. 550; No. VI., 1898, p. 567; No. VIL., 1898, p. 1016. I regret that various
circumstances, mostly beyond my control, have caused so great a lapse of time
between this paper and No. VII.
2 Chun: ‘ Die Ctenophoren der Plankton-Expedition,’ p. 26.
3 Van Beneden: ‘ Anthozoaires de la Plankton-Expédition?
118 DR. G. HERBERT FOWLER ON THE [ Feb. 3,
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1903.]
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buoy opruyepe | + it i i+? ay ee Ret er ed oe te SGrararse foe cpaean il) op
*yooog,
igi bebcow traaonyy Dba: ee ees re be ee eee erceep a et pelts ge geye
snqvNuagIy snuv)oINGT Janae Ror arg ae ar Pap ge. RB Bar ape Pane Gee eadc
nee ee ee a 2 a eee eee
|
eaT Cajon fy “vue | a ee gine a = Be ges |
ees |
120 DR. G. HERBERT FOWLER ON THE [ Feb. 3,
PHYLLOPODA,
Popon INTERMEDIUS Lilljeborg.
My friend the Rev. T. R. R. Stebbing, F.R.S., was kind enough
to identify and count the specimens of this species. It would no
doubt have occurred in more of the surface hauls had not my
finest tow-net been devoted to the collection of Diatoms for the
Scottish Fishery Board. It is no doubt a purely epiplanktonic
form in the Faeroe Channel; a single specimen only was taken at
20 d (500-400 fathoms) as against about 106 specimens in 7 hauls
at the surface ; the single deep specimen was probably a sinking
corpse.
This species 1s, I believe, known only from the surface’; it
ranges over the Baltic, Norwegian coast up to Vadsé, Denmark,
Boulogne, Concarneau, Trieste.
OSTRACODA.
The representatives of this order belonged exclusively to the
Halocypride, and were mostly taken in the mesoplankton. Only
one species, Conchacia maxima, occurred sufficiently often to allow
of a generalivation as to its horizon.
The identification of a Halocypridis rarely satisfactory without
dissection of the mouth-parts, which means destruction of the
specimen. I have, however, dissected a considerable number, and
feel at all doubtful only in the case of C. porrecta; I have pre-
ferred, however, to leave the specimens under this species rather
than create a new species on the strength of slight differences
in the armature of the mandible.
The table following shows those surface and deep-water hauls,
made by Professor Chun between Finistére and the Canary Islands
with open vertical nets*, which contained the same species as
5S 8
> ‘~
Ss R cy
SE ii aes Res PS Ss
= =| 8 S38 aS = 8
S | Ss S&S 2 Ss
: : SS SS Ss aS =
Haul. | Horizon in fathoms. 2 = 2/3) SS 3 SES
S S : J S
S S cw sy S
JOG 546 to O * L
S
Il. 819 to 0 3 * aS) %
m
VE 546 to O % % ate B %
VII. 873 to O fe * >
|
Surface. iis as % 8
1 J. de Guerne: Bull. Soc. Zool. France, xii. 341 (1887). W. Lilljeborg:
“Cladocera Suecix,’ Upsala, 1900, 4to. F
2 Chun: SB. kén. preuss. Akad. Wissensch. (1889 xxx. Claus: ‘Die Halocy-
priden,’ Wien, 1891, 4to.
1903. ] PLANKTON OF THE FAEROE CHANNEL, 121
were captured by the ‘Research’ in the Faeroe Channel; they
are cited in the text by Roman numerals,
Concuacra MAximA Brady & Norman,
As I have previously pointed out’, this species appears to be
purely mesoplanktonic, in the latitude at any rate of the Faeroe
Channel. It occurred in 50 per cent. of the mesoplankton hauls,
and in three hauls which began at or over 300 fathoms and finished
at the surface ; it was not captured once in hauls between 100
fathoms and the surface (cf. table, p. 118). The species was
fairly common.
The record of previous captures was cited in the second paper of
this series }, and also indicated a mesoplanktonic habit in subarctic
regions, but it is not surprising that in yet colder waters it should
appear at the surface. At 84° 32’ N., 76° E., it was captured
with a surface-net by the ‘ Fram’ *,and is recorded as abundant in
most of the samples of Crustacea from this voyage *.
ConcHa@CIA HYALOPHYLLUM Claus.
Twelve specimens in haul 137, 100 fathoms to surface. Five
specimens too small for satisfactory identification, but perhaps
referable to this species, occurred in hauls 20¢ and 20 d.
Claus * records as other occurrences Chun’s haul LV., Ischia at
492 fathoms, Orotava at the surface.
2 ConcHacIA PORRECTA Claus.
Numerous specimens from 20a 6¢d and one from 137%. Claus
(op. cit.) records it from Chun’s hauls IL, I1., PV., VII.
CoNCH@CIA BOREALIS G, O. Sars.
A single specimen in haul 19 a, 480 to 350 fathoms. Recorded
previously from 250 to 300 fathoms at the Lofoten Islands’, and
from Trondhjem Fjord at 150 fathoms®, This appears to be a
purely cold-water form.
PARACONCH@CIA OBLONGA Claus,
Six specimens in haul 20¢, four in 20d. Claus (op. cit. p. 64)
cites this species as from Chun’s hauls III. and IV.; and states
that it also occurs at the surface, but without giving authority or
details. He remarks on the probable identity of this species with
G. W. Miiller’s variabilis’, a suggestion with which Miiller seems
to agree®. This would extend the distribution considerably, as
Miiller? records it, from the ‘Vettor Pisani’ collections of
1 Proc. Zool. Soc. 1897, p. 523.
2 ¥. Nansen: Norwegian North Polar Expedition. G. O, Sars: Crustacea, p. 11.
3 Td. ibid. p. 137.
4 C. Claus, op. cit. p. 61.
5 G.O. Sars: Forh. Vid.-Selsk. Christiania (1865), vol. 1866, p. 120.
G.S. Brady & A. M. Norman: Trans. Roy. Dublin Soe. (2) v. p. 686.
G. W. Miller: “ Ueber Halocypriden,” Zool. Jalirb. Syst. v. p. 273.
G. W. Miller: ‘ Ostracoden des Golfes von Neapel,’ p. 229.
ara
oe)
122 DR. G, HERBERT FOWLER ON THE [ Feb. 3,
Chierchia, as occurring at various points in the tropics at depths
between 382 and 546 fathoms; it occurs also in the Gulf of
Naples. Brady! records it as having been taken by the ‘ Chal-
lenger’ Expedition off Kandavu, Fiji, and between Marion and
Crozet Islands at unrecorded depths.
HALOCYPRIA GLOBOSA Claus.
Six specimens in haul 137.
This species is known from the surface and at various depths
in the Atlantic’, and is recorded from Gibraltar as taken by the
‘Vettor Pisani’*. Of specimens taken by the ‘Challenger,’ the
record was in one instance lost *; other specimens were captured
at the surface between Api and Cape York’. It seems to be a
form widely distributed both vertically and horizontally.
CONCH@CILLA DAPHNOIDES Claus.
Only three complete specimens, and one empty carapace, refer-
able to this genus were obtained. The specimens on which Claus
founded the genus (with this single species) were all young males :
larger specimens of the genus, including females, were obtained by
Sir John Murray on H.M.S. ‘Triton’ in 1882, from the Cold
Area of the Faeroe Channel, and were described by Canon
Norman and Dr. Brady under the specific name of lacerta, not
without the “suspicion that they may perhaps belong to the adult
form of C. daphnoides”°. My own specimens were too few to
settle the point ; but as the two smaller specimens most resembled
in outline the figure of Claus, and the two largest that of Brady
and Norman, I have left them provisionally under the older specific
name.
Tn addition to Chun’s hauls IIT. and IV., it has been captured
at 200 fathoms off Achill Head (daphnoides), the Faeroe Channel
as above (lacerta), and off Kandavu, Fiji, at an unrecorded depth”
(daphnoides).
CoPEPODA.
Mr. I. C. Thompson was kind enough to report on the
Copepoda in No. IV. of this series of papers*. Since that date,
the arrangement of three then doubtful hauls has required modi-
fication: 12a, which was suspected at the time of capture to have
remained open too long, proves to have no apparent contamination
of undoubted surface forms, and has been moved to the Meso-
plankton; 13e was also suspected, in this case with justice, as
1G. 8S. Brady : “ Myodocopa of the ‘Challenger’ Expedition,” Trans. Zool. Soc.
Xiv. p. 95.
C. Claus, op. cit. p. 79.
G. W. Miller: Zool. Jahrb. Syst. v. p. 270.
G.S. Brady & A. M. Norman: Trans. Roy. Dublin Soc. (2) v. p. 705.
G. 8. Brady: Trans. Zool. Soc. xiv. p. 97.
G.S. Brady & A. M. Norman: Trans. Roy. Dublin Soc. (2) v. p. 697.
G.S. Brady: Trans. Zool. Soc. xiv. p. 95.
Proc. Zool. Soc. 1898, p. 540.
bo
onan } w
1903. | PLANKTON OF THE FAEROE CHANNEL, 123
containing several undoubtedly epiplanktonic species (e. g. Arach-
nactis albida), and has been relegated to the “ doubtful” category,
closure of the net not having taken place at the proper time ;
12 was known to be epiplanktonic, although there was some
doubt as to the exact depth at which it had been towed. I have
therefore reprinted in the table (pp. 118, 119) the captures of the
seven forms which were taken at least six times, sufficiently often
to give approximate data for an estimate of their vertical distri-
bution. In discussing this question, I gave a short table on
p. 546 showing the occurrences of these seven species expressed
in percentages of those Epiplankton and Mesoplankton hauls
which contained Copepoda; this can now be amended as follows,
omitting the four doubtful hauls from the calculation :—
Total hauls containing Copepoda.
ER CRETE
SS Ta ae
Epiplankton. Mesoplankton.
Calanus finmarchicus occurred in 83 9/5 and in 91 /p
1 Bucalanus attenuatus ry) 22 4; 5 41 ,,
Bucheta norvegica _ iit. = Us op
Metridia longa 3 22 ,, & 1555
1Pleuromma abdominale ,, bys 55 58 ,,
Acartia clausii ie 33, z 25 4,
Temora longicornis 5 33 4, 25 OF
The amended table is in harmony with the conclusions drawn
from the former, as to the vertical distribution of these forms,
except in the case of Acartia clausii, the question of which was
expressly reserved (op. cit. p. 549).
Since the publication of Mr. Thompson’s report, I found that
Dr. R. Norris Wolfenden was making an exhaustive study of the
fauna of the Faeroe Channel, and naturally placed my collection
at his disposal. He has been kind enough to furnish the following
notes on new and other species, with some of which he has
already dealt briefly elsewhere *?. Exact data of depth &c. were
not always available, as by the time that Dr. Wolfenden received
the specimens all the epiplankton hauls of a station had in many
cases been put together in one bottle, all the mesoplankton hauls
in another, for economy of space.
PLEUROMMA ROBUSTUM Dahl, Zool. Anzeig. v. p. 16 (1893).
“This is the common Plewromma of the Faeroe Channel, and
was found to be present in considerable numbers in Dr. Fowler's
collection, Pl. abdominale occurring much more rarely. In these
northern latitudes it almost entirely replaces the latter species.
It is readily distinguishable by the horseshoe-shaped mass of red
pigment which is present in the anterior and inferior portion of
the head at the hase of the mouth-organs, and in the male by
1 Dr. Wolfenden points out that as, according to his wide experience, Mucalanus
attenuatus is nowhere met with in the Faeroe Channel, unless perhaps quite
exceptionally, these figures probably refer to Hucalanus elongatus. Similarly, for
the species Pleuromma abdominale should probably be substituted Pl. robustum.
2 R. N. Woltenden: Journ. Marine Biol. Assoc. (1902), vi. p. 344.
124 DR. G. HERBERT FOWLER ON THE [ Feb. 3,
the clasping antenna being on the left side, and the pigment-
spot invariably on the right side. The second feet in both
male and female have the characteristic notch and hook on both
limbs. Length 3-4mm. Haul 20a (200 to 100 fms.) and several
others.”
HererocH#ra ZETEsIos ¢, Wolfenden, op. cit. p. 367.
“The head is like H. papilligera Gbt. Though the end joints
of both anterior antennze were broken off, the 19 joints left had
a length of over 4 mm., with the geniculation between the 18th
and 19th segments. The anterior antenne were therefore much
longer than the whole animal, which was 35mm. There was
considerable asymmetry of the furcal segments, that on the left
being much the longest and broadest. The anterior foot-jaw had
one thick hooked bristle on the 5th lobe, but no ‘“ tooth-comb”
bristle, and the 5th feet were peculiar and unlike those of any
other Heterocheta, displaying an upright and stiff process of the
2nd basal joint, armed with fine stiff hairs on the inner aspect
(like a “tooth-comb ”), and the proximal inner margin of the Ist
joint of the exopodite with a protuberance armed with 4 teeth.
The 2nd basal joint of the foot of the opposite side is armed distally
with short stiff bristles. It could be only the male of 7. grimaldiu
or of H. longicornis, neither of which is yet known, or of H. major
(Dahl). The latter and H. grimaldii ave very large (5-10 mm.),
and though H. zetesios resembles H. longicornis in some points, it
is perhaps better for the present to distinguish it as a new species.
Only one example was met with in Dr. Fowler’s collection, in
haul 20q@ (200 to 100 fathoms).”
ANGISTHUS ATLANTICUS Wolfenden, op. cit. p. 364.
“The occurrence of an example of this genus in the Faeroe
Channel is remarkable. This specimen was found in the collection
made by Dr. Fowler as 20a (200 to 100 fathoms). It had a
total length of 1:45 mm., a 6-segmented anterior antenna with
very long and peculiar sensory processes. It has distinct differ-
ences from dg. mucronatus or Ag. aculeatus Gbt., and also from
the species described by Scott from the Gulf of Guinea as
Lgisthus longirostris.”
Lucicutia MAGNA Wolfenden, sp. n. 3.
“‘ A single specimen found in Dr. Fowler’s collection from 19 @
(480 to 350 fathoms), of 3°54 mm. length, was apparently new.
The anterior antenne were larger than the whole body, by the
terminal one and a half joints. The endopodite of the Ist foot
was two-jointed. The right 5th foot has a strong spiny process
on the inner side of the 2nd basal and the exopodite of two seg-
ments ; the endopodite and exopodite of the left 5th foot being
each of three segments. The size alone distinguishes 1t from the
males of any other known species, only LZ. grandis being larger.”
or
1903.] PLANKTON OF THE FAEROE CHANNEL. 12
AUGAPTILUS zETESIOS Wolfenden, op. cit. p. 369.
“One specimen only was found, in the bottle marked 19 a.c.
Another specimen occurred in the sample marked 20,”
Evucananus crassus Gbt. 1888, Atti Acc. Line., and 1892, Fauna
u. Flora Neapel, v. p. 19.
“This species was of not infrequent occurrence, especially in
the bottles marked B 1 and 13 & (2 to 0) and 137 (100 to 0).
“The writer also has frequently noted its occurrence in the
Faeroe Channel.”
GAETANUS MAJOR Wolfenden, sp. n.
“Two examples of this genus (Gaetanus) were found in
Dr. Fowler's collection marked 19 @ (480 to 350 fathoms). The
copepod very greatly resembled Gaetanus armiger Ght., but the
anterior antenne were longer, reaching beyond the furca by the
length of the last joint; the spines of the last thoracic segment
were comparatively shorter, the lst abdominal segment and the
anal segment shorter, and the fureal segments only as long as
broad (longer than broad in G@. armiger), and each abdominal
segment had a row of pectinations on the posterior border. The
saws of the swimming-feet possessed more teeth; the abdomen
was not nearly half the length of the cephalothorax, and the
whole length of the animal was 5:3 mm. In all these points it
differed from the typical G. armiger, the size of which reaches
only about 3 mm., and justifies its being made into a separate
species,”
Garpius Gbt. 1895, Bull. Mus. Harvard.
“A good many examples of this genus occurred in the deep-
water collections of Dr. Fowler, e.g. in the bottle marked Meso-
plankton 20 (500 to 100 fathoms).
There is no doubt that in the Faeroe Channel there are two
kinds of Gaidius—one agreeing in every particular with the
Gaidius pungens of Giesbrecht; the other, a larger species, also
differing in the segmentation of the Ist and 2nd feet. Gaidius
pungens Gbt. has the exopodite of the Ist foot with only two
segments and the endopodite of the 2nd foot with only one seg-
ment; whereas the northern species has a 3-jointed exopodite
of the 1st foot and a two-jointed endopodite of the 2nd foot.
There are other minor differences. In size the northern species is
much larger; Gaidius pungens Gbt. being 45 mm., as compared
with about 3mm. The Chiridius tenwspinis of G. O. Sars is the
Same species, all being characterized by the peculiar series of
lamellar appendages of the basipodite of the 4th foot.”
PSEUDAETIDEUS ARMATUS Wolfenden.
“Some examples of this species occurred in Dr. Fowler’s col-
lection. It was drawn and described (in MSS. only) before the
126 DR. G. HERBERT FOWLER ON THE | Feb. 3,
writer became acquainted with the recent work of Prof. G. O. Sars,
who figured and described the species as Chiridius armatus. The
writer has published reasons why this generic name should not
be used (see Rep. of the Brit. Assoc. 1892, ‘‘ A proposed Revision
of the Subfamily Aetidtine”), as it is not a Chiridius.”
EucuIRELLA CARINATA Wolfenden, op. cit. p. 366.
“‘One example of this species, measuring 3°54 mm. in length,
was found in Dr. Fowler’s collection in the bottle marked 20.
‘These new species will be described in full, along with the
drawings, in the writer’s monograph which is in hand.”
AMPHIPODA.
T am indebted to the Rev. T. R. R. Stebbing for help in the
determination of some of these forms. Only one species occurred
in sufficient quantity and with sufficient frequency to enable
deductions as to its habitat in the Faeroe Channel being drawn,
namely
PARATHEMISTO OBLIVIA Kroéyer=abyssorum Boeck.
I have already discussed the distribution of this form at some
length’; to the records there given must be added two stations
of the ‘National’ expedition—N. of the Hebrides, and S.W. of
Iceland “—and 12 stations along the route of the ‘Fram’*. Canon
Norman also cites Bonnier as having taken it at 950 metres in
the Bay of Biscay *.
It is apparent from the table (p. 119) that the species is a
true member of the Mesoplankton in this locality, having been
captured in 66 per cent. of the deep hauls; it rises to the surface
at midnight, the only occasion out of 26 Epiplankton hauls being
at that hour (haul 15d).
CyCLOCARIS GUILELMI Chevreux.
Two specimens of a Cyclocaris were obtained in haul 20d,
between 500 and 400 fathoms. Mr. Stebbing informs me that
they agree undoubtedly with the above species, captured in a net
sunk to 600 fathoms near the Lofoten Islands by the ‘ Princesse
Alice’’, the largest of the six specimens being about 12 mm. in
length. My largest specimen, a female, would have measured
about 20 mm. if straight.
Another species of this genus, Cyclocaris tahitensis Stebbing,
was described ® from a single specimen taken by the ‘ Challenger’
off Tahiti, apparently at the surface ’.
1 Proc. Zool. Soc. 1898, pp. 583-585.
2 J. Vosseler: Amphipoden der Plankton-Expedition, p. 80.
3 ¥, Nansen: Norwegian North Polar Expedition. Crustacea, by G. O. Sars,
p. 14.
4 A.M. Norman: Ann. Mag. N. H. (7) v. p. 131.
5 KB. Chevreux: Bull. Soc. Zool. France, xxiv. (1899), p. 148.
6 T. R. R. Stebbing: Chall. Rep. Zool., Amphipoda, p. 661, pl. xvii.
4 J. Murray: Chall. Rep., Summary of Results, p. 1077.
1903.] PLANKTON OF THE FAEROE CHANNEL. 127
Mr. Stebbing points out “that this species has been again
figured in great detail by Professor Sars ', and that Dr. Norman 2
has with almost equal fulness represented his Cyclocaris faroensis.
In Mr. Stebbing’s opinion the differences between the two sets of
figures are purely casual, depending on individual or accidental
‘conditions of the specimens examined, Further, while accepting
provisionally the distinction between the boreal form and the
‘Challenger’ C. tahitensis, he agrees with the view thus expressed
by Dr. Norman, ‘so remarkable is the resemblance, that the
differences seem scarcely varietal; but I hesitate to unite a form
found in the Faroe Channel with one from so distant a locality as
Tahiti.’ ”
Canon Norman does not appear to have known of M. Chevreux’s
name (which apparently has priority); his specimens were captured
by the ‘Triton’ in the Faeroe Channel in 1882, at a depth of
640 fathoms. Professor Sars’s specimens were apparently from
various depths along the course of the ‘ Fram,’ the smallest
specimens at the least depths.
This species (omitting tahitensis as specifically distinct) is
evidently a Polar and deep-water form,
TRYPHANA MALMI Boeck.
A single specimen, from haul 13 7, 100 fathoms to the surface,
was identified by Mr. Stebbing. The previous records of its
occurrence are: Hardangerfjord, 100 fathoms’; from Nansen-
fjord to Hardangerfjord *; Folgeré, Sunde, Foldenfjord, 80 to
100 fathoms °. Sars’ regards 7’. nordenshivldi Boeck and 7’, boechi
Stebbing as being the males of 7’, malmi. Of these the former 4
was described from the Sofia Expedition as off the Faeroe Islands
at 65° N., the latter ® from 18° 8’ N., 30° 5’ W., at the surface.
EvTHEMIsTOo sp. indet.
A number of specimens too much broken for recognition
occurred in the haul 15d at midnight.
EUTHEMISTO BISPINOSA Boeck.
Occurred in 15 (530 to 0 fathoms) and 15d (surface, at mid-
night). The records of this species were exclusively Arctic, till
it was captured by the ‘ National’ expedition in the Sargasso Sea
between 218 fathoms and the surface.’
EvUTHEMISTO COMPRESSA Goés.
A single specimen at 12 ¢ (450 to 320 fathoms). This also was
believed to be exclusively an Arctic form, but has now been
1 F. Nansen: Norwegian North Polar Expedition. G. O. Sars: Crustacea, p. 20,
pls. ii. & ili.
2 A. M. Norman: Ann. Mag. N. H. (7) v. p. 197, pl. vi.
3 Boeck: ‘Skandinaviske og Arktiske Amphipoda,’ 1872, p. 92,
4 C. Bovallius: Vega-Exped. Vetensk. lakttag. iv. p. 573.
® Sars: Crustacea of Norway ; Amphipoda, i. p. 17 (1890).
6 T. R. R. Stebbing: Chall, Rep. Amphipoda, p. 1539.
128 DR. G, HERBERT FOWLER ON THE [ Feb. 3,
recorded from the Antarctic region, as well as from the Gulf-
Stream proper, the Sargasso Sea, and South Equatorial Drift’.
EUTHEMISTO LIBELLULA Mandt.
A single specimen in haul 15d at the surface at midnight. It
is widely distributed over the Arctic Seas, but, unlike the two
foregoing species, was not taken in southern waters by the
‘ National.’
SCINA BOREALIS G. O. Sars.
A single specimen from 13g (465 to 335 fathoms) was acci-
dentally included among the Copepoda sent to Mr. Thompson
and identified by Mr. A. O.Walker. According to Canon Norman ’,
the distribution of this species is mesoplanktonic, mostly northern,
but also in the Bay of Biscay (960 metres). It reaches to about
SOUUINE
SCHIZOPODA.
The Schizopoda captured belong exclusively to the Huphausiacea,
and are referable to only three species. Several forms which
might have been reasonably expected among the captures were
absent. <A list of the British species with their distribution is
given by Canon Norman, in his paper on British Lophogastridee
and Euphausiidee *. :
TTHYSANOESSA LONGICAUDATA Kroyer.
A considerable number of specimens of this species were
captured: it appeared to be the commonest Schizopod of the
Faeroe Channel at the time. The synonymy appears to be
Thysanopoda longicaudata Kroyer=Thysanoessa tenera Sars=
Thysanoessa longicaudata of Hansen, Norman, Ortmann, We.
The species ranges from the West Coast of Norway right across
to Greenland and into the Labrador current; the ‘ National’ ’ ceased
to take it (after almost daily captures up to that moment) from
the date of entering the warm water of the Gulf-Stream (‘ Florida-
strom’). Sars*, who described the species from deep water in
the Varanger Fjord, records it also as from the surface at four
stations between Norway and Jan Mayen on the cruise of the
‘Vormgen’". It appears, therefore, to be essentially a cold-
water species, an Arctic type-form, and was captured by the
Siirenotdes |
It has, however, been recorded twice from British coasts *.
Vosseler: Amphipoden der Plankton-Expedition, p. 86.
A. M. Norman: Ann. Mag. N. H. (7) v. p. 185.
F.. Nansen, op. cié. p. 19.
4 A.M. Norman: Ann. Mag. Nat. Hist. (6) ix. p. 454.
5 A. Ortmann: ‘ Decapoden und Schizopoden der Plankton-Expedition,’ p. 14.
6 G.O. Sars: Forhandlinger Videnskabs-Selskabet (Christiania), 1882, no. 18, p. 58.
7 G.O.Sars: Norwegian North Atlantic Expedition (Crustacea), pt. 1. p. 18.
8 F. Nansen: Norwegian North Polar Expedition. G. O. Sars: Crustacea, p. 14.
9 A. M. Norman: Aun. Mag. Nat. Hist. (6) ix. p. 463 ; and the papers there cited.
a
1903. ] PLANKTON OF THE FAEROE CHANNEL. 129
Once it occurred in enormous quantity in St. Andrew’s Bay in
company with WVyctiphanes norvegica, once at Redear with a
similar swarm of Huthemisto compressa, the latter being also an
Arctic type-form'. In both these cases it is probable that the
creatures had been driven down the North Sea by a strong southerly
current, in the manner which I have already suggested’ for
Parathemisto oblivia; and it has therefore no more right to be
regarded as a “ British” species than an occasional Velella or
Tanthina brought up by the North Atlantic Drift to our shoves.
According to Ortmann (op. cit.), the ‘ National’ hauls gave no
indication of the vertical distribution of this species.
Tt will appear from the table (p. 119) that the ‘Research’ was
more fortunate, and the hauls point to its having a distinct
preference for the mesoplankton in the Faeroe Channel. Like
(at any rate some) other mesoplanktonic species, it rises to the
surface at night. Specimens with adult characters were captured
in 19 per cent. of epiplankton hauls, but in 66 per cent. of meso-
plankton hauls. On the other hand, larval and post-larval stages,
apparently referable to this species, were obtained in 38 per cent.
of epiplankton hauls, but only in one mesoplankton haul, and
that one terminating near the 100 fathoms. The species, there-
fore, appears to be epiplanktonic when young, mesoplanktonic
when adult, so far as these observations go and in the Faeroe
Channel at this time of year. In seeking deeper (colder) water
in this locality, it follows what appears to be the practice of other
Arctic type-forms when they meet the warmer water of the North
Atlantic Drift. That this was not apparent from the results of
the ‘ National’ is probably due to the fact that from the Hebrides
almost up to the moment of coming into the Gulf-Stream the
vessel was in far colder surface-water than that of the Faeroe
Channel in summer.
The larvee mentioned above ranged from an early Calyptopis
stage up to the adult condition. It was not, of course, possible to
derive them all with certainty from Thysanoessa longicaudata ; but
the majority may be safely referred to this species, not only
because the adults captured were far in excess of any other
Kuphausid, but also because the larvee could be traced gradually
through successive stages back to the Calyptopis. The meta-
morphoses of this species follow the lines indicated by Sars * for
Nyetiphanes, Huphausia, and Thysanopoda.
As Paul Mayer* has shown, the spination of the telson of
Malacostracan larvee yields a character important both for phylo-
geny and for diagnosis. It has not as yet, I think, been pointed
out that the condition of the telson’ in Huphaustid: affords a
further argument for the view maintained by Boas’ and others,
1 ©, Chun: ‘ Beziehungen zwischen dem arktischen und antarktischen Plankton,’
Stuttgart, 1897, 8vo, p. 30.
2 Proc. Zool. Soc. 1898, p. 583.
3G. O. Sars: Chall. Rep. Zool. xiii. (Schizopoda).
4 P. Mayer: Jenaische Zeitschrift, xi. (1877), p. 246 et seqq.
® J. E. V. Boas: Morphologisches Jahrbuch, viii. p. 485.
Proc. Zoot. Soc.—1903, Vou. I. No. TX. 9
130 DR. G. HERBERT FOWLER ON THE [ Feb. 3,
that this family takes its origin very near to the root of the
Decapodan stem, and that it has far closer affinities with the
latter than with the Myside. In Wyctiphanes’ and Huphausia *,
and possibly in other genera also, the youngest Calyptopis-larvee
show seven spines on each side of the telson; unlike all other
Schizopoda, so far as is known, except perhaps Lophogaster, they
are thus in absolute accord with the ‘“ urspriingliche Borstenzahl
7+7” which Mayer attributes to the primitive Macruran and
Brachyuran. In Huphausia, Nyctiphanes, Thysanopoda, and
Nematoscelis according to Sars (op. cit.), and in Thysanoessa,
the number is increased at later stages by a median terminal
spine, which, like the others, is jointed to the telson. Accepting
Mayer’s enumeration of the spines from the middle line outwards,
and styling the median azygos spine of the EKuphausiide as 0,—
spines 7 are found in the adult Thysanoessa about one-third of
the length of the telson from the root; spines 6 at about two-
thirds of its length from the root; spines 5 are lost; spines 4
persist as the large lateral jointed spines near the end of the adult
telson *; and spines 3, 2, 1, 0 disappear altogether in the course
of development. On page 131, I have illustrated four stages in
this reduction omitted by Sars, of which fig. 15 does not quite
bear out his description: these show the disappearance of the
median spine 0, and the commencement of a new wnjointed
growth of the telson backwards, to form the lanceolate tip of the
adult. The character of the telson and the presence of this
median spine will apparently form a good criterion for the
separation of Kuphausidan larvee (at stages later than the Meta-
nauplius) from other Schizopodan and from Decapodan larvee.
The earliest Calyptopis-larvee captured by the ‘ Research’
resembled closely those figured by Sars (op. cit.) for other genera,
except for the facts that the carapace was much more globular
anteriorly and was devoid of spines or processes.
NYcTIPHANES NORVEGICA M. Sars.
This form was captured on only six occasions. Although a
North Atlantic type, it is not an essentially Arctic type like
Thysanoessa longicaudata : it is of constant occurrence in certain
localities on our own coasts, and has been recorded from as far
south as Portugal. The various records of its occurrence are cited
by Canon Norman *, but unfortunately the size of the individuals
and the depth from which they were derived are only rarely noted.
I am informed by Sir John Murray that, in his experience, large
adult specimens are taken only in deep water.
1 G. O. Sars: Chall. Rep. Zool., xiii. Schizopoda, pl. xxvii. fig. 6.
2 C, Claus: Untersuch. Crustaceen-Systems, pl. i. fig. 2, Wien, 1876, 4to.
3 With regard to these, Boas (op. cit. p. 623, note 5) has suggested that they may
be homologous with the long caudal appendages of Nebalia and many Phyllopods.
This possibility is rendered considerably more remote by their being merely two
persistent spines out of a series which is not represented in the forms cited by him.
4 A.M. Norman: Ann. Mag. Nat. Hist. (6) ix. pp. 459-460 (1892).
1903. ] PLANKTON OF THE FAEROE CHANNEL, Le
Text-fig. 13. Text-fig. 14.
5
Yi
Text-fig 15. Text-fig. 16. Text-fig. 17.
+
Telsons of larvae attributed to Thysanoessa longicaudata, showing reduction of the
primitive seven pairs of spines and formation of the median unjointed spine
of the telson.
Fig. 13. Early Calyptopis, 15 mm. long.
14. Furcilia, 4°5 mm. lone.
15. Cyrtopia, 5 mm. long.
16. Late Cyrtopia, 5 mm. long.
17. 4mm. long.
Q*
132 ON THE PLANKTON OF THE FAEROE CHANNEL. [ Feb. 3,
The same distribution was noted by Vallentin and Cunning-
ham ':—‘“ The adult, so far as our information allows of a
decision, lives on the bottom, and never swims far from the
ground [90- —95 fathoms in this ‘instancel, while the young, up to
half or three-quarters the size of the adult, occur abundantly at
the very surface and at all intermediate depths. As mentioned
above, Mr. Murray found swarms of individuals at the surface in
the Farée Channel, but none of these were full-grown, and very
few more than half the adult size.”
Thave no doubt that this generalization will prove true for
greater depths: I took adult specimens, over 35 mm. in length,
only between 350-220 fathoms, 400-300 fathoms, 500-400
fathoms: the remaining specimens varied from 9 to 17 mm. in
leneth. In other words, Vyctiphanes norvegica is apparently
mesoplanktonic when adult
A few larve, larger for their stage of development than those
attributed to Thysanoessa, were taken at the surface, and may
perhaps belong to Vyctiphanes: they have not been included in
the tables.
THYSANOPODA MICROPHTHALMA Ortmann. (? = Thysanopoda
microphthalma G. O. Sars.) .
Three specimens, recognizable as young forms by the character
of the second maxilla and gills, and by the spination of the telson,
of about eight, twelve, and fourteen mm. in length, appear to be
referable to the same species as specimens recorded in quantity
by the ‘ National’ in 60° 3’ N., 27° 0' W., at a probable depth of
between 218 and 328 fathoms. These were referred by Ortmann”
to Thysanopoda microphthalma of Sars *, a species founded on two
specimens from the surface at 26° 21' N., 33° 37’ W., and 7° N.,
23° W. respectively. The identity of the ‘National’ specimens
with those described by Sars seems to me rather doubtful: firstly,
because it is not very likely that a rare form such as this should
occur as adult both at the surface near the Equator and also at
218-328 fathoms in the Greenland Sea, or at 500-400 fathoms
in the yet colder water of the Faeroe Ohammel ; secondly, because
Ortmann himself indicates some points of difference between his
specimens and those of Sars. My own specimens agree with
Ortmann’s figure, and differ from Sars’s description, in “the shape
of the antennal scale, and in the absence of a spine from the
second joint of the first antenna. The telson was not hispid,
probably owing to immaturity ; the eye was somewhat flatter than
in Ortmann’s figure, and showed slight signs of a constriction
such as is characteristic of Thysanoessa. The matter cannot be
settled in default of further specimens, owing to the fact that
Sars gave only a woodcut of the entire animal, and no figures of
the detailed anatomy.
1 R. Vallentin & J. T. Cunningham: Quart. Journ. Micr. Sci. xxviii. pp. 325-6.
2 A. Ortmann: ‘ Decapoden und Schizopoden der Plankton-Expedition,’ p. 9.
3 G. O. Sars: Chall. Rep. Zool., xiii. Schizopoda, p. 106.
1903. ] ON THE ELK IN NORWAY. 133
Of the three ‘ Research’ specimens of this species, two came
from the Mesoplankton, one from a haul of 480 to O fathoms:
no examples were captured at the surface, unless some of the
larvee attributed to Thysanoessa belonged to this species; this is
unlikely, because in that case the small size of the eyes would
probably have betrayed them.
9, On the Present Condition and Habits of the Elk in
Norway. By H. J. Exwss, F.R.S.
[ Received January 19, 1903. ]
(Text-figures 18-26.)
So little seems to be known by naturalists concerning the actual
condition of this remarkable animal, that I think some of the
observations I have made during six Septembers, which I have
spent entirely in the fascinating sport of Elk-hunting, may be
worth recording in the Proceedings of this Society.
My experience has been gained entirely in the provinces of
North and South Trondhjem, where the Elk is more numerous
than perhaps in any other part of Europe, and where it seems to
attain a greater size and vigour, if one may judge by the develop-
ment of the horns (see text-fig. 18, p. 134) than anywhere else in
Scandinavia. ‘
I have also derived much valuable information from Capt. Gerard
Ferrand, who was, I think, the first Englishman to hunt regularly
in the same districts, and whose experience continued almost
without a break for 20 years from 1865, This has been invaluable
in confirming, and to some extent modifying, my own observations
and what I have been able to learn from the natives of these
provinces.
Forty years ago the Elk was a much scarcer animal in Central
and Northern Norway than it is now, or rather was ten years ago,
since which time it has probably decreased in numbers. At that
time it was hunted for its meat alone by the proprietors of the
regions where it occurred, and in the large tracts of forest
belonging to the Government it was not allowed to be shot until
1880. Up till that time very few foreigners knew what magni-
ficent sport Elk-hunting was, and as the natives preferred the
meat of cows to that of bulls, which, after the rutting-season
begins, is extremely rank and hard, the bulls increased in numbers
and were able to grow to a size and age which they rarely
attain now.
The legal season for killing Elk was formerly much longer
than it is now, and in North Trondhjem lasted for three mone:
Then it was cut down to 6 weeks, from the lst September till the
15th October, and now has been still further reduced to a month
or 3 weeks in some provinces, whilst in Sweden only 15 days are
allowed during which the animals can legally be killed.
(azIs juu #) ‘BIABUIPUBOG UWOAF Uoas aarty J “qsosiey ayy Jou Yonoyy
‘QsopiM oy ‘ueds UT SeyOUL PE +N PY Jnoqe ‘KeMio yy ‘way puory, WosZ YL Fup AT[NF Jo susopyy
[Feb. 3,
MR. H. J. ELWES ON
‘BI SU-9x0],
134
1903. THE ELK IN NORWAY. 135
Text-fig. 19.
Horns of fully adult Elk killed in Bjorndal, 65° N., Sept. 41895; 18 points; a fair
average size for this district, but much larger than usually seen in the south.
(4 nat. size.)
Text-fig. 20.
Horns of a full-grown but not adult Elk, killed Sept. 1896 in Upper Namdalen, 65° N.
Supposed by natives to be 4 or 5 years old. Probably this type would develop
into such antlers as shown in text-fig. 18 if conditions were favourable.
(1, nat. size.)
Text-fig. 21.
Horns of young Elk, supposed to be 3 years old. These would probably develop
into such a head as that shown in text-fig. 19. (;'; nat. size.)
136 MR. H. J. ELWES ON i ebra;
This rapid increase of an animal which, on account of its great
size and conspicuous tracks, cannot escape unnoticed in an
inhabited country, must be entirely attributed to the wise game-
laws made by the Norwegian Government, and in most parts
honestly observed by the most law-abiding and _ well-governed
people I have ever met with. In former times the Elk, in Scandi-
navia, as In North America, was hunted down in winter on ski
(the Norwegian form of snowshoe), and slaughtered for its meat
by every peasant farmer, till it had almost been exterminated.
When, however, a law was made that it could be hunted only in
the month of September, which period was for a time somewhat
extended in North Trondhjem, its numbers soon increased, and
about 20 years ago attained such proportions that English and
German sportsmen began to visit Norway to hunt Elk. The
landowners in some districts then discovered that, instead of
hunting themselves or paying Swedish hunters, who, from long
experience, were more expert, to kill thew Elk for them, the
right of killing Elk, which is limited to one animal on each farm,
had a letting value; and when many such rights are united so
that a large tract of country can be reserved to the lessee, this —
value was worth a little trouble to maintain. The consequence
has been that, though poaching and killing Elk out of season is
not entirely unknown, yet it cannot be carried on extensively ; and
I have little doubt that the Elk will continue to thrive wherever
the country is suitable.
The statistics which I append show the numbers of bull and cow
Elk which are known to have been legally killed in the various
‘“« Amts” or provinces of Norway in 1894, and the average for the
previous five years, and may be taken as very nearly exact, though,
according to some, the returns for the southern provinces are not
so accurate as for the northern ones, and this record takes no
account of those illegally killed. It will be seen that only four
provinces of Norway (excluding Finmark), namely, Stavanger,
North and South Bergenhus, and Romsdal, all of which are on
the south-west coast and exposed to the warm and wet influence of
the Gulf-Stream, are without Elk; and it may be added that in
those four provinces wild Reindeer are most numerous.
North Trondhjem is before all the rest in numbers; and in this
province I believe the size of the horns is or was also much larger
on the average than in any of the southern districts (text-fig. 18,
p. 134).
It is said that the Elk is gradually extending its range north-
wards, and has appeared in the southern parts of the province of
Nordland only in the last few years; and there seems to be no
reason why it should not go still further, as in North America the
Moose (which is so nearly allied to the Elk, that I do not think
it ean be looked on as more than’‘a variety of that animal) is
found in regions where the climate and food is certainly not more
favourable to its habits than they seem to be in some parts of
Nordland and Finmarken.
The greater part of North Trondhjem and a large proportion of
1903.] THE ELK IN NORWAY. 137
South Trondhjem are mountainous, barren, and thinly populated
except on the coast and in some inland valleys and fiords, and
are covered with forests of birch, spruce, and Scotch fir up to an
elevation of about 2000 feet, wherever the ground is not too rocky
or swampy for these trees to grow. Above that elevation there
are fjelds or bare mountainous uplands, the lower slopes and
sheltered dells in which are more or less clothed with birch, willow,
mountain-ash, and alder, mixed with stunted firs. There are
many large lakes and large areas of peat-bog; but these bogs are
rarely so deep and soft as to be impassable, and even after long
periods of rain a man can cross them by picking his way.
The country is divided into so-called farms, most of which con-
sist of small patches of oats and potatoes, with from 10 to 30 acres
of meadow-land which is mown for hay, and the produce of which
is eked out by small ricks which are put up wherever a sledge-
load of grass can be got together in the forest. A large area of
forest and mountain is attached to each farm, and there are usually
one or two sxeters' on the mountain, to which the cows are driven
for about three months in summer. Though most of these farms
belong to the occupiers there are some large private properties,
belonging to timber companies and public institutions, and much
of the higher fjeld and barren mountain remains in the hands of
the Government.
In the wilder and less populated districts the Elk are hardly
disturbed during eleven months of the year except by the occasional
attacks of bears and wolves. Though the bear has become rarer
of late years, the wolf, on the contrary, has appeared in districts
where it was until recently almost unknown; but at present
they have confined their attacks rather to the semi-wild rein-
deer, which are kept in some numbers by the Lapps all along
the Swedish frontier, from Roros northward, and to the sheep, of
which every farmer has from 10 to 30. It seems to be doubtful
whether bears can kill full-grown bull Elk’, but during my last
hunting-season I found the remains of no less than three Elk calves
which had been killed by them.
The favourite food of the Elk in summer consists almost entirely
of the twigs and leaves of birch, willow, and mountain-ash, and
in winter of the branches and bark of the same shrubs. Wherever
there is a grove of mountain-ash, Elk will live almost entirely on
it so long as they can get it, and in districts where Elk are abun-
dant the tree is constantly eaten down, so that it seems likely to
become much scarcer than at present.
Scotch-fir twigs are also largely consumed during the winter,
but do not seem to be eaten so long as mountain-ash can be
easily procured. Though I have never actually seen Elk eating
grass, | am assured by the native hunters and by Capt. Ferrand
that they do so in summer to some extent, and also bite off the
flower-heads of Hpilobiwm and other plants. Water-lily roots,
1 “Szeter” is the Norsk term for a shieling.
2 Capt. Ferrand has known a bull Elk beat off the attack of two bears in company.
138 MR. H. J. ELWES ON [ Feb. 3,
which are said to be the favourite summer food of the Moose in
America, are also eaten, though the plant is rare in the districts
where I have hunted.
The Elk seems to spend the greater part of the day in feeding,
though it lies down for some hours to chew the cud. Some
hunters say that it has a regular time for lying down, and
will not hunt between 11 and 2 o’clock, because of the great
difficulty of approaching the animal when it is resting. I have,
however, seen them lying at all hours between 9 a.m. and 3 P.M.,
and have found them up and feeding at midday and later, so that
there is evidently no rule beyond the appetite of the individual
animal.
From its great size and the nature of its food, the Elk requires
a much larger extent of feeding-ground than any animal I am
acquainted with, except the elephant; and I should suppose that
at least three square miles of suitable forest would not support
more than one Elk continuously, judging from the number I have
found in places where they were at home and undisturbed.
It is an animal of extremely solitary habits, and in summer
more than three will hardly ever be found together, and more
often only two. <A family-party usually consists of a cow, a calf,
and a yearling; very often the cow and the calf are alone, and
two bulls are frequently found together before the rutting-season
begins. In winter, however, they are said to be somewhat more
gregarious, but I have never heard of more than nine actually
being seen in company in the autumn.
In some Swedish forests, which are strictly preserved for shooting,
it is said, however, that the Elk associates in larger herds, and has
become so numerous that much damage is done to the forest by
their biting off the shoots and tops of the young Scotch pines; but
in such localities they do not attain the size and vigour that they
do where they have a wider range and a greater choice of food, as
in the two provinces where I have observed them.
The Elk is commonly supposed to be essentially an inhabitant
of forests, and though this is to some extent the case, and in winter
it no doubt almost invariably keeps to the shelter of the forest,
yet I have lately become convinced that it is found on the higher
fjelds to a much greater extent than is generally known. As
high as the birch is found, Elk may certainly be seen in summer ;
and the old solitary bulls in districts where open fjelds exist
rarely descend into the pine-forest until the rutting-season begins.
From 1500 to 2500 feet is a very common range in summer, and
on the high mountains of Upper Tydal I have seen tracks much
higher than this, far out in the open fjeld on the Swedish frontier.
And in North Trondhjem, where the timber-line is not so high, the
open hill-tops in some places are covered with Elk tracks and dung,
as though the animals had remained there during the whole of
the hot weather. Capt. Ferrand assures me that he has killed a
bull on the mountain above Lake Feemen at an elevation of at least
3700 feet, on regular reindeer-ground, where arctic willows were
1903.] THE ELK IN NORWAY. 139
the only shrubby plants. When found in such situations, they
may be stalked in exactly the same way as red deer, but when
disturbed usually make for the shelter of the forest ; and even on
the fjeld they nearly always lie in or close to a small grove of
dwart birch, and are in consequence not so easy to find with the
telescope as red deer.
The Elk is supposed to be monogamous, and I have never seen
more than one cow in company with a bull during the rutting-
season; but Iam assured by a Lapp, whose knowledge of their
habits is very great, that in districts where cows are more
numerous than bulls, the older bulls will change their mates two
or three times during the season. This seems to be confirmed by
the fact, which I have often noticed, that at the end of September,
when the rutting-season is approaching its height, cows are often
found without bulls, and the tracks of solitary bulls are found,
travelling presumably in search of fresh cows.
Col. Walker, of Tykillen, Co. Wexford, who has had great experi-
ence in Elk- hunting, relates the following event in a letter to me :—
“Thomas and I saw a magnificent bull with a splendid head in
company with a cow and calf ; they were quite in the open on the
edge of a small lake about half a mile from us.
“T noticed that whenever the bull went to the cow she ran at
him, and butted him very hard in the ribs; he then each time ran
from her, and browsed on the trees. Thomas then told me that
it would be useless to stalk this bull as he would be gone in a few
minutes. His story was that the bull only remains with the cow
for three days, and then she beats him off, and he has to go and find
another. Thomas had hardly done telling me this when the bull
made his last appeal to the cow. She gave him a rough reception,
and he at once started off at a fast trot. She remained grazing,
and we could see the bull going away very fast for nearly two miles
over a long stretch of open mountain, which took him across our
boundary. We never saw this bull again.”
Another proof of their polygamous habit is that on such ground
as that on which I hunted last year, where during eight years only
one cow has been killed for every five or six pale ‘and where in
consequence the cows were far more numerous, I found no more
barren cows than in districts hunted by Norwegians only, where
more cows than bulls had been killed.
A point upon which I have never been able to get any certain
information in Norway is whether the sexes attract each other
by calling at night, as the American Moose do. The art of calling
is unknown to Scandinavian hunters so far as I can learn, though
it is said to be practised in North Russia and Kurland. The
weather in Norway is usually so wet and stormy at the end of
September that there is no inducement to lie out in the forest ;
and though I have spent many nights in remotely situated huts
and seters, and have listened at dark and daylight, I have never
heard such a call asis made by the Moose. The cows do, however,
call to their calves, and I have heard bulls uttering a low grunting
140 MR. H. J. ELWES ON [ Feb. 3,
noise when alone, and apparently looking for cows. On this
point, however, | quote Capt. Ferrand’s experience, which is that
he has on several occasions attracted bulls whose whereabouts
he knew, especially on a still frosty evening, by imitating the
erunting of the bull, which can be heard when they are looking
for cows at a distance of from 500 to 1000 yards, or possibly
more. Moreover, he has heard the cows make a somewhat
similar noise, and the calves make a bleating noise when they
have lost their dam.
Col. Walker also writes to me on the subject of the cow calling
for the bull, as observed by him at Storvand, on the property of
Mr. Collett near Mo, in North Trondhjem, in 1890 :—
‘“One day my hunter and I came on the spor of a bull Elk and
cow ; after some time we looked over a small hill and saw, in the
valley below it, the bull and cow about 150 yards from us. Seeing
that he was paying her great attention I waited to see what would
- happen; after one or two attempts he jumped on her, they having
their backs tome: I aimed at him, but waited until he had finished
his performance ; as he came off her I fired, and shot him stone
dead. He rolled off on to the ground. She did not appear to
have noticed the shot or that he was dead, but began to graze
close to him, and once or twice went over and smelt him, then
grazed again. I then walked up to the bull; she did not seem to
mind me, only stared at me and stood close to the bull. When I
was within twenty yards of her she got alarmed and went off,
very slowly at first, constantly looking round for him to follow
her. Shortly afterwards, while we were cutting up the bull, we
heard a very loud harsh roaring noise, just like the noise made by
a badly wounded bear when pinned in a corner. Both my hunter
and I thought it was a bear. I then went to the top of a small
hill, and on looking over saw the cow in the middle of a large open
bog. She was roaring, I could see her quite plainly through my
glasses ; she then got wind of me and bolted off.
“Qn another occasion I had killed a bull which was in company
with a cow and two calves; these bolted, the cow one way, calves
the other. About 7.30 p.m., when I was in my hut, I heard a cow
calling just like a domestic animal in this country. I asked
Thomas whether we were near a farm; he then told me it was
the Elk cow calling for her calves.”
Though the scent and hearing of the Elk are unusually acute,
their sight seems to be by no means so quick as that of deer: if it
were, the difticulty of shooting them, which is already very great,
would be much increased. The usual method adopted in Norway is
to use a trained dog in a leash, which can scent the Elk at a distance
of a mile or more in the forest; but owing to the difficulty of
approaching without noise in the thick forest, and the cunning
which the Elk almost invariably display in lying down so as to
get the wind of anyone following their tracks, only those who have
great patience, caution, perseverance, and an intimate knowledge
of the habits of the Elk and the ground, are successful in
1903. ] THE ELK IN NORWAY. 141
getting more than a glimpse of a frightened Elk disappearing
among the trees. But if you can suc ceed in seeing the Elk before
he has heard or smelt you, you may take liberties with his eye-
sight which would not be possible in deerstalking. I once
approached a cow Elk, which was lying with her calf on an open
hillside, within 10 yards by keeping her body between me and
her head, and might have got nearer if the calf had not seen me.
On another occasion I rowed in a boat, just as it was getting
daylight, within 150 yards of a bull Elk, which had been lying on
the shore of a lake, and had not yet got up from his lair, and
shot him from the boat before he had t: iken alarm.
When bulls are rutting they sometimes run towards a man whom
they have heard but not seen, thinking that it is another Elk, and
in consequence stories are told of their attacking human beings,
But these stories will rarely bear investigation, and though there
are undoubted instances of wounded Elk attacking human beings,
I never but once saw one turn upon me, and then only because
he could not get away.
Besides the system of hunting them with a dog in a leash, they
are also hunted to some extent with loose dogs which bring them
to bay ; and though thissystem requires a man of great endurance
in order to follow fast enough to keep the hounds within hearing,
sometimes for many hours, it is perhaps more exciting and de: adly
than still hunting. When Elk are so disturbed by dogs and
have not seen, heard, or smelt man, they will sometimes come to
bay very soon, but more often run many miles, and always try
to throw the hounds and hunter out by crossing large rivers or
lakes. The Elk is an extremely strong and good swimmer, and
quite at home in the water, and though he usually crosses a large
river where it is shallow, he can, when pressed, pass a roaring
torrent, where no boat could live. In the very wet autumn of
1893 I lost several of the bulls I hunted, in consequence of their
crossing rivers which we could neither ford nor swim; when
severely wounded an Elk usually takes to water.
They may sometimes be driven successfully in places where the
ground is very steep or confined, but as a rule Elk do not follow
particular paths so much as most large animals, and can go up and
down very steep ravines and rocky places which would be thought
impossible for so large and heavy an animal. They usually feed
up wind and run down wind, but there is no rule about this, and
after a long hunt they sometimes come back nearly to the same
ground they started from.
On the whole, I consider Elk among the most difficult animals
to hunt that I have ever had experience of, and, even where they
are quite numerous, | have more than once spent ten days without
getting a fair chance at an animal worth shooting.
~ The bulls are said by some to shed their horns as early as
Christmas and not later than January, as Mr. Meade-Waldo tells
me they have done in our Gardens. Capt. Ferrand, however,
has been informed by woodcutters who were working all winter
[ Feb. 3,
MR. H. J. ELWES ON
142
(‘azis ‘qeu 4) “pojetouedap YONUL pey ase WOIF SUIOY SITY ynq
‘sprvMioye Svat dULOS JOIIJSIP 94} UL “AIO SIY} paliawo savy oF posoddns sva Worm Inq B pel[ty [—'elavuipuvog
Ul UWees T9Ae BABY YT ULOY ysostel oy, “JaSuTWI0g “FY Ag Aq pardnodo “44e][09 “zy Fo esnoy oy} Ul Mou pue ‘oy ye dn peyord us0y pegs Jo surov.y,
‘GG SG-9X9,
THE ELK IN NORWAY. 143
1903.]
“Unasn yy Torawrg ur eravnyywy MOF ALG OS.1v] B Jo susr0 zy
'€3 “SY-9xa7,
144 MR. H. J. ELWES ON | Feb. 3,
in the forest, that March is a more usual time, and this seems
more probable having regard to the time at which the horns of
other deer are shed. The new ones begin to grow in April
or May, and are fully developed about the middle of August. In
September the velvet is rubbed off against a young fir-tree,
which is usually destroyed in the process, and from the middle to
the end of September the rutting-season commences. At this
season the bulls have a very strong, rank, musky smell, and
scrape shallow round holes in the ground like stags’ wallows, which
retain the scent of their urine for some days. Occasionally the
bulls fight desperately, and I have seen places where the ground
was tor up over many yards and sprinkled with hair and blood,
but few hunters have had the good fortune to witness such an
encounter.
The development of the horns of the Elk in the Namsos district
of Norway is apparently greater than in South Norway and
Sweden, or in the districts which it still inhabits in Kast Prussia
and Kurland, though I cannot speak with the same certainty as
to Russia.
On my return from Siberia, four years ago, I saw semifossil horns
of the Elk from the district of Perm, which were larger than
any I have seen in Hurope; but on visiting the best collections I
could hear of in St. Petersburg, I saw none which appeared to me
better than the best Norwegian heads, or so good as the one
figured (text-fig. 23, p. 143), which is that of a Lithuanian Elk
in the Branicki Museum at Warsaw.
The bull calf has no horns the first year. The second year he
has a small spike on each side, sometimes, but rarely, two spikes,
and I have seen one on one side and two on the other. As he
grows older it is supposed that the number of points increase
annually, one on each side; but I think this is not at all invariably
the case, and that the size and strength of the individual has much
more influence than the age, and probably, as in the case of other
deer which shed their horns annually, food and climate have a
good deal to do with it. As a rule the horns inerease in size and
number of points up to 10 or 12 years old and possibly more, and
in old animals have a tendency to diminish in size and in the
length and strength of the points. From 18 to 22 points are
perhaps the average of adult males in Northern Norway, and 10
to 14 points in Southern Norway.
The widest span which I have ever seen is 54 inches (text-
fig. 18, p. 134), and the greatest number of points 16 on one side,
as on the shed horn of which I show a tracing (text-fig. 22, p. 142).
The breadth of the palm in this instance and the number of points
are quite exceptional, and I very much doubt whether there are
any such Elk now alive in Norway, as Herr Bruun of Trondhjem,
who has the opportunity of seeing all the finest specimens which
are procured, says that he has seen nothing equal to them.
Moose heads of as great or greater size are, however, often
killed, and though the largest which I have ever seen is 65 inches
1903. ] THE ELK IN NORWAY. 145
across, | have heard on fairly good authority of one over 6 feet in
span, (This was written before anything was known of the
gigantic heads found in Alaska.)
With regard to the comparative size and weight of the Elk and
Moose, I cannot speak positively, as it has never been possible to
weigh an entire animal in the places where I have killed them.
Mr. Abel Ch: apman, however, gives me the following weights of
a bull, of fair but not unusual size, killed by him on Sept. 1 15th :-—
Kilogrammes. Kilogrammes.
Head with skin of neck ...... 4() = 40
Shoullderssk cheat Secs 42 each = 84
Haunches with feet ............ 5A eS
Sidlesk nce eee eee ene A (ee)
IWiealke ermal eevee soosnacoocasonadec 60 = 60
Skeimy, Sanya saat easement ans 3 = 30
Motalleeee 402
This would make the weight of the living animal at, least
1000 pounds or over, as the intestines are enormous.
Capt. Fervand’s best bull, a 10-year old, weighed 1240 English
pounds without the intestines.
The pace of Elk, when undisturbed, is a slow walk, and their
movements are very deliberate, but they can trot for many
miles over boggy and rough eround at a pace of 6 or 8 miles an
hour, and oceasionally when much frightened break into a
lumbering canter.
The female Elk has her first calf at three years old in the month
of May, which makes the period of gestation about 7 months.
Usually she has only one, but not unfrequently two calves, which
erow very rapidly, and by the end of September are as big as a
red-deer hind. They suck thei dams until late in the winter,
and keep company with them until another calf is dropped, and
sometimes longer. They have occasionally been tamed in Sweden
and taught to go in harness, but owing to the difficulty of feeding
them they are not easy to keep in confinement. I have seen,
however, in the Zoological Gardens at Rotterdam a cow Moose
which is said to have been about 20 years in confinement, and
T hope that we shall be equally successful with the one which was
captured by Mr. Nickalls and presented by him to our Gardens.
Statistics of the number of Elk killed in Norway in the season
of 1894. (Translated from the ‘Tidsskrift Norsk. Jeger og
Fisker Forenings,’ Heft 3, 1895.)
Name of Amt. No. of Bulls. No. of Cows. Total.
Smaalenene............... 8 3 Tl
INEGTESINUE scabcoseuesscocsce 67 60 937/
Hedemarken ............ 74 a2, 146
Konishiansry. «cast saseemee 9() 87 177
Proc. Zoo. Soc.—1903, Vou. I. No. X. 10
146 MR. H. J. ELWES ON | Feb. 3,
Name of Amt. No. of Bulls. No. of Cows. Total.
Buskerud@eseses-ceeee 69 73 142
Jarlsberg og Larvik ... 9) 8 17
iBrabsbergesn erent. D4 48 102
INedemes! (o.-erccrisccee oes 3 7 10
Lister og Mandal ...... 1 — 1
Sondre Trondhjems ... 50 dO 100
Nordre Trondhjems ... 212 194 406
Nordland Reece 9 4 13
SSIEANENGVEREIO nqooogodonssane = a =
Sondre Bergenhus ...... == — —
Nordre Bergenhus ...... — — —
Romsdal eereeeries co: — — a=
IMM AKEM sonososcogoooce — — —
646 606 1252
Average for the 5 years, 1889-93. 610 512°6 1122-6
To give a better idea of the numbers of Elk found in what was
perhaps, at that time, the best forest in Norway, | give the
following statement copied from my diary in 1895. The number
actually seen may possibly represent from 3 to 3 of the number
actually on the ground at the time :—
Cows &
Bulls. yearlings. Calves.
Sept. 2. Haende ...... aS 1 Re Barren cow shot for meat.
3. Storen ...... ie 1 1
4. Strommen... oat 1
5. Strommen... 1 11 ai 1 bull killed: 18 points.
6. Blank day.
BMG ca tenons 1 1 1
Oeis Mo reescascee: sa 1 oe
10. Oplo ......... ae 1 1
11. Synnees ...... Bao 1 ie
TAS ie Were seopsaceae) eld aL 1
INS sen A ey oscene esence 11 1 bull killed: 16 points.
15. Blank day.
1G: Mo ssicse: 1 1 1
17. Storengen ... 66 11 a
18. Storengen ... 1 1 1
O° JE), sbsedanas a3 1 1
QO M0 aie eases aoe 11 1
21. To Storvand, blank day.
22. Vennevik ... 1 11 1 1 bull killed: 6 points.
23. Synnes...... 11 111 11
24. Brotten...... 11 fg ae 1 bull killed: 14 points.
25. Synnes ...... 1 1 1 1 bull killed: 13 points.
26. Quisten ...... 1 ee te 1 bull killed: 11 points.
Dae Oplowess-rae ae 1 1
29. Mo : 1 1 1 1 bull killed: 18 points.
Total ...... 17 25 14
This number of Elk was found on about 12 farms out of 24
over which I had the right of hunting. These farms might
average 10,000 acres more or less in extent.
On the same ground as this Col. Walker kept the following
record of the Elk which he saw himself during the four seasons
1887-1890, between Sept. Ist and Oct. 10th. It tends to show
a
1903. ] THE ELK IN NORWAY. 147
how rapidly Elk increase where they are protected and the cows
not killed :—
| |
Foe | Olde Monn lyre een gee
Year. Balle Balle Cows. Calves.
| 1887.} 3 —_ 9 10 | Two of these cows had two calves each.
1888.| 7 2 8 7 | One of these cows had two calves.
1889.| 6 6 17 | 14 | Six of these cows had two calves each.
1890.| 5 10 18 20 | Nine of these cows had two calves each.
Most of these observations were written at the request of
our late President, Sir W. Flower, six years ago, but were
withheld from publication until this year, when the question
of a new species of Elk existing in Siberia was raised by Mr.
Lydekker and the Hon. W. Rothschild. I then added the following
appendix.
Text-fig. 24.
Horns of a very old Elk showing degeneration. (About 51; nat. size.)
APPENDIX.—ON THE VARIATION OF THE ELK.
Dr. Lénnberg’s paper on the variation of the Elk appears to
me to confirm the opinion I expressed at a recent meeting of the
Society (P. Z. 8. 1902, vol. ii. p. 144), that the Siberian Elk
described by Mr. Lydekker (P. Z. 8. 1902, vol. i. p. 207) as Alces
bedfordie was nothing more than an inconstant variety of the
European Elk. The Hon. Walter Rothschild has since expressed
a contrary opinion, for which, however, I am not aware that he
has any further evidence than the fact that other horns similar
10*
148 ' MR. H J. ELWES ON [ Feb. 3,
to those of the so-called A. bedfordie have been received by him
from Siberia. I stated that the only horns of the Elk which I
succeeded in obtaining in the Altai Mountains in 1898, which
are now in the St. Petersburg Museum, were palmated in pre-
cisely the same manner as those of the Norwegian Elk, and others
which I have seen in Russian collections had all well-developed
palmation. Dr. Linnberg’s figures go to show that the develop-
ment of the antlers in those parts of Sweden from which they
come is nothing like so fine as in the districts of North and South
Text-fig. 25.
Tracing of a cast horn found at Solem in Bangdal by Capt. Ferrand.
(About =; nat. size.)
Trondhjem, where most of my hunting has been done. When I
first began Elk-hunting in Tydal, a mountain valley in South
Trondhjems-amt running up to the Swedish frontier, I should have
considered the horns figured by Lénnberg in fig. 3 as a fair repre-
sentation of an adult Elk in that district; but I have seen them with
as many as eleven points on each horn, and believe that horns with
even more points have been obtained. Further north, in North
1903. ] THE ELK IN NORWAY, 149
Trondhjems-amt, where the Elk a few years ago was extremely
numerous, much larger heads occurred, owing, I believe, to the much
greater quantity of mountain-ash, which seems to be the favourite
winter food of the Elk, and which has been to a great extent
destroyed by the vast number of animals constantly devouring it.
Out of the whole number of bulls I have killed, about twenty,
only four or five had horns of the type shown in Liénnberg’s
Text-fig. 26.
One-horned Elk in Ipswich Museum.
figures 1, 4, 6, and 9; and all of these, with one exception,
appeared to be quite young animals.
Among the hunters who accompanied me in different years
were a Swede, a Norwegian, and a Lapp, all of whom knew the
Elk of those districts most intimately, and were considered the
most experienced hunters of their districts. None of them ever
even suggested the possibility of two races of Elk existing ; and I
150 ON THE ELK IN NORWAY. [ Feb. 3,
attribute non-palmation of the horns entirely to imperfect
development of the animal, caused by insufficient food in their
earlier years, or degeneration caused by old age, wounds, or other-
wise. We did not reckon an Elk to be adult until he had at
least seven points on each horn, and the oldest and largest bull
that I ever killed, which had been well known in the district, and
continually hunted for at least ten years, had, when I killed him,
well-palmated horns of about one-half the size of what they ought
to have been in an adult animal of his size, showing that in the
Elk, as in the red deer, the horns degenerate in size and number
of points in old age, which may be fifteen or twenty years or less.
Two such instances of degeneration are figured. Text-fig. 24,
p- 147, is the head of an old Elk killed by Thomas Bate, Hsq., in
Lurudal Namdalen on Sept. 25, 1890. The horns measure 43 inches
in expanse, with ten points on one side and eleven on the other ;
the points are, however, not arranged in a uniform series on the
edge of the palm, and the development of the brow-antlers is very
abnormal.
The shed horns of what was probably the same animal were
picked up on the same ground in the year previous by Col. Sullivan,
and are of the same type with the same number of points.
An Elk displaying still more remarkable abnormal degeneration
was killed by Capt. Ferrand, and is shown in text-fig. 26, p. 149.
This animal was supposed to be 25 years old or more, and
the incidents of his death have been most graphically described in
the ‘ Badminton Magazine’ for March 1901 by Capt. Ferrand. The
horns are now in the Ipswich Museum.
The largest horns I have seen from Norway, belonging to an
animal which I unsuccessfully hunted for many days, but which
was afterwards killed by a farmer, and sold to me by Mr. Bruun
of 'Trondhjem, were 54 inches in width, with nine points on each
side (see text-fig. 18, p. 134); but there is a pair of shed horns in
Sir Henry Pottinger’s house at Mo, of one of which I send a tracing
(see text-fig. 22, p. 142), showing sixteen points on each side. It
is well known in Norway that the Elk of the southern districts,
which are much more fully timbered, and where there is nothing
like the same extent of open fell and good feed as in North
and South Trondhjem, and where both sexes are much more
constantly hunted and the calves frequently deprived of their
mother’s milk in September, do not, in modern times at least,
produce anything like such fine heads as those of the wilder
districts of the north, the conditions being probably very similar to
those deseribed by Dr. Lonnberg as in the southern provinces
of Sweden’. Taking his nine figures, I should be inclined to say
that all except 2 and 3 might, if they had come from Siberia,
have been considered as belonging to Alces bedfordiw ; and form,
to my mind, ample proof of that being (if the horns belong to
1 Mr. Percy Godman informs me that a well-known Elk-hunter in South Norway
considers that on his property Elk have diminished by two-thirds in the last ten
years, and attributes this decrease to their having eaten and destroyed all the “ leaf-
trees,” z.e. willow, mountain-ash, and aspen.
7
i if
ay iH
‘
My
'
dea/Aays) a enOSe weil J, Jed. IN),
950°
OG
10.
Pickard €ambridge del et lith. West, Newman imp,
SPIDERS OF THE FAMILIES PISAURIDA| AND SENOCULIDA®,
Piso, 1903-0 1a aye
Newman ump.
est,
Wi
i Pickard..Cambridge del. ec lith.
OF THE FAMILY PISAURIDAS.
SPIDERS
1903. | MR. F. PICKARD-CAMBRIDGE ON NEW SPIDERS. 151
adult animals) nothing more than a local variety produced by com-
paratively unfavourable conditions. In considering the probability
of the existence of local races of Elk, it must be remembered
that no northern mammal, except perhaps the reindeer, has such
wandering habits; and though a race might become temporarily
isolated in an area which was entirely surrounded by a large
extent of country providing no suitable food, yet there is no
doubt, in my mind, that when their favourite food had become
partially exhausted, they would migrate many hundreds of miles,
and thus prevent the establishment of local races.
A very interesting paper on the former existence of the Elk in
the Thames Valley in England, with a plate showing the great
similarity of its horns to those now existing, is published by
Mr. E. T. Newton, F.R.S., in the Quarterly Journal of the
Geological Society, vol. lix. (1903), and I am much indebted to
that gentleman for sending it to me.
February 17, 1903.
Dr. Henry Woopwarp, F.R.S8., Vice-President,
in the Chair.
Mr. R. EK. Holding exhibited and made remarks upon an adult
skull of a Collie Dog, indicating a displacement of the incisors
caused by the closing of the left lower canine upon the third
incisor of upper jaw, and which apparently retarded the develop-
ment of the second incisor of that side. The skull also showed a
supernumerary canine which could in no way be mistaken for a
retained milk-canine.
Mr. Holding also exhibited portions of three Rabbits’ skulls
having ‘“ over-shot” incisor teeth. In one specimen the abnor-
mality had caused a deviation from the median line of the anterior
portion of the skull; and in another the incisors, after leaving the
premaxilla, had formed a complete circle and grown into the palate,
causing starvation and death of the animal.
The following papers were read :—
1. On some new Species of Spiders belonging to the
Families Pisauride and Senoculide ; with Characters of
a new Genus. By Freperick PicKARD-CAMBRIDGE,
B.A., F.Z:S.
[ Received January 10, 1903. ]
(Plates XIV. & XV.")
While working out the Spiders of the two families Pisauride
and Senoculide of Central America for the ‘ Biologia,’ oppor-
tunity was kindly offered by Mr. R. I. Pocock for an examination
of the material in the British Museum Collection, with the result
1 For explanation of the Plates, see p. 168.
152 MR. F. PICKARD-CAMBRIDGE ON NEW SPIDERS. | Feb. 17,
that a new genus and eight new species were found, several of
them occurring amongst those collected by myself on the Lower
Amazons in 1895-6.
The Spiders belonging to the Pisawride are interesting from
the fact that they run freely and with great rapidity over the
surface of the water, even in astrong current. One of the genera,
Dolomedes, and another, Thalassius, have both been credited with
capturing and devouring small fish of various kinds; and of the
latter genus Mr. A. N. Stenning, himself formerly a gamekeeper,
whose observations are likely to be trustworthy, declares that he
has found members actually devouring the small fry of a variety
of trout which occurs in South Africa, some of the culprits bemg
now in the Museum Collection. Other notes on the habits of
these interesting Spiders will be found under the genus 7'rechalea.
Species described or figured below.
Thaumasia velow KW. Simon, @, p. 154, Pl. XIV. fig: 1.
a annulipes, sp. nov., 2, p. 154, Pl. XIV. fig. 2.
Dossenus marginatus E. Simon, @ 5 [Oe 155, Pl. XIV. figs. 3-5.
Paradossenus nigricans, sp. nov., &, p. 155, Jel, SD figs. 6-9),
Thanatidius spinipes, sp. nov., Q, p- 156, PL. XIV. figs. 1O=12.
Senoculus parallelus KE. Simon, 2, p. 166, Pl. XIV. fig. 13
es albidus, sp. nov., 2, p. LOSS PEP Xan; fies 142
Trechalea longitar sis C. L. Koch, GO Qo jo MOO, IA, Nave figs. 13
‘ keyserlingt, sp. nov., ° , p. 163, PL XV ericslee 2:
5 urinator K. Simon, ¢ Q, p. 161, Pl. XV. figs. 3-5 a.
we ellacombei, sp. nov., 2, p. 161, Pl. XV. fig. 6.
macconnells Poc., 3, p. 162, Pl. XV. figs. 7, 8.
connexa O. P.-Cambr., 3, p. 162, Pl. XV. figs. 9, 10.
- extensa O. P.-Cambr., 3, p. 162, Pl. XV. figs. 11, 12.
amazonicd, Sp. NOV., 3 ©. im 163, PEP XaVeniest 18- 20.
Hes nyt us palustris KW. Simon, 3 @, p. 165, Pl. XV. figs, 22-25.
» habilis O. P.- Cami... Es [Oo WOE, LI, LOWY, tee, Al
Fam. PISAURID..
Synopsis of Genera dealt with below.
A. Tibia i. with four (2-2-2-2) or five (2-2-2-2-9)
paired spines beneath, the last pair very small and
apical. Protarsi 1. with three or four pairs beneath.
A!, Tibia i. with four pairs of spines beneath. Protarsi
i-iv. with a single small central apical spine beneath.
Sternum produced into a long narrow conical point
between coxe Iv.
1. Tarsi shorter, not flexible. Protarsi i. with three
(2-2-2) paired spines beneath.
aa. Lower margin of fang-groove with 3 teeth.
a. Tibial and protarsal spies comparatively short,
only two or three times as long as the diameter
of the segment.
* Anterior row of eyes wider than that formed
by the posterior centrals. Central posterior
eyes nearer to each other than to the
laterals.
a), Teeth on lower margin of fang-groove not
equidistant, the 3rd more remote............__ Thawmasia Perty.
1903. ] MR. F. PICKARD-CAMBRIDGE ON NEW SPIDERS. 153
b!. Teeth on lower margin of fang-groove
GTAP eco Asdare oammes nonendeaabeonoe Tinus ¥. P.-Cambr.
** Anterior row of eyes not wider than that
formed by the posterior centrals. Cen-
tral posterior eyes much further apart
than each is from the lateral ............. Hana O. P.-Camby.
b. Tibial and protarsal spines comparativel
very long, the longest being quite two
thirds the length of the whole segment... Dossenus Ki. Simon.
bb. Lower margin of fang-groove with 4 teeth,
HEN SILAAEILID cagcosmsncdmearito eon onSe0s600§06009
ii. Tarsi long and flexible. Protarsi i. with four
(2-2-2-2) paired spines beneath.
a*, Legs subequal in pairs, 4—2 and 1—3. Man-
dibles conspicuously gibbous above, glabrous,
more distinctly carinate on the outer side
(males). Lower margin of fang-groove with
b*. Legs unequal, 4, 2, 1, 3. Mandibles not
conspicuously gibbous above, hairy, less
carinate (males). Lower margin of fang-
groove with 3, 4, or 5 teeth ..................... Trechalea Thorell.
B!. Tibia i. with five pairs of spines beneath. Pro-
tarsi i-iv. with two apical spines beneath.
Sternum very broad between cox iv., not
1 VOTH TADS TEP FD] | LO)D ITs dao opsmod apnmanEbe donc odeceReSooaEsToe ? Thanatidius 1. Simon.
B. Tibia i. with 9-10 pairs of spines beneath, the last
pair not apical. Protarsi i. with 8 or 9 pairs of
spines beneath .......0..0.eccccceccccscseucectsreesreeseese Syntrechalea ¥. P.-Cambr.
Parvadossenus, gen, nov.
Hesydrus Simon.
If the above genera be separated by the perhaps more easily
observed character furnished by the eye-formula, then the table
would run as follows :-—
A. Anterior row of eyes not or only a little wider than the posterior
central row. CTR ith
a. Anterior vow of eyes not wider than the posterior central row... 5 ;
0 Dossenus.
b. Anterior row of eyes a little, but distinctly, wider than the
posterior central row ...... Paradossenus. Tinus. Thaumasia. Hesydrus.
Trechalea. Syntrechalea.
B. Anterior row of eyes much wider than the posterior central row,
as wide as the posterior lateral row ................:0eeeeeeeeeeeeeeeees Thanatidius.
Nore.—(1) The number of protarsal spines in some species of Trechalea ave
difficult to locate, and the character is thus rendered less valuable.
(2) One cannot be quite certain that Thanatidius has been correctly identified.
(3) Tinus is closely allied to Drances Simon, but the latter has, according to this
author, no apical pair of spines beneath tibia i. and ii., while they are present in the
former. The name Drauces (not Drances) was used by Champion for Coleoptera
in 1889.
(4) In the Biologia Centr.-Amer., Arach. Aran. ii. p. 312, I have referred the
species Hesydrus jullient Simon to Enna O. P.-Cambr. on the ground that they
have the characters, the tarsi flexible and central posterior eyes 2 diameters apart, in
common. This genus therefore comprises Simon’s Hesydrus Sect. 2a, while
Hesydrus, as here diagnosed, comprises his Hesydius Sect. 1 a.
Genus TuAumAsiA Perty.
Del. Anim. Art. Bras. p. 192 (1833). Type, 7’. senilis Perty.
Species recorded.
I ME RSeTUIS: POY y OC MEL amr. ce ou cicces cn sere. ee Nae ta Brazil.
2. 7’. marginella (C. L. Koch) ..... . (Sub Dolomedes.)
154 MR. F, PICKARD-CAMBRIDGE ON NEW SPIDERS. [ Feb. 17,
By MS SGC MEUG (18, SIMON) oseosccgsees (Sub Saltwinus.)
4. ?7. scapularis (C. L. Koch)...... (Sub Dolomedes.)
5. ? 7. binotata (C. i. Koch)......... (Sub Dolomedes.)
6. Thawmasia velow Simon, Ann. Soc. Ent. Belg. xli. p. 18
fie iol) Rac. san ope Mon aaa nH aRrA ane RAM ohanadhuod as Brazil.
THAUMASIA VELOX Simon. (Plate XIV. fig. 1.)
Loe. cit. Type ¢, in coll. EK. Simon. Total length 8 mm.
Thaumasia velox F. P.-Cambr. Biol. Centr.-Amer., Arach, Aran.
u. p. 309, t. 30. figs. 5, 5 a-c (¢).
The type of this species (¢) was taken on the Amazons, and,
short of first-hand evidence, I do not doubt that the male and
female, taken by myself amongst the foliage on the margin of a
small “‘ furo” in the Parana Buyassu, near Breves, Lower Amazons,
must be referred to this species.
The male identified as belonging to this species from Guate-
mala may possibly be not identical, but it is impossible to be
certain with but one example from each locality.
The female has not been before described, but a figure of the
vulva is given on the Plate. For figures of the male see Biol.
Centr.-Amer. loc. cit. above.
Hab. LowErR AmAzons: Santarem; Breves, Parana Buyassu
(fF. P.-C., 1895-96); Rio Tocantins, 8. Paulo de Olivenga (de
Mathan); Guatemala (Sarg).
7. THAUMASIA ANNULIPES, sp. nov., 9. (Plate XIV. fig. 2.
Type 2, in coll. Brit. Mus. Total length 5°5 mm.
2 .—Colowr. Carapace deep brown, with pale yellow narrow
central band, slightly dilate behind eyes. Abdomen brown,
shoulders and anterior lateral portion deep black. Legs pale
yellow, femora annulated with dusky brown.
Structure. Carapace horizontal above, convex and abruptly
deflected behind. yes: posterior row recurved, subequal, equi-
distant; laterals on a stout black tubercle. Anterior row straight
or slightly procurved, wider than width occupied by posterior cen-
trals. Anterior eyes equidistant, centrals larger. Clypeus more
than three times the height of the diameter of anterior centrals.
Mandibles long and stout. Fang-groove with three denticles on both
margins. Sternwm cordiform, posterior angle attenuate, produced,
its apex extending as far as the pedicle. Cox of pedipalp long,
straight, slightly dilate and rounded at apex. Labiwm half the
length of maxille, broad, squarely truncate at apex. Legs
4, 1, 2, 3; iii. the smallest, extending as far as apex of tibia iv.
Tibia 1. and 11. with 2—2—2 spines beneath ; 1—1 lateral on apical
half ; 1—1 at base and apex above; 2 at apex beneath. Pro-
tarsus 1. and 11. with 2-2-2 spines beneath and | at apex; 1—1
on each side; none above. For vulva, see Pl. XIV. fig. 2.
Several females of this small Spider were taken by myself in the
neighbourhood of Manaos, Lower Amazons, 1895-6.
1903. ] MR. F, PICKARD-CAMBRIDGE ON NEW SPIDERS. 155
Genus Dossenus Simon.
Hist. Nat. Ar. 2, i. p. 314. Type, D. marginatus EK. Simon.
Lower Amazons.
DossENUS MARGINATUS E. Simon. (Plate XIV. figs. 3-5.
ta)
Ann, Soc. Ent. Belg. xlii. 1898, p. 19. Type 9, and androtype
¢ in coll. EK. Simon. Total length, 9, 7-7 mm.
Two adult females of a Spider, which I feel pretty certain must
be referred to this species, were taken by myself at Breves and
Para, and are now in the British Museum Collection. The spines
on the legs are very long in comparison with those of allied
genera.
Hab. Isu. Trinrpap; Braziu: Amazons; Matto Grosso ((rer-
main); Lower Amazons, Para, Breves (Parana Buyassu, /. P.-C.,
1895-96).
PARADOSSENUS, gen. nov.
Type, P. nagricans, sp. n. Lower Amazons.
Lower margin of fang-groove with 4 teeth, the 5rd being
smaller. ‘Tarsi short, straight, not flexible. Protarsi 1. with
3 paired spines beneath; protarsi of all four pairs with a single
canta apical spine beneath. Tibia i. with four paired spines
beneath, the last pair apical. Sternum produced into a conical
point between cox lv. Spines on legs short, two to three times
the diameter of the segment. Posterior row of eyes recurved ;
eyes subequal, centrals one diameter apart, two and a half from
the laterals, which are set on a low tubercle. Anterior row
straight or very slightly procurved, slightly wider than the central
posterior row. Clypeus one and a half times the height of the
diameter of an anterior central eye.
Legs 1, 2, 4, 3—i1. very short, extending as far as two-thirds
of tibia iv.
PARADOSSENUS NIGRICANS, sp. nov. (Plate XIV. figs. 6-9.)
Type ¢, 8 mm.; gynetype 9, 85 mm.—carap. 3°75 mm.
Tn coll. Brit. Mus.
3 2 .—Colouwr. Carapace dull yellow-brown, with two longi-
tudinal dark brown bars on each side of the pale central line, the
whole being almost entirely clothed with yellow-grey scale-like
hairs. Margins of carapace reticulated with fine dusky lines.
The anterior portion of the brown bars presents within their
margin a narrow elongate accent-like pale mark. Abdomen pale
yellow ; ; dorsal avea with a broad longitudinal dark brown (some-
times paler) foliated band, becoming narrowed towards the
spinners, from the centre of whose lateral margin, on both sides,
issues an oblique interrupted lateral bar. The centre of the
foliated band is occupied by a pale lanceolate band, much attenu-
ated towards the spinners, its basal portion again occupied by a
broad lanceolate dark band, scarcely reaching the centre of the
156 MR. F. PICKARD-CAMBRIDGE ON NEW SPIDERS. [ Feb. 17,
abdomen. Lateral area mottled with dusky brown; ventral
surface pale yellow, unicolorous. Legs entirely yellow: 1. and i.
very faintly annulated and speckled with dusky brown, ili. and iv.
annulated, in some cases very richly, with black or brown.
Mandibles ved-yellow, with a large brown patch on each in front.
N.B.—These colours are very variable and in some cases the
central dark band within the inner pale band on the abdomen
is itself pale.
The vulva consists of two parallel plates, which have a promi-
nent angle on their inner margin. Between these, in front, lies
a deep transverse semicircular depression, and a convex central
piece, with a few shallow longitudinal grooves, is situated between
the plates.
For figures of tibial spur of male palpyus and the vulva, see
Plate XIV. figs. 7, 9.
These spiders are very abundant on the Lower Amazons, racing
over the surface of the water with legs extended and body flat,
and are exceedingly difficult to catch. None of the females had
egg-cocoons with them, but they probably carry them in the same
position as Z'rechalea, for in habits and general appearance these
spiders are very similar to those belonging to the latter genus.
Hab. Lowrr Amazons: Breves, Parana Buyassu (/. P.-C.,
1895-6).
Genus THanatipius EK. Simon.
Hist. Nat. Ar. 2, ii. p. 293. Type, 7. dubiws (Hentz).
N. America.
The following three species are referred to this genus by
Simon :—
1. 7. dubius (Hentz), Bost. J. N. H. v. p. 449 (1847).
(Sub Zhomisus.)
he TUE ueeqooIs (Valentws)), Oe, Gittis oho ocassosacc09 080007 (Sub Zhomisus.)
3. 7’. undulatus (Keys.), Verh. z.-b. Ges. Wien, p. 486 (1887).
(Sub Zetragonophthalma.)
4, THANATIDIUS SPINIPES, sp. nov. (Plate XIV. figs. 10-12.)
Type @ in coll. Brit. Mus. Total length 6 mm.
Hab. Parana Buyassu, Breves, Lower Amazons.
? .—Colour. Carapace, mandibles, sternum, and legs bright,
unicolorous orange. Abdomen olive-green, with a central band
of cretaceous-white pigment-cells.
Structure. Carapace nearly circular, ocular region produced,
parallel-sided, slightly raised. Thoracic region gibbous, abruptly
deflected behind. Hyes: posterior row strongly recurved, eyes
subequal, less than one diameter apart, equidistant ; laterals on a
stout conical tubercle. Anterior row strongly procurved; eye-
tubercles subequal, less than one diameter apart, equidistant, much
wider than the width occupied by lateral posteriors. Clypeus,
in centre as high as four diameters of anterior central eyes.
1903. | MR. F, PICKARD-CAMBRIDGE ON NEW SPIDERS. 157
Mandibles long, slender, more than twice as long as height of
clypeus. Mang-groove with three denticules on lower margin,
Maxille long, parallel, dilate at apex; labiuim broad, half the
length of maxille, truncate at apex.
Sternum longer than broad, cordiform, produced posteriorly
between coxie iv. into a broad rounded plate, the cox being their
own diameter apart. Legs 1,4,2,3. Femora with two long
dorsal spines and several others on the sides and apex. Femur i.
with two long spines in front in apical half. Tibia i. and ii. with
2—2-2-2-2 long spines beneath, the longest (the basal) being half
as long as the segment itself, cial lateral spines; 1—1 dorsal.
Protarsus i. and ii, with 2-2—-2—2 long spines beneath (basal pair
longest): 1—1 lateral, 1—] ae Spines on tibia and pro-
tarsus lil, and iv. numerous, but less regular. Patellee 1-4 with
a single long apical spine. Scopula absent. Tarsal claws 3,
superiors with three or four denticules beneath. Spinners 6,
posteriors longest. Pedipalp with large tarsal claw, having 3
very long denticules beneath towards base. Vulva, see Pl. XIV.
fig. 12.
Hab. Asingle 9 from the Parana Buyassu, near Breves, Lower
Amazons.
Genus TrecHaLeA Thorell, 1869 (nom. nov. for Vriclaria).
Type, 7. longitarsis (C. L. Koch), Colombia. Sub 7rielaria
(1848), nom. preeoce. by Wagler for Aves (1832).
The following characters are common to all the species men-
tioned below :—Teeth on upper margin of fang-groove 3, on the
lower, 3, 4, or 5. =Tarsi long and feible! Tibia i. with four
pairs of spines beneath, the last pair small and apical, besides two
lateral spines. Protarsii. with four pairs of spines beneath (often
irregularly situated); protarsi of all four pairs with a small
central apical spine beneath. Anterior row of eyes slightly
recurved, extending laterally slightly beyond the posterior
eentrals. Lateral anteriors smaller than the central anteriors.
Legs variable in length, 4, 2, 1, 3. Clypeus variable in height,
porrect, and the eyes also variable in their relative positions.
As regards the type-species of the genus, two distinct forms
have been identified as longitarsis of Koch—one by Keyserling,
now in coll. Brit. Mus., a female, having 5 teeth; the other by
Simon (Ann. Soc. Ent. Belg. xlii. p. 20, 1898), male and female,
having 3 teeth. Since they cannot both be the true longitarsis,
I describe Keyserling’s identification as a new species, for it is
totally distinct from examples from Colombia, now before me,
whence the type of the genus originally came. It forms, indeed,
the type of a distinct group, so entirely different is the form of
the vulva from that of the typical Trechalea from Colombia.
At present, however, although the identity of 7’. longitarsis
cannot be settled ial absolute certainty until an examination
of the type-specimen (if it still exists in the Imperial Museum at
158 MR. F, PICKARD-CAMBRIDGE ON NEW SPIDERS. [| Feb. 17,
Berlin) has been made, the examples from Colombia are regarded
and here described as 7’. longitarsis C. L. Koch. All of the
species described and recorded below seem to be good and distinct,
though of course much more material from all parts is required
for comparison and the confirmation of characters.
The habits of these spiders are very interesting, for although
they are not amphibious in the true sense of the term, as are
Argyroneta and Desis, they are quite at home on the water, and
speed along over the surface with great ease and rapidity, the
hairy clothing of the legs being waterproof. When racing away
over the water, they are not easy to distinguish from the large
Hemipterons (probably a species of Ranatra) of similar habit,
when the two creatures happen to be together in the same locality.
On being pursued over the water, the spiders will rush up the sides
of the boat and take refuge under the boards, but I have never
observed them dive below the surface as do Pirata, Dolomedes,
and some other Pisauridee. Whether, as has been reported of
certain species of Dolomedes and Thalassius, the spiders actually
prey upon small fish of various kinds, I cannot say from actual
experience, though the long flexible legs and exceedingly sharp
claws are well adapted for the pursuit and capture of such prey
in their native element. J may add that McCook’s records of the
fish-catching capacity of Dolomedes (Amer. Spid. vol. i. p. 236 and
ii. p. 66) has recently received confirmation by Mr. A. N. Stenning
in South Africa. He tells us that 7’alassius, a genus represent-
ing Dolomedes in the Ethiopian Region, has been often observed
by himself in the act of devouring the small fry of a species of
trout, and calls the attention of pisciculturists in those regions to
the fact, and begs them to keep an eye on these spiders.
On a swampy island in the large largo opposite Santarem on
the Amazons, I have taken examples of one beautiful species
squatting flat, like Sparassids, on the trunks of the trees, where
their hoary-white hairs and mottled legs and body afforded them
excellent protective colouring. Nor do they resemble Heteropods
merely when at rest, for when disturbed they dash round to the
opposite side of the tree-trunk with all the rapidity of a Selenops.
The egg-cocoon, as in many Pisaurids, is carried by the female
attached to the central pair of spinners at the tail-end of the
abdomen ; and it is curious to note how, when these females stop
suddenly in headlong flight, the weight of the cocoon swings them
round on the water, and carries them along backwards for some
distance.
The species recorded and described below may be recognized by
the following characters :—
Males.
A. Lower margin of fang-groove with 4 teeth, the third
smaller. Legs unicolorous ................................. wrinator BE. Simon.
1903. | MR. F, PICKARD-CAMBRIDGE ON NEW
B. Lower margin of fang-groove with 3 subequal teeth.
Legs more or less spotted or annulated.
I Legs longer in proportion, leg iv. being about 8
ene longer than the carapace.
. Third tooth on lower margin of fang-groove more
anne from the others.
. Tibia of pedipalp from one-half to twice longer
than broad; tibial spur distinctly unculate at
apex, not concave on inner side. Embolus of
bulb with a long conspicuous spur at its base.
. Tibia of pedipalp not longer than broad; tibial
spur not unculate at ‘apex, concave on inner
side, the inner margin having a prominent
angle. Kmbolus of bulb without any long spur,
but having two chitinous ridges .. “a
b. Third tooth on lower margin of fang- aroove not
remote, the 3 teeth almost equidistant.
a2. Tibial spur not uneulate at the apex.
Embolus of bulb without any long spur ......
b2, Tibial spur unculate at its apex. Embolus
of bulb with a long spur at its base :
II. Legs shorter in proportion, leg iv. being 5-6 times
longer than the carapace .............0.00c00e0eeeeeeeeeee
SPIDERS. 159
longitarsis C. Li. Koch
(sec. F. P.-C.).
macconnelli Pocock.
connexa O. P.-Cambr.
extensa O. P.-Cambr.
amazonica, sp. 0.
An alternative table for Males.
A. Embolus of bulb with a long conspicuous erect black
spur in its basal cavity.
a. Lower margin of fang-groove with 4 teeth
Lower margin of fang-groove with 3 teeth.
a!. Central posterior eyes one diameter apart ; apical
portion of embolus shorter, more distinctly
flanged on both sides; 3rd tooth more remote ;
tibial spur less broad and flattened..................
. Central posterior eyes less than half a diameter
apart; apical portion of embolus longer, less
conspicuously flanged on the lower margin; 3rd
tooth not remote; tibial spur broader and more
flattened .
B. Embolus of bulb naihonta any erect "iba spur in its
basal cavity.
a*, Tibial spur straight, blunt at apex.
a, Tibial spur forming a conspicuous angle on its
inner margin. ‘looth 3 remote from the others ;
apex of embolus less broadly pacer on its upper
margin .
. Tibial spur without ‘angle on inner margin.
Teeth equidistant; apex of embolus more lose
flanged on its upper margin ............
. Tibial spur strongly curved, pointed at apex
Females.
A. Lower margin of fang-groove with 5 teeth, the 4th
very small, situated on the inner side between the
3rd and 5th. Vulva without any central tongue-like
sclerite, but with two small blunt cusps in the middle
of the anterior margin, and an oblique elongate
lateral lobe on each side, converging behind the
cusps 55 daa bon Ee: eat ROR Cac eB Cae
B. Lower margin] of fang-groove with 3 (or 4) teeth.
Vulva with a well-detined central tongue-like sclerite.
a. Lower margin of fang-groove with 4 teeth, the 3rd
smaller
urinator.
longitarsis.
extensa.
macconnelli.
connexa.
amazonica.
heyserlingi, sp. 0
urinatory BE. Simon.
160 MR. F. PICKARD-CAMBRIDGE ON NEW SPIDHRS. _[ Feb. 17,
}. Lower margin of fang-groove with 3 subequal teeth.
a, Size much larger, 25 mm. Leg iv. at least ten
times longer than the carapace. Third tooth on
lower margin of the fang-groove slightly more
remote from the second than the latter is from
the first.
a*. Central tongue-like sclerite much narrower,
dumb-bell-shaped, about pally dilate at each
end . vesseees. €llacombei, sp. n.
. Central | ‘tongue- like sclerite much ‘broader,
almost as broad in the middle as posteriorly,
much broader anteriorly but somewhat vari
Plt) (aiernctsresnarcoacauacc Lona ae eee ee auee RAR UeEec aASps longitarsis C. Li. Koch.
61, Size much smaller, 11 mm. leg iv. not more
than 5-6 times longer than the carapace. Teeth
on lower margin of fang-groove equidistant.
Central cavity of vulva long and narrow, straight
on the imner margins; central tongue elongate,
not dilate anteriorly, but narrowed.................. amazonica, sp. Nn.
1. TrecHALEA Lonerrarsis (C. L. Koch). (Plate XV. figs. 13-17.)
Die Arach. xv. p. 65, t. 522. fig. 1462 (sub 7’riclaria).
? Trechalea longitarsis K. Simon, Ann. Soc. Ent. Belg. xlu.
1898, p. 20.
Type 2, in coll. Mus. Imper. Berlin. Hab. Colombia.
Although the identity of this species cannot be satisfactorily
settled without reference to the type-specimen, if it still exists,
T give a description and figures of the characters of a form with
annulated legs which was taken in Colombia. If the type no
longer exists, and no more exact locality than ‘“ Colombia” can
be ascertained for the type- specimen, then the examples here
described stand as “ ¢opotypes” —examples from the same locality
in which the type was taken—and will serve as a standard of
comparison for the species in the place of the original type.
Had Simon given more definite characters (for the majority of
the species of 7’rechalea possess 3 teeth) which would enable us to
recognize his identification of 7’. longitarsis with any certainty,
and if these (for he makes no mention of the locality) were taken
in Colombia, then his examples would be the recognized “ topo-
types” to which we should have to refer for an identification of
T. longitarsis (C. Li. Koch).
Present identification of 7’. longitarsis (sec. F. P.-Cambr.).
Total length, ¢ 20, 9 25 mm.—®. Leg i. 61; 11.68; 11. 55;
Tio (CH ummm, Wile, i, Wiss wy, Geer, Leow, in, All aman,
eee in coll. Brit. Mus. from Colombia.
3 2 .—Colowr. Carapace rich brown-black, with a pale orange
A-shaped spot behind the eyes, having a central dusky line, and
one on each side; a pale rufous-white, narrow, submarginal
band, and a narrow black velvety marginal band. Abdomen
brown, with a black central foliated band: a more exact description
is impossible from dried and wrinkled examples. Sternum and
ventral area dull yellow-brown. Legs brown, very distinctly
annulated with rufous-white hairs, protarsi less distinctly, tarsi
1903. | MR. F, PICKARD-CAMBRIDGE ON NEW SPIDERS, 161
not at all. Mandibles black, clothed with hoary-white hairs in
front.
Structure. General characters similar to those of other species of
the genus. Central posterior eyes one diameter apart (less in ¢ );
clypeus six times as high as the diameter of an anterior central
eye (in ¢ four times).
Vulval area broader than long, rounded, rectangular ; central
tongue distally broad, widely dilate basally; variable in different
examples.
Palpus. Tarsus twice as long as broad; tibia two-thirds the
length of the tarsus; one-fourth longer than the patella, Tibial
spur broad basally, curved apically, terminating in a small hook,
having on the inner side a slightly serrated ridge. The embolus
of the bulb has a stout erect spur springing from the cavity below
the apex of the lamina of the bulb, while the embolus itself is
short, strongly curved, and conspicuously flanged on each side.
Hab. Couompra, dried examples (¢ & 2); Venuzuena, dried
examples (¢ & @); also many examples in spirit, in coll. Brit.
Mus.
2. TRECHALEA URINATOR E. Simon. (Plate XV. figs. 3-5 a.)
Ann. Soc. Ent. Belg. xlii. p. 20 (1898).
Type ¢, gynetype Q, in coll. E.Simon. Total length (3, 2)
90-25 mm—(d¢). Carap. 11. leg i. 745 i. 82; m1. 62; iv,
85 mm. Tib.i. & iv. 19°5 mm. Prot. iv. 22 mm.
Both sexes with 4 teeth on the lower margin of the fang-
groove. Legs 4, 2, 1, 3. Clypeus 3-3} times in height the
diameter of an anterior central eye.
Carapace, abdomen, and legs entirely without dark bands or
annules, or these are at least very faint; entirely clothed with
dusty olive-brown hairs and pubescence.
Palpus. Tibia 14 times longer than broad ; tibial spur slender,
its apex bent over, forming a single small hook. The inner
apophysis deeply impressed across the middle, forming two blunt
cusps.
Vulva with a short broad central tongue, broadly dilate basally.
Tf I am correct in my identification of Simon’s species, it is a
common Spider throughout Ecuador.
Hab. Ecuapor: Guayaquil, Loja; Rio Durango; Bulun ;
Cachavi; Cavondelet ; Paramba; Salidero (Rosenberg).
3. TRECHALEA ELLACOMBEI, sp. nov. (Plate XV. fig. 6.)
Type @, in coll. Brit. Mus. Total length 25 mm. Carap.
10mm. Leg i. 68; ii. 75; iii. 60; iv. 80mm. Tib. i. Les hive
17-5 mm. Prot. iv. 22°5 mm.
Lower margin of fang-groove with 3 teeth. Legs 4, 2, 1, 3.
Clypeus 4 times the height of the diameter of an anterior central
eye. The whole of the central area of the carapace black, with
margins and striz flavous. Abdomen too much shrivelled to admit
of the coloration being recorded. Legs very deeply streaked and
Proc. Zoou. Soc.—1903, Vou. I. No. XI. 11
162 MR. F, PICKARD-CAMBRIDGE ON NEW SPIDERS. [FF eb. 17,
annulated with black, except the tarsi. Vulva witha long central
tongue, dilate basally, and its apex incurved.
This species is nearly allied to 7. urinator, but may be recognized
by the annulated legs, the possession of only 3 teeth on the
mandible, and by the form of the vulva.
Hab, Surinam, Bergen Daal (collected by Mr. C. W. Ellacombe).
4, 'TRECHALEA MACCONNELLI Pocock. (Plate XV. figs. 7, 8.)
Trans. Linn. Soc. Lond., Zool. (2) viii. 2, p. 67, Sept. 1900.
Type S, in coll. Brit. Mus. Total length 16 mm. Carap.
Primm Wee G4 - wii O mua o:D)> ava vie lt Dente cost yan lumen
Prot. iv. 18 mm.
Lower margin of fang-groove with 3 teeth. Legs 4, 2, 1, 3.
Clypeus twice the height of the diameter of an anterior central
eye. Carapace with a large black patch in the centre, its margins
scalloped by the flavous margin round the carapace and the pale
strie. Abdomen dusky with a flavous pattern and two flavous
spots above the spinners. Palpus: Tibia not, or scarcely, longer
than broad. ‘Tibial spur slender, concave lamelliform, its lower
edge slightly serrate; convex and incurved at the apex; seen from
above it has a A-shaped emargination. The inner apophysis is
cylindrical, low and impressed across the top.
Hab. Brivisn Guana: Mt. Roraima (collected by Messrs.
Quelch & McConnell).
5. 'TRECHALEA CONNEXA (O. P.-Cambr.). (Plate XV. figs. 9, 10.)
Triclaria connexa O. P.-Cambr. Biol. Centr.-Amer., Arach.
Aran. 1. p. 233, t. 30. figs. 1, 1 a-e (3), 1898.
Trechalea connexa F. P.-Cambr. loc. cit. i. p. 313, t. 30. fig. 17
(3), 1902.
Type <o, in coll. Godman & Salvin. Total length 14 mm.
Carap.7 mm. Leg i. 40; un. 45; in. 39;iv.48 mm. Tib. i. 10;
i. 11mm. Prot. iv. 13°5 mm.
Lower margin of fang-groove with 3 teeth. Legs 4, 2, 1, 3.
Clypeus as high as two diameters of an anterior central eye.
Carapace flavous, streaked and mottled with brown. Abdomen
dark brown. Legs decidedly annulate. Palpus: Tibia half longer
than broad ; spur, seen from below, without any hook at its apex.
Embolus of palpus without two ridges as in 7’. macconnelli, shorter
and more broadly flanged on its anterior margin.
Hab. Mexico, Atoyac (H. H. Smith).
6. TRECHALEA ExTENsA (O. P.-Cambr.). (Plate XV. figs. 11, 12.)
Triclaria extensa O. P.-Cambr. loc. cit. 1. pp. 174, 233, t. 22.
figs. 10, l0a—f( 3), 1896.
Trechalea extensa F. P.-Cambr. loc. cit. ii. p. 313, t. 30. figs. 16,
16 a,6 (6), 1902.
Type 3, in coll. Godman & Salvin. Total length 20 mm.
Carap. 10. mm. Wee i. 693 ii. 77: ii. 58; iv. 82 mm. ibs
175 iv. 19 mm, Prot! tv. 20'mm:
1903.] MR. F. PICKARD-CAMBRIDGE ON NEW SPIDERS, 163
Lower margin of fang-groove with 3 teeth. Legs 4, 2, 1, 3.
Clypeus 3 times as high as the diameter of an anterior central
eye. Carapace unicolorous flavous, margined with dusky black.
Abdomen unicolorous brown, with a short central anterior pale
dorsal bar. Legs faintly annulate. Palpus: Tibia twice as long
as broad; spur, seen from above, stout, projecting, curved, with
a stout sharp cusp or hook on the inner side at the apex, very
similar to but much stouter than in 7’. wrinator. Inner apophysis
deeply impressed in the middle. Embolus as in 7". longitarsis,
but less broadly flanged.
Hab. GuaveMaua (Sarg).
7. TRECHALEA AMAZONICA, sp. nov. (Plate XV. figs. 18-20.)
Type 3, gynetype 2, in coll. Brit. Mus. Total length ¢ 10;
(2? 15mm.) Carap.6mm. tegi. 31; 1. 32; mi. 29; iv. 37°5
mm. Tib.i. & iv. 7-8 mm. Prot. 1. & v. 10 mm.
Lower margin of fang-groove with 3 teeth. Clypeus in height
equal to 13 diameters of an anterior central eye. Legs 4, 2, 1, 3.
Sarapace dull straw-yellow; margins and strize shghtly suffused
with dusky brown, produced by fine dark hairs; mottled with
patches of white hairs, angular on margins; clypeus fringed and
clothed with hoary-white hairs. Mandibles, pedipalps, and a spot
on ocular area behind the eyes clothed with hoary-white hairs.
Abdomen olive-green mottled with brown and _hoary-white.
Ventral area clothed with silver-white hairs. Sternum and legs
dull yellow beneath, the latter brighter above ; femora indistinctly
annulated with dusky brown above ; base of patella black. Tibi
and protarsi with two broad dark annulations. Pedipalps also
annulated with brown.
Vulva with a narrow oblong central cavity, its lateral margins
straight and parallel, with an elongate central tongue, not dilated
at either end, not incurved at apex, without any black shining
bosses on each side.
The male and female here described were taken from the same
trunk of a tree on aswampy island in the largo opposite Santarem.
They were crouching on the bark like Sparassids, and when dis-
turbed moved with great rapidity.
Hab. Lowrr Amazons: Santarem; Breves, Parana Buyassu;
Para (f. P.-C. 1895-6).
8. TRECHALEA KEYSERLINGI, sp. nov. (Plate XV. figs. 1, 2.
Type 2, in coll. Keys. Brit. Mus. Total length 20 mm.
Carapace 9°75 mm. Leg i. 44; il. 46; ni. 36; iv. 49 mm.
ibe! bis ie, bed mm.) sProteavol3 mm.
Lower margin of fang-groove with 5 teeth. Clypeus five times
as high as the diameter of an anterior central eye. Legs 4, 2,
1, 3. Carapace with two broad central bands, widely separate
behind the eyes, connivent behind; the clear space behind the
eyes marked with lateral and central fine streaks. Abdomen with
a broad deep-brown foliate band above, constricted just behind
ile
164 MR. F, PICKARD-CAMBRIDGE ON NEW SPIDERS. [Feb. 17,
the middle and again slightly above the spinners, broadly margined
with pure white pubescence. Femora of legs deeply blotched and
streaked above with black; tibiz distinctly, protarsi indistinctly
annulated. Vulva without any central tongue, but with two
small cusps in the middle and two separate posteriorly converging
Jateral sclerites, quite distinct in form from that of any other
species here recorded (see Plate XV. fig. 2).
This specimen was identified by Keyserling as 7’. longitarsis
CliaiKtochea Seekps lane
Hab. Braztu, Rio Grande do Sul (Keys. Coll.).
Genus Hesyprus Simon, 1898 (June 30).
Simon, Hist. Nat. Ar. 2, u. p. 315. Type, H. palustris Simon.
Ecuador.
The species belonging to this genus are very closely allied to
Trechalea, but form a small group chiefly distinguishable by the
glabrous, gibbous mandibles, with a well-marked carina on the
outer side, in the male sex. The legs in both sexes are subequal
in pairs 4-2, 1-3; being also much shorter in proportion than in
Trechalea. Both margins of the fang-groove with 3 teeth. Eyes
as in 7'rechalea, central posteriors slightly over one diameter
apart; clypeus low and porrected. Tibia and protarsus i. with
2—2—2-2 spines beneath; protarsi 1-4 with a small central apical
spine beneath.
In Biol. Centr.-Amer., Arach. Aran. ii. p. 305, I have referred
the species given as the type of Hesydrus (sec. 2a, H. jullient)
to Hnna O. P.-Cambr., and the name Hesydrus i is applicable to the
species under sec. la with H. palustris as the type. It appears
that Simon has somehow confused the characters of these two
groups. In Hnna (including H. jullieni) undoubtedly the clypeus
is more vertical; the central posterior eyes are two diameters
apart, or more, and the anterior row is not wider than the
posterior central row ; while the tarsi are straight and not flexible.
Bubypume vez palustris (for if not identical, the form before me
is certainly congeneric with this species) the clypeus 1s porrect,
the central poster lor eyes one diameter apart, the anterior row
wider than the posterior central, and the tarsi flexible; the eyes
of the posterior row are certainly not equidistant, a character
given for distinguishing Hesydrus from Trechalea. Fortunately,
however, the citation of a definite type species enables one to
rectify the confusion.
The two species known to me may be recognized by the
following characters :—
a. Tibial spur of palpus, seen from above its apex, not bifid,
slightly emarginate and prominent at its angles, but
not incurving in the form of two teeth ............ . habilis O. P.-Cambr.
b. Tibial spur of “palpus, seen from above its apex, deeply
bifid, its angles forming two sharp teeth curving in-
wards and downwards agucneanqudisengHe speadaooo Geb aanbbe conan. | VAMIAINTET UN KorveaKayel,
1903.] MR, F, PICKARD-CAMBRIDGE ON NEW SPIDERS. 165
1. Hesyprus PAtusrris Simon, (Plate XV. figs. 22-25.)
Ann. Soc. Ent. Belg. xlii. 1898, p. 21(¢, 2)'.
Type d, gynetype 9, in coll. E. Simon, Total length ¢ 11
(2 16 mm.).—Carap. 6 mm. Leg i. 25; ii. 30; iii, 24:5;
iv. 30 mm. Tib.i.6; iv.7 mm. Prot. iv. 10°5 mm.
Lower margin of fang-groove with 3 teeth. Legs 4-2, 1-3.
Clypeus porrect, as high only as the diameter of an anterior
central eye. Carapace dull flavous with dusky hairs. Palpus:
Tibial spur short, concave lamelliform, deeply bifid at its apex,
forming two small, sharp, incurving teeth. Inner apophysis elon-
gate, not impressed across the middle, produced at its anterior
apical angle to form a small cusp. Vulva with a broad central
cavity, more than two-thirds the transverse width, concave on its
lateral margins, having a long narrow central tongue, not dilate
at either end, with a shiny black boss on each side, on the floor
of the cavity ; very variable in form.
Hab. Kevavor: Paramba (osenberg). Loja, Zamora '.— VENE-
ZUELA: Merida.
It is not certain that my identification of this species is correct
but it is highly probable.
2. Hesyprus HaBILis (O. P.-Cambr.). (Plate XV. fig. 21.)
Triclaria habilis O. P.-Cambr. loc. cit. i. pp. 173, 233, t. 22.
figs. 9, af (3), 1896.
Trechalea habilis F. P.-Cambr, loc. cit. ii. p. 313, t. 30. fig. 15
(3), 1902.
Type 3 incoll.Godman and Salvin. Totallength 10mm. Carap.
5mm. Tib, 1. 5°5 mm. (other legs broken off or mutilated).
Lower margin of fang-groove with 3 teeth. Legs 2, 4, 1, 3
(sec. O. P.-C.). Clypeus as high as 14 diameter of an anterior
central eye. Legs dull yellow-brown, slightly annulate. The
colours of the carapace and abdomen appear to have faded.
Mandibles glabrous shiny and very gibbous dorsally. Palpus :
Tibia one-third longer than broad. Tibial spur, seen from above
its apex, not bifid, its angles not forming sharp incurving teeth.
Hab. GUATEMALA: Costa Rica (Sarg, Rogers).
?
3. HESYDRUS ESTABANENSIS KE. Simon.
Ann. Soc. Ent. Belg. xii. 1898, p. 20(¢, 9).
Type ¢, gynetype 2? ,incoll. H. Simon. Total length’ ¢ 9,7mm.
One would suspect, from the descriptions, that this species is
congeneric with H. palustris, the males having the mandibles
convex and carinate on their outer apical margin.
Hab. VENEZUELA: San Esteban.
4, H&rsyDRUS BUCCULENTUS EK. Simon.
Ann. Soc. Ent. Belg. xlii. 1898, p. 20 (4).
Type 3, incoll. E. Simon. Total length 10 mm.
The remarks made under the last species apply also to this.
Hab. Brazit: Rio (Gounelle).
166 MR. F, PICKARD-CAMBRIDGE ON NEW SPIDERS. [Feb. 17,
Fam. SENOCULIDS,
Genus Senocuuus Taczanowski.
1872. Senoculus Taéz. Hor. Soc. Ent. Ross. ix. p. 106.—Type,
S. maronicus. ;
1873. Labdacus O. P.-Cambr. Proc: Zool. Soc. Lond. p, 118.—
Type, LZ. monastordes.
1880. Platyctenus Keys. Verh. z.-b. Ges. Wien, xxix. p. 338,
note. Nom. nov. for Senocwlus (nom. inappro. sec. Keys.).
1880. Stenoctenus Keys. Verh. z.-b. Ges. Wien, xxix. p. 340.—
Type, S. gracilis.
1883. Weothereutes Holmb. Bol. Acad. Nac. Ciene. Cordoba, v.
p- 35.—Type, V. darwint Holmb.
In a former paper, ‘‘ Cteniform Spiders from the Lower Amazons
&e.,” Ann. Mag. Nat. Hist. (6) xix. p. 90, Jan. 1897, I have tabu-
lated the characters of the known species of this genus so far as
I could draw reliable characters from the existing descriptions.
The table is here re-modelled and includes additional species
from Central America, published in the ‘ Biologia Centvrali-
Americana,’ and a new species occurring in the British Museum
Collection, of which a diagnosis is given below. I have not seen
S. parallelus Simon, but it is certain that my example from the
Lower Amazons belongs to that species, since M. Simon has kindly
compared a figure of the vulva with that of his own type.
The following is a list of the species already published :-—
1872. S. maronicus Tacz., 2, St. Laurent de Maroni. Hor. Soe.
_ Ent. Ross. rx. p. 106, t. in. fig. 4.
1873. S. monastoides (O. P.-Cambr.), 9, Brazil, Rio Grande.
P. Z.8. Lond. p. 118.
1879. S. rubromaculatus Keys., 9, Peru. Verh. z.-b. Ges.
Wien, p. 339, pl. iv. fig. 30. ey
1880. S. plumosus (K. Sim.), 9, Brazil, Para. Bull. Soc. Zool.
Fr. p. 154.
1880. S. purpureus (EK. Sim.), 9, Panama. Bull. Soc. Zool. Fr.
5 UD)
1880. S. parallelus (E.Sim.), 9, Brazil, Teffé. Bull. Soc. Zool.
Fr. p. 156.
1880. S. rujficapillus (HE. Sim.), ¢, Brazil, Para. Bull. Soe.
Zool. Fr. p. 154.
1880. WS. tricolor (Ki. Sim.), ¢ 2, Brazil, Teffé. Bull. Soc. Zool.
Fr. p. 153.
1880. S. gracilis (Keys.), 9, Peru, Aimable Maria. Verh. z.-b.
Ges. Wien, p. 341, t. iv. fig. 29.
1883. S. darwint (Holmb.), 9, Formosa, Chaco, Arg. Rep.
Bol. Acad. Nac. Ciene. Cordoba, v. p. 35.
1896. S. prolatus (O. P.-Cambr.), ¢ 92, Mexico, Atoyac. Biol.
Centr.-Amer., Arach. Aran. i. p. 218, t. 28. figs. 3, 3a—/( @ ).
1897. S. pallidus (F. P.-Cambr.), ¢, Brazil, Rio Janeiro. Ann.
Mag. Nat. Hist. (6) xix. p. 92, pl. iv. fig. 5 (sub
Stenoctenus).
1903. ] MR. F. PICKARD-CAMBRIDGE ON NEW SPIDERS. 167
1902. S. canaliculatus F. P.-Cambr., ¢ 9, Panama, Bugaba.
Biol. Centr.-Amer., Arach. Aran
figs. 3(¢), 4, 4a (9).
ii, p. 350, 6. 33.
The characters by which the species may be distinguished, so
far as I can gather them from the descriptions, are as follows :—
Males.
A. Carapace and abdomen clothed with plumose hairs.
B. Carapace and abdomen clothed with simple hairs.
I. Tibia i. and ii. with 9—10, protarsi with 8—9 long
spines beneath on each side .
II. Tibiai. and ii. with 4—4 long spines Deneath, pro-
tarsi with 5—6.
a. Anterior portion of ocular area almost vertical .
6. Anterior portion of ocular area porrect, hori-
zontal.
1. Apex of lamina round the inner anterior por-
tion of the bulb of the palpus much broader
and more deeply and broadly bifurculate ...
2. Apex of lamina round the inner anterior
portion of the bulb of the palpus much
narrower and less deeply bifurculate (simply
Ditid and canaliculate) 22. <2. ........00.o0cc ees one
Females.
A. Carapace and abdomen clothed with plumose hairs.
Ir Cane deeply striate, central stria oval-elon-
gate; tarsi i. and ii. with 4—4, protarsi i. and ii.
with 3—3 long spines beneath..
2. Carapace scarcely striate, central stria short, sub-
punctiform ; tibia i. and ii. with 5—5, protarsi
i. and ii. with 4—4 long spines beneath..
BG. Carapace and abdomen clothed with simple hairs.
a. Legs very long; tibia i. and ii. with long spines
beneath, 10 on inner side, 9 on outer side.
a, The lateral lobes of the vulva bearing a rounded
boss on the outer margin just behind the
middle (see Keyserling’s figure) ..
. The lateral lobes of the vulva without any
rounded boss ........
6. Legs shorter, tibia i. and ii. with ‘only 4—4 or
5—5 long spines beneath.
aa. Tibia i. and ii. with 5—5 long spines beneath.
a. Posterior eyes very large, centrals less than a
diameter apart, scarcely 1} diameters from
the laterals ; anterior Eee of ocular area
almost vertical aeaias
. Posterior eyes moderate in size, , centrals over
a diameter apart, at least two diameters
from the laterals. Anterior portion of
ocular area porrect ...
oe au we i, and ii, with 4—4 long spines beneath.
. Protarsi i. and ii. with 3—3 long spines
beneath, besides lateral and apical spines...
68, Protarsi 1. and ti. with 5—é long spines
heneath, besides lateral and apical spines.
a’. External sclerites of vulva eyenly rounded
on their outer margin, sinuously concave
on their inne. margin, their apices
(anterior portion) sharp, with a minute
cusp on their outer margin.
b4, External sclerites of vulva emarginate on
their outer sides, evenly convex on their
ner margins, "their apices forming a
broad hook, not very sharp at the point,
without any minute CUSD.........000-00cee es
tricolor Simon.
pallidus I. P.-Cambr.
ruficapillus Simon.
prolatus O. P.-Cambr.
canaliculatus FP. P.-Cambr.
plunosus Simon.
iricolor Simon.
gracilis Keyserling.
parallelus Simon.
ruficapillus Simon.
monastoides O. P.-Cambr.
albidus ¥. P.-Cambr.
prolatus O. P.-Cambr.
canaliculatus F. P.-Cambyr.
168 MR. F, PICKARD-CAMBRIDGE ON NEW SPIDERS. [Feb. 17,
SENOCULUS ALBIDUS, sp. nov. (Plate XIV. fig. 14.)
Type 9,in coll. Brit. Mus. Total length 14 mm.
In general appearance this species is similar to others of the
genus, the legs being clothed with a lateral fringe of curving
simple silky white hairs. The abdomen is also clothed at the
apex on each side with long pure-white hairs. The vulva consists
of two large oblong convex sclerites convergent forwards, and
somewhat concave at this point, see Plate XIV. fig. 14, where
also the vulva of S. parallelus Simon is figured (fig. 13).
Hab. Brazil; Rio Janeiro.
Nore.—Of S. purpureus (KE. Simon) and S. darwint (Holmberg)
1 cannot give any reliable characters. S. maronicus Tacz., the
type of the genus, is, according to the author, immature, and no
mention is made of the number of spines on the legs.
EXPLANATION OF THE PLATES.
PratE XIV.
Vig. 1. Thaumasia velox, 9, vulva, p. 154.
xD annulipes, 2, ee | p. 154.
| Dossenus marginatus, 9, p. 155.
” oF 2 p) vulva.
2 2 » €yes.
Paradossenus ni igréicans, 2, p. 155.
fs 35 ©, vulva.
” ” 2, eyes.
e 6, tibial spur of palpus.
Thanatidius spinipes, &, eyes, p. 156.
9p a ditto, in front.
Bs 9, vulva.
| Senoculus parallelus, 2, vulva, p. 166.
a albidus, 2, vulva, p. 168.
fa Sd lt
PER ooesaaer aos
PratEe XV.
Fig.1. Trechalea keyserlingi, 9, p. 168.
2. A 5 vulva.
3 urinator, 6, tibial spur of palpus, p. 161.
A. i Hs 2, vulva.
5. Bs 6 3, embolus of palpus.
BOR x 6, 5 from below.
6 ie ellacombei, ©, vulva, p. 161.
7 5 macconnelli, 6, tibial spur of palpus, p. 162.
8. ie oe 6, embolus of palpus.
9. 55 connexa, 6, tibial spur of palpus, p. 162.
10. ms 5 3d, embolus of palpus.
il. 36 extensa, 6, tibial spur of palpus, p. 162.
12. 3 6, embolus of palpus.
13. a) longitarsis, 6, embolus of palpus, p. 160.
14, 55 a 6, ditto, apex enlarged.
15. es % 6, tibial spur of palpus.
16. an 6, ditto, example from Venezuela.
WW) aA By ©, vulva, example from Colombia.
18. ne amazonica, 6, tibia of palpus and spur, p. 163.
9} op a 3, embelne of palpus.
20. ©, vulva.
21. . Hesydrus habilis, 6, tibia of palpus and spur, p. 165.
op palustris, 8, tibia of palpus and spur, p. 165.
AY 55 6, embolus of palpus.
a e 6, right mandible, from outer side.
25. oD ee 2, vulva.
7 ‘ . a
Pos ar ina
fi 1@
pies
h 7 s 4 7
| fi ul
5. Gal Bale & Danielsson L® bth.
: 8. j
bsiodleea cen ots), PeNAbL OC si4n, MOP WARS ALLOW,
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EGS SOS vol IPN SOV
Ciemdelk Bale & Danielsson [,°4 hth.
IZAENARILIONG Aa WOME S, EOL
1903. ] ON THE MARINE FAUNA OF ZANZIBAR. 169
2. On the Marine Fauna of Zanzibar and British East Africa,
from Collections made by Cyril Crossland in the Years
1901 and 1902.—Polycheta. Part I. By COvynix
CrossLanp, B.A., B.Sc.1
[Received January 15, 1903.]
(Plates XVI. & XVII.)
Introduction.
The collections referred to were made under the following
circumstances :—Sir Charles Eliot, K.C.M.G., C.B., late Fellow
of All Souls College, Oxford, at present H.M. Consul-General at
Zanzibar and Commissioner for British East Africa, took me out
with him as his private assistant in his researches on Nudibranch
Mollusca. I made collections not only of this, but of the other
marine groups; accounts of which, by various specialists, will
appear in these ‘ Proceedings’ from time to time. The largest
collections are those of the Nudibranchiate Mollusca and Polychete
Annelids, of which groups about 150 species in each are to be
described. Sir Charles Eliot has already published one part of the
results of his examination of the Nudibranchs (P. Z. 8. 1902, vol. ii.
p- 62); other papers by various authors will shortly be ready.
For the benefit of other possible workers in this region, I may
mention that the greater part of my collections was made in two
localities, viz. Chuaka Bay, on the east coast of Zanzibar, and
Wasin Harbour, near the Anglo-German boundary on the main-
land. The former locality is extremely rich in shore forms, and
dredging in 3 fathoms of water at the north side of the mouth of
the bay was often very productive. Wasin Harbour averages a
depth of 10 fathoms, and here I collected almost entirely by
dredging. The bottom is extensively covered by a species of
elesto, the branches of which are overgrown by an encrusting red
sponge. Among this a great variety of the smaller Polycheta
and Nudibranchiata are found.
My mention of these two localities is of the more importance to
future workers because large stretches of the coast are extremely
barren. Unfortunately this applies to the two principal towns of
Kast Africa, and especially to Mombasa. A few miles from
Zanzibar, e.g. round the islands and sandbanks which surround
the harbour, and on one portion of the shore about a mile to the
south of the town, at low spring-tide only, ave rich collecting-
grounds, and near this latter area dredging is profitable, especially
at a depth of 5 fathoms. The greatest portion of the Zanzibar
channel is, however, extremely barren.
More detailed descriptions of the reefs of East Africa, with
maps, will be found in my two papers: “On the Coral Reefs of
Zanzibar,” and “ Pemba and British East Africa,” in the Proc.
Camb. Phil. Soc. vols. xi. & xii. (1902).
1 Communicated by Pref. W. C. McInrosn, F.Z.S.
2 For explanation of the Plates, see p. 176.
}
170 MR. CYRIL CROSSLAND ON THE Feb. 47,
The gratitude of all zoologists is thus due to Sir Charles Eliot,
whose generosity and scientific zeal have enabled these collections,
the first of considerable size from this region, to be made. I
wish also to express my best thanks to Mr. Stanley Gardiner, who
helped me by his advice and in every other possible way, both
during my residence in Hast Africa and in the working out of the
collections on my return to England.
CHAHTOPTERIDS,
Genus PHYLLOCH ©TOPTERUS.
The position of the genus Phyllochetopterus among the lower
Cheetopteridx is shown by the following table of the genera :—
A. Notopodia of the second body-rezion not foliaceous
(z. e. two body-regions only) ................. RANZANIA.
B. With foliaceous notopodia posteriorly.
1. Body divided into two regions ..................... LELEPSAVUS.
2. Body divided into three regions.
; One pair of tentacles ........................... .. SPIOCHETOPTERUS.
AUG JORNERS Cue WIUEVOTES |... cs,cponon ondcoo ganado sue boo PHYLLOCH ETOPTERUS.
The most characteristic feature of the Chetopteride is the
adaptation of certain parapodia for the production of a respiratory
current, which modification, completed in the genus Chetopterus,
makes the latter one of the most remarkable forms of animal life.
This pecuhlarity is not developed in Ranzania, whose only
Chetopterid features are its general build of body, which is
divided into an anterior flattened muscular and glandular portion
bearing long notopodial setz only, and a rounder weaker posterior
portion with delicate notopodia and neuropodia of uncinigeral
tori; in addition it is tubicolous in its mode of life, and procures
food by the ciliated grooves of its tentacles and dorsal surface.
To these features are added notopodial gills in Telepsavus, in
which case they are developed on every segment of the hind-body.
In the remaining two genera this modification is restricted to
more or fewer of the middle segments, to two only in Spiocheto-
pterus and several species of Phyllochetopterus, but up to 25 in
other species of the latter.
Tn all the eight species of Phyllochetopterus yet known the
body is vermiform, the notopodial gills comparatively small and
simple, bifid, and containing capillary sete. All the sete are
characteristically Cheetopterid, in the first region long with leaf-
like ends, and arranged in a row as in Chetopterus, though the
notopodia of P. aciculigerus alone approach in their long pointed
shape those of the former genus. In the fourth notopodium of
the first body-region one or more sete are enlarged and of a dark
brown colour, but are not flattened as in Chetopterus. The uncini
are of one kind only throughout the body, but their form is more
specialized than in Cheetopierus, and is that which occurs in, for
example, the Sabellidee. The parapodia are alike in all the species,
differing only in the proportions of the parts, excepting the hind-
body notopodia of P. aciculigerus (6).
1903.] MARINE FAUNA OF ZANZIBAR. 171
The prostomium is, in this family, usually more or less reduced.
In Chetopterus itself it is completely merged with the oral
funnel, and the same condition is described for some Phyllocheeto-
pterids (e. g. P. claparedi McIntosh). In all the members of this
genus it is normally scarcely distinguishable externally, both pairs
of tentacles being carried by the peristomium. In certain specimens
of P. elioti, however, as described below, it is quite prominent,
and so was discoverable in all the four species I have examined.
The first body-region is an almost solid mass of muscular and
glandular tissue in all species, but posteriorly musculature is
confined to the ventral surface, except at the level of the parapodia,
where muscle-bands surround the body. Between these the body-
wall is excessively thin dorsally and laterally, and always black
from the colour of the gut-walls.
Up to the present six species have been described, viz. :—
P. socialis Claparéde. Mediterranean and Atlantic (1) (4).
P. major Claparéde. Mediterranean (1).
P. gracilis Langerhans. Canary Islands (2).
P. claparedi McIntosh. Japan (3).
P. aciculigerus Crossland. The Maldive Islands (6),
P. gardineri Crossland. The Maldive Islands (6).
To these I now add :—
P. elioti. Zanzibar.
P. pictus. British East Africa.
Claparéde’s P. fallax was founded on such characters as the
numbers of segments composing the three body-regions, the
ringing of the tube, &c.; these have been shown by Roule (4) to
be variable in members of this genus.
The principal distinctions of systematic importance are :—
(1) Presence or absence of eye-spots, and of the development
of the prostomium.
(2) The number of the strong setz in the notopodium of the
fourth setigerous segment.
(3) The proportionate sizes of the parts of the parapodia of
the regions B and C.
(4) Ringing and other such details of the tubes are variable,
but (1) in some species they are straight and occur
singly, and (2)in others they are of a U-shape and occur
in numbers twisted together.
These characters are distributed as follows :-—
(1) Eye-spots present in P. socialis, P. major, P. elioti, P.
gardineri, P. pictus; absent in P. claparedi, P. aciculigerus,
P. gracilis.
(2) A single strong seta in 4th notopodium in P. socialis',
P. major, P. claparedi, P. pictus. More than one seta
strengthened. PP. gracilis with three, P. gardineri three,
P. elioti two, and P. acieuligerus eight.
1 In P. socialis two sete may exceptionally occur, but the extra one is always
smaller than the one it accompanies.
172 MR. CYRIL CROSSLAND ON THE [Feb. 17,
(3) The parapodia of the region C consist of double neuro-
podial tori and a clavate papilliform notopodium containing
a single seta in all the species except P. aciculigerus and
P. claparedi.
(4)* Tubes straight, occurring singly in P. claparedi, P. elioti,
P. major. Twisted together, in numbers, P. socialis and
P. pictus.
PHYLLOCHATOPTERUS ELIOTI*, sp.n. (Plate XVI. figs. 1-3 & 8;
Plate XVIT. figs. 10-13.)
Of this species numerous specimens occur in Chuaka Bay,
Zanzibar. The straight, stiff, opaque black tube, 6 to 9 inches
long, 1s buried in the sand at low tide, one or two inches only
projecting. Its presence is usually rendered conspicuous by the
growth of a tuft of bright green enteromorpha on the projecting
portion.
The colour of the living animal is milk-white anteriorly, and
black posteriorly. There is no pigmentation, and the brilliant
blue and yellow colours described by Claparéde and Lo Bianco in
the Naples species are not found in either of the two species I
have seen alive. The black colour of the gut gives the usual
green solution in alcohol.
The peristomium is very mobile, the whole shape of the head
being thus very different in different specimens. When expanded,
its shape is as in Pl. XVI. fig. 1. In a few cases only, where the
head is much contracted and bent back, does the prostomium and
its eye-spots come into view as in fig. 2.
The first pair of tentacles are very slender and long, attaining
a length of 9 mm.
The fore-body measures, in a large specimen, 6 mm. in length
by 2°5 mm. in breadth, and consists of about 14 setigerous
segments. (There were 13 in three specimens, 14 in three more,
and 15, 16, and 17 each in three more. The average is thus 14
in nine specimens.) The notopodia are very short and stumpy,
but the setz and their arrangement, as in the other species of the
genus, recall Chetopterus. The fourth foot has two strong sete
on each side, rarely three on one side or the other, though in one
specimen there were three on one side and four on the other.
The ordinary sete are all straight.
The region B (fig. 3) consists of from 20 to 25 segments, the
parapodia of which are very like those of P. gardineri, except that
the notopodial flap is slightly smaller, and there is a long space
between it and the ventral portion of the neuropodium, and the
parts of this latter are of more nearly equal size. The neuropodium
of the first segment of the gill-region is not divided. There are in
each notopodium five long setze whose flexible heads project. The
uncini (fig. 8, c) are triangular with very fine teeth, just visible
under a 4-inch objective.
The hind-body has more than 25 segments, but none of my
1 The tubes of Gardiner’s two species were unfortunately not obtained.
2 Thus named in honour of Sir Charles Eliot, K.C.M.G., C.B., H.M. Consul-
General at Zanzibar, and Commissioner for British Hast Africa.
1903.] MARINE FAUNA OF ZANZIBAR. 173
specimens are quite complete. The notopodia are the usual
clavate papille containing a single seta. The neuropodia are as
in the gill-bearing region, but much smaller.
The two species P. eliott and P. gardineri are closely related,
differing only in the reduction of the peristomial collar in the
latter and the proportions of the mid-body parapodia and setee.
The only work on the internal anatomy of a member of this
genus is the section of the fore-body of P. claparedi given by
McIntosh, although examination of the anatomy of the species
of Phyllochetopierus would be extremely interesting, both for
comparison among themselves and with the other genera of the
family’, especially Chetopterus. Having neither time nor full
opportunity for this work, to which my inclinations are most
strongly drawn, I can only give here a few deductions from the
examination of a series of sections of a specimen of this genus,
which appear to differ in many minor details from the above-
mentioned section of P. claparedn.
Body-wall.—The region A is, as shown by the sections figured
(BD Xa ies. & 12), an almost solid mass of glandular
and muscular issues in marked contrast to the delicacy of the
regions Band C. The whole of the ventral epithelium is strongly
glandular beyond the limits of the well-marked ventral shield, and
these glands extend anteriorly to the dorsal surface and even on to
the prostomium. The cuticle is of extreme delicacy, if really
present. The musculature consists of weak circular and very
strong longitudinal muscles, but the division of the latter into
two dorsal and two ventral bundles cannot be made out. In the
regions B and C the only muscle occurring dorsally is a very
delicate circular layer (Pl. XVII. fig. 13). Diagonal fibres can be
traced in the region A between the nerve-cords and the notopodia,
which become more definite muscles in the region B (fig. 13),
The nervous system is in contact with the skin. The two
ventral cords le at a distance from one another in the region A,
but approach one another posteriorly (cf. figs. 12 & 13), their
arrangement corresponding thus to that in Chwtopterus. Trans-
verse commissures, portions of one of which are shown in Pl. XVII.
fig. 12, connect the two cords.
The brain (Pl. XVII. fig. 10), which in Cheetopterus is a narrow
band dorsal to the mouth, not differentiated from the circum-
cesophageal commissures, is here more distinct, in correspondence
with the presence of the prostomium in this genus and its absence
in the case of Chetopterus. Itis a perfectly simple swelling of the
circum-cesophageal commissures. The continuity of the nervous
system with the epidermis is shown by a comparison of the
sections of the prostomium given in figs. 10 & 11. The former
shows the circumeesophageal connectives, that on the left as it
crosses from the ventral to the dorsal side, that on the right, near
the eye-spot (e) passing into the prostomium. The replacement
of the ordinary glandular epithelium of this area by the densely
1 For anatomy of Telepsavus, see pl. xiii. and its explanation in Claparéde’s
‘ Annélides Sédentaires,’
174 MR. CYRIL CROSSLAND ON THE [ Feb. 17,
staining nerve-nuclei of the brain (which are similar to those
seen on the outer side of the ventral nerve-cords &ec.) is shown
distinctly, this replacement being complete in fig. 10, where the
epidermis is composed entirely of nerve-cells.
The eye-spots are a pair of groups of cells, each containing
numerous minute granules of black-brown pigment. These are in
continuity with the nerve-cells of the brain, but are anterior and
dorsal to its fibrous part.
The celom is small and broken up into several distinct portions.
The largest of these are a pair of spaces lying laterally and
ventrally to the gut (fig. 12), bounded dorsally by the powerful
muscles of the seta-sac. Dorsally and medianly, commencing as a
space in the prostomium, is a third part which posteriorly becomes
a mere canal surrounding the dorsal blood-vessel. In the region B
the eelom is more normal, though reduced in size. It remains
divided into right and left halves by the dorsal and ventral
mesenteries of the gut.
The vascular system consists of dorsal and ventral vessels, both
running in the gut-mesenteries. The former breaks up at the
base of the prostomium into three branches (fig. 10), and large
connecting-vessels are found in the anterior segments. Posteriorly
the dorsal vessel forms a large sinus covering the dorsal wall of
the gut (fig. 13).
Alimentary canal.—The mouth is richly ciliated, and the outer
ends of its columnar epithelial cells contain a few minute specks
of the black-green pigment so characteristic of the family. These
are absent from the narrow triangular gut of the region A, but
reappear in great quantities in the larger thick-walled alimentary
canal of the regions B and C (fig. 13).
The gut-lining here consists of long crowded cells, the swollen
distal ends of which are crowded with minute specks of the
pigment, indicated by dark dots in the figures.
The transverse sections of the notopodial gills, shown in fig. 13,
are interesting. The space which they contain is ccelomic, and at
its centre is a bundle of five or six setee wrapped closely together
by fleshy tissue. The transverse section of this resembles that of
a telegraph-cable, the wires being represented by the sete. At
each end of the oval section is seen a sharply-cut groove, the
sides of which are of granular, rather deeply-staining protoplasm,
without nuclei, supporting externally very long cilia. Laterally
is a more deeply-stained area of the epithelium, perhaps of nervous
function.
PHYLLOCHAHTOPTERUS PICTUS, sp. n. (Plate XVI. fig. 5.)
The tubes of this species are brown and translucent, quite free
from sand or mud. They were found clustered together in con-
siderable numbers on the under side of a large stone at low-water
level on Pungutiayu islet, Wasin. Each tube has two openings,
each limb of the U being about 4°5 cm. long. The bending is
irregular, and they are twisted and fused together, so that 1t 1s
usually almost impossible to separate out any one tube.
1903. MARINE FAUNA OF ZANZIBAR, 175
The contained worms are the smallest of the species described
here, or in (6), being slightly smaller every way than in P. elioti.
The specific name refers to the abundance of brown colour
found on their tentacles and fore-body, Pigmentation, except of
the gut, is rare in the Cheetopteride. Of the other species, it is
described in only two, viz. P. claparedi (8) and P. socialis (a): ari
both these it is comparatively slightly developed.
The ground-colour of the anterior region, tentacles, and para-
podia of P. pictus is, in life, creamy white. The greater part of
the posterior regions is, as usual, black, by reason of the pigmen-
tation of the gut.
On either side of the groove which runs along the upper surface
of the larger tentacles are regularly arranged squarish blotches
of brown, as shown in Pl. XVI. fig. 5. Brown dots are scattered
also over the pro- and peristomium and the anterior segments of
the fore-body. There are two dark bands with indefinite edges
along the ventral bases of the notopodia, and a broad crescent
across the ventral surface of the end of the region A.
The mouth is not at all funnel-shaped, but rather slit-like, the
two small peristomial lappets coming together from either side.
The prostomium is large, flattened from side to side, and projects
above the dorsal surface of the fore-body. The very slender second
setigerous pair of tentacles lie close on either side of it, being thus
very inconspicuous. Two elongated, but distinct, eye-spots, of a
dark brown colour, occur one on each side of the prostomium.
The regions A and B consist of the following numbers of
segments in different individuals :—
* % *
Doren OM UG.) Sls SMe eaieye CaS
Bese uh 9 9) 7 9) 3 8 —
* These specimens were below the average size.
The fourth notopodium has but one large seta of the shape
shown in Pl. XVI. fig. 9,a. The bending of this seta is remarkable
as occurring only in this species. Its three teeth, when seen from
one point of view, give an explanation of Langerhans’s figure of
the corresponding seta in P. gracilis, which is not otherwise
intelligible.
The other setz are all straight, and present no peculiarities
except those of the last notopodium, which are bent, and the head
is finely toothed (fig. 9, c). .
The parapodia of the region B (figs. 5 & 6) are small, the
neuropodial ridges especially so. In correspondence with this the
uncini are very minute and delicate (03 mm. long), and their
teeth barely visible under a 1-inch objective (fig. 9, d). The
neuropodia do not extend far up the side of the body, and the
space between them and the notopodial gills is not filled up by
the triangular membranous gill-flap as in the other species (fig. 6).
The notopodia contain two or three thin sete which do not project.
In the region C the notopodia are reduced to little clavate
papille, containing one seta, as in all the other species except
176 ON THE MARINE FAUNA OF ZANZIBAR. | Feb. 17,
P. claparedt and P. aciculigerus (see fig. 6, of P. elioti). The
neuropodia also are small.
List of the Literature.
(1) Ciuaparrpr.—Annélides du Golfe de Naples, 1868.
(2) LancerRHANS.—“ Ueber einige Canarischen Anneliden.” Noy.
Act. der Leop.-Carol. Deutschen Akad. Bd. xliii., 1881.
(3) McInroso.— Challenger’ Reports, vol. xii., 1885, p. 374.
(4) Route.‘ Campagne du Caudan.” Ann. d. l'Université de
Lyon, 1896.
(5) Lo Branco.—Ann. Tub. nel Golfo di Napoli, 1893.
(6) Crossnanp.—J. Stanley Gardiner’s ‘Fauna and Geography
of the Maldive Archipelago.’ In the press.
EXPLANATION OF THE PLATES.
Prate XVI.
Fig. 1. Normal condition of the head of Phyllochetopterus elioti (p. 172), as seen
from dorsal side. X 10.
Side view of a specimen whose prostomium and eye-spots are visible. > 10.
. Side view of three segments of the region B. > 10.
Ventral view of the same. ’
P. pictus (p. 174). Dorsal view of anterior part of body. x 10.
. Side view of three segments of the region B. X 10.
A notopodium of the region C in either species. 100.
. Sete of P. elioti (p. 172).
(a) Thickened seta of fourth parapodium. x 83.
(6) Normal seta from third foot. x 90.
(c) An uncinus. > 1000.
9. Setee of P. pictus (p. 172).
(a) Thickened seta from the fourth foot. > 100.
(b) Head of a normal seta from the same bunch. 200.
(c) From last foot of region A. The shaft is bent and part of the head
finely toothed. x 100.
(d) One of the uncini. > 1000.
Puate XVII.
Fig. 10. Section of Phyllochetopterus elioti (p.172) through the brain, bases of
short tentacles, &c.
11. Section a little anterior to fig. 10, through mouth, first feet, &c.
12. Section at level of fifth foot.
13. Section near beginning of region B. All sections x 42.
DAD HH oo wo
The meaning of the lettering on the figures of the Plates is as follows :—
b.v.=blood-vessel. n.=nerve,
cire.m.=circular muscle. n.c.=nerve-cells.
ce.=ceelum. n.cd.=nerve-cord.
d.bv.=dorsal blood-vessel. n.f.=nerve-fibres.
d.cil.=dorsal ciliated groove. neph.=nephridium.
d.coe.=dorsal division of ccelom. neur.=neuropodium or ventral division
d.neur.=dorsal division of the neuro- of the foot.
podia. noto.=notopodium or dorsal division
e.=eye. of the foot.
g-=gut. peri.=peristomium.
gill.=gill- flap between noto- and ppd.=parapodium or foot.
neuropodia. pro.=prostomium.
gl.=glandular areas or epithelia s.=seta.
&e. ¢, & ¢,=first and second pairs of ten-
Zm.=longitudinal muscle-layer. tacles.
m.=ruscle. v.bv.=Vventral blood-vessel.
mo.=mouth. v.neur.=ventral division of the neuro-
wm. of ppd.= muscles of parapodium. podia.
PA S419 085) vol lela leE Qvale
K.Broom del.
MP Parker lith. Parker & West imp.
AXIS, ADLAS: i PROAT WAS Or
GOMPHOGNATHUS & TRIRACHODON.
1903.] ON THE VERTERR® OF THERIODONTS. Lie
3. On the Axis, Atlas, and-Proatlas in the Higher
Theriodonts. By R. Broom, M.D., B.Se., C.M.Z.8.
[Received January 20, 1903. }
(Plate XVIIL.')
Hitherto, though something has been known of the structure
of the axisin at least two of the Dicynodonts and in Cynognathus,
we have been practically ignorant of the structure of the atlas in
any of the Theriodonts or Anomodonts. In fact, Prof. Seeley (1)
in describing the axis of Cynognathus, seems to be in doubt
whether he is really dealing with an axis, or with an axis and
atlas combined. On page 100 of his paper he says, ‘The first
vertebra appears to be anchylosed to the second”; while a little
further on he states that “this vertebra is remarkable for its
form being exactly like the odontoid process of the vertebra in
many animals, and suggesting the idea that the atlas is lost” ;
and in a note he adds: ‘ This appears to be confirmed by the sub-
sequently to be described condition in Z'ropidostoma dunni.” In
counting the vertebra, however, he counts the axis as the Ist
cervical, and states that “there are 6 cervicals.” Gadow (2), in
his recently published work on Reptiles, possibly misled by Seeley,
definitely states that “the atlas is fused with the axis.” Con-
sidermg how mammal-like the higher Theriodonts are, and that
the axis bears a very marked resemblance to the mammalian axis,
having a large spime and a well-developed odontoid process, it
seems remarkable that the idea should have arisen that the atlas
was anchylosed to the axis, more especially as there is no part
of the axis in Cynognathus that bears any resemblance to any
kniown atlas.
When in Grahamstown recently, I had an opportunity, through
the kindness of Dr. Schonland, of examining the Theriodonts in
the Albany Museum, and especially the very fine specimens of
Gomphognathus and Trirachodon, which have been described by
Seeley (3), and which have been so marvellously developed under
his direction. In both the type specimens of Gomphognathus
kannemeyert and Trirachodon kannemeyeri the anterior cervical
vertebra are most beautifully shown, but Seeley in his description
ot the specimens makes no reference to this most important
region, beyond stating (p. 53) that “ the occipital plate [in 7’rira-
chodon]| is not completely exposed owing to some of the vertebre
being in contact.” Twoof his figures of the skull of 7'rivachodon
show the atlas and part of the axis; but as no distinction is shown
between matrix and bone, it is difficult to make anything of the
drawing.
The beautiful specimen which forms the type of Gomphognathus
1 For explanation of the Plate, see p, 180.
Proc. Zoou. Soc.—1903, Vou. I, No. XII. 12
178 DR. R. BROOM ON THE [| Feb. 17,
consists of two complete mandibles in articulation with the
posterior half of the skull. The lower jaw lies almost at right
angles to the axis of the skull, and the upper cervical vertebrae
have been to some extent protected by passing down between the
two rami. The skull consists of the practically complete occiput
with both temporal arches in almost perfect condition, and with
most of the parietal and sphenoidal regions well shown. Though
the sutures unfortunately are not distinct, the occiput is so well
preserved, practically without distortion, that I think it worthy
of being figured.
The vertebrze so far as preserved consist of the proatlas, atlas,
axis, and the 3rd and 4th cervical vertebre.
The atlas (a¢.) consists of a well-developed arch and a distinct
hypapophysis (A.at.). The arch is partly broken in the middle, but
must have borne considerable resemblance to that of the mammals.
It differs in having had a distinct posterior zygapophysis (z.at.)
for articulation with the small anterior zygapophysis of the axis
z.ax.). The zygapophysis of the left side is well preserved, but
that of the right has been broken off. On the right side the
outer part of the arch is continued backwards as a well-developed
bony process or rib (7.a¢.). It is probable that the process is
really a rib, but I have been unable to find a suture between it
and the arch. On the under surface, what appears to be part of
the rib of the left side is seen lying by the side of the body of the
axis (7.at.). The well-developed hypapophysis is also well seen.
It appears to have had only a ligamentous connection with the
arch of the atlas.
In front of the arch of the atlas, and lying in the hollows formed
above and outside of the occipital condyles, are the two portions
of the proatlas. Each consists of a short bony arch with a
well-developed process passmg upward and outward from its
outer end. e
The axis is well developed, and bears considerable resemblance to
that of Cynognathus. The spine forms a very large flattened crest.
In the specimen it is somewhat damaged, but there is probably
not a great deal missing. In front, the spine passes forwards
over the arch of the atlas, and possibly forms an articulation
withit as in the Monotremes. Between the anterior part of the
spine and the odontoid process is the small anterior zygapophysis
(z.ax.). The odontoid process (0.p.) is well developed, and on the
right side it is seen articulating with the arch of the atlas. On
the under surface the body of the axis is seen, asalso a part of the
odontoid process in articulation with the hypophysis of the atlas.
On the upper surface, near the posterior end of the centrum, on
the right side is a portion of the rib of the axis.
The 3rd and 4th cervicals present no features worthy of any
special remark. Remains of ribs are seen in connection with
each.
In the type specimen of Zrirachodon kannemeyert there are
preserved and well shown the two portions of the proatlas, the
1903. ] VERTEBRE OF THERIODONTS, 179
arch and hypapophysis of the atlas, and the odontoid process and
part of the body of the axis. Though the structure of the
vertebra is essentially similar to that in Gomphognathus there are
many points of difference.
The proatlas (p.a.), though occupying the same relative position.
as in Gomphognathus, is less specialized. It is present as a pair of
curved, moderately thin, bony plates lying in front of, and probably
somewhat overlapping, the arch of the atlas, There is no bony
process developed as in Gomphognathus.
The atlas (a¢.) consists of a well-developed arch and a distinct
hypapophysis. ‘The arch is not so complete as in Gomphognathus,
and must have differed considerably in regard to the arrangement
of the anterior articular surfaces. In Gomphognathus the occi-
pital condyles (0.c.) are small and moderately close together,
whereas in Z'rirachodon the condyles (0.c.) are very large and as
wide as the atlas. The arch, perhaps on this account, does not
close in inferiorly to meet the hypapophysis, with which it can
only have been attached by hgament. The ‘zygapophy sis (z.at.)
for the axis is very small. “The hypapophysis (/.at.) is very
sunilar to that in Gomphognathus.
Of the axis only the odontoid process, imperfectly displayed,
and a part of the body remain.
The structure of the upper cervical vertebre is so imperfectly
known in the majority of fossil reptiles, that there is some diili-
culty in satisfactorily dealing with the aftinities of the structures.
It is difficult to avoid being struck by the close resemblance of
the axis, atlas, and proatlas in the Theriodont to those structures
in the Crocodiles, more especially as there is no close aflinity
between the groups. The explanation is probably to be found in
the fact that both the Crocodiles and the Theriodonts, though far
removed from each other, have retained with but slight modifi-
cation the type of structures met with in their common ancestor.
In Sphenodon, though the proatlas is small, the type is practically
the same; and I have recently discovered that Procolophon has a
well-developed proatlas. Procolophon has usually hitherto been
associated with the Dicynodonts, Theriodonts, and Pareiasaurians,
but as it has a persistent notochord, abdominal ribs, a plate-like
pubis and ischium, and 2, 3, 4, 5, and 4 phalanges in the digits, it
is manifestly much more closely related to Palwohatteria than it is
to the Theriodonts and Dicynodonts. In most of its structures it is
probably as primitive as the common ancestor of the Crocodiles
and Theriodonts, so that there is good reason for believing that
the common ancestor had a well-developed proatlas and probably
an atlas and axis very similar to those found in later forms,
but almost certainly with the odontoid process as a distinct
element, as seen in Jcehthyosaurus. No proatlas has yet been
detected in Pareiasaurus, but I have found one in Lystrosaurus
(= Ptychognathus Owen). It will probably yet be found in most
of the primitive reptilian types,
12*
180 MR. C. TATE REGAN ON THE (ctor
Works referred to.
(1) H. G. Srerey: “On the Skeleton in new Cynodontia from
the Karroo Rocks.” Phil. Trans. vol. 186 B. p. 59 (1896).
(2) H.Gapow: ‘ Amphibia and Reptiles.’ London, 1901.
(3) H.G.Sreney: “On the Gomphodontia,” Phil. Trans. vol. 186 B.
p. 1 (1896).
EXPLANATION OF PLATE XVIII.
Fig. 1. Occiput and upper cervical vertebra of Gomphognathus kannemeyeri. Nat.
size.
Fig. 2. Under view of atlas and axis of Gomphognathus. Nat. size.
Fig. 3. Posterior view of atlas &c. of Drivachodon kannemeyeri. Nat. size.
Fig. 4. Under Baill ess 55 a Be Nat. size.
Fig. 5. Side Penne 3 # Ss Nat. size.
at., atlas; ax., axis; e.0., exoccipital; h.at., hypapophysis of atlas; 0.c., occipital
condyle; o.p., odontoid process; p.a., proatlas; 7°, r4, bs; 7.a¢., atlas rib; r.aa.,
axis rib; sq., squamosal ; z.a¢., zygapophysis of atlas; z.aa., zygapophysis of axis;
8c, 4c, 3rd and 4th cervical vertebra.
4. A Revision of the Fishes of the Genus Triacanthus.
By C. Tarn Reean, B.A
[Received January 20, 1903.]
Although six species of 7riacanthus were described and figured
by Bleeker in the ‘ Atlas Ichthyologique,’ Giinther, in his Catalogue,
recognized only three (viz.: Zr. strigilifer Cantor, biaculeatus
Bloch, and brevirostris Schlegel), and did not accept either of the
four described by Dr. Bleeker as new, but placed three of them
(viz.: Tr. macrurus, blochi, and oxycephalus) in the synonymy of
Tr. biaculeatus, and the fourth (77. niewhoft) in that of 7%. brevi-
rostris. Since then this arrangement has not been challenged, nor
has any new species of this genus been described.
Subsequent to the reading of my paper on the Plectognathi?, I
examined the specimens of Zriacanthi in the British Museum
Collection, which include Bleeker’s types, and I have come to
the conclusion that all six species described by Bleeker are valid,
although the one he called 7'r. macrurus is certainly identical with
Tr. biaculeatus Bl., a specigs not recognized by him; whilst a
seventh species, which has been generally confused with 7'r. brevi-
rostris Schleg., is now described for the first time as 7’r. indicus.
A complete revision of the synonymy has thus become necessary,
and the need for more complete diagnoses of the various species is
obvious. In the descriptions given below, which are in each case
based on several specimens, the total length is measured to the
base of the first caudal ray, the length of head to the gill-opening,
the length of the caudal peduncle from the base of the last dorsal
to that of the first caudal ray, the length of the snout from its tip
to the vertical from the anterior margin of the eye, that of the
1 Communicated by G. A. BouLENGER, F.R.S., V.P.Z.S.
2 P.Z.S. 1902, ii. p. 284.
1903.] FISHES OF THE GENUS TRIACANTHUS. 181
postorbital part of the head from the vertical from the posterior
margin of the eye to the gill-opening. Young specimens have a
shorter snout, a deeper body, and longer dorsal and ventral spines
than adults, and have been excluded from the diagnoses. In each
case the total length of the largest example here described is given.
TRIACANTHUS Cuvier.
Body compressed, covered with small rough scales; caudal
gees more or less elongate, t apering ; mouth small; teeth in
2 series in each jaw, 10 in each outer series, incisor-like ; Enos of
the inner series obtuse, rounded, 6 in the upper jaw, 2 in the
lower. Branchiostegals 6; pseudobranchize present. Spinous
dorsal with 5 rays, the first a long and strong spine; soft dorsal
with 20-25 rays; anal with 15- 20 rays ; ventral fins each repre-
sented by a strong spine, without soft rays; caudal widely forked.
Vertebree 20.
In all the species the colour is strikingly similar, being bluish
grey above, silvery below. The only tangible differences in colour
are that the membrane of the spinous dorsal fin is in some specimens
immaculate, in others wholly or partly blackish.
Key to the Species.
I. Second ray of spinous dorsal more than half as
long as the first, membrane of spinous dorsal
immaculate ; length of yest of anal half that of
soft dorsal ; D. V, 20-22; A. 16-16 ....... . 1. Dr. strigilifer Cantor.
II. Second ray "of spinous dorsal less than half as
long as the first.
A. Snout concave; pelvis between ventral spines
tapering to a distinct point posteriorly.
a. Membrane of spinous dorsal immaculate;
length of caudal peduncle 43-5}; times in
total length ; length of base of anal 1{-14
times in that of soft dorsal; D. V, 22-2
A. 16-17 .. : 2. Ti. blochi Bleeker.
6b. Membrane between first two rays ‘of spinous
dorsal blackish; length of caudal peduncle
4-5 times in total ‘Tength ; length of base
of anal 12-18 times in that of soft dorsal;
DV 23 —2ai ARGS O\ ee ed sas hdee adhe tsees 3. Tr. biaculeatus Bloch.
c. Membrane of spinous dorsal immaculate ;
length of caudal peduncle 63-7 times in
total length; length of base of anal 13-
1} times in that of soft dorsal; D. v,
21 25; A. 17-19.. 4, Tr. oxycephalus Bleeker.
B. Snout straight ; pelvis between ventral spines
scarcely narrowed posteriorly ; D. V, 24-25;
A. 18-20.
a. Membrane of spinous dorsal blackish; snout
half as long as the head; postorbital part of
head not shorter than eye-diameter ......... 5. Fr. brevirostris Schlegel.
6. Membrane of spinous dorsal blackish; snout
more than half as long as the head; post-
orbital part of head shorter than eye-diameter. 6. Tr. indicus, n. sp.
c. Membrane between first two rays of spinous
dorsal blackish ; snout Jess than half as long
as the head ; postorbital part of head shorter
than eye- -diameter De ee ee de ie mieunofu Bleeker.
182 MR. C. TATE REGAN ON THE [ Feb. 17,
TRIACANTHUS STRIGILIFER.
Triacanthus strigilifer Cantor, Mal. Fish. p. 363, pl. ix. (1847);
Bleeker, Atlas Ichth. v. p. 89, pl. cexxix. fig. 3 (1865); Giinther,
Cat. vill. p. 211 (1870).
Triacanthus longirostris Hollard, Ann. Sci. Nat. (4) 1. 1854,
p. 46, pl. 11. fig. 3.
Depth of body about equal to the length of head, about 3
times in the total length, length of caudal peduncle 43—5 times.
Snout very slightly concave, its length 13-12 times in that of the
head, eye-diameter 3-4 times, interorbital width 4-5 times.
Interorbital space concave, without distinct median ridge. Upper
edge of occipital crest in the same straight line as that of the
snout ; distance to base of first dorsal spine from posterior margin
of eye about 14 times the eye-diameter. D. V, 20-22; A. 15-16;
the first dorsal spine longer than the head, the second considerably
more than half as long as the first, the others much shorter; the
base of the anal fin about half as long as that of the soft dorsal
fin; pelvis between the ventral spines moderately broad, tapering
to a point posteriorly. Membrane of spinous dorsal immaculate.
Total length 185 mm.
Hab. Seas of Arabia and India; East Indian Archipelago.
TRIACANTHUS BLCCHI.
Lriacanthus blochi Bleeker, Nat. Tijds. Ned. Ind. ii, 1852,
p- 81; Atlas Ichth. v. p. 89, pl. ecxvil. fig. 1 (1865).
Triacanthus biaculeatus (part.) Giinther, Cat. viii. p. 210
(1870).
Depth of body about 3 times in the total length, length of head
33-3+ times, length of caudal peduncle 43-5} times. Snout
concave, its length 14-2 times in the length of head, eye-diameter
3-3? times, interorbital width 4—5 times. Interorbital space
concave, with median ridge scarcely, if at all, distinct. Upper
edge of occipital crest forming an angle of about 160° with that
of the snout; distance from posterior margin of eye to base of
first corsa! spine about 17 times as long as the eye-diameter.
D. V,-22-23; A. 16-17; the first dorsal spine longer than the
head, the others short; length of base of anal fin 17-14 times in
that of the base of soft dorsal fin. Pelvis between ventral spines
narrow, tapering to a slender poimt. Membrane of spinous
dorsal immaculate.
Total length 150 mm.
Hab, Kast Indian Archipelago; China.
TRIACANTHUS BIACULEATUS.
Balistes biaculeatus Bloch, Ausl. Fische, pl. 148. fig. 2 (1785).
Triacanthus biaculeatus Cuv. Régne An. 11. p. 152 (1817).
Triacanthus angustifrons Hollard, Ann. Sci. Nat. (4) 1. 1854,
p. 40, pl. n. fig. 2.
1903. ] FISHES OF THE GENUS TRIACANTHUS, 183
Triacanthus macrurus Bleeker, Atlas Ichth. v. p. 91, pl. cexxii.
fig. 3 (1865).
Triacanthus biaculeatus (part.) Giinther, Cat. viii. p. 210 (1870).
Depth of body about 24 times in the total length, length of
head about 33 times, length of caudal peduncle 44-5 times.
Snout concave, its length 12-14 times in the length of head, eye-
diameter 33-4 times, interorbital width 4-5 times. Interorbital
space concave, with median ridge scarcely, if at all, distinct.
Upper margin of occipital crest forming an angle of about 170°
with that of the snout; distance from posterior margin of eye to
base of first dorsal spine 12-14 times as long as the eye-diameter.
D. V, 23-24; A. 18-19; the first dorsal spine longer than the
head, the others short ; length of base of anal fin 12-14 times in
that of the base of the soft dorsal fin; pelvis between ventral
spines rather narrow, tapering posteriorly to a slender point.
Membrane between first and second rays of spinous dorsal fin
blackish.
Total length 180 mm.
Hab, Kast Indian Archipelago; Australia; China.
TRIACANTHUS OXYCEPHALUS.
Triacanthus oxycephalus Bleeker, Verh. Bat. Gen. xxiv. 1852,
p. 27, pl. v. fig. 10; Atlas Ichth. v. p. 90, pl. ecxx. fig. 3 (1865).
Triacanthus biaculeatus (part.) Giinther, Cat. viii. p. 210 (1870).
Depth of body 24-23 times in total length, length of head
about 3 times, length of caudal peduncle 64—7 times. Snout
slightly concave, its length about 14 times in the length of head,
eye-diameter 3-4 times, interorbital width 3-3} times. Inter-
orbital space flat. Upper edge of occipital crest convex ; distance
from posterior margin of orbit to base of first dorsal spine about
12 times as long as the eye-diameter. D. V, 24-25; A. 17-19;
first dorsal spine longer than the head, the others short ; length of
base of anal fin 13—1? times in that of the base of the soft dorsal
fin ; pelvis between the ventral spines broad anteriorly, tapering to
a point posteriorly. Membrane of spinous dorsal fin immaculate.
Total length 140 mm.
Hab. Kast Indian Archipelago,
TRIACANTHUS BREVIROSTRIS.
Triacanthus brevirostris Schlegel, Faun. Japon., Poiss. p. 294,
pl. exxix. fig. 2 (1846); Bleeker, Atlas Ichth. p. 94, pl. ccxxxi.
fig. 3 (1865); Giinther, Cat. vii. p. 209, part. (1870).
Triacanthus biaculeatus Bleeker, Verh. Bat. Gen. xxii. 1849, p. 6.
Triacanthus rhodopterus Bleeker, Verh. Bat. Gen. xxiv. 1852,
p. 25, pl. iv. fig. 8.
Triacanthus russellii Bleeker, t. c. p. 25.
Depth of body 24-2? times in the total length, length of head
34-32 times, length of caudal peduncle 47-54 times. Snout
straight, its length about twice in that of the head, eye-diameter
33~5 times, interorbital width about 34 times. Length of post-
184 ON THE FISHES OF THE GENUS TRIACANTHUS. [ Feb. 17,
orbital part of head equal to or greater than eye-diameter. Inter-
orbital space with a more or less distinct median ridge, with a
groove on each side of it. Occipital crest convex, moderately
elevated, becoming nearly horizontal in front of the base of the
first dorsal spine. D. V, 24-25; A. 18-20; first dorsal spine
shorter than the head, the others short; length of base of anal fin
12-12 times in that of soft dorsal fin; pelvis between ventral
spines broad, scarcely narrowed posteriorly. Membrane of spinous
dorsal fin blackish.
Total length 250 mm.
Hab. Kast Indian Archipelago, Seas of China and Japan.
TRIACANTHUS INDICUS, 0. Sp.
Russell, Indian Fishes, p. 14, pl. xxi. (1803).
Balistes biaculeatus Bennett, Fishes of Ceylon, pl. xv. (1830).
Triacanthus biaculeatus Cantor, Mal. Fish. p. 360 (1847); Day,
Fishes of Malabar, p. 260 (1865).
Triacanthus brevirostris Hollard, Ann. Sci. Nat. (4)1. 1854, p. 45,
pl. ii. fig. 1; Giinther, Cat. vii. p. 209, part. (1870); Day, Fishes
of India, p. 685, pl. clxxv. fig. 1°(1878).
Depth of body 23-2 times in the total length, length of head
31-34 times, length of caudal peduncle 47-5 times. Snout
straight, its length about 1? times in that of the head, eye-diameter
3-43 times, interorbital width 3-3? times. Length of post-
orbital part of head distinctly less than eye-diameter. Interorbital
space with a more or less distinct median ridge with a groove on
each side of it. Occipital crest elevated, its upper edge nearly in
the same straight line with that of the snout. D. V, 24-25;
A. 18-20; first dorsal spine nearly as long as the head, the others
short ; length of base of anal fin 14-14 times in that of soft dorsal
fin; pelvis between ventral spimes broad, scarcely narrowed
posteriorly. Membrane of spinous dorsal fin blackish.
Total length 220 mm.
fab. Coasts of India from Kurrachee to Penang; Ceylon ;
Andaman Is,
This species differs from 7. brevirostris chiefly in the longer
snout, the shorter postorbital part of head, and the shorter and
more elevated occipital crest. The figures given by Russell and
Bennett are excellent.
TRIACANTHUS NIEUHOFI.
* Nieuhof, Gedenkw. Zee en lantr. p. 272, fig.
Willoughby, Ichthyology, Appendix, p. 5, pl. x. fig. 2 (1686).
? Gronow, Mus. 1. p. 52 (1754-6) & Zoophyl. p. 53 (1763-81).
Triacanthus nieuhofi Bleecker, Verh. Bat. Gen. xxiv. 1852, p. 26,
pl. iv. fig. 9, & Atlas Ichth. v. p. 92, pl. cexvii. fig. 3 (1865).
Triacanthus brachysoma Bleeker, Nat. Tijds. Ned. Ind. tv.
1853, p. 128.
1 T have not been able to verify this reference.
1903. ] ON VARIATIONS OF THE SAND-VIPER, 185
? Balistes bipes Gronow, Cat. Fish, p. 37 (1854),
Triacanthus brevirostris (part.) Giinther, Cat. viii. p. 209 (1870).
Depth of body about 2} times in the total length, length of
head 33-33 times, length of caudal peduncle 41~4} times. Snout
straight, about 21 times in the length of head ; eye-diameter about
37 times and equal to interorbital width. Length of postorbital
part of head less than eye-diameter. Interorbital space with
median ridge with a groove on each side of it; occipital crest
strongly elevated, the distance from the base of first dorsal spine
to the upper angle of gill-opening rather more than the distance
from the base of the first dorsal spine to the anterior margin of
eye. D. V, 24-25; A. 18-20; first dorsal spine longer than the
head, the others short ; length of base of anal fin about 12 times
in that of the base of soft dorsal fin; pelvis between the ventral
spines broad, not much narrowed posteriorly ; membrane between
first two rays of spinous dorsal fin blackish.
Total length 126 mm.
Hab, Kast Indian Archipelago.
This species has a shorter and more declivous snout than
Tr. brevirostris, and also occipital crest more elevated, postorbital
part of head shorter and body deeper.
5. On the Geographical Variations of the Sand-Viper, Vipera
ammodytes. By G. A. BoutencEr, F.R.S., V.P.Z.S.
[Received January 30, 1903.1]
(Text-figures 27 & 28.)
The variations of Vipera ammodytes in connection with the
distribution of the species have not received sufticient attention.
Having succeeded in bringing together and carefully comparing a
series of 55 specimens from various localities, J have convinced
myself that the South-eastern specimens (Greece, Archipelago,
Syria) can be distinguished from the typical form from Austria-
Hungary, Dalmatia, Bosnia, and Montenegro, not by means of
any single absolute character, but by a combination of characters,
as shown by the following definitions :—
Forma typica (text-fig. 27 @)—Naso-rostral shield usually *
reaching the canthus rostralis, and extending considerably higher
up than the upper border of the rostral, which is usually broader
than deep (text-fig. 28 a); rostral “horn” with 3 (rarely 2 or 4)
transverse series of scales between the rostral shield and the
apex. Ventral shields 145 to 163. The dark shade on the lower
lip, if present, broken up by light bars separated by 2 to 4 labial
shields. Lower surface of end of tail usually red?, Grows to
80 centimetres.
Var. meridionalis (text-fig. 27 6)—Naso-rostral shield never
1 5 exceptions out of 30 specimens examined
2 Yellow in one specimen from the Dinaric Alps, Bosnia.
186 ON VARIATIONS OF THE SAND-VIPER. [Feb. 17,
reaching the canthus rostralis, and but rarely extending higher
up than the upper border of the rostral, which is often as deep as
broad or a little deeper than broad (text-fig. 285); rostral “ horn”
with 3 to 5 transverse series of scales between the rostral shield
and the apex. Supraciliary edge usually more prominent than in
the typical form, sometimes slightly angular, foreshadowing the
condition in V. raddit. Ventral shields 133 to 147. A more or
less distinct dark blotch on the lower lip, involving 5 or 6 labial
shields without interruption. Lower surface of end of tail yellow.
Grows to 60 centimetres.
Text-fig. 27.
Side views of heads of a, Vipera ammodytes, f. typica (Feldkirchen,
Carinthia), and 6, var. meridionalis (Athens).
Front view of end of snout, showing the lepidosis.
a. Feldkirchen, Carinthia; 6. Athens; c. Cocosu, Roumania.
T submit these definitions to the consideration of herpetologists
who may have the privilege of examining large series of speci-
mens from any single district. The material at my command is
still deficient in examples from Bulgaria, Turkey, and the Caucasus,
and I may mention that the only two specimens from Roumania
which I have been able to examine differ from both forms as here
characterized, and are on the whole intermediate between them.
The rostral shield is deeper than broad, and the naso-rostral does
not reach the canthus rostralis (text-fig. 28 ¢); one has two series
of scales on the rostral “ horn,” the other has three ; supraciliary
edge not very prominent; ventral shields 155 and 153; no light
bars on the lower lip; lower surface of tail yellow.
1903.] ON THE HABITS OF THE HOOLOCK. 187
6. Notes on the Habits of the Hoolock.
By Geo. Canpirr, M.B.Cantab.?
[Received January 20, 1903.]
The Hoolock (Hylotates hoolock) is one of the most interesting
of the family Simiidee, and is perhaps not so familiar to naturalists
at home as are some other members of the family, as, owing to its
extreme delicacy and the great difficulty experienced in keeping it
alive in confinement, it does not often find its way into European
collections. Even in the Calcutta Zoological Gardens it is difficult
to keep Hoolocks alive for any length of time. They often
succumb to pneumonia, or if they escape actual disease they mope
and die from the effects of confinement, or possibly from depri-
vation of some article of diet which in the wild state they have
been accustomed to. I have ventured therefore to submit to the
Society these short notes, made from the point of view of a field-
naturalist rather than from a scientific aspect.
The Hoolock is clothed all over with a fine soft hair, which,
perfectly black in the male, in the female shows a greyish tint,
especially over the back. This uniform dark colour is only
relieved by narrow horizontal streaks of white hair above the eyes.
The face, palms of the hands, and soles of the feet are devoid of
hair, and here the black skin is smooth and finely wrinkled and as
soft as the finest kid.
The hallux and pollex have a flattened nail, the remaining digits
have the nail laterally compressed and resembling a claw.
There is no tail. Ischial tuberosities and cheek-pouches are
absent.
When the Ape is sitting, the vertebral column presents a single
marked curve with the convexity backwards. On the ground the
Hoolock has a very characteristic gait. He goes along in a sort of
shambling waddle, with legs bowed and knees bent, the soles of his
feet applied flat to the ground with the hallux widely abducted, both
arms being carried upwards and extremely abducted as if to balance
himself. He cannot get up any speed, and invariably swings up
into the first tree he comes to, where his movements are suddenly
changed from extreme awkwardness to extraordinary grace and
agility.
He swings along to the thinnest part of a bough, or to the
slender end of a bamboo, until it bends to his weight, then with a
swing and a sort of a kick-off he flies through the air, seizing
another bough and swinging along it with the unerring accuracy of
a finished trapeze performer. I fancy he does very little walking
in the wild state, for I have never seen a wild Hoolock on the
ground. Moreover, they are only found in the dense jungle where
the ground is everywhere covered by tangled vegetation. It is
puzzling to me why these anthropoids, being so entirely arboreal
1 Communicated by F>G. Parsons, F.Z.8,
188 MR. G. CANDLER ON THE [ Feb. 17,
in habit, should be lacking in such a useful appendage as a tail.
I think, at any rate, that it points to the fact that the apes have
been developed along a line distinct from the monkeys, the earlier
traces of which line are yet to be discovered.
The Hoolocks are extremely shy, and it is most difficult to
watch them, as they are concealed by leaves high up in the tops of
the bamboo-clumps or forest trees. You may hear their cries all
round you as you ride quickly along a jungle-tract, but the
moment you leave the path or look up at them there is a dead
silence and scarcely a leaf stirs, until, tired of waiting, you move on
again.
The cry of the Hoolock is a characteristic sound in the Cachar
jungle. It is a very pleasing note, risimg and falling in intensity,
and reminding one somewhat in its rhythm of a pack of beagles
giving tongue on a scent which is waxing and waning in strength,
as a larger or smaller number of the band join in the chorus.
Tt is heard chiefly in the early morning, then all through the heat
of the day there is silence, but towards evening, as the sun sinks,
you may hearitagain. Hooloo! Hooloo! Hooloo! with the accent
on the Hoo syllable, is supposed to describe the sound, but it is
really quite indescribable in writing.
As in other species of apes, there is a special modification of
the larynx, which acts as a sort of resonating-box, and helps (I
suppose) to make the sound carry, as it does, long distances. There
is also a peculiar arrangement of the upper aperture of the larynx,
with its small and inadequate looking epiglottis, which more
resembles the arrangement in birds than the leaf-like epiglottis in
man.
As, day after day, I have ridden through the jungle, it has seemed
to me that the Hoolocks work their ground systematically in
their search for food, just as the planter plucks one section of
his tea to-day and another section on a distant part of the garden
to-morrow. For I have found them filling the air with their cries
along a particular stretch of jungle-road one day, whilst the next
day not one was to be heard; then, perhaps, a week later they are
back again in the same place. Living as they do in communities,
they are constantly on the move, and from what we know of their
great intelligence, it seems to me highly probable that their move-
ments are guided by very definite plans, and that very probably
they have some sort of government system.
There is a point about the Hoolock that strikes me as very
extraordinary, and that is the fact that he cannot swim. I had been
told this by both natives and Kuropeans, but I confess I was
somewhat sceptical about it until I tried experiments myself. We
put a full-grown Hoolock into a big tankin 10 feet of water. He
struggled helplessly, as a boy would before he learns to swim,
He sank twice, with head thrown back and arms waving franti-
cally, and we were obliged to rescue him almost asphyxiated and
choking in the most human way.
1903. ] HABITS OF THE HOOLOCK. 189
This weakness he shares with man, but I do not know whether
(or not) it has been noted in the other anthropoids.
It is a significant fact that the range of the Hoolock is bounded
by two vast rivers, the Br ahmaputra on the north and the Irawaddi
on the south. It may well be that, with his natural aversion to
water, these rivers have confined him to the comparatively limited
stretch of country he occupies. Travelling high up in the jungle,
he could swing easily across the ordinary streams which would
come in his path without having to take to the water. The
monkeys of India take readily to water, and it is a pretty sight to
see them spring out froma lofty overhanging bough and drop, one
after another, with a splash into the stream, and strike out boldly
for the further bank.
In Cachar, where these notes were written, the tea-planters
often keep Hoolocks for years, allowing them to run. loose about
the compound, and they are certainly the cleanest and most
interesting pets imaginable, offering a very marked contrast in
this respect to the red monkeys, which, chained to a pole, are so
common a feature in Indian compounds.
A Hoolock, to be tamed in this way, must be caught quite young
and not tied or shut up in any way. A native boy is generally tolid
off to watch him for a few days, and to prevent him i om bolting,
but he soon learns to come down from the trees for a a plantain,
and he will in most cases settle down to a solitary life, remaining
about the same compound for years. But chain him or restrain
his liberty in any way, and he inevitably begins to mope and pine,
and invariably dies in a few weeks. It is strange that the calls of
the wild Hoolocks, which he must hear almost daily all round him,
do not tempt him to revert to his natural life as a member of
wandering community. I imagine a Hovlock, who attempted to join
a strange band, would meet with a rough reception, anyway they
never ti ‘y to retur n to the jungle after they become tame.
Several such tame Hoolocks I have had the opportunity of
observing for some months past. Often they will be away up in
the tree-tops for days together, when nothing will tempt them
down, but when one chooses to be sociable he will come and sit
on the arm of your chair at breakfast, and never reach or snatch
things off the table: in fact his manners are unexceptionable, and
he keeps his skin beautifully clean without that exaggerated
parade of flea-hunting which makes the monkey tribe so objection-
able as pets. At sunset you may see him settle down to sleep,
jammed tight in a fork in a squatting position. In this semi-
domesticated state I notice that the Hoolock seldom uses his
voice. I suppose, leading a solitary bachelor life, he finds no
necessity for chattering or calling. With regard to the diet of
the Hoolock, Dr. Blanford, the Indian naturalist, gives a long
list, including fruit, leaves, young shoots, spiders, insects, birds’
eges and young birds. But, it seems to me, the diet of such shy
creatures must be largely a matter of conjecture, for no certain
190 ON THE HABITS OF THE HOOLOCK. (Pebuli,
conclusions can be drawn from the habits of captured specimens,
nor can we recognize as a rule substances in the stomach of shot
specimens, as we can in the crop in the case of birds. My own
observations lead me to believe that fruits and the succulent shoots
of young bamboos and other trees form the bulk of their diet.
They will certainly catch and eat certain spiders; but I have
invariably found them to refuse such insects as moths or butter-
flies, perhaps because many such insects have a bitter taste. Hggs,
too, I found they would not eat. If you give an insect or a small
bird to a Hoolock he will certainly pull it to pieces, and possibly
taste or bite it, but it by no means follows that it is one of the
regular dishes he enjoys in his wild life.
The following list of leaves and shoots which are eaten by the
Hoolock is given by Anderson :—Morugo pterygosperma, Spondias
manifera, Kicus religiosa, Beta vulgaris, Ipomea reptans, Canna
imdica.
I hope later on to supplement these short notes with some
anatomical observations on weight of brain relative to body, and on
the number and depth of the convolutions. But this is a matter
of time, for specimens are not very readily obtained. The Hindoo
coolies, who form the bulk of the population in the tea-districts
of Cachar, will never kill a Hoolock. The Kuki tribes in the
Cachar Hills, on the other hand, kill and eat them, and regard
them as somewhat of a delicacy, I believe. But even a Kuki finds
it difficult to get a shot at these creatures, so shy are they and so
active in their movements.
1903. ] ON CERCOCEBUS ATERRIMUS AND LUTRA CAPENSIS. 191
March 3, 1903.
G. A. Boutencer, Esq., F.R.S., Vice-President,
in the Chair.
The Secretary read the following report on the additions made
to the Society’s Menagerie in February 1903 :-—
The registered additions to the Society’s Menagerie during the
month of February were 57 in number. Of these 8 were acquired
by presentation, 48 were received on deposit and 1 on approval.
The total number of departures during the same period, by death
and removals, was 105.
Amongst the additions special attention may be directed to :—
1. An example of Cuvier’s Gazelle (Gazella cwvieri), deposited
by the Hon. Walter Rothschild, M.P., F.Z.S., on Feb. 9th. No
example of this rare North-African Gazelle has been exhibited in
the Gardens since 1867.
2. A Tamandua Anteater (Vamandua tetradactyla) from South
America, received on approval on Feb. 12th. This is a very
healthy and lively specimen of an animal which we have not had
living in the Gardens for some years.
3. A young male Chimpanzee (Anthropopithecus troglodytes)
about a year old, deposited by Mr. J. C. Lamprey, of the West
African Regiment, on Feb. 18th. This animal is said to have
been obtained at Kronko in the French Soudan, and was brought
home from Sierra Leone by the depositor. It is in very good
health.
4, A Frilled Lizard (Chlamydosaurus kingi) from Australia is
the second example of this species received at the Gardens, the
first having been presented by Mr. Saville Kent some seven years
ago. The present specimen was presented by Mr. H. W. Fawdon
on February 18th.
The Secretary read the following extracts from a letter addressed
to Mr. P. L. Sclater by Major C. Delmé Radcliffe from Uganda,
and exhibited the skins of a Monkey (Cercocebus aterrimus) and
an Otter (Lutra capensis) obtained by his collector Mr. Doggett,
and sent home by parcel post, at the same time :—
“YT am sending you the skins of two monkeys—one completely
prepared with bones, and the skin of the other, also skulls of both.
The animals were both female. Doggett got them for me a few
days ago, as I had sent him collecting by road here from Entebbe.
This monkey is quite new tome. I shall be glad to hear from you
if it is a new species. It is very interesting, and the skull has some
interesting features, for instance the small canines. The hair is
curiously like a Chimpanzee’s. Doggett tells me he thought from
the noise they made that they were Chimpanzees at first. Is it a
true Cercopithecus? It interests me very much, and I shall be
Proc. Zoo. Soc.—1903, Vou. I. No. XIII. 13
192 MR. W. E. DE WINTON ON [ Mar. 3,
glad of your opinion. In the same parcel I enclose a Lake Otter
skin,”
Mr. F. KE. Beddard, F.R.S., exhibited the mounted skin of the
Greater Bird of Paradise (Paradisea apoda) that had lately died
in the Society’ Gardens.
Mr. J. L. Bonhote, F.Z.8., exhibited a photograph of two tame
Hlephants in Ceylon accompanied by a baby one, which latter
showed a considerable amount of hair, especially on the forehead.
One of the old Elephants in the photograph, presumably the
mother, also showed a certain amount of hair,
With reference to a recent paper by Col. C. K. Stewart, dealing
with the real home of the Tiger, Mr. J. L. Bonhote said that he
had asked a friend of his, a Sanskrit scholar at Cambridge, as to
whether there was a word fox the tiger in Sanskrit ; the reply was
in the affirmative, the word being vyaghra (or in the Pali form,
vyaggho). This note was not brought forward against the fact
of the original home of the tiger being in the north, but to support
Mr. Thomas’s contention that it had not spread south so recently
as Col. Stewart was inclined to believe.
Professor F. Jeffrey Bell, F.Z.S., exhibited a specimen of a
Holothurian of the genus Actinopyga from shallow water off
Zanzibar, in which there was not only an oral extremity with
tentacles, as in the Cucwmaria planci described some ten years
since by Prof. Ludwig, but also an anal extremity; these additions
(whether the result of gemmation or of fission cannot at present
be said) do not occur in the same radius of the Holothurian.
The following papers were read :—
1. On a new Species of Pigmy Antelope of the Genus
Neotragus from the Cameroons District, W. Africa.
By W. E. pz Winton, F.Z.8.
[Received March 3, 1903. ]
(Plate XIX. & Text-fig. 29.)
In a collection lately received from Mr. G. L. Bates from the
Cameroons are two specimens, male and female, of a new form of
Pigmy Antelope, adding a second species to the genus Neotragus.
I propose to name this very interesting new form in honour of the
collector, who has added so much to our knowledge of the fauna
of the Cameroons country.
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1903. ] A NEW PIGMY ANTELOPE, 193
NEOTRAGUS BATESI, Sp. n. (Plate XTX.)
Size nearly half as large again as the Royal Antelope, V. pyg-
nugus. The colour darker and slightly more smoky, the feet not
so rufous, and the whitish markings of the underparts not so
pure.
The front of the face, crown of the head, and neck are dark
smoky-brown, the back and upperside of tail more rufous or bay-
coloured. All the fur annulated with dark brown and bright bay.
The type of this new species, a male, bears the following notes
on the label attached by the collector :—‘ Kfulen, Bulu Country,
Kamarun, 1500 ft. above sea. Caught in trap on edge of garden,
15 Sept. 1902.” Measurements: head and body 540 mm., tail 65,
hind foot (without hoof) 160, ear (inside from notch) 50.
Dimensions of skull of male (text-fig. 29):—Gyreatest length
107 mm.; basal length 95; greatest breadth 51 (below orbits) ;
breadth of brain-case 37:5; orbit to points of premaxille 48;
length of nasals (middle line) 33; length of frontals 33; length
of parietals 30; length of interparietals 13; greatest breadth of
Text-fig. 29.
Skull of Neotragus batesi.
nasals 20; breadth in narrowest part 9; length of horn (broken)
27, the perfect horn could not be less than 35 mm. in length ;
distance between bases of horns 25; distance between tips
(circa) 35; length of orbit 23; height of orbit 22; leneth of
upper tooth-row 32; breadth of palate between pms.' 17, between
ms.’ 20, between ms.’ 21°5,
The dimensions of the skull of the female are practically the
same as those of the male.
The discovery of this new species extends the range of the genus
Neotragus very considerably, for the Royal Antelope (V. pygmeus)
is found only on the Gold Coast from Liberia to Lagos.
13*
194 MR. E. R. SYKES ON | Mar. 3,
An examination of the skull somewhat modifies the definition
of the genus as laid down in ‘The Book of Antelopes’ by Sclater
and Thomas; for in this new species are found extra maxillo-
premaxillary vacuities 9 mm. long and 3 mm. broad, very similar
to those found in the genus Wesotragus of Kast Africa. The
horns again of the members of this genus cannot be said to be
“less than the diameter of the eye” in length; for the horns
of the type of this new species, in their broken state, are longer
than the diameter of the orbital cavity, and would undoubtedly
measure half as much again in their perfect state. These horns
are not ‘perfectly smooth,” but show slight ridges or rings in
their basal portion. The skull is otherwise fairly similar to that
of the Royal Antelope.
There can be no doubt that Bates’ Pigmy Antelope somewhat
bridges over the differences between this genus and the Kast-
African Nesotragus, and practically reduces the distinguishing
characters to those of the horns alone.
The horns of Veotragus are very small, practically smooth, and
laid back on the head in a plane with the forehead; while those
of Vesotragus may be half as long as the head or more, strongly
and closely ridged and directed upwards.
2. On the Land Operculate Mollusca collected during the
“Skeat Hxpedition” to the Malay Peninsula in 1899—
1900. By E. R. Syxgs, F.Z.S.
[Received February 2, 1903. ]
(Plate XX,")
The species of Land-Mollusks collected by the “Skeat Expedi-
tion,” though not very numerous, are of considerable interest on
account of the welcome addition made to a fauna which is as yet
but little known. They include :—
LEPTOPOMA ASPIRANS Benson.
Leptopoma aspirans Benson, Ann. Nat. Hist. ser. 2, vol. xvii.
p. 229.
Hab. Biserat, Jalor. It has been recorded from Bukit
Pondong and the Kinta Valley, having been originally described
from Tenasserim.
LAGOCHILUS KOBELTI, sp. nov. (Plate XX. figs. 13-15.)
Testa modice wmbilicata, turbinata, solidula, lirulis numerosis
cincta, periostraco brunneo leviter induta; spira conica ;
anfr. 54, conveai ; apertura modice obliqua, subcircularis, pert-
stomate duplici, incrassatulo et subrefleco, juxta insertionem
breviter sed distincte inciso. Alt. 6:8; diam. max. 7:0 millim.
Hab. Biserat, Jalor. i
1 For explanation of the Plate, see p. 199.
P.Z.S .1903 ,vol.I.P1.XX.
ee
J.Green del.et hth. The} <
Mmtern Bros imp.
MOLLUSCA FROM THE MALAY PENINSULA.
1903. ] MOLLUSCA OF THE “ SKEAT EXPEDITION.” 195
The species of Zagochilus recorded from the Malay region are
very puzzling and I think Dr. Moellendorff was quite right in
describing as Z. rollei the form that I noted (under the name of
L. townsendi) from Kelantan. The nearest ally of Z. kobelti
appears to be LZ. townsendi Crosse; I have not seen an authentic
specimen of that species, but have compared the form now described
with the description and figures given by Crosse and with a
specimen collected by Herr Grubauer, from whose collections
Dr. Moellendorff has recorded Z. townsendi as the only species
found. The shell I now name is a trifle smaller and more
elevated in proportion to the breadth, and the umbilicus is
narrower. It may be noted that the reference to Crosse’s original
paper in the ‘ Journal de Conchyliologie’ should be to p. 200 and
not p. 208 as given by De Morgan and Dr. Moellendorff in their
papers on the Perak fauna.
T have named the form after Dr. Kobelt as a trifling recognition
of his recent study of the Cyclophoride.
DITROPIS CAVERNS, sp. nov. (Plate XX. figs. 17-19.)
Testa depresso-conoidea, late wmbilicata, olivacea, tenwis, glabra ;
spira mediocriter elevata, apice eroso, sutura impressa ;
anfr. 4 (2), convent, ultimus antice vin descendens, carinatus,
carinis duabus supra peripheriam, unica ad peripheriam, et
plurimis in regione umbilicali ; apertura subovalis, peristomate
incrassatulo. Alt. 2°2; diam. max. 1:7 millim.
Hab. In a eave, Biserat, Jalor.
A single specimen only.
CyCLOPHORUS MALAYANUS (Benson).
Cyclostoma malayanum Benson, Ann. Nat. Hist. ser. 2, vol. x.
p- 269.
Hab. Gunong Inas, Perak.
Recently when cataloguing (J. Malac. ix. p. 61) a collection of
shells from Kelantan, I gave the names of Cyclophorus saturnus Pfr.
and borneensis Mete.: both the forms then recorded have occurred
in the present collection, and I have therefore again considered
the identifications. Both, according to my present view, are
erroneous, and the group is a very difficult one. The present
species, which I regard as a form of C. malayanus, was then named
C. saturnus, and the next species was called C. borneensis.
CycLoPHORUS TUBA (Sby.).
Cyclostoma tuba Sowerby, Proc. Zool. Soc. 1842, p. 83.
Hab. Gunong Inas, Perak.
See remarks under the last species; probably the C. borneensis,
recorded by De Morgan from Perak, also belongs to this species.
PYrEROCYCLOS SUBALATUS, sp. nov. (Plate XX. figs. 1, 2.)
Testa late umbilicata, orbiculato-depressa, lineis increments notata,
196 MR. E. R. SYKES ON | Mar. 3,
brunnea, stregis corners picia, fascia unica nigro-brunnea ad
peripheriam ornata ; anfr. 44, mediocriter crescentes, convext,
sutura bene notata separati; apertura subcircularis, peri-
stomate mdistincte duplicato, ala parva antice angulato.
Alt. 8; diam. max. 16 millim.
Hab. Gunong Inas, at 5000 feet.
I thought at first that this might be a form of P. blandi Bens.,
but 1t appears to be smaller, more elevate and not so widely
umbilicated, and to differ in colour. In the two specimens that I
have seen the lip is duplicated only on its outer margin, and the
wing is small and thin.
RHIOSTOMA JALORENSIS, sp. nov. (Plate XX. figs. 6-8.)
Nearly related to A. houset but larger, slightly more depressed,
and with the whorls not so tightly coiled. The tube is long and
large, reaching the body of the shell, and is bent slightly back-
wards at the junction. The disjoined portion of the last whorl is
much longer than is the case in 2. housei, as will be seen from
the figure, and is more descending. The colour resembles that
of A. houset, and a dark band is usually present at the periphery.
Operculum as usual. Diam. max. 29 millim.
Hab. Limestone Hills and Caves, Biserat, Jalor.
1 have been in some doubt as to whether this is not a local race
of R. housei, but the differences are constant in the specimens
examined and, I think, are of specific value. The porcellaneous
appearance of the operculum, so often seen in this group, appears
to be due to the wearing down of the outer layers.
RHIOSTOMA, sp.
Hab. Kwala Aring, Kelantan.
A single specimen which agrees well with &. jousseawmer
De Morgan, save that the tube is placed a little further back from
the mouth, and is thinner and more cylindrical. It appears safer,
however, to await further material ere describing this form.
OPISTHOPORUS PENANGENSIS Stol.
Opisthoporus penangensis Stoliczka, J. As. Soc. Bengal, vol. xl.
pt. 2, p. 265, pl. x. fig. 7.
Hab. Kwala Aring, Kelantan.
A single specimen, which I refer to this species with some doubt.
RHAPHAULUS ASCENDENS, sp. nov. (Plate XX. figs. 11, 12.)
Testa pupoidea, anguste umbilicata, fusco-purpurea, dense cost-
lato-striata, spira bene elevata, apice obtusulo; anfr. 6, plano-
convert, penultimus .gibbosus ; apertura subcircularis, intus
pallide fusca; peristoma pallide corneum, expansum et reflecum,
marginibus callo junctis ; tubulus brevis, incrassatus, w sutura
oblique ascendens. Alt. 18:5; diam. maa. 9°5 millim.
Hab. Patalung.
1903.] MOLLUSCA OF THE “‘ SKEAT EXPEDITION.” 197
A single specimen “from rotten wood.” Recalling in shape
A. perakensis Smith, the present species is larger and stouter, and
the tube slants obliquely upwards instead of descending ; as com-
pared with 2. lorraini Pfr., which also has an ascending tube, the
form is not so cylindrical, the whorls are flatter, and the size is
much greater.
RHAPHAULUS PERAKENSIS Smith, var. JALORENSIS, n. var.
(Plate XX. figs. 9, 10.)
Shell very similar to 2. perakensis Smith, but a little thinner
and paler in colour, and the tube is bent along the outer lip, being
attached to the lip, and not diverted behind it as in typical
R. perakensis. The tube is broad and short. This form may prove
to be a distinct species, but bearing in mind the great variation
shown by Col. Godwin- Austen (Moll. India, vol. i. p. 202, pl. xlvii.)
to exist in A. blanfordi Benson, and also considering how little we
know of the group, most forms having been described from very few
specimens, I have thought it wiser to give only a varietal name.
Hab. Bukit Bisar, on the borders of Jalor, altitude 2000 feet.
A single specimen.
SCHISTOLOMA ANOSTOMA (Benson).
Cyclostoma anostoma Benson, Ann. Nat. Hist. ser. 6, vol. x.
p. 269.
Hab. Belimbing, Ligeh.
SCHISTOLOMA SECTILABRUM (Gould).
Cyclostoma sectilabrum Gould, Boston Journ. Nat. Hist. vol. iv.
p. 459, pl. xxiv. fig. 10.
Hab. Ula Selama, Perak.
Pupina Lowr De Morgan.
Pupina lowi De Morgan, Bull. Soc. Zool. France, vol. x. 1885,
p. 414, pl. vii. fig. 3 (lowrt on plate).
Hab. Gunong Inas, Perak.
PUPINA AUREOLA Stoliczka.
Pupina aureola Stoliczka, J. As. Soc. Bengal, vol. xli. pt. 2,
p. 267, pl. x. figs. 11, 12.
Hab. Jalor, a single specimen.
Agrees well with Stoliezka’s figure 12, but his figure 11 looks as
if it might belong to a different species.
ALyc2&us THIEROTI De Morgan.
Alyceus thieroti De Morgan, Bull. Soe. Zool. France, vol. x.
1885, p. 403, pl. vii. fig. 6; Moellendorff, Proc, Zool. Soc. 1891,
p. 342.
Hab. Belimbing, Ligeh, a single specimen.
198 ON MOLLUSCA OF THE “ SKEAT EXPEDITION.” | Mar. 3,
Atyca{us DIPLOCHILUS Moellendorff.
Alyceus diplochilus Moellendorft, J. As. Soc. Bengal, vol. lv.
pt. 2, 1886, p. 313; Proc. Zool. Soc. 1891, p. 342, pl. xxx. fig. 8.
Hab. Cave near Biserat, Jalor.
ALYCHUS CONFORMIS Fulton.
Alyceus conformis Fulton, Ann. Nat. Hist. ser. 7, vol. 1x. 1902,
p. 68.
Hab. Kwala Aring, Kelantan.
ALYCHUS PERAKENSIS Crosse.
Alyceus perakensis Crosse, J. Conchyl. vol. xxvii. 1879, p. 206,
pl. xii. fig. 7.
Hab. Biserat, Jalor, amongst rocks.
OPISTHOSTOMA ANNANDALEI, sp. nov. (Plate XX. figs. 4, 5.)
Testa conico-pyramidalis, perforata, pallide rufescens, spira bene
elevata, apice obtuso; anfr. 7, conveai, primi leves, reliqui
distante lamellati, ultimus constrictus, retroversus, ascendens,
conspicue solutus ; apertura rotundata, peristomate leviter
incrassato. Alt. 2°2; diam. max. 2°8; diam. min. 1°6 millim.
Hab. Jalor.
A single specimen, found by Mr. Annandale in débris on the
floor of a cave. Related to O. laidlawi Sykes, from Kelantan, but
the present species is larger, and the spire is much more produced,
thus becoming more cylindrical in appearance.
DIPLOMMATINA SKEATI, sp. nov. (Plate XX. fig. 3.)
Testa sinistrorsa, vix rimata, ovato-fusiformis, aurantio-rufa,
solidiuscula ; anfr. 64, conveat, primi leves, reliqui costulis
parvis subremotis regulariter sculpti ; sutwra bene impressa ;
apertura fere circularis, peristomate incrassatulo, bene reflexo,
subalato ; lamella columellaris parva. Alt. 3:1; diam. max.
1:9 millim.
Hab. Gunong Inas, Perak, at about 5000 feet.
Only a single specimen of the shell, the most salient features of
which are the inflated whorls, deeply cut suture, and fine, regular
costulations.
DIPLOMMATINA LAIDLAWI, sp. nov. (Plate XX. fig. 16.)
Testa sinistrorsa, rimata, ovato-fusiformis, corneo-albida, apice
rubello ; anfr. 54, convext, primi leves, reliqui costulis parvis
remotis regulariter sculpti, interstitiis sub lente dense spiraliter
striatis; apertura quadrato-circularis, peristomate duplica,
expanso, margine columellari sinwato, subalato; lamella
columellaris minima, indistincta. Alt. 2; diam. maz.
1 millim.
Hab. Gunong Inas, 3000-4500 feet, Perak.
Found “ on the under surface of dead leaves, young with adults.”
1903.] ON THE CALLOSITIES ON THE LIMBS OF THE EQUID. 199
Much smaller than the last species, the costule more distant, and
densely marked with minute spirals. I can find no trace of these
latter in D. skeati, but the single specimen was not taken alive.
GEORISSA MONTEROSATIANA Godwin-Austen & Nevill.
Georissa monterosatiana Godwin-Austen & Nevill, P. Zool. Soc.
1879; p2729) pl. xs fie) 6:
Hab. Perak.
EXPLANATION OF PLATE XX.
Figs. 1,2. Pterocyclos subalatus, p. 195.
3. Diplommatina skeati, p. 198.
4,5. Opisthostoma annandalei, p. 198.
6, 7,8. Rhiostoma jalorensis, p. 196.
9,10. Rhaphaulus perakensis, var. jalorensis, p. 197
11,12. Rhaphaulus ascendens, p. 196.
13, 14, 15. Lagochilus kobelti, p. 194.
16. Diplommatina laidlawi, p. 198.
17, 18,19. Ditropis caverne, p. 195.
3. The Significance of the Callosities on the Limbs of the
Equde. By R. LypexKer, F.Z.8.
[Received February 5, 1903. ]
The question as to what structures in other mammals are
represented by the callosities on the inner sides of all the limbs of
the Horse, and those of the hind-limbs of the Kiang, Ass, and
Zebras, 1s one which has attracted the attention of many naturalists,
nearly all of whom appear to be in accord in regarding them as
vestigial structures. The late Sur W. H. Flower, for instance, in
his work ‘ The Horse’ expressed the opinion that these structures
are degenerate glands; pointing out at the same time that the
so-called ergot on the hinder aspect of the horse’s pastern appears
to represent one of the pads, or cushions, which are still functional
in the foot of the 'Tapi.
In his volume on ‘ Mammalia” in the ‘Cambridge Natural
History’', of which the preface is dated February 1902, Mr.
Beddard gives a qualified support to this gland-theory ; stating in
one passage that the equine front callosities probably correspond
to the carpal glands of several other mammals, although on
another page their glandular nature is questioned. In an
apparently later communication? the same gentleman suggests
that the callosities on the fore-limbs of the Hguide may represent
a carpal sense-organ, vestiges of which he believes to survive in
the carpal bristles of the Dassies. The degeneration of such an
organ would, it is urged, very likely result in the formation of
structures resembling those under consideration.
1 Pp. 12, 13, and 240.
2 Proc. Zool. Soc. 1902, i. p. 135.—I am indebted to Mr. Beddard himself for
directing my attention to this passage.
200 MR. R, LYDEKKER ON THE CALLOSITIES [ Mar. 3,
There has, however, long existed an idea that the equine callo-
sities are remnants of a vanished toe. Somewhat analogous to
this idea is a theory, which has been advanced in a paper lately
read before. the Royal Society of Edinburgh by Prof. Ewart’.
That gentleman, it appears, has based his investigations to a
very great extent on the condition obtaining in the fetus; and
has arrived at the conclusion that the callosities in question have
nothing to do with glands, but that they represent certain foot-
pads of polydactyle mammals. According to Prof. Kwart’s view,
the fore-callosity of the Horse is homologous with the supplemental
pad on the fore-foot of the Dog, while the hind callosity (which is
wanting in all existing Hquidw save the true Horse and the so-
called Hguwus przewalskii) corresponds to the hinder plantar pad
of the Banded Anteater (A/yrmecobius fasciatus). Such correlations
will, I venture to think, scarcely be taken seriously by the great
majority of zoologists ; and I shall therefore confine my attention’
to the question whether the identification of these callosities with
foot-pads generally is well founded. In all cases my observations
are confined to the condition obtaining in adult animals. It may
be added that I purposely brought these observations to the
notice of the Society before the publication of Prof. Ewart’s paper,
since I treat the subject from a different standpoint.
In the first place, the callosities on both limbs of the Horse are
situated on the inner surface, whereas, if they represented
vestigial foot-pads, their position should be, primd facie, on the
hinder. aspect, as is the case with the ergot. It might, indeed,
be argued that they have changed their original. position, but
of such a shifting there is no evidence in the adult. A second,
and perhaps more important, objection to the foot-pad theory may
be drawn from the fact that the callosities in the fore-limb are
situated above the so-called knee-joint (carpus), and are therefore
altogether higher up than any of the foot-pads of plantigrade
mammals. Unless, therefore, another shift of position has taken
place, the fore-callosities do not represent foot-pads. This argu-
ment, it may be mentioned, was used by Sir W. H. Flower to
disprove the theory that the callosities are remnants of the lateral
digits.
The hind-callosities, on the contrary, are situated a short distance
below the joint of the hock (tarsus), and are therefore on a part
of the limb, albeit on its inner side, which is included in the foot
of a plantigrade mammal. If, however, the front callosity be
regarded as serially homologous in a general sense with the hind
one—and this is an integral part of Prof. Ewart’s theory,—it will
be evident that in the event of the former not being a foot-pad,
the same will hold good for the latter.
A third, and perhaps stronger, objection may be urged against
the foot-pad theory. On the assumption that the callosities of the
existing Hquide are vestiges of foot-pads, it is clear that these
1 See ‘ Nature,’ vol. lxvii. p. 239 (1908).
1903. | ON THE LIMBS OF THE EQUIDA. 201
structures must have existed in the ancestors of that family ever
since the time when these ancestors were plantigrade. But, so far
as I know, no ungulate was ever wholly plantigrade in both feet; the
nearest approach to this condition obtaining in the Lower Eocene
Coryphodon, in which the hind-limb was wholly plantigrade, while
the front one was partially digitigrade. It has thus to be assumed,
on the foot-pad hypothesis, that the front callosities of the Horse
have been functionless structures from a period antedating the
evolution of the Ungulata. Such a persistence, on exposed parts
of the body, of a wholly functionless structure seems very im-
probable, especially when the modifications are borne in mind
which, on this hypothesis, the horse-line must have undergone
since the time when the callosities were functional str uctures.
Perhaps the case of the ergot may be cited against this argument ;
but it should be remembered that this structure certainly acted
as a functional pad at a much later stage of evolution than could
possibly have been the case with the callosities.
Having now stated what appear strong objections, so far as the
adult is concerned, against correlating the callosities of the Horse
with the foot-pads of polydactyle mammals, it remains to consider
whether they can be identified with any other structures. Those
familiar with the morphology of the Cervide will be aware that a
certain number of representatives of that family—notably the rein-
deer, the White-tailed Deer, the Mule- Deer, and, in a rudimentary
condition, the Elk,—are furnished on the inner side of the hock
with a glandular tuft corresponding very closely in situation with
the hind-callosity of the Horse. In fact, the only difference in the
position of the two structures is that the tarsal tuft of the Deer
in question is placed rather lower on the hock. From the fact of
its oceurrence in Deer so widely separated from one another as
are the species mentioned, it seems evident that the tarsal gland
(which is doubtless a scent-organ) is a very ancient structure,
which was present in all the ancestors of the group, but has been
lost, probably from disease, in the great majority of Old World
forms.
Judging from their position, there would seem to be a certain
probability that the hind-callosities of the Horse and the tarsal
gland of the Deer are homologous structures.
With regard to the homology of the fore-callosity of the Zquide,
it may be mentioned that many Gazelles have tufts of hair
(“‘knee-brushes”) at the knee (carpus), which are probably
glandular in origin. And it is possible (if the suggestion with
regard to the hind-callosities hold good) that these may represent
‘he fore-callosities of the Horse, for there seems no good reason
why the position of a gland should not have somewhat shifted in
two widely separated groups of mammals. Then, again, we have
the carpal bristles of certain mammals, such as the Coatis and
Dassies, already referred to as being regarded by Mr. Beddard as
the remnants of a “scent-organ,’—a structure probably not far
removed in its nature from a gland. The occurrence of these
202 ON THE CALLOSITIES ON THE LIMBS OF THE EQUIDA. | Mar. 3,
bristles in the Dassies (Procavia) is very important. Mr. Beddard
states that these are the only ungulates in which he has found
these bristles. Carpal callosities are, however, described by
Dr. W. Leche’ as occurring in Wart- Hogs (Phacocherus); although
they are stated by their deseriber to be acquired, and not primitive
structures. Whether the latter statement is calculated to modify
Mr. Beddard’s opinion with regard to the nature of the carpal
bristles in the Dassies, I am, of course, unable to say. Of special
importance is the occurrence of bristles in these structures, since,
even if hairs be found to exist on the callosities of foetal Hquide,
this would be no bar to the supposition of their glandular nature.
As regards the structure of the callosities themselves, it may be
noted that in the Horse both pads are of a distinctly warty nature,
and that the hind pair are certainly in a more decadent condition
than the other, being in fact on the verge of disappearing. In
the Zebras, on the other hand (in which the hind one has been
lost), the fore-callosity is larger and much less warty and also
situated higher up. In dried skins it is, in fact, much more like
the pale glandular patch of skin below the ear of a Reedbuck’.
In this connection we have to bear in mind not only Mr. Beddard’s
observations alluded to above, but likewise others by Mr. Bland
Sutton’, in which it is pointed out that in certain Lemurs decadent
glands are actually converted into bunches of spines, which are
practically almost the same as warts; that is to say, they are
hypertrophied growths of somewhat abnormal dermal tissue.
Hence there seems no primd facie reason why the callosities of
the Hquide should not be decadent glandular structures, the
decadence being more marked in the two pairs of callosities of the
Horse than in the single pair of the Asses and Zebras.
There is, however, another point which may have an important
bearing on the subject. From the presence of a depression in
the skulls of Wipparion, Hippidiwm, &e., it is evident that
primitive Horses were furnished with face-glands comparable to
those of Deer; such glands probably having a function somewhat
analogous to that of the scent-glands on the limbs of the latter.
If, then, the existing Hquwide have got rid of their face-glands, as
being (perhaps on account of change of habit) useless, it is con-
ceivable that, for the same reason, they may have also discarded
their limb-glands.
If these suppositions (and they are but suppositions) be well
founded, it follows that a tarsal and a carpal gland must have
existed in the common ancestors of the Horses and Deer ; that is
to say, in the common stock of all modern Ungulates save the
Elephants and perhaps the Dassies. And it may be urged that if
this were the case, traces of such glands ought to be met with in
Tapirs, Rhinoceroses, Pigs, Hippopotamus, &. So far as I am
1 Biol. Centralblatt, vol. xxii. p. 79 (1902).
2 It would be important to examine the histological structure of .the callosity in a
Zebra.
3 Proc. Zool. Soc. 1887, p. 369.
1903.] ON A FOSSIL OSTRICH FROM THE ISLAND OF SAMOS, 203
aware, the only instances of structures which can be regarded as
at all approaching this nature are the carpal bristles of the
Dassies and the carpal callosities of Wart-Hogs, the latter of which,
as already mentioned, are considered to be of modern origin. No
trace of any structure comparable with the hind-callosity of the
Horse has, so far as I am aware, ever been detected on the tarsus
of any of the above-mentioned Ungulates.
Tf an objection of this nature be regarded as fatal to the gland-
theory (or sense-organ-theory, for I regard the two as practically
identical) of the origin of the equine callosities, it will, I think,
be still more so to the foot-pad hypothesis, since short-limbed and
polydactyle mammals ought to have retained traces of ancestral
foot-pads for a greater period than long-limbed monodactyle forms
like the Hqwde.
In conclusion, I may state that it has been my object, not so
much to attempt to show what the equine callosities represent, as
to demonstrate, from paleontological considerations, the impro-
bability of their being vestigial foot-pads.
P.S.—I am informed that if a callosity be pared down, and a
finger moistened with the resulting exudation held to a horse’s
nose, the animal will follow anywhere. If this be true, it affords
strong testimony in favour of the gland-theory.
4. Note on some Remains of Struthio karatheodoris Maj. of
the Island of Samos. By Rupotr Martin, of Basel
University.’
[Received February 11, 1903}.
(Text-figures 30-34.)
In the Catalogue of the collection of Mr. Barbey at Valleyres
s./Orbe (Switzerland), published in 1894 by Dr. Forsyth Major *,
besides a great number of mammals there is mentioned the femur
of a ratite bird, which, no doubt, belonged to the genus Struthio
(Dr. Major could scarcely find any difference), Dr. Major,
considering the geological age of the deposits in which the bone
had been found (Upper Miocene), and recognising that it belonged
toa form different from the recent Ostrich, created the new species
Struthio karatheodoris.
Some time ago, Dr. Major received the fragment of a pelvis
from the Museum of the Vienna University (found in the same
place as the femur), which probably belonged to the same species,
and which is the subject of the following remarks.
I have to thank Dr. Major for having entrusted me with the
study of this pelvis and for having placed two photographs of the
1 Communicated by Dr. C. I. Forsyrm Maysor, F.ZS.
2 Le Gisement ossifére de Mitylini et Catalogue d@’ossements fossiles recueillis a
Mitylini ete., Lausanne, 1894.
204 MR. RUDOLF MARTIN ON A FOSSIL [Mar. 3,
femur (rostral and caudal aspect) at my disposal, on which the
following description is based.
Evidently this femur (text-figs. 30, 31) belonged to a large
struthious bird, as it differs little from the same bone of a modern
Ostrich, The size of the femur is somewhat larger than that of
the same bone of recent Ostriches at my disposal, but very probably
there would be no difference between a large specimen of Struthio
camelus and the fossil.
Text-fig. 30. Text-fig. 31.
+ nat. size.
Text-fig. 31.— _,, i u a Rostral view. 4 nat. size.
Text-fig. 30.—Right femur of Struthio karatheodoris. Caudal view.
The following table gives the results of a comparison between
the fossil in question and that of a medium-sized modern
ostrich :—
Length of Length ofcaput Smallest Width of
Species. femur. + collum. width. trochlea.
Str. karatheodoris 36°0 10°5 5D 12:4 mm.
Str. camelus ...... 31-2 10:0 52 11:5 mm.
The measurements of the caput + collum have been taken
1903. ] OSTRICH FROM THE ISLAND OF SAMOs. 205
from the apex of the caput to the margin of the trochanter, and
that of the trochlea in the direction of its greatest expansion,
A close study of the bone, however, revealed some features in
which the fossil differs from the recent Ostrich, on the sup-
position that the photograph is not distorted.
In the fossil, the neck supporting the head of the femur is much
stouter than in the recent species and is much less constricted.
This is best seen in the form of the distal border of the collum.
Although measurements may include an error of, say, ;/5, they
give an idea of that feature, because the difference between the
two species exceeds that error. The only measurable line is the
plummet in the apex of the head to the linea aspera, dividing
the rostral surface of the neck from that of the bone itself.
Width of the neck = 1.
Struthio camelus _
3 haratheodoris = :
It seems to me that the groove in which the ligamentum teres
is inserted is shallower in the fossil than in Struthio camelus. It
may be that the photograph gives a wrong impression, and I think
it would be better not to regard this feature as a specific one.
Co|1c00}~1
Text-fig. 32.
Pelvis of Struthio karatheodoris. Ventral view.
a & 6 = true sacral vertebra. i., ii., & iii. = first three postsacral vertebra.
+ nat. size.
206 MR. RUDOLF MARTIN ON A FOSSIL | Mar. 3,
The trochanter and the line aspere of the proximal end of the
femur are not to be distinguished from the same parts in the
recent Ostrich ; even the pneumatic foramen in the caudal surface
is found in the same place.
The middle part of the bone is just as in S. camelus, nd the
distal end differs only in one particular: the fossa intercondy-
loidea is much elongated, so as to form a slight valley in the distal 4
or 2 of the rostral surface of the femur. In the caudal surface,
immediately above the proximal border of the trochlea, there is
also a funnel-shaped pneumatic foramen which enters the bone
obliquely (from the mesial border of the bone, and is directed
towards the middle of the axis of the trochlea). As in the recent
Ostrich, there is a tuberosity between the wide orifice of that
foramen and the mesial edge of the caudal surface.
The pelvis (text-figs. 32-34) differs in some respects rather
considerably from that of the recent Ostrich; but the individual
Text-fig. 33.
Pelvis of Struthio karatheodoris. Worsal view.
variation met with in the recent form, and very probably also in
the fossil, shows that the difference is much less than it seemed to
be at first sight, and very probably the extreme variations of
the two forms closely approach.
1903. ] OSTRICH FROM THE ISLAND OF SAMOS. 207
The pelvis at my disposal is represented only by a fragment
from the last presacral to the third postsacral vertebra (6 vertebrae,
the first and the last incomplete), and in connection with it the
corresponding parts of the iia, i.e. the acetabular and immediately
postacetabular region. The acetabula themselves are not entirely
preserved, but only the caudal border of that on the left is quite
intact, and the antitrochanter is broken away on both sides, so
that on the left side only its outlines can be determined.
Text-fig. 34.
Lateral view of the pelvis of Struthio karatheodoris. + nat. size.
In the principal and most characteristic features, the fossil
does not differ from S. camelus, and there can be no doubt that
the ratite bird of Samos belonged to the same genus,
In the following features the fossil pelvis agrees with or differs
from that of the recent Struthio :—
The pelvis is compressed from both sides in the same manner
as in Struthio camelus (though the compression is a little less
marked), so that the dorsal aspect of it is much as in the
recent form.
li. The sacral vertebre of the fossil are much stouter. The
same irregular sculptures occur, forming rough longitudinal
ridges.
iii, The centra of the two “ true sacral vertebre” are, compared
with the following centra, narrower, but better rounded
than in the recent Ostrich.
iv. The acetabulum has quite the same position as in SS. camelus.
v. The plane between the ventral border of the antitrochanter
and the dorsal edge of the foramen obturatorium is—in
comparison with S. camelus—turned forward, i.e. it slopes
more rapidly towards the acetabulum.
vi. The outlines of the antitrochanter ave just as in S. camelus,
and its size is also the same.
vii. The ilia of the fossil and the recent Ostrich are closely
similar. There is no difference in the dorsal view of the
pelvis exceeding possible individual variation. The vertical
Proc. Zoot. Soc.—1903, Vou. I. No. XIV. 14
208 MR. RUDOLF MARTIN ON A FOSSIL [ Mar. 3,
distance between the antitrochanter and the lateral edge
of the area dorsalis is relatively longer in the fossil than
in most of the recent medium-sized Ostriches; but on one
side this distance cannot exactly be determined in the fossil
(because the antitrochanter is broken away and the edge
of the area dorsalis is much rounded), and on the other side
the individual variation in this regard in S. camelus is very
great. The largest individuals of this species differ but
little from the fossil, and very probably the fossil pelvis at
my disposal must be referred to an adult and rather large
animal.
The distance between the dorso-mesial borders of the ilia is
somewhat shorter in the fossil than in most of the pelves of
S. camelus with which I have compared it. But it is in this respect
that individual variation has the greatest amplitude, and in the
largest pelvis of S. camelus at my disposal this distance is equal
to that of the fossil.
S. karatheodoris. S. camelus.
Costal process of the Ist “true sacral
vertebra” to the distal border of the
2nd postsacral vertebra ..............-..+.-- 78cm. 9-2 cm.
Width of the centra of the “true sacral
VOTED IAA Ase ae Lek A sea AL aa ate 2:5 1:8
Width of the centrum of the 2nd postsacral
Aifer Mee] OV ee AN) Mec AMC A eer a RMR a ey 2°8 Died
Greatest width of the dorsal surface of the
WOLVAS S58 Yc arthur ae eatc i enenmen URINE Birr 12 12°2
Plummet in the dorsal border of the anti-
trochanter to the median plane ......... ca. 12 12
Plummet in the dorsal border of the anti-
trochanter to the plane of the dorsal
Suistace iofibhenpelvisiees ae nsaee-eeeeeeeaaee ca.6°5 4
The results show that there are great differences in the pro-
portions of the pelvis of the two species which justify a
separation of the fossil from the recent Ostrich. The individual
variation, however, does not allow us to place the two forms in
different genera, because it greatly reduces the differences above
mentioned.
I have said that the separation of the two Ostriches in different
species was justified. On the other hand, the question arises,
whether or no the differences between the two forms be not the
consequences of a pressure acting during the great tectonic trans-
formations in the region between Asia Minor and Greece. A
study of the other fossils of Samos, however, gives no evidence of
such a force ; there is only a lateral compression to be seen, a result
of the pressure of the weight of the more recent strata.
These considerations add to the importance of the features which
I have selected for special comment, whilst on the other hand other
1903. ] OSTRICH FROM THE ISLAND OF SAMOS, 209
features lose their systematic value. But I have shown that the
latter had really already lost their importance because of their
individual variation.
The species Struthio karatheodoris Maj. is therefore to be kept
separate.
What are the relations between S. karatheodoris and S. asiaticus
from the Siwalik Hills? I can now give further information
on S. asiaticus. A comparison of the fossils with the figures on
the unpublished plate R of Falconer’s ‘ Fauna Antiqua Sivalensis,’
and those accompanying Davies’ ' paper and Lydekker’s” deseription,
did not reveal any differences, except that the drawings in
Falconer’s Atlas are much better than the others. Davies found
that the principal difference between S. asiaticus and S. camelus lies
in the greater stoutness of the cervical vertebre in the former ;
and Lydekker kept the two species separate only on account of
this feature. The answer to the question, whether the greater
stoutness of the sacral vertebrae on one side and that of the cervical
vertebree on the other indicates a special relationship, cannot be
given now. There is no evidence for such relations; and tor
my part, consider this character in the fossil forms (compared with
the modern Ostriches) only as more primitive.
There may be reason to unite the two forms in one species ; but
considering their different geological ages’, I think it will be better
not to do so. But there can be no doubt that Struthio asiaticus
is in direct relation with S. karatheodoris, i.e. that the former is
a descendant of the latter, as Dr. Major‘ supposes the whole
Siwalik fauna to be a later and transformed generation of the upper
Miocene fauna of Pikermi and Samos.
It is noteworthy therefore that S. karatheodoris and S. asiaticus
give us no evidence for a specialisation of the Struthionids in
Southern Eurasia’, and a consequent emigration into Southern
Kurope, Syria, and Africa, but support rather the view that the
order of migration took place in the opposite direction. However,
the genesis of the Struthionid cannot, as Burckhardt © is inclined
to suppose, be associated with the Miillerornithide; for the
geological age of the former is opposed to such an hypothesis.
By the discovery of a Sérauthio in the island of Samos, the ege
on which the species of Struthio chersonensis Brandt™ has been
based is of some interest. Though the circumstances surrounding
its discovery may be somewhat obscure, yet the size and proportions
show that it cannot be the egg of a modern Ostrich; that it really
1 Davies, Geol. Mag. 1880, 2 Lydekker, in Paleontologia Indica, 1884-86.
3 Lydekker, Fossil Vertebrata of India: Records Geol. Survey, India, vol. xx. 1887.
4 Op. cit. Comptes rendus des Séances de l’Acad. d. Sci., Oct. 1888. (An account of
the fauna of Samos.) Atti della Societ’ Toscana di Scienze Naturali, Proc.-verbali,
vol. v. 3 July, 1887, p. 272. (Letter addressed to Prof. Meneghini, in which
Dr. Major shows the difference between the mammalian remains of Samos and tho
of Kos.) be
® Burckhardt, “ Uber Aepyornis,”’ Pal. Abh., Jena, 1893, p. 21.
6 Burckhardt, “Das Problem des antarktischen Schépfungscentrums,” &c., Zool.
Jahrbiicher, 1902, p. 26.
7 Bull. Ac. Imp. Se. St. Pétersbourg, vol. xviii. 1873.
14*
210 MR. F, E, BEDDARD ON | Mar. 3,
is an egg of a Struthio has been proved by von Nathusius. Its
geological age cannot certainly be determined because it was found
floating ina river. Brandt supposes that it had been embedded
in the bottom of the river and had been worked out by the water.
The age of the strata forming the bed of that river is supposed to
be that of the strata in which von Nordmann? discovered the
mammalian remains in 8, Russia (environs of Odessa), and therefore
that of the breccia of Pikermi; but it may be much younger,
because Nordmann* does not separate the Tertiary from the
Pleistocene. In the paper above mentioned, he attributes the
bone-beds of 8. Russia to the “‘offenen Diluvium,” and says that it
has the same geological age as the bone-beds of the Val d’Arno.
Therefore we have no evidence of the exact geological time during
which a Struthio lived in South Russia, and the egg 18 in con-
sequence of little importance in regard to this question. Moreover,
it seems to me more than doubtful to assign this egg to the
modern Ostrich, and “very likely to the species occurring at
Samos,” as is done in Nicholson and Lydekker’s ‘Manual of
Paleontology’ (p. 1228).
5. On a new Genus and two new Species of Harthworms of
the Family Mudrilide, with some Notes upon other
African Oligocheta. By Frank EH. Bepparp, M.A.,
ID Ja sSioy JUAN
[Received March 3, 1903. ]
(Text-figures 35-38.)
The first set of specimens referred to in the following descriptions
form a part of the collection in the British Museum; I am greatly
indebted to the kindness of the Director for allowing me the
opportunity of examining them. They belong to two species,
both of which are undescribed. The first is a third species of the
genus Stuhlmannia, which I call
Stuhlmannia michaelseni, n. sp.
I have examined so many individuals of this form of Stuhl-
mannia, and the agreement between them is so close, that I have no
hesitation in regarding it as a distinct species, which I name
after the founder of the genus, Dr. Michaelsen °.
The worms were collected by Mr. 8. L. Hinde in the Mt. Kenya
district. There were thirty mature examples.
The general appearance and proportions of this new species are
quite the same as in S. variabilis. The length reaches 100 mm.
and the diameter 2-2°5 mm. The colour is a yellowish grey; the
clitellum is greyer.
1 Nordmann, in ‘ Jubileum semisaeculare Fischeri de Waldheim,’ Moscou, 1847.
2 Nordmann, Palaontologie Stid-Russlands, 1858.
3 JB. Hamb. wiss. Anst. vii. (1890), p. 24.
1903. ] NEW SPECIES OF EARTHWORMS. 211
The sete are strictly paired and the ventral pair of segment xvii.
appear to be missing. The clitellum occupies segments xiv.—xvil.
The median orifices of the spermatheca and of the spermiducal
glands are very obvious, and frequently lie wpon projecting papille.
The spermathecal pore opens far back upon the xiiith segment.
The male pore opens far back upon the xviith segment, or even
intersegmentally (xvii./xviii.). The penial setze can often be seen
to protrude from it. Among the external characters, however,
there is one which distinguishes the present species from its allies,
and that is the total absence of the penial process of the body-wall.
There was not the least trace of this organ observable in any of
the individuals. It is true that this penis is not always to be
seen in Stuhlmannia variabilis; but I do not think that it would
be likely to be absent from thirty individuals selected at random ;
and this absence is therefore one of the reasons which lead me to
create a new species.
Text-fig. 35.
Dissection of Stuhlmannia michaelseni.
xi, xii, segments eleven and twelve ; L.V., intestino-tegumentary trunks ; o.d., ovidueal
apparatus; Sp., spermathecal sac; Sp.gl., spermidueal glands.
The accompanying drawing (text-fig. 35) illustrates a general
212 MR. F. E. BEDDARD ON [ Mar. 3,
view of the viscera seen on cutting open the body-wall in the
dorsal median line. No such figure of the anatomy of this genus
has been published up to the present; and it is convenient to
show the relations of the different organs and their comparative
sizes.
There are points in the anatomy of the genus Stuhlmannia
which are illustrated by that drawing, and to which attention does
not appear to have been called. The last pair of hearts lies in the
eleventh segment, as in so many other Eudrilids, for instance in
the genus Polytoreutus ; and the septum which should separate
segments xiii, and xiv. is nearly missing. I am disposed to
associate this latter fact with the presence of the ccelomic sacs
surrounding the gut, which may have been developed at the
expense of the septum, as in Hudrilus', in which genus the
corresponding septum is also much reduced. Another matter to
which I would wish to direct attention, is the existence of intestino-
tegumentary trunks, which have not yet been recorded in this
genus. They are, of course, of wide occurrence among earth-
worms.
A further point of some little interest is visible in the sketch
exhibited herewith. In my ‘Monograph of the Oligocheeta,’ * I
pointed out the existence of at least occasional asymmetry of the
female reproductive organs shown in the presence of only a single
receptaculum ovorum, that of the opposite side of the body being
absent. In other specimens*® I found precisely the same state of
affairs. The species that I examined was, I believe, Stuhlmanmea
variabilis.
In the present species of Stwhlmannia there is exactly the same
asymmetry, the receptaculum being only present upon one side,
and that the right. Or, to be more accurate, the receptaculum 1s
possibly present also on the left side, but is quite rudimentary,
and, I imagine, functionless. The “ Hitrichterblase,” as Michaelsen
has termed it, is present on the left side, and is simply a loop of the
oviduct, the two sections of the tube running in close contact side
by side; just opposite to it is a small spherical projection. A
lumen is present in this, which is the rudimentary funnel, but
the lumen is no wider than that of the oviduct elsewhere, and
there is no question of any free communication with the body-
cavity outside; there was no break to be detected in the muscular
wall of this projection, which perhaps should be regarded as funnel
+ receptaculum.
A series of sections through the rudimentary funnel and the
adjoining parts of the oviduct confirms the appearances displayed
in a preparation mounted in glycerine. The rudimentary funnel
(or funnel + receptaculum) is but a slight protuberance, which is
traversed up to its very end by a blindly ending branch of the
1 Beddard, “On the Gonad Ducts and Nephridia of Hudrilus,’ P. Z.S. 1902,
vol. 11. p. 89.
2 Oxford, 1895, p. 602.
3 “On some Harthworms from British East Africa, &c.,” P. Z.S. 1901, vol. 1.
p- 355.
1903.] NEW SPECIES OF EARTHWORMS. 213
descending limb of the oviducal loop, which constitutes the
“ Hitrichterblase” of Michaelsen. No communication between
this short diverticulum of the oviduct and the general eccelomic
cavity could be detected. Evidently, however, it is the forerunner
of the funnel which opens into the fully developed receptaculum,
or, possibly, of both. It is noteworthy that the oviduct
is divisible into two regions. ‘The section lying nearest to the
spermathecal sac is of a wider calibre than that lying nearer to the
external orifice. Furthermore, the lumen of this wider section
is a straight tube, while the lumen of the narrower section of
the oviduct is sinuous in its course within the muscular sheath.
It is possible that the wide straight section is really to be looked
upon as a greatly drawn-out funnel, drawn out by reason of the
growth of the spermathecal sac.
The walls of the oviduct, as is the case with other Eudrilide,
are very stout and muscular; and attention has already been
called to the fact that part of the oviducal tube runs a sinuous
course, independent, therefore, of its muscular covering. ‘These
facts, coupled with the conditions obtaining in Hudrilus ', where
the oviduct runs for a considerable space actually within the
septum, lead me to consider that the muscular wall of the oviduct
in Stuhlmannia may be a purely adventitious sheath, not belonging
to the oviduct at all, but consisting chiefly of the otherwise
missing septum between segments xiii./xiv. Its continuity with
the receptaculum can at any rate be explained on this view, which
is in any case not at variance with the other facts to which I have
just called attention.
The present species, in correlation, it is to be presumed, with
the entire absence of a penial process, does not possess the median
unpaired bursa propulsoria which characterizes S. variabilis. In
the former point possibly, but not certainly, it agrees with
S. gracilis. Dy. Michaelsen states, only as a possibility, since he
only examined two specimens, the absence of a copulatory process
in S. gracilis; furthermore, a thickening of the integument of
segment xv. appeared to him to be possibly intelligible as a not
fully developed penis. With regard to the presence or absence of
a bursa propulsoria he is silent. Stuhlmannia michaelseni, however,
cannot be confused with S. gracilis by reason of the characters
of the penial sete. They are provided at the end with a row of
sharp denticles on either side as in S. variabilis. In short, I
think that a case has been made out for the creation of a new
species of this genus Stuhlmannia.
Bettonia lagariensis, n. g. & sp.
Of this apparently new genus and species [ have examined but
a single specimen, complete and sexually mature. It measures
90 mm. by 4-5 mm. in breadth. The colour is bluish violet. It
1 Beddard, “On the Gonad Ducts and Nephridia of Eudrilus,’ P. Z.S. 1902,
vol. ii. p. 89.
7
214 MR. F. E. BEDDARD ON [ Mar. 3,
was collected by Mr. Stuart Betton, at Lagari, British Central
Africa,
The prostomium is continued by grooves over about half of the
buccal segment.
The sete are wider apart in the case of the ventral couples than
in the case of the lateral.
The limits of the clitellum are a little obscure; it appears to
embrace segments xiv.—xvii. inclusive.
The chief, indeed practically the only, reason which leads me
to separate this Hudrilid generically, is the condition of the repro-
ductive apertures. The rule in the family is that the apertures
of the sperm-duct and of the spermathece are unpaired and
median in position.
Text-fig. 36.
ste
Ventral surface of Bettonia lagariensis. §, spermathecal pores; g, male pore.
There are, however, a few exceptions, such as Hudrilus itself,
and an apparently close ally of the present genus, viz. H’mznoscolex.
Betionia offers a third arrangement of the reproductive pores.
4
1903.] NEW SPECIES OF EARTHWORMS. 215
The spermathecal pores are paired, while the male pore is single
and median (text-fig. 36, p. 214).
The two former pores are upon the boundary line of segments
xil./xill.: each is somewhat eye-shaped in contour and corresponds
exactly in position to the outermost of the ventral couple of setz.
The single and median male pore lies on the boundary line of
segments xvil./xvill. It is very large and conspicuous, with
radially folded margins, indicating, perhaps, the possibility of the
extrusion of the bursa propulsoria,
The internal organs, unfortunately, were much softened, and
therefore the female reproductive apparatus was rather difficult to
decipher. The nature and relations of the other organs of the
body were not so difficult to ascertain.
As in many, if not in all, Kudrilide, the last pair of hearts
occurs in segment xi. In front of these were four distinct pairs.
The dorsal vessel is single throughout. I have not been able to
study the alimentary system in detail, but I have ascertained
that the present species is a member of the section Eudrilacea by
virtue of the existence of an unpaired ventral median alimentary
gland in segment x., which showed on a microscopical examination
the usual laminate structure of these glands, not at all to be
confounded, even in a badly preserved specimen like the present,
with the totally different though corresponding organs in the
other subfamily of the Eudrilide. There are also a pair of these
glands in segment xuli., of an oval form, which contain abundant
crystals, such as are met with in other genera.
Text-fig. 37.
Termination of the male efferent apparatus of Bettonia lagariensis.
B.p., Bursa propulsoria; Sp.gi., spermiducal glands.
The only conspicuous parts of the male generative system are
216 MR. F. E. BEDDARD ON | Mar. 3,
the sperm-sacs and the terminal apparatus, which opens on to the
exterior in the xviith segment. The sperm-sacs are in segments
x1. and xil., and occupy a considerable space in those segments.
They are simple, solid, sac-shaped structures, not racemose in form.
The spermiducal glands are paired. Hach of them (see text-fig.
37, p. 215) is shortish and rather thick, tapering somewhat towards
the tip. The last third of the gland is bent forward and lies above,
parallel to and in contact with the anterior section of the gland.
This arrangement occurred on both sides of the body. Hach
gland is sharply constricted at its opening into a large median
bursa propulsoria lying below the ventral nerve-cord. This latter
sac, however, presents obvious signs of having been produced by
a fusion of two sacs; for posteriorly it is completely double. It
is into each of these posterior lobes that the spermiducal glands
open.
My description of the female apparatus must unfortunately be
incomplete. The organs, as already stated, are paired. The
spermathece, near to where they open, have very thick muscular
walls; and this region at least is enveloped in a ceelomic sac, as 1s
the base of the spermatheca in the species of Pareudrilus (?), with
which I deal later in the present communication (see below).
How this ccelomic sac is otherwise related to the spermatheca
and to the receptaculum ovorum I am unable to state. The latter
organ presents no noteworthy peculiarities, and the oviduct leads
from it to the exterior, on the fourteenth segment.
On «a Species of PAREUDRILUS.
I believe that a number of individuals belonging to this genus,
which were collected by Mr. Crossland, may represent a new
species. But I am unable to speak with absolute certainty on the
matter, since the material was not in good order for investigation,
and since the specimens of P. papillata examined by Michaelsen *
were likewise much softened by evaporation of the alcohol ; if the
worms upon which I report here are not referable to P. papillata,
then the species is unquestionably new.
But whether the species be new or not, I have something to add
to what is known about the structure of this genus.
The dimensions of my specimens agree apparently with those
given by Michaelsen for his species. The length wassome 100 mm.
and the diameter about 3 mm. The dark purple colour, turning
to yellow below and in the clitellar region, is like that of my
Pareudrilus stagnalis. The setz are closely paired, and I observed
a tendency in the neighbourhood of the genital pores for one seta
of a pair to be lost or not developed. I do not refer to a mere
dropping out ; on examining the cuticle, occasionally no pore was
to be noticed in the place where such a pore (through which the
seta is extruded) should be. The irregularity of this state of affairs
1 “Die Regenwtirmer Ost-Afrikas,” in Deutsch-Ost-Afrika, p. 11.
1903. ] NEW SPECIES OF EARTHWORMS. 217
does not incline me to describe in detail the cases observed, which
were, moreover, not many. I can eonfirm Michaelsen’s statement
that there are no genital or any specially modified sete in any
region of the body.
In the external sexual characters I find some little differences
from the description of Michaelsen, and it is partly on this account
that I am disposed to regard my specimens as belonging to a new
species. The clitellum is perhaps a little more extensive, xili.—
Xvill. or even xix.; but that is a slight difference. I find that the
pores of the pair ed spermathece are situated definitely on the
border-line of segments xiv./xv. and not upon xiv. They are,
however, in the line of the ventral couple of sete. On the other
hand, I agree with Michaelsen in placing the pores of the spermi-
ducal glands upon segment xvii. just in front of the ventral sete
of that segment. As to the location of the genital papille, Dr.
Michaelsen and I have not found the same conditions in the
specimens studied. In my specimens, the constant rule appears to
be the presence of two pairs of papille of which the most anterior
are on segment xvii., just behind the ventral setee but on a line
with the ene most setze of the couple ; on the following segment,
the xviith, there is a precisely similar pair also behind the ventral
sete, but in this case corresponding to the innermost seta of the
couple.
The papille are fairly conspicuous, and are to be noted upon the
cuticle when stripped off. The area is slightly raised and is
studded with the mouths of large glands. Corresponding to the
papille internally, are glands to which I shall recur in describing
the anatomy. These papille were never absent.
I could discover no nephridial pores; but I imagine that this
species, like Pareudrilus stagnalis and some other Pareudrilacea,
will turn out to possess a ramified system of nephridial end-
tubes in the integument. In any case the nephridia, viewed
internally, appear to be paired structures. As to the alimentary
tract, I find a gizzard far forward, in the sixth or seventh segment,
and I have not been able to find calciferous glands, The last
heart is in xi. The funnels of the sperm-ducts are in x., xi. The
two pairs of sperm-sacs, in xi., xii., are tongue-shaped.
As to the female generative system, my observations do not
altogether agree with “those of Michaelsen ; but it is no disparage-
ment to that excellent and accurate observer to suggest that the
condition of his specimens may possibly have led him into some
slight error. Otherwise I must place my specimens not only in a
new species, but in a new genus. The ovary I have not been able
to find at all. This in an Kudrilid is not of course surprising
for as a rule the ovary disappears early, having transferred its male
to the egg-sac. On the other hand, while it is unlikely that I
could have missed so conspicuous an object as the large ovarian
sac figured by Michaelsen, it is equally unlikely that he was misled
by bulging septa or other structures into stating the presence of
such a sac communicating with the egg-sac. It seems to me
218 MR. F. E. BEDDARD ON | Mar. 3,
therefore to be possible that, after all, in spite of undoubted
resemblances, Michaelsen and I have examined different species
and indeed different genera.
The female organs of the species which I have examined show
a number of not uninteresting features. The two most salient
parts of that system, which are visible on a dissection, are the two
spermathece and the egg-sacs. The spermathece are sausage-
shaped, and distinctly divisible into two regions. The proximal
part, ¢. e. that nearest to the external orifice, is strongly muscular,
and indeed is enveloped with stronger muscular bands than is the
bursa propulsoria of the spermiducal gland. Its epithelium is
perfectly continuous with the general epidermis of the body-wall,
and it has every appearance of being formed as an ingrowth from
the exterior. The distal region of the spermatheca has very thin
muscular walls, much thinner than the walls of the glandular part
of the spermiducal gland.
The interior has an epithelium which is raised into folds. I
cannot speak of the histological characters of the cells, as the
material was not sufficiently good. There is a very close resem-
blance, on a superficial view, between this spermatheca and the
spermiducal glands. Indeed, on a cursory inspection, they
might be taken for consecutive pairs of either spermathece or
spermiducal glands. The next most obvious part of the female
reproductive system is a very large mushroom-shaped body closely
adherent to the septum dividing segments xiii./xiv. This body is
stalked, and appeared, on dissection, too large to be identified
with a receptaculum ovorum (or egg-sac). Nevertheless it is the
egg-sac, and by virtue of its large size it appears to be precisely
like the egg-sac of P. papillata described by Michaelsen. On a
closer inspection, a fine tube, apparently leading from the stalk
of the egg-sac to the muscular part of the spermatheca, was
apparent; this seems to correspond to the narrow tube (sq.) figured
by Michaelsen.
A series of longitudinal sections through the body showed more
accurately the relations of these diverse organs to each other. I
find that the spermatheca is entirely independent of the rest of
the female apparatus, and that its cavity does not communicate
with the narrow tube arising from the ege-sac. That narrow tube
exists, as I have already mentioned; but on reaching the base of
the spermatheca, 2. e. the muscular end portion, it dilates into a sac
which entirely surrounds the muscular part of the spermatheca,
but does not, so far as I could ascertain, open into it anywhere.
The conditions, therefore, are those of such a genus as Hyperio-
drilus or Helhiodrilus, where a true spermatheca is invested by a
ceelomic sac. Now, though the difference may appear to be slight,
I am disposed to think that it is important, and that a sperma-
theca which has no communication with the egg-conducting
apparatus is essentially different from a spermatheca which has
such acommunication. It seems to me, for example, to be wrong to
compare the spermathecal sac of Lybiodrilus or Stuhlmannia with
1903. | NEW SPECIES OF EARTHWORMS. 219
the spermatheca of Heliodrilus. For this reason I cannot agree
to Michaelsen’s placing of my genus dAlvania within the genus
Hyperiodrilus. The former has a true spermatheca, homologous
with that of other non-Eudrilid earthworms, while the latter has
not. ‘This also is the case with the genus or genera with which we
are dealing now. Pareudrilus stagnalis has a spermathecal sae
which communicates with the ccelomic sacs envolving the ovary ;
“ Pareudrilus” papillata has not. It is possible, therefore, that we
should revive Michaelsen’s Unyoria for the latter species. The
stalk of the egg-sac, as might be expected, lodges the funnel of the
oviduct, or, to speak more accurately, the greater part of the funnel.
The exact conditions obtaining are the following. In a series of
sections it may be seen that the narrow tube communicating with
the ccelomic sac surrounding the proximal end of the spermathec:
runs forwards and opens into the cavity of the xilith segment bya
wide orifice ; its walls are therefore continuous with, and no doubt
developed from, the septum bounding segment xiii. posteriorly,
The upper ‘ lip” of the ostium is covered by the cubical cells of
the oviduct, which here opens freely into the cavity of the xiiith
segment. Further on in the series of sections, the mouth of the sac
surrounding the spermatheca is closed, and the tube opens into the
egg-sac through a wide tube which is entirely lined by the cells of
the oviducal funnel ; these are, of course, perfectly continuous with
those cells which he in the xiiith segment. The part of the
oviducal funnel which les in the xivth segment appears to be
divided into two, and to open by as many mouths into the huge
egg-sac, which is so kidney-shaped as to be nearly divided into two
sacs; I cannot pretend to an accurate description of the funnel and
its various foldings. It is clear, however, that the conditions which
obtain are those of the more typical EKarthworms, where the funnel
opens partly freely into the xiith segment and is partly reflected
so as to open within the ege-sac. Now, the simpler forms of
Kudrilidee, such as the genus Hudriloides, are distinguished by the
fact that the ovary is unenclosed in any sac, and that the funnel
of the oviduct opens precisely as has been just described in Pareu-
drilus papillata. On the other hand, in the more complicated
forms, such as Séuhlmannia, the sacs containing the ovaries
envelop also the oviduct-funnel and communicate with the
spermathecal sac. The species which forms the subject of the
present remarks is plainly intermediate between these two
extremes; and for that reason, as I think, deserves generic
separation from Pareudrilus. i
§ On the Spermatophore.
The spermatophores of the Eudrilide have not been much
studied, and, so far as I am aware, are known only in the genera
Stuhlmannia and Polytoreutus, in which I have myself described
them. Many, if not most, of the individuals of the present species
which I examined had a single spermatophore in both the
spermathecze,
220 MR. F. E. BEDDARD ON | Mar. 3,
As will be seen from the accompanying drawing (text-fig. 38),
the spermatophore consists of a globular swelling followed by a
long thin tube. The sperm, which appears blackish in glycerine
preparations, is limited to the spherical or nearly spherical recep-
tacle at the distal end of the case. The spermatophore, as will be
Text-fig. 38.
Spermatophore of Pareudrilus sp.
also apparent from the figure referred to, is of an elegant form,
not precisely corresponding to that of the spermatheca in which
it lies. The swollen and globular receptacle of the sperm is not
more than one-third of the length of the “ stalk,” which reaches
down to the very mouth of the spermatheca. At its termination the
walls of the spermatophore project in a ring-like fashion ; a state of
affairs exactly recalling the spermatophores of the Tubificidee, and
of the genus Stuhlmannia among the Eudrilide. No doubt the ~
shape is due to a moulding upon the walls of the spermatheca ;
but the state of preservation of the specimens does not enable me
to give details. The walls of the spermatophore appear to be firm
and thick, and rather brittle in consistency. In teased prepara-
tions the rupture of the stalk was invariably a clean fracture.
The walls are fibrous in appearance, and of the usual pale brown
colour that is generally associated with chitimous membranes. The
extremity of the tube, 7. e. that which is nearest to the mouth of
the spermatheca, is open; the other end is quite blind. The
1903. | NEW SPECIES OF EARTHWORMS. 221
shape of the whole spermatophore is sufficiently elucidated by the
drawing referred to.
§ Note on the Clitellum of Alma stuhlmanni, and on a possibly
new species of the genus Alma.
I believe that a note by myself’ upon the clitellum and
spermatophores of a West-African species of the genus Alma is
the first record of the extent of the clitellum in that genus. Like
other aquatic forms, Alma seems to be characterised by a seasonal
development of the clitellum; and hitherto, with the exception
just mentioned, no one appears to have seen or at least described
this organ in that Geoscolecid. In the species to which I have
just referred, the clitellum was found to extend from segment xlv.
to Ixxxv., a position which is quite unlike that found in any other
Geoscolecid and present only in a few Lumbricids. This is an
additional reason for associating the genus dima more particularly
with Criodrilus, asis done by Michaelsen; for both these genera,
though referable to the family Geoscolecidee, have many points of
kinship to the Lumbricidze. As, however, up to the present time,
but one species of the genus Alma has been described in the fully
mature condition, it is possible that the position and extent of the
clitellum characterising that species are not normal but exceptional
in the genus. Therefore I do not hesitate to describe the con-
ditions occurring in a second species of the genus, which I owe to
the kindness of Mr. Cyril Crossland, who collected specimens on the
shore of Victoria Nyanza, among weeds cast up by the waves. I
have two fully mature examples of a species which I believe to be
identical with Dr. Michaelsen’s Alma stuhlmanni. The dimensions,
however, are rather less; only one of the two examples was quite
intact —the other had lost the hinder end of the body; in the
complete example, measurements showed a length of 120 mm.
The other might have been slightly longer, as it was rather thicker.
In both, the penial appendages were rather longer than those of the
original specimens deseribed by Michaelsen. I found them to be
10 millimetres long; in one example the two were unequal in
size, one penial appendage only measured 6 mm. The structure
of these appendages is usually characteristic of the species. The
worms which T have examined agreed in almost every detail with
the description of A. stuhlmanni as given by Michaelsen °.
I may remark, however, that there were only two sete at the
free end of the penial process, and, indeed, one of these had
dropped out. The two papille upon which these sete are placed
were partly encircled by a horseshoe-shaped region of specially
glandular epidermis, which was conspicuously marked out from
the rest of the integument covering that process. It 1s conceivably
this region which secretes the spermatophore. In addition to this,
1 “On the Clitellum and Spermatophores of an Annelid of the Genus A/ma,” Proc.
Zool. Soc. 1901, vol. i. p. 216.
2 “Die Regenwitrmer Ost-Afrikas,” in Deutsch-Ost-Afrika, 1896, p. 4,
222 PROF. NEWTON ON THE WHITE RHINOCEROS. [Mar. 17,
I fancy that the single papilla at the base of the penial process is
rather nearer to the base than is figured by Michaelsen, and much
nearer than in the West-African form, and, moreover, it appeared
to me to be not symmetrical in its position. The clitellum was
not seen by Michaelsen at all. In my specimens the clitellum
occupied the same segments; but, as in the species described by
myself from Western Africa, that generative region was somewhat
undefined in its beginning and ending. The greatest number of
segments referable to the clitellum in the West-African species
are from xlv.—lxxxv.; but the clitellum was only fully developed
upon segments xlvii—Ixxxii. Its general appearance was precisely
like that of the species which I have just mentioned. In the
present species—and the observations apply to more than one
specimen, and are therefore all the more reliable as an expression
of normal conditions—the clitellum was much shorter, only
extending from segments xl.-Ixxi. This rather leads me to the
inference that the species with which I am concerned here is in
reality different from that which I described from McCarthy Island
on the Gambia, and referred to A. stuhlmannt. The present worm
is undoubtedly A. stwhlmanni; and it seems to be necessary, on
account of the difference in the clitellum, to use another name for
the West-African form. I would propose, therefore, to call the
latter A. budgetti. It is, however, clearly a very close ally of
A. stuhlmanni. It is interesting to note that the species of this
genus go more or less in couples. The East-African A. nilotica
corresponds to my A. millsoni from West Africa, while A. stuhlmanni
seems to be nearest to the form which I propose to name A. budgetti.
At present more information is wanted about A. emint; but it
appears to be formed rather after the plan of A. mllsoni and
A. nilotica. For in those species there are special setze on the penial
processes, while in A. stuhlmanni and S. budgett: there are sete of
the same pattern as those on the body generally.
March 17, 1903.
G. A. Boutencer, Esq., F.R.S., Vice-President,
in the Chair.
The Secretary exhibited, on behalf of Prof. Newton, F.RS.,
three photographs of the White Rhinoceros (Rhinoceros simus),
sent to him with the following letter by Mr. C. R. Saunders,
C.M.G., Chief Magistrate and Civil Commissioner in Zululand :—
Kshowe, Zululand,
DEAR Sir, 6th January, 1903.
I received a letter from you in August 1900, following
on an account, written by me in ‘ The Field,’ of an interview I had
with White Rhinoceroses about that time. I did not answer
your letter at the time, hoping I should be able before long to
send you a photograph of the living animal. This, however, I
1903. | PROF. NEWLON ON THE WHITE RHINOCEROS. 223
have not yet been able to procure, although I visited the reserve
in which they lived for this special purpose last winter. Their
traces were abundant, but my time was limited, and they could
not be found. There are, I believe, about ten of these animals
living in that reserve, ond I do “Age despair of yet obtaiming
a photogy: iph of them in life, in which case | shall be pleased
to send you a copy of it.
Karly last December two of the animals (both bulls), one a very
old one and the other not full-grown, strayed out of the reserve
into one of the native locations and were killed. J obtained
three photographs of one of them, the old bull, taken by an
amateur two or three days after it had died. Lam forwarding by
the same mail as this a copy of each of these photographs, which
you ave welcome to, and which, I think, demonstrate the fact that
they were taken from a specimen of the White Rhinoceros, although
the carcass was a good deal distended,
The killing of these two Rhinoceroses was most unfortunate.
They suddenly appeared among some native kraals, and the men
went out and attacked them with spears. The young one was
see outright ; that of which the photograph was taken travelled
a long fierce after being wounded, and was not found until
some days had elapsed. Yours faithfully,
Alfred Newton, Esq. C. R. SAUNDERS
As these photographs were probably the only representations
Recently-killed Rhinoceros simus, adult ¢. Dee. 1902.
Proc. Zoou. Soc.—1903, Vou. I. No. XV. 15
224 MR. OLDFIELD THOMAS ON A NEW MONKEY. [Mar. 17,
ever taken of this animal in the flesh, it seemed well worth while
reproducing two of them (text-figs. 39, 40).
Text-fig. 40.
Recently-killed Rhinoceros simus, adult g. Dec. 1902.
Mr. Oldfield Thomas exhibited the skin of a Chinese Monkey,
which had been obtained from a hunter by Mr. Henry Brelich,
and presented by him to the National Museum. It appeared to
represent a new species, the third known, of the remarkable genus
Rhinopithecus, and was described as follows :-—
RHINOPITHECUS BRELICHI, sp. n. (Plate X XT.)
Size very large, apparently larger than either /. roxellane or
R. bieti; for the skin, though that of a female, is as large as the
male of either of the other species. Fur not abnormally elongated
in any region, the longest being on the flanks, where the hairs
may attain to about 90 mm. in length ; those of the back 50-60 mm.,
and those across the shoulders 70-80. General colour of back
glossy slaty grey, the hairs grey to their roots, with shining tips.
A prominent oval white patch, 5 inches long by 2 broad, present
in the middle line between the shoulders, its hairs white to their
roots. Crown suffused with yellowish, its hairs yellow at base,
whitening terminally, but with broad black tips; hairs of cheeks
PZ). 1908 vol. Pike
J.Smit del.et lth.
=
Mantern Bros imp.
REINO PITON CUS BRE LICHE,
1903. ] MR, OLDFIELD THOMAS ON A NEW ANTELOPE. 225
yellow with black tips; nape between the yellowish crown and the
white wither-patch pale brownish with black tips to the hairs;
neck both on sides and below blackish grey, the hairs dull whitish
basally, with black ends. Ears white, markedly contrasting with
the head. Front of shoulders and inner aspect of forearms deep
yellow, which shades into whitish along the under aspect of the
latter, and contrasts markedly with the dark slaty of the outer
side of the forearm, this colour darkening to black on the wrists.
(The hands are lost in the specimen, but are presumably black.)
Hind limbs light greyish, more or less suffused with yellow behind
and blackish in front, but the colour contrasts are not sharp and
defined as they are in the other species. Belly uniformly grey
(about grey No. 5 of Ridgway). Tail very long, conspicuously
longer than in the other species, its hairs, which average about
40 mm. in length, curiously curved on each side downwards and
away from the centre line, along which there is an irregular
parting; in colour the tail is black throughout except at the
extreme tip, where there is a small white pencil; on each side of
its base there is a small yellow patch, outside of which there is a
blackish line passing round across the anal region: but owing to
the condition of the skin, the exact situation of these lines and
patches is not quite certain.
Approximate dimensions of the type, measured on the skin,
which has been made up from a flat native pelt :—Head and
body 730 mm., tail 970 (with hairs 1040).
Habitat. Mr. Brelich states that, ‘as far as I could gather, this
monkey inhabits a range of mountains known as the Van Gin
Shan Range, about 108° E., 29° N., in the north of the province
of Kwei-chow, Central China.”
Type. Female. B. M. No. 3.3.14.1. Collected and presented
by Henry Brelich, Esq.
This magnificent Monkey, one of the largest in the world apart
from the anthropoids, is a very remarkable discovery, and one on
which we may congratulate Mr. Brelich, who obtained and sent
it to the Museum on the suggestion of Mr. Herbert Ingram,
himself a frequent contributor to the National collections.
As may be seen from the above description and from the figure
(Pl. XXI.), the differences between this monkey and its only near
allies are so numerous as to render any detailed comparison un-
necessary. Good figures have been given of &. roxellance by
Milne-Edwards'! and De Winton’, and of 2&. bieti by Milne-
Edwards and Pousargues’.
Mr. Oldfield Thomas also exhibited adult and young examples
of a Bush Duiker, which had been sent to the British Museum
by Mr. F. W. Isaac, from Eldoma Ravine, British East Africa,
1 Rech. Mamm., Text, p. 233, Atl. pls. xxxvi. & xxxvii. (1874). P
2 P. Z.S. 1899, p. 572, pl. xxxi.
3 N. Arch. Mus. (8) x. p. 121, pls. 9-12 (1898).
15*
226 MR. OLDFIELD THOMAS ON A NEW ANTELOPE. [Mar. 17,
{t appeared to belong to a new species allied to the Congolese
Cephalophus weynsi Thos., and was described as follows :—
CEPHALOPHUS IGNIFER, Sp. n.
Size medium. Fur close, fine and glossy, hairs of back about
an inch in length. General colour of back bright rufous or bay
(nearest to ochraceous-rufous of Ridgway), darkening forwards on —
the neck and shoulders to dull brownish. Forehead mixed rufous
and black; crown and occiput bright rufous like back, coronal
tuft a deeper and more chestnut or vinaceous rufous. Hairs of
occiput reversed upwards to the crest as usual, those of nape all
directed backwards. Muzzle blackish ; lips and chin white; ears
dark brown behind, with white edges and inner surfaces. Throat
rufous. _ Belly brown mesially, grading into rufous laterally.
Inner side of forearms, inguinal region, and inner side of thighs
white. Outer side of forearms and thighs rufous; feet brown,
darkening almost to black above the hoofs. Tail rufous above,
white below, proximally, with a mixed brown and white terminal
GulGHene
Skull of normal proportions; premaxille just reaching nasals,
frontal convex as usual; posterior palate variable, that of the
male cut out some way in front of the lateral notches, while in
the female the median notch is posterior to the lateral ones.
Horns laid back just in the line of the face, those of an adult
male 98 mm. long, with a basal diameter of 31 mm.; the same
dimensions in an immature female 49 and 20.
The young specimen (skull 105 mm. long, the last milk premolar
only just up) is nearly black all over, the coronal region and the
posterior back alone being rufous.
Dimensions of the type, an adult male, measured in skin :—
Head and body 810 mm., hind foot, with hoof, (c.) 240, ear 81.
Skull—basal length 162, greatest breadth 79°5, muzzle to orbit .
94, nasals 73 x 32; palate length 97; length of upper tooth-row
53, of three upper premolars 23:4.
Hab. Eldoma Ravine, British East Africa, alt. 7200 feet.
Type. Male. B.M. No. 2.11.17.6. Collected and presented by
F. W. Isaac, Esq. Three specimens.
Native name ‘“ Meindet.”
This Duiker is allied to C. weynsi of the Eastern Congo and
C. johnstoni of Toro by its colour and general characters, but
differs from them by the hairs of its nape being all directed back-
wards in the usual way. ©. harveyi of Kilima-njaro, also a
member of this group, has a blackish forehead and no chestnut
corona] tuft.
, The following papers were read :—
1903.] ON JAPANESE LONG-TAILED FOWLS. 227
1. Observations and Experiments on Japanese Long-tailed
Fowls. By J. T. Cunninenam, M.A., F.Z.S
[Received January Ist, 1903.]
(Text-figures 41 & 42.)
The enormous length of the tail-feathers in Japanese ‘“ Long-
tailed Fowls” has been known to zoologists in this country for many
years from the stuffed specimens in the hall of the Natural History
Museum in London. These specimens were figured by the late
G.J. Romanes in his ‘ Darwin and After Darwin,’ Part I., published
in 1892. The figures represent fairly accurately two males of the
breed, but the females are not represented at all, unless by indistinet
figures in the background; and the accessories, instead of being
dvawn from those actually present in the Museum case, are entirely
imaginary, showing the birds in a state of perfect freedom in open
country, a state which they never enjoy in Japan. Romanes’s
figures are among a large number given as “ typical proofs of the
efficacy of artificial selection ” : and it is the object of the present
paper to show that this involves an assumption, with regard to
this particular breed, which is by no means justified by the facts.
The tails of the male specimens in the National Museum are
6 to 9 feet in length, that is to say, the longest feathers are of
that length. I have endeavoured, but without success, to obtain
information concerning the history of these specimens. I believe
they were reared in Japan, and they were probably sent from that
country after death, either as skins or as stuffed specimens,
There are some female specimens also, but in these the tails are
scarcely longer than in the hens of ordinary breeds,
The breed has been known to poultry-fanciers fora considerable
time, and the following is the account of it given in ‘ The Book of
Poultry’ by Lewis Wright, published in 1885 :—
“ About the year 1878 there appeared in Germany, and a year or
two later in England, fowls imported from Japan, whose principal
peculiarity consisted in an immense length of tail and hackle-
feathers. Some of these were exhibited as Yokohamas ; others,
said to be superior in these points, were called Pheenix fowls.
The tails of these specimens averaged about a yard in length, and
the general appearance was not only that of a Game-fowl, but all
the colours were Game colours, Whites, Piles, Duckwings, and
later a few Black-Reds. The long plumage was, however,
unique, and a fair idea of it may be gathered from the illus-
tration.
“Correspondence in the poultry journals brought out the fact
that such birds had been occasionally exhibited as Japanese game
so far back as about 1872. But it further appeared that in the
Japanese Great National Museum at Tokio there were preserved
two specimens. of an allied race in which the tail-feathers measure
228 MR. J. T. CUNNINGHAM ON [ Mar. ale
134 feet and 17 feet respectively, and a feather has been actually
sent to France which measures 2 metres 85 centims. in length
(say 94 feet). In 1884 Mr. Gerald Waller, of Twywell, imported a
pen of these birds ; and from his statements we gather that they
are known in Japan as Shinowaratao, Shirifuzi, or Sakawatao
fowls and various other names. He says the very long-tailed
ones are kept in high, narrow cages, always sitting on a perch
covered with straw-rope, with no room to turn or get down, but
with a food- and water-tin at each end of the perch. Three times
daily they are lifted down for a few minutes’ exercise, their tails
being carefully rolled up in paper cases to keep them from injury.
The Japanese state that a tail has been measured 23 feet in length,
and that the birds only moult the tail once in three years. This
last statement is highly interesting. It is obvious that if a tail
23 feet long were grown in one year, it must be at the rate
of nearly three-quarters of an inch per day; and though
Madame Bodinus states that she could see the tails grow daily,
it is difficult to realise this; but experience will soon decide the
oint. The birds which have reached Europe have never yet
exceeded 5 or 6 feet in length of feather, which is not beyond the
possibility of a single season, though it appears of an enormous
length. The saddle-hackles of Mr. Waller’s birds are 16 inches
in length; but it is manifest that such enormous feathers as
reported from Japan could never be preserved under the ordinary
conditions of an English poultry-yard. The feathers are not only
long but extremely narrow and flexible, trailing low after the
birds.”
Mr. Wright does not mention the comb, but the illustration
which he gives represents the male bird with what is called a pea-
comb of small size and with small wattles, whereas the specimens in
the National Museum have single vertical serrated combs and large
wattles. The pea-comb is a rounded mass, with small rounded
tubercles projecting from it.
Mr. Frank Rice, of Acton, Suffolk, who breeds Yokohamas,
gives in his circular the same illustration which appears in
Mr. Wright’s book, and which therefore is certainly not a new
figure from a living specimen. But in his description he states
that the head should be neat and small, with evenly-set pea-comb.
Tt would thus appear that the long-tailed fowls comprise varieties
which differ in comb as well as in colour, though they seem to be
all similar in the excessive growth of the tail, and probably are
all grown in Japan under the same artificial treatment.
The principal purpose of this paper is to discuss the causes by
which the elongation or excessive growth of the tail has been
produced. About two years ago, Prof. Lankester, in a letter to
‘Nature,’ referred to the specimens in the Museum of which he is
Director as ‘‘a magnificent sport,” comparing their exceptional
character to what is called genius in human beings. On the
other hand, in the ‘ Dictionary of Birds’ by Newton and Gadow,
article “ Fea ther,” the length of the tail-feathers is attributed to
1903. ] JAPANESE LONG-TAILED FOWLS, 229
continuous growth, and it is stated that the moult is checked or
prevented by some means unknown to Europeans. We have seen
that in Wright’s book the statement is cited that the feathers are
only moulted once in three years, but this might be a congenital
peculiarity, whereas the expression used in the ‘ Dictionary of
Birds’ implies that the moult is prevented by artificial treatment.
Very definite statements on the question are made in a paper
by Mr. Basil Hall Chamberlain (“ Note on a Long-tailed Breed
of Fowls in Tosa,” Trans. Asiatie Soc. Japan, vol. xxvii. 1900),
for my knowledge of which I have to thank Mr. Frank Finn.
My. Chamberlain made enquiries on the subject in the country
from which the breed is obtained, and where presumably it
originated, but nevertheless his statements are based on the
assertions of breeders and not on any investigations of his own.
The following is a summary of his paper :—
The origin of the breed is not known, but it is believed to be at
least a hundred years old. It has been produced simply by
selection of the best specimens; one highly-prized variety, the
Haku, was produced in this way within the last few years.
The proper general name for the Long-tailed fowls is Shino-
wara-td, derived from the village of Shinowara in the province of
Tosa, east of Kochi the capital. Some are still bred in that place,
put most now in Kochi itself, whence the majority are exported
to Kobe and some of the finest to Tokio, but the very finest are
retained by the producers.
The following varieties were described to Mr. Chamberlain :—
Shira-fuji: white head- and body-feathers, tail black as in the
other varieties. He saw one specimen of this two years old, and
measured its tail-feathers, which were 73 feet long. Another
specimen, fourteen months old, had tail-feathers 4 feet long.
Others have black bodies. Another variety is the Haku, white
all over with yellow legs; another, Totenko, has red neck- and
pbody-feathers ; another, Dokiri, has reddish colour mixed with the
white of the body. All these except the Haku have black tail-
feathers.
As great a length as 18 feet has been reached in the tail-
feathers, but even 12 feet isa rarity. From 7 or 8 to 11 feet is
the usual length. The feathers grow about 4 inches a month,
and continue to grow while the bird lives, which may be eight or
nine years. The beautiful body-feathers growing from the
shoulders reach a length of 4 feet. (Vote.—This evidently refers
to the saddle-hackles, which grow not from the shoulders but from
the top of the rump.)
Some of these saddle-hackles may fall off in moulting, but the
tail-feathers never do so. He saw the birds in October 1898
when moulting, and only the ordinary feathers were gone or
going, not the long ones.
He also saw the hen, which was a handsome bird, distantly
reminding one of a hen-pheasant, with fawn-coloured breast and
white quill to the delicately-coloured feathers of the back. She,
230 _ MR. J. T. CUNNINGHAM ON | Mar. 17,
too, has longer tail-feathers than an ordinary hen, sometimes as
long as 8 inches. One, or at most, two hens are allowed to each
breeding-cock. The latter’s tail-feathers are cut to allow of his
walking about freely. He lives a little longer than the others
which must be kept shut up; but all are hardy, bearing both heat
and cold. The ordinary number of long tail-feathers is 15 or 16,
some cocks have as many as 24,
The tail-feathers must not be wound up, as people ignorantly do
away from Kochi, but must always be allowed to hang free; for
which reason the cocks are kept in high, narrow cages, quite dark
except at the top, for light at the bottom would attract them.
When the tail-feathers become too long and touch the ground in
the cage, a bamboo is put a little way back so as to form an arch
and make more distance. The birds sit all day on a flat perch
3 inches wide, and are only taken out once in two or three days
and allowed to walk about for half an hour or so, a man holding
the tail all the while to prevent its getting torn or soiled.
The high, narrow cages may be made of any wood; they are
64 feet high, 3 feet deep, and 6 inches wide. The wonderful
feathers both on tail and body come from quills much stouter than
any possessed by ordinary fowls.
The price in Kochi was 15 dollars for a cock with tail under
10 feet, 25 dollars over that length.
There is absolutely no artificial method of making the feathers
grow. Allis done by selection. Any failure is due to not having
a hen or parents of the proper breed. Also one must know how
to treat the birds.
At Kobe in November 1898 Mr. Chamberlain saw three speci-
mens, one with tail-feathers 133 feet long. He also saw a splendid
white tail 103 feet long, which had been pulled out from a white
bird owing to its falling off its perch and fluttering about. The bird
was five years old, and the feathers were growing again. Thefancier
said that the feathers in young birds grow about 4 inchesa month,
in older birds more, up to 7 inches a month.
Two photographs are given with the paper, but no reference to
them is made in the text. The tails are very long, but there are
no long feathers from the shoulders, only tail- and saddle- or
rump-hackles.
It is evident that Mr. Chamberlin, although his observations are
of considerable value, was not an experienced naturalist, and that
he is simply reporting what he was told. He says nothing about
the combs of the fowls he saw. With regard to his assertions
about selection as the sole means by which the breed has been
produced, it is to be noted that he is evidently referring in the
case of the Haku chiefly or entirely to colour. The Haku isa
white variety, and Mr. Rice, above mentioned, also has a strain
of this colour. Nearly all domesticated birds and animals vary in
colour, and nothing is easier-than to separate a white variety in
fowls, horses, pigeons, dogs, &c. These fowls, like others, vary
in colour; and the question before us now is not the separation of
1903. ] JAPANESE LONG-TAILED FOWLS. 231
colour varieties, but the excessive length of the tail which occwrs
in all the varieties. Mr. Chamberlain states that a specimen
fourteen months old had a tail 4 feet long. At the rate of
4 inches a month, this would mean a period of growth of twelve
months, and therefore the chickens must have begun to acquire
their permanent feathers at two months of age, and the feathers
must have grown without intermission afterwards. This is
perhaps not impossible, but according to my observations, recorded
below, it is very improbable. It may further be pointed out that
the statement “ there is absolutely no artificial method of making
the feathers grow,” is difficult to reconcile with that which follows
it, “also one must know how to treat the birds,” unless proof is
offered that the artificial treatment has no effect on the growth of
the feathers.
The birds in my possession, which have formed the subject of
my own observations and experiments, were descended from a
pair which were imported direct from Japan by Mr. John Sparks,
of London, and purchased by Mrs. J. C. Williams, of Caerhayes
Castle, Cornwall. Before proceeding to my own observations, I
wish to discuss the question whether the moult occurs in the
specimens kept in this country, and also some evidence I have
obtained as to specifie treatment of the feathers. We have seen
in Mr. Wright’s account that the feathers of birds in this country
had not been known to exceed 5 or 6 feet in length. This
does not throw much light on the question of the moulting,
because it may be that, the birds not being kept on perches as in
Japan, the feathers get broken when they have reached a con-
siderable length. I enquired of Mr. Rice what his experience
was, and he replied that his birds usually moulted their tail-
feathers each year, but he had had some cocks omit this operation
in theix second year. He said that his cockerels at eight months
of age had tails from 2 ft. 6 in. to 3 ft. 6 in, in length, and that
he had one cock, three years old, that had a tail 53 feet long.
Mrs. J. C. Williams was also kind enough to answer my
enquiries. She informed me that the original male bird which
she obtained from Mr. Sparks had a tail just 5 feet long. At first
its plumage suffered from the change of climate, but in 1901 its
tail was considerably better than it had been. (The length at
this time was not stated.) She had a young bird whose tail had
measured 42 feet, but he broke it in getting about. The young
birds usually moulted about a year after hatching. This must have
been the first moult of the adult plumage, which begins to appear
in the autumn after hatching. In reply to a request for further
particulars about the moult, Mrs. Williams stated the birds do
certainly cast their long tail-feathers. The bird that had some
feathers 43 feet long was then (March 1902) nearly three years
old, and she thought it was at the second moult that a feather of
that length was measured.
There is evidence here that, when left to themselves, the long
tail-feathers are moulted in the ordinary way, at any rate for two
232 MR. J. T. CUNNINGHAM ON [ Mar. 17,
successive seasons after the first year in which the birds are
hatched.
It will be seen that the absence of the moult, or the occurrence
of continuous growth, if true for the birds in Japan, is not true
for those reared in this country; and thus we have reason to
doubt that uninterrupted or continuous growth of the tail-feathers
is a fixed congenital peculiarity in the breed.
Mr. John Sparks in May 1901 supplied me with the following
information concerning the method of treatment applied to the
birds in Japan :—‘‘ In order to ensure very great length of tail,
the cocks ought to be kept on a perch as much as possible after
they are six months old; and the tail-feathers should be pulled
gently every morning, grasping the centre bone-like part firmly
with the finger and thumb and pressing steadily downwards
towards the tip, each feather being done several times. This
softens the quill and causes it to lengthen. The birds do not
moult the tail-feathers, but if one or more come out others
immediately grow in their places.
“The Japs themselves, those who take great pride in their birds,
always roll the long feathers up like a lady rolls up her hair, and
tie them, whenever the birds are let off their perches to walk
about, which is about twice a day for an hour at a time.
““T have often seen them thus treated in Japan, and those
which Mrs. Williams and the Hon. W. Rothschild had from me
were so treated on the voyage by the man in charge of them, and
T sent them down to St. Austell in their regular perch-cages.”
There is here a detail in the treatment of the feathers which,
so far as I can discover, has never been mentioned in any
published account of the matter ; and my own experiments, which
I now proceed to describe, so far as they have yet gone, tend to
show that this mechanical treatment of the feathers is the whole
secret of the mystery.
My own EXPERIMENTS.
In May 1901, I received from Mr. John Sparks twelve eggs,
laid by the fowls of this breed in the possession of Mrs. J. C.
Williams. I afterwards ascertained that some of these were the
produce of the pair originally imported from Japan, others were
from the offspring of this pair, but all the eggs were of perfectly
pure breed with no cross or admixture whatever. The eggs were
incubated by an English hen of mixed breed, and on June 13th
ten healthy chicks were hatched. Of the other two eggs, one
contained a dead chick, the other was addled or not fertile. This
shows that the eggs were remarkably fertile and of great vitality.
The chicks were small, the eggs being not much more than half
the size of ordinary table-eggs. Their colour was fawn, with a
broad dark brown stripe down the middle of the back, a narrower
stripe on each side of this, and a thin stripe of the same colour
running from the outer corner of the eye. They were very
1903. ] JAPANESE LONG-TAILED FOWLS. 233
pretty, very active, and very healthy, and throve well on ordinary
food, consisting of oatmeal, chopped meat and vegetables, and
mixed grain of various kinds. One of the chicks was accidentally
killed when I turned the hen out of the nest, having got beneath
its mother’s feet as she was scratching the earth. I examined
this specimen, and found that the primaries and secondaries of
the wings were present as short black quills with a little down at
the tip, but that there was no trace of tail-feathers or tail-coverts,
nothing but down over the rest of the body. The comb was
visible at the back of the beak as a slight yellow ridge with six
teeth. The toes were four in number; the skin on the legs
was yellow.
June 23rd. Age 10 days.—Another chick had been lost up to
this date, having escaped and probably fallen a victim to a cat.
The feathers of the wings now reached nearly to the end of the
body, and were chequered i in colour, being marked transversely
with dark colour and grey alter nately, ‘The tail-feathers, i. ¢.
rectrices, had begun to sprout in some of the chicks, elsewhere
there was still only down. The combs appeared no larger.
July 6th. Age 23 days.—The chicks were now half-fledged:
another had been lost by escaping, so that only seven survived.
Feathers were growing on the shoulders and sides of the breast,
but the head, back, and front of the breast were still downy, with
the original markings. Tail-coverts also appearmg. Feathers,
except tails, all barred with dark and grey, so that the general
appearance was speckled and very inconspicuous against the soil.
The tail-feathers were dark, nearly black, but not steel blue-black
like those of the adult. It was interesting to see the chicks scatter
in all directions, and then crouch down whenever the hen uttered
her special warning cry, as she did often when she heard a jackdaw
croak. In four of the chicks the tails were more developed than
in the rest; in these also the combs were beginning to grow
higher and to get red. These four proved to be cocks, so that the
sexual difference begins to show itself at this early age.
July 28th. Age 1 month 15 days.—The four cocks showed
their sexual characters a ttle more distinctly, the wattles and
ear-lobes being indicated by a tinge of red. Tn the three hens
the combs had not begun to enlarge and showed no red.
One of the cocks was darker than all the other chicks, and had
a slender tail, not very long and bent downwards. The hens were
light-colour ed, with white breasts, their tails being as long as those
of some of the cocks. Two of the cocks had dark breasts.
Aug. 4th. Age 1 month 3 weeks.—Chicks now fully fledged.
In the four cocks reddish tints were appearing in the feathers of
the back and wings, while in the hens only neutral tints were
present. The combs i in the cocks were a little more dey eloped, in
the hens not developed. The tail-coverts in the cocks were growing,
but not longer than the rectrices, and the sickle- feathers not
conspicuous.
The red feathers mentioned above are the beginning of the
234 MR. J. T. CUNNINGHAM ON [Mar. 17,
adult plumage. There is at least one complete change of feathers,
from the chicken plumage to the adult, both in cocks and hens.
Whether the chicken-feathers are changed before this I do not
know, but there were indications that the wing-feathers at least,
in the chicks, had been changed before the assumption of the adult
plumage.
Aug. 12th. Age 2 months.—I examined one of the cocks, and
could distinguish two steel-blue sickle-feathers on each side, not
longer than the rectrices and rather narrow. These were the first
of the adult tail-feathers to appear, succeeding others of dark,
lustreless, brownish black.
Aug. 20th. Age 2 months 1 week.—I counted the tail-feathers
in some of the chicks. In one of the hens I found 14, or 7 pairs,
of rectrices. In one of the cocks, the dark one previously men-
tioned, the saddle-hackles were almost black with a grey stripe
down the centre, instead of reddish. In this bird I found also
seven pairs of rectrices, and only four pairs of sickle-feathers
which were neutral brown. The other three cocks could also now
be individually distinguished, and I give the peculiarities to show
the amount of variation :—
A. The largest: more advanced in plumage than the rest, more
red on the back, breast nearly all black except at the sides;
a kink in the comb.
B. Similar to A, but not quite so far advanced.
C. Similar to A and B, but lighter in tone; back rather
yellowish than red.
D. The dark bird above mentioned.
Aug. 24th. Age 2 months 11 days.—Examined Cock A and
found a number of steel-blue sickle-feathers, or tail-coverts, all
with very long horny sheaths at the base indicating vigorous
growth. In Cock B similar feathers were only just beginning to
sprout.
Aug. 26th.—In Cocks C and D very few new sickle-feathers
showing. In B the rectrices were being replaced.
Sept. 6th. Age 2 months 3 weeks,—In Cock A all the chicken-
rectrices had been moulted, and the new steel-blue ones were
growing out with long horny sheaths at the bases. In all the
cocks the spurs had begun to show as very slight blunt knobs.
None of the cocks had crowed yet.
Sept. 10th.—Examined one of the hens. Saw several tail-
coverts growing with long sheaths but no new rectrices.
In Cock D, the dark one, the two central rectrices were long,
slender, and curved, and had sheaths at the base, also sheaths to
some of the other rectrices. I thought at the time that these
central rectrices had not replaced others in the chicken plumage,
but grown continuously. I afterwards concluded that they had
moulted earlier than in the other cocks.
In Cock A the longest tail-covert measured 19 cm. or about
7? inches.
1903. ] JAPANESE LONG-TAILED FOWLS. 935
Sept. 19th. Age 3 months 1 week.—Condition of plumage :—
Cock A. Breast, thighs, and belly all black. Neck-hackles
very light grey, with thin dark stripe down the centre of
each feather. Back with some steel-blue feathers behind
the hackles, the rest red. Saddle-hackles developing,
yellow, long and thin. Nearly all the rectrices and tail-
coverts growing with long sheaths, colour steel-blue.
Cock B. Nearly the same as A, but not quite so far advanced.
Cock C. Much less advanced, white on sides of breast, speckled
brown feathers mixed with red of back, only the outer-
most rectrices showing sheaths.
Cock D. Very little white on breast, hackles of neck steel-blue,
back black with a little red at tips of feathers. Saddle
whitish. Rectrices 7 pairs, only the central and outer-
most pairs with sheaths, the rest apparently not yet
moulted.
Hens. Breasts almost white, the sides of breast buff; neck-
hackles dark, with whitish stripes down centres of feathers.
Back and tail greyish brown, speckled, 7. e. with white
quills. Two hens with black heads, one with head lighter,
grey and speckled.
T now decided to stroke and pull the tail-feathers in Cock B and
to leave Cock A untouched, to see if any difference would result.
These two, as I have said, were closely similar except that A was the
larger, finer bird and slightly more advanced in development. I
chose, therefore, the one which was congenitally inferior, so that
if any superiority in growth of feather appeared in it, it could
only be due to the artificial treatment. In both there were seven
pairs of rectrices, the central pair slender and curved, the rest
broad and_ str aight. The outermost pan had not yet been
moulted, I fixed up a sort of cage with a round perch at the
bottom, and put Cock B into it w hile I stroked his feathers, but
did not keep him in it.
Oct. 1st.—Found that the hens had shed most of their rectrices
and were producing new feathers, as well as new tail-coverts, but
without change of colour.
Oct. 7th.—The upper tail-covert in Cock A measured 25 em. or
about 10 inches, not including the basal sheath.
At this time I changed my residence, and the fowls were
installed in a place divided into tworuns; into one I put Cocks B,
©, and D, in the other Cock A with the three hens.
On Oct. 12th Cock A crowed for the first time, another proof
that he was a little more precocious than the others.
Oct. 16th. Age 4 months 3 days.—Longest feather, a tail-
covert on left side, in Cock B 27 ecm., about the same as the
longest in Cock A.
Oct. 27th. Age 4 months 2 weeks.—Tried tying the cocks by
one leg on ordinary perches about 3 feet from the ground, and
thus was able to measure the feathers better.
236 MR, J. T. CUNNINGHAM ON [ Mar. 17,
Cock A. Longest feather 13 inches from skin.
Cock B. The same. But there were more long feathers in
Cock A, the outer ones being more nearly the length of the
middle.
As I found I had not time to pay attention to more than two
cocks, and as, moreover, they soon began to fight, I got rid of
Cocks C and D; and what I have to say hereafter refers only to
Cocks A and B, which were undoubtedly the best of the four, and
had the most beautiful colours. I began tying these two cocks
on ordinary perches in the daytime by means of a piece of tape
tied to one leg and round the perch.
Dec. 1st. Age 5 months 18 days.—Longest feather in Cock A
18 inches, in Cock B the same. This shows a growth of 5 inches
in five weeks, or 1 inch per week. Perhaps the feather in Cock A
was really a little longer, as nearly all the feathers in this bird
had lost their tips by friction against the ground. ‘There is thus
no evidence up to this time that pulling the feathers mi Cock B
has increased the rate of growth.
Dec. 8th.—I had been stroking the feathers of Cock B regularly
at morning and evening. Usually I tied the birds on the perches
at night, and generally tied the feathers up in paper when the
birds were free. I used tissue-paper, and rolled the tail up trans-
versely, fastening it with tape. I tried a rough narrow cage with
wooden bars for Cock B, but the bird turned round and escaped
through the opening behind the perch.
T noticed now that one of the long feathers in Cock B seemed
to have stopped growing, the sheath having peeled off and left a
dry stalk. This was one of the outer tail-coverts, but the rest
were still growing vigorously.
Dec. 27th.—As my first attempt at a cage did not succeed, and
LT was unwilling to confine the birds so completely as the Japanese
do, I made a perch of about 9 imches in length supported on two
uprights. This apparatus was movable, not fixed to the ground.
T used this for Cock B, tying him to it by a piece of tape fastened
round one leg. This answered very well, although he once upset
the perch and slightly wounded his head. No permanent injury
resulted from this accident.
On this date, when I was stroking the feathers, one of the
smaller tail-coverts on the right side came out of the socket, with
the epidermic sheath attached to it. This feather was 14 inches
long. This seemed to indicate that the effect of stroking the
feathers was, as suggested by a correspondent of ‘ Nature,’ to pull
the growing feathers out, instead of increasing their growth ; but
my experiments were not finished yet.
Dec. 31st. Age 6 months 18 days.—Measured the feathers as
accurately as possible on a flat wooden measure.
Cock A longest two feathers, the central feathers of the tail,
22 inches. ;
Cock B longest feather scarcely 22 inches, also a central
rectrix.
1903. ] JAPANESE LONG-TAILED FOWLS. 237
It seemed, therefore, that the growth of the feathers in Cock B
was barely keeping pace with that in Cock A, although only in
the former were the feathers pulled or stroked. In Cock A the
longest feathers were the two central rectrices ; in B only one of
the central rectrices was as long, the adult feathers having mostly
started earlier in A than in B.
Up to this time Cock A had been with the three hens, and B
with the other cocks until I had got rid of these.
1902, Jan. 8th.—On this date one of the hens laid the first egg.
It was small and of light brown colour, like those the birds were
hatched from.
The hens have a low serrated comb and scarcely any wattles,
only ved skin about the cheeks.
Jan. 12th. Age 7 months.—In Cock A the 2nd pair of rectrices,
counting from the middle, seemed to be coming to the end of their
growth, the sheaths drying up and the quills forming. This pair
were not so long as the corresponding pair in Cock B which
had been pulled. The five outer pairs of rectrices had ceased
growing in both cocks. These were broad and stiff and only about
6 or 7 inches long, in fact like the rectrices of ordinary fowls.
All the hens were now laying. One of the hens was put with
Cock B, the other two with A.
Feb. 4th. Age 7 months 22 days.—
Cock A, longest feather......... 251 in. = 64 cm.
GockeBe war Reacts 252 in, =65'7 cm.
It is thus evident that the growth in B had been greater than
in A. The feathers in B were pulled once or twice a day when-
ever I was at home, and B was tied on the perch with his feathers
loose at night, while A was usually left free with his tail rolled
up in paper. The sheaths of the central rectrices, which were
the longest feathers in both birds, were in Cock A shorter, and
seemed as though growth were about to cease.
It is to be noted here that so long as the feather is in full
growth, the horny sheath which surrounds the base is milky
white, or bluish white, and soft, while when growth ceases the
sheath dries up and appears black because the feather is seen
through it; then it scales off, and leaves the quill of the feather
bare down to the surface of the skin. The quill at the base of a
full-grown feather is only formed at the end of growth, and most
of it is contained in the skin-socket. Thus I learned to recognise
the approaching cessation of growth and formation of the quill by
the appearance of the horny sheath.
Feb, 10th.—One of the feathers was accidentally pulled out in
Cock A while the tail was tied up in paper. I do not know how
this occurred; I only saw the root of the feather hanging free
while the length of it was held by the paper. When a growing
feather is thus pulled out, the horny sheath comes out of the
dermal socket and remains attached to the base of the feather,
238 MR. J. T. CUNNINGHAM ON [Mar. 17,
and the end of the sheath is quite soft and exudes a little blood
and moisture when pressed. This feather was 52°8 cm. or about
1 ft. 9in. long. It was one of the anterior tail-coverts, and was
the only feather ever pulled out of Cock A.
Feb. 11th. Age 8 months nearly.—On this date, when I was
stroking the feathers of Cock B, another came out. It was one
of the anterior lateral coverts of the right side, and its length was
152 inches (38 cm.). The feather had not ceased growing, but
showed signs that it was about to cease, as the rhachis at the base
was stout and thick, and the barbs emerging from the sheath were
free and downy, as they are at the base of a full-grown feather,
and never at its more distal part
Feb. 14th.—Examined the bases of the feathers in both cocks.
In A the central rectrices had nearly or quite ceased growing, and
also nearly all the tail-coverts, only two or three still showing the
basal sheaths. In B two or three of the posterior coverts had
ceased growing on each side, but a number of the others still
showed vigorous growth.
Feb. 15th.—The longest feather in Cock B came out when I
was stroking the feathers, though I was not pulling at all hard.
The feather was the central rectrix of the left side, and was in
growing condition, with bluish pulpy sheath at the base. It came
clear out of the socket and was not broken off at all. The extreme
length was 2 ft. 44 in., but about 1 inch of this was in the socket
before it was pulled out. The total length of the sheath up to
the point where the feather proper emerged was 3} inches (8 cm.).
Feb. 16th.—Ascertained that in Cock B three tail-coverts on the
right side had ceased growing and two on the left. The 2nd
rectrix on right side had ceased to grow, that on the left nearly so.
In Cock A the 2nd pair of rectrices had ceased growing, the
1st or central pair had nearly ceased. Two central coverts, which
were nearly as long as the central rectrices, had nearly ceased, and
there were only two others on each side with growth-sheaths.
There were two rows of coverts in front of the rectrices, anterior
and posterior, with four or five feathers in each row on each side.
When the feathers are specified individually they are counted from
the middle line.
Feb. 17th.—One of the posterior lateral coverts on right side in
Cock B was pulled out today. I was really trying to see if it
would come out when pulled, as it had completely ceased to grow.
T found that it required much more force to pull it out than is
required in the case of a growing feather. The quill was com-
pletely formed. It was evident that the attachment of the feathers
was most feeble when growth was ceasing, not when the feather was
growing vigorously ; while after the quill was completely formed,
the feather was very firmly held in the socket. The feather was
192 inches long.
March 2nd.—In Cock B another feather was unintentionally
pulled out. It was in full growth, and was one of the lateral
anterior coverts on the right side. Total length 163 inches.
1903. ] JAPANESE LONG-TAILED FOWLS. 239
March Ath.—Measuvred feathers, exactly four weeks since last
measurement :—
Cock A, longest feather............ 2 ft. 34 in.
Cock B pe Peer SEE Soterarior 2 43 i
The longest feathers in A were the two central rec trices, and
these had almost entirely ceased growing since last measurement.
They had grown 23 inches in four weeks.
In B, on the other hand, the longest feather was the central
rectrix on the right side, <a was still erowing and was | inch
longer than the rectrices ‘of A. The left rectrix of B, which was
pulled out, would have been somewhat longer.
In Cock A there were now only two feathers with growing
sheaths, both on the right side, all the rest having ceased to grow.
In Cock B, on the other hand, there were sev eral coverts with
growing sheaths, as well as the central rectrix of the right side.
Mar ch 9th.—The two feathers still growing in Cock A were the
central posterior covert on the right side and one next to this on
the same side. They seemed to be about to stop growing. There
were only five long feathers in Cock A, all the rest being shorter
than the corresponding feathers in B.
March 10th.— Another feather came out from Cock B today when
the feathers were stroked ; it was the Ist anterior covert on the left
side, and, although still growing, showed signs of cessation of growth.
It was evident that the feathers, as noted abov e, were easily
ulled out at this stage. Feather measured 1 ft. 97 in. (55°3 cm.).
March \\th.—Still another feather pulled out of Cock B, Ist
anterior covert of right side. It measured 1 ft. 8 in.
March 16th. Age 9 months—The two feathers which were
still growing in Cock A seemed now to have both ceased to grow,
one of them certainly. The only other growing feather in this
bird was the one on the left side which had sprouted i in place of
the one pulled out.
In Cock B four of the old feathers were still growing, namely,
the right central rectrix, and three coverts on the left side, besides
the new feathers sprouting in place of those pulled out.
Many of the golden saddle-hackles in B had growing sheaths.
One came out when they were being stroked the other day, and
measured 7 inches in length. A few of the feathers of the same
kind were growing in A, but most have stopped, and these hackles
altogether were about 2 “inches shorter in A than in B.
Between the saddle-hackles and the long tail-coverts there were
a number of feathers which may be called transition feathers.
They were flexible and projected upwards first and then drooped
in an arch; the outer ends were steel-blue, the bases carried pure
white down, which was exposed and was very ornamental to the
bird.
April 1st.—Measured the feathers, exactly four weeks since last
measurement :—
Longest feathers in A, central rectrices, ee 4} in.
Longest feather in B, right central rectrix, 2 ft. 8 in.
Proc. Zoo. Soc.—1903, Vou. I. No. XVI. 16
240 MR. J. T. CUNNINGHAM ON | Mar. 17,
The rectrices in A had therefore grown | inch in four weeks,
the rectrix in B 32 inches. The former had ceased to grow some
time before this date.
One covert on the right side in A had still a little of the blue
sheath at the base; it had almost finished growing, and was still
shorter than the four longest central feathers.
In B the next longest feather was the central covert of the left
Text-fig. 41.
Japanese Long-tailed Fowl.
Cock A, photographed April 1903.
side, which was 1 ft. 11 in. long, or 9 inches shorter than the central
rectrix. The left central rectrix, which had been pulled out on
Feb. 15th, was growing again rapidly and was now about 4 inches
long.
In Cock B the left central covert was still growing vigorously,
with long sheath. It had a twist in it and hung im a spiral. Of
the other two growing coverts on the left side, the outer had
nearly ceased growing, the other still had a blue sheath.
1903. | JAPANESE LONG-TAILED FOWLS. 241
I found that the feathers which had ceased growing for some
time were very firmly attached. It was impossible to pull them
out by moderate force, and I did not wish to pull them out by
violence. There were 8 long coverts on each side in both cocks.
April 7th.—There were signs that more of the feathers in Cock B
were coming to an end of their growth. The sheath of the right
central rectrix was diminishing, The outer of the two upper
Text-fig. 42.
Japanese Long-tailed Fowl.
Cock B, photographed April 1903,
posterior coverts on the left side had, I believe, ceased to grow,
and the inner, though still growing, was apparently about to cease.
The only feather which appeared to be in full growth was the
central covert on the left side.
April 13th. Age 10 months.—The right central rectrix in
Cock B came out today when I was pulling the feathers. I had
pulled it rather hard and rather frequently in the hope of stimu-
16*
242 MR. J. T. CUNNINGHAM ON [ Mar. 17,
lating its growth, with this result. I am sure it would not have
grown much more, as its growth was evidently ceasing.
Total length of the feather 2 ft. 9% in., of which 2 in. was
embedded in the socket.
April 20th.—On this date the light-coloured hen being broody,
I put 9 of her own eggs under her to be incubated.
April 29th.—Longest feather in Cock B now was the central
covert of-the left side, which was 2 ft. 2 in. long and still growing.
The covert next to this was also growing. The feathers which
had been pulled out were all regrowing vigorously, the longest
being the left central rectrix, which was 7 or 8 inches long.
In Cock A there was no further growth, except in the anterior
covert which had been pulled out.
May 12th.—Allof the eggs incubated by the light-coloured hen
were hatched on this date, and all the chicks were vigorous and
healthy. My experience of the breed has been in direct opposition
to the statements in Lewis Wright’s account, with regard to their
vigour and fertility: I have found them exceptionally hardy,
vigorous, and fertile.
There was no new variation observable in the chicks, except in
one which was a rich golden-brown on the head and shoulders,
instead of light buff colour. All were striped as described in the
original chicks of the preceding season.
May 27th. Age 11 months 14 days.—Left central covert in
Cock B 2 ft. 5 in. long, so that it had only grown 3 inches in four
weeks. The covert next to this came out on this date when I
stroked it. It was 2 ft. 24 in. long. It had evidently nearly
ceased to grow, and was beginning to form downy barbs at the
base.
June 2nd.—First signs of moulting noted in the hens; a good
many of the breast- and body-feathers on the ground from day to
day, and one primary wing-feather from the light-coloured hen
found. There was, however, no sign of moulting in the tail-
feathers of either cocks or hens. Cock A had lately been left
at liberty with tail free, and none of the feathers had been
broken or lost.
June 8th.—Hens moulting a good deal, more primaries shed,
and in some cases the new primary was half developed. The
primaries in each hen were shed a single pair at a time.
In the cocks none of the wing-feathers, saddle-hackles, or tail-
feathers had been shed.
The chick of this season’s brood which was browner than the
others in the down, showed the same peculiarity in the immature
plumage, having a yellowish-brown colour instead of the neutral
brown of the others; the marking was the same.
June 24th. Age | year 11 days.—Left central covert in Cock B
2 ft. 8 in. long, so that it had grown only 3 inches in four
weeks.
July 13th.—The left central covert in Cock B was accidentally
pulled out when J was away from home; the paper in which the
1903. | JAPANESE LONG-TAILED FOWLS. 243
feathers were tied up having come undone, the feather got caught
in a door and was pulled out. There was some indication that
the feather was ceasing its growth, as, although the sheath was
long and soft, the bar bs at the base were beginning to be slightly
downy.
The total length of the feather was 2 ft. 10} in., or 2 ft. 92 in.
beyond the socket.
As this completes the history of the feathers of the first season,
I will here give a table showing the history of the feathers which
were pulled out, as it is the fur ther history of these feathers
which constitutes the most important result of my experiment.
Feathers of first adult plumage in Cock B pulled out.
Right side of Tail. Hae oe
Small lateral covert, growing.................. Dec. 27.
Anterior lateral covert, nearly ceased ...... Feb. 11.
Posterior covert, ceased) .s..c.5-s.c-sc-.er esse Feb. 17.
Anterior lateral covert, growing ............ Mar. 2.
Anterior Ist covert, STOWING .................. Mar. 11.
Central "rectrix, STOWE... ...--.¢--c-c-e ssc ee. Apr. 13
Left side of Tail.
Central rectrix, growing......0.0iececss cesses es Feb. 15.
Anterior central covert, growing ............ Mar. 10.
Anterior 2nd covert, nearly ceased ......... May 27.
Posterior central covert, growing ............ July 13.
Of these it is important to note that all but the last two
sprouted again almost immediately, and continued to grow,
though not very fast, till the general moult. The last two did
not appear again until the month of September, that is to say in
the moulting-season. It is evident, therefore, that the recrescence
of a feather is influenced by the season of the year. After the
moult, that is to say in winter and spring, there is a general
tendency to feather-growth, and if a feather is pulled out ‘during
this time a successor at once begins to grow in its place. After
the end of April, at any rate in ‘the second year of the bird’s life,
the activity of feather-growth slackens or ceases, and feathers
pulled out after this time are not replaced till the following
moulting-period.
July 31st. Age 1 year 1 month 18 days.—Found today that
Cock A had begun to moult bis wing-primaries: there were three
or four new feathers growing, and one of them was already half
its full length. Cock B was in the same condition, but had not
shed so many feathers. The wing-secondaries were moulting also.
Aug. 4th.—Found one of the outermost short tail-coverts in
Cock A growing again, the old feather having been moulted.
This was the beginning of the moult in the tail-feathers.
244 MR. J. T. CUNNINGHAM ON [Mar. 17,
T believe there were 10 tail-coverts on each side in each cock, but
the two outermost on each side were small and unimportant. It
will be seen from the list given above that only eight of the
coverts in Cock B had been pulled out, so that eight feathers were
left, apart from the rectrices, which had completed their growth,
and could not grow any longer till they were moulted ; while in
Cock A all the feathers except one had completed their growth,
and must necessarily have been moulted before they could have
grown again.
Aug. 11th.—The longest feather in tail of Cock B now was the
central vectrix of the left side, which I measured today and
found to be | ft. 83 in. long.
The old feather was pulled out on Feb. 15, so that 1ts successor
had been growing nearly six months, and had grown at the rate
of little more than 3 inches per month.
Aug. 16th.—One of the dark-headed hens had moulted all the
rectrices as well as some of the tail-coverts, so that the rectrices
were shed almost simultaneously.
Aug. 24th.—Found that Cock A had shed another short tail-
covert, one of the transition-feathers, and also three of the outer
rectrices: the latter were the 6th and 7th on right side and 6th
on left, counting from the centre.
Cock B had shed none of the rectrices or old tail-coverts yet,
but this morning when I was stroking the feathers the Ist anterior
covert on the right side came out. The predecessor of this had
been pulled out on March 11th, and it was in full growth. It was
138 inches in length, and this was the growth of five months.
There was no reason to regard the loss of this feather as connected
in any way with the moult, as growing feathers are not moulted.
The feathers growing in Cock B in place of those which were
pulled out in the spring were as follows :—
Three anterior coverts on right side.
One posterior as .
One anterior covert on left side.
Two central rectrices.
Sept. 1st.—The outermost rectrix on each side in Cock B came
out when I was stroking the tail: they were loose and just about
to moult. This may be regarded as the beginning of the moult
of the tail, although one of the saddle-hackles and of the inter-
mediate feathers had been moulted occasionally for some time.
Sept. 2nd.—The 2nd posterior covert on right side in Cock B,
a full-grown feather, was moulted. It was 1 ft. 77 in, without
the imbedded quill, 1 ft. 83 in. in extreme length.
Sept. 3rd.—6th rectrix on right side shed in Cock B; it was
74 inches long without the quill.
Sept. 4th.—\st (nearest centre) posterior covert in Cock B
moulted; it was 1 ft. 73in. long. Also one of the outer anterior
coverts in Cock A moulted.
Sept. 5th.—In Cock B three rectrices and one covert on left side
1903. ] JAPANESE LONG-LTAILED FOWLS. 245
came out when I stroked the feathers: these were all feathers of
completed growth, which came out in consequence of the moult.
In the light-coloured hen, nearly all the rectrices and some of the
coverts came out when I tried them. The two central rectrices
in the hen were longer than the others, slightly curved downwards
at the tip, and speckled with brown at the borders: length
22°5 em. (9 inches) without quill. The other rectrices were broader
and black, length 19°5 em. (8 inches) without quill: the quill was
1:8 em. in length.
Sept. 6th.—More tail-feathers moulted from Cock B_ this
morning. The 2nd anterior covert on left side pulled out on
May 27th had just begun to appear, so that renewed growth was
deferred till the moulting-season.
Sept. 7th.—Nearly all the rectrices and longer tail-coverts which
had completed their growth had now been moulted in Cock B.
T had pulled many of them out without violence when they were
loose. By looking at the base of the feathers, I could see when
they were loose, because the transparent hollow quill was pushed
out and was visible beyond the socket.
In Cock A only four of the rectrices and none of the longer
coverts have been moulted spontaneously.
The left central rectrix in Cock B, which was still growing,
measured today 2 ft. 04+ in., or had grown not quite 4 inches in
four weeks.
Sept. 18th. Age 1 year 2 months.—Today, when taking up
Cock A to examine him, I accidentally trod upon the growing
feather the predecessor of which had been pulled out on Feb. 10th.
It was one of the anterior coverts on the left side, possibly the
first. The feather broke off at the top of the sheath, the latter
remaining in the socket. It was nearly 18 inches long (44°5 em.),
and would probably continue to grow from the same base.
This cock had still six long feathers not moulted, namely: the
central rectrices, the 2nd pair of rectrices, and two posterior
coverts.
In Cock B the fourth anterior covert on the right side, the
outer of the three growing again, was coming to an end of its
growth, showing downy barbs at the base, and it would be curious
to see if it were moulted.
This fact is worthy of particular notice, for it proves that a
feather which replaces one pulled out before the moulting-season
does not necessarily continue to grow through that season, and
it raises the question whether feathers which are growing when
the moulting-season begins, necessarily grow longer or for a
longer time than those which are moulted in the natural way.
This question is answered to some extent by the subsequent
observations.
At the end of September I ceased to reside at Penzance and
came to London, and having obtained permission through my
friend Mr. F. KE. Beddard, F.R.S., to deposit the fowls at the
Society's Gardens, | took them there on Oct. Ist. They were
246 MR. J. T. CUNNINGHAM ON [ Mar. 17,
accommodated in two of the pheasants’ aviaries. I brought them
from Cornwall in hampers in a night train, with the tails tied up
in papers, and none of the feathers were injured. While they
were at the Gardens I kept the tail of B tied up, while that of A,
in which there were no long growing-feathers, was left free. Both
the birds were left at liberty. During the fortnight after their
arrival, all the remaining old feathers in both tails were shed, and
the moult was thus completed, while in B there were left the seven
long feathers which had grown in place of those pulled out earlier
in the year.
This observation disposes, at any rate for these specimens, of the
supposition that the tail-feathers of the breed are not moulted
when left to themselves.
Oct. 11th.—Took away Cock B and the light-coloured hen from
the Gardens and placed them in a private fowl-house. The left
central rectrix measured 2 ft. 53 in., showing a growth of about
5 inches in five weeks. I fixed up a short perch on a stand, and
at first allowed both birds to fly up to it in the afternoon, then
later in the evening tied the cock to the perch and unfastened its
tail till the morning.
Oct. 22nd.—1 intentionally pulled out the fourth of the anterior
coverts on the right side which had stopped growing, in order to
make the new feather grow again. Thus there were left only six
long feathers, all growing.
Oct. 29th.—In the morning found the Cock B hanging from the
perch by the leg which was attached by the piece of tape.
Generally, if he tried to fly off when he was tied, he was able to get
back again, but this time the tape slipped down the vertical sup-
port, and he was unable to do so. In his struggles he had injured
the longest feather, the left central rectrix, at the base, and it
broke off near the top of the sheath. The detached part was
2 ft. 54 in. long. The root of the feather was uninjured, and the
basal part continued to grow.
Nov. 2nd.—Measured the central rectrix of the right side, which
was now the longest feather: it was 1 ft. 11 in. in length.
Nov. 15th.—Examined Cock A at the Zoological Gardens. The
rectrices were nearly full-grown, except the two central pairs
which were shed last, and which were only a few inches in length.
The coverts were only a few inches in length, and were not likely
to reach the ground for some weeks.
Nov. 17th.—This evening when I removed the paper from
Cock B, I found that another of the long feathers had come out.
The bird had been free all day, and must have pulled out the feather
by catching it in some projection when wallowing in the earth, or
must have accidentally pulled it out when preening himself. It
was the second anterior covert of the right side, and was 1 ft. 93 in.
long (53°8 cm.).
There were now only four long growing-feathers besides the
left central rectrix which was not lost but broken off.
Dec. 6th.—I brought away Cock A and the other three hens
1903.) JAPANESE LONG-TAILED FOWLS. 247
from the Gardens. The new feathers in A had grown no faster
than the new feathers in B, in fact they were not quite so long.
The longest feather in B hung about 2 ft. 3 in. from the perch,
and all four of theilong feathers were still growing. In Cock A,
none of the feathers touched the ground yet.
Dec. 9th.— Another of the feathers in Cock B pulled out today,
namely, the central anterior covert on the left side. The loss was
due to accident when the bird was free, not to manipulation.
Since this date I had kept Cock B tied up nearly every day in
a separate house, to prevent any more accidents. The feathers
that were pulled out were all sprouting again, and there were three
long feathers still growing, namely, the right central rectrix, one
of the posterior coverts on the right side, ‘and the third anterior
covert on the same side. The first two were about the same length,
that is about 2 ft. 6 in.
In Cock A, the 3rd pair of rectrices had ceased to grow, and
were a little longer than the outer rectrices. The 2nd _ pair of
rectrices were. still growing, and were not much longer than the
3rd pair. The central pair were growing, and were only about
4inches long. The tail-coverts were growing, but still very short.
In Cock B, the 2nd pair of rectrices were growing, and about
9 inches long; ; the 3rd rectrix on the right side had ceased to
grow, and the basal sheath was peeling “off, the corresponding
feather on the left side was still growing.
Dec. 21st.—The third anterioun covert on right side, which was
pulled out last spring and grew again, appeared to have almost
come to the end of its growth: the basal barbs were downy, and
the sheath at the base had almost all peeled off. I had not pulled
it out.
History of Feathers of Second Growth in Cock B,
to Dec. 28, 1902.
Right side of Tail.
lst Anterior Covert: the preceding feather was accidentally
pulled out while growing on March 11th, when it was 1 ft.
8 in. long. It was accidentally pulled out a second time
while growing on August 24th, when it was 1 ft. 13 in. long.
A successor immediately sprouted in its place.
4th Anterior Covert: stopped growing, and formed a complete
quill a little before October 22nd, on which date it was pur-
posely pulled out, and its successor sprouted in its place: it
was not measured. Originally pulled out December 27th,
1901, when it was 1 ft. 2 in. long.
2nd Anterior Covert: preceding feather pulled out Mareh 2nd
when | ft. 44 in. long: accidentally pulled out again while
growing on November 17th (1 ft. 93 in. long), and its
successor sprouted in its place.
3rd Anterior Covert: predecessor pulled out February 11th, when
248 MR. J. T. CUNNINGHAM ON [ Mar. 17,
1 ft. 22 in. long: had almost entirely ceased to grow, but had
not been pulled out: it was a little over 2 feet long.
Posterior Covert pulled out February 17th, 1902, when it was about
2 feet long. Now still growing, and about 2 ft. 6 in. long.
Central Rectrix: predecessor pulled out while growing on
April 13th, when 2 ft. 9in. long: was still in full growth, and
about 2 ft. 6 in. long.
Left side of Tail.
Central Rectrix: predecessor pulled out while growing on
February 15th, when 2 ft. 41 in. long: was accidentally
broken off on October 29th, but continued to grow from the
base, and the basal portion is now about 7 inches long. The
total length of the feather, if it had not been broken, would
have been 3 ft. 04 in.
Ist Anterior Covert: predecessor pulled out growing on
March 10th, when 1 ft. 97 in. long: accidentally pulled out
growing on December 9th, when 2 feet long. Its successor
immediately sprouted again, and was just showing.
There were thus in Cock B only three feathers left of the eight
which had begun to grow before the moulting-season. The two
longest of these were about 2 ft. 6 in. long, while the third was
shorter and had ceased growing.
The other feathers which began to grow after the moult were on
the whole longer than the feathers in Cock A; and the difference
between the tails in the two birds, the one artificially treated and
the other left to nature, was sufficiently striking.
SUMMARY AND CONCLUSIONS.
These observations extended only to the 2nd moult, or the first
shedding of the adult plumage.
They show that when the feathers are not pulled or artificially
treated in any way, care being taken to protect them and prevent
them from being accidentally pulled out, the growth continues
till about the end of March, when it ceases, the quills of the feathers
are formed in the normal way and the feathers are moulted nor-
mally in the following autumn.
On the other hand, when the feathers are pulled out in the
spring, successors immediately sprout in their places, continue to
grow till the following season, when they go on growing without
moulting, except in some cases when growth may come to an end
in the moulting-season.
In the cock whose feathers were stimulated by pulling, growth
did not go on at a more rapid rate, but continued for a longer time
and produced a longer feather. Thus in Cock A, no growth took
place after April Ist, and the maximum length was 2 ft. 43 in.;
while in Cock B, growth continued till July 13th, and the
maximum length was 2 ft. 94 in.
1903. ] JAPANESE LONG-TAILED FOWLS. 249
I think it is tolerably certain that these experiments explain
the method adopted by the Japanese in the production of tail-
feathers of extreme length, and that much that has hitherto been
mysterious and inexplicable in the matter is now explained. The
statement that the feathers do not moult or moult only once in
three years, or that the Japanese have a secret method of pre-
venting the moult, is explained if we assume that the Japanese
fancier strokes the feather as Mr. Sparks stated, and that he either
deliberately pulls a feather out when it shows signs of diminishing
growth, or that the feather is automatically pulled out in the
process when its growth diminishes ; because, as I have shown, the
attachment of the feather is very feeble when the growth is ceasing.
As arule, when a feather is pulled out its successor immediately
sprouts again, and its growth is afterwards continued regardless
of the moulting-season. The results of my experiments in fact
are in complete agreement with the statement of the matter
furnished by Mr. Sparks.
The long-tailed cock in its perfection, therefore, is neither a sport
nor a br eed, but a product of artificial cultivation ; and the excessive
growth of the feathers is the result of stimulation applied to the
individual. The most important part of the stimulation is not
the mere pulling of the feather, but the extraction of it which
causes the growth of its successor.
On the other hand, the method of treatment is applied not to
any breed at random, but to a particular breed which includes
several varieties of colour, and apparently two varieties of comb.
It can scarcely be supposed that the same treatment applied to
another breed would produce results at all comparable, and we
therefore may conclude that in this special breed there is a special
and extraordinary tendency to growth in the tail-feathers and
saddle-hackles. The congenital peculiarity is evidenced in the
case of Cock A in my experiment. The question therefore arises,
whether this congenital peculiarity has been developed entirely by
spontaneous variations and selection, or whether it has been
influenced by the excessive growth artificially induced in every
generation. It is probable enough that in most cases a cock which
showed the most rapid and most prolonged growth was used for
breeding, but it is by no means certain that the cocks so used
were never subjected to the artificial treatment. The probabilities
ave rather the other way, that specimens which had been found
to respond to the treatment were used for breeding. It is a
significant fact that this is the only breed of long-tailed fowls in
existence; and that the method of treatment applied to it is so
elaborate and so absolutely artificial, requiring daily attention for
months and years. If a similar result could have been obtained
by selection alone, it is difficult to understand why poultry fanciers
in some part of the world have not made the discovery.
The results of this inv estigation are-remarkably in agreement
with the theory advocated in my book ‘Sexual Dimorphism in the
Animal Kingdom.’ In that book I pointed out that wherever
250 SIR CHARLES ELIOT ON NUDIBRANCHS [ Mar. 17,
hypertrophy of organs or parts occurred in one sex only, the parts
affected were subjected by the habits of the animal, as known by
observation, to special irritation and stimulation.
Postscript, added April 25th.—On March 9th, as the left central
rectrix in Cock B seemed to be ceasing to grow, I tried to pull it
out, but it broke off at the top of the sheath. The total length
from the surface of the skin was 4 feet. The right central rectrix
in the same bird-on March 16th was 3 ft. 4 in. long and still
growing. At present it shows no sign of cessation of growth,
and has now been growing for more than a year. The appearance
of the two birds at the present date is shown in the two photo-
graphs (text-figs. 41, 42, pp. 240, 241).
2. On some Nudibranchs from Hast Africa and Zanzibar.
Part I. By Sir C. Huot, K.C.M.G., H.M. Commis-
sioner for the Hast Africa Protectorate, F'.Z.S.
[ Received February 20, 1903. |
CERATOPHYLLIDIA AFRICANA, gen. et sp. nov.
One specimen from near Wasin, E. Africa, in 9 fathoms.
The living animal was described by Mr. Crossland, who dredged
it, as of a light greenish-yellow colour on the upper surface, but
with the foot, branchiz, and under side of mantle, white. The
back was very hard and smooth, but its most remarkable character-
istic was the presence of a number of papilla, consisting of round
or pear-shaped bodies set on stalks. The stalks as well as the
base and tip of these globes were white, but the middle part was
black, owing toa dense aggregation of black spots, which, however,
can be seen to be separate under a lens. The globes were quite
soft and the stalks flexible; they shook when the animal was
moved, but were not observed to execute any spontaneous move-
ments. The mantle-edge was wavy.
The alcoholic specimen is of a uniform pale lemon-yellow, the
black bands of the globes being, however, preserved. The breadth
across the middle of the back is 1:9 centim. Unfortunately the
animal is contracted almost into a circle, but apparently the
length, when stretched out, must have been about 2°2 centim.
The consistency of the body is like hard wax, and fragments of
the mantle, which is ample, could easily be detached with the
forceps. The whole dorsal surface is a mass of closely packed
spicules. It bears about a hundred of the stalked globes. They
are of very varying size; many are quite minute, but the largest
is about 3 millim. high including the stalk, and about 2 millim.
across the ball, which is quite soft and can easily be pressed flat.
They are distributed over the whole of the back irregularly, and
not in any pattern, but are perhaps thickest round the mantle-
edge, including the space in front of the rhinophores. Both the
1 For Part I. see P. Z. S. 1902, vol. ii. p. 62.
1903. ] FROM EAST AFRICA AND ZANZIBAR. 951
rhinophore-pockets and the anal papilla project ; the edges are
smooth. In the preserved specimen the rhinophores are grey.
The branchiz ave arranged in a circuit interrupted only by the
head and genital papilla. They vary in size, but though in places
long and short branchi seem to alternate, this cannot be said to
be the general rule.
Over the mouth are two tentacles each about 1:5 millim. long,
and 1 millim. broad at the base. They are not directed sideways
but straight forward, and being set close together so that the
division is not visible, they appear to form a sort of head. They
are united at their bases. The mouth is larger than is usual in
this order, and though it is suctorial is hardly poriform. Though
the animal was dissected only three months after capture, the
internal organs were already much dried and shrivelled, the spirit
having apparently been unable to penetrate the hard integument.
It was clear, however, that the buccal organs are of the type of
Phyllidiopsis rather than Phyllidia. The buccal opening led into
a sausage-shaped tube about 6 millim. long and 2 millim. broad,
with muscular walls transversely striped. This passed into a long,
narrow, coiled tube, which preserved the same calibre until it
dilated into the stomach. ‘Two ample glands (salivary ?) entered
the larger part of the tube on either side, but were not in any
way fused with it. The liver was large and undivided behind.
The central nervous system was enclosed in a thin capsule and
somewhat concentrated, the cerebral and pleural ganglia being
hardly distinguishable and the pedal ganglia lying beneath them.
The eyes were large, black, and distinct. The genital mass was
much hardened, but the two spermatothecs, one white and empty
and the other black and full, were quite distinct. It was impossible
to ascertain whether the glans was armed with hooks and whether
the folds on the dorsal wall of the pericardium (sometimes called
the pericardial gill) were present, but it is highly probable that
the species possesses these family characteristics.
In virtue of its buccal apparatus this animal belongs to Bergh’s
genus Phyllidiopsis, although the tentacles are not attached
through their whole length and are rather larger than is usual in
the Phyllidiade. It is remarkable that the genus Phyllidiopsis
contains one species, Ph. papilligera, which has also black papille
on the back. To me, the presence of these dorsal papille seems a
peculiarity sufficiently marked to merit generic rank. If Hchino-
doris is a genus, why should not Phyllidiadze which have the same
peculiarity enjoy the same distinction? I would propose to call
the genus Ceratophyllidia, and its characters will be :—Back
studded with papille ; buccal apparatus in the known species
similar to that of Phyllidiopsis.
PLEUROPHYLLIDIELLA HORATII, gen. et sp. nov.
One specimen from Wasin, East Africa. Mr. Crossland, who
captured it, gives the following notes on the living animal :—
“Three inches long. Mantle edged with light salmon-colour; it
252 SIR CHARLES ELIOT ON NUDIBRANCHS [ Mar. 17,
ends in two ridges near rhinophores, similarly edged; front edge
of velum similarly edged. Back grey, mottled with a darker
shade, the top of the numerous longitudinal ridges being sprinkled
with clear black spots. Underside white. Gills also white.
Rhinophores longitudinally lamellated, grey in colour, and can be
contracted or retracted, though the pockets do not seem very
definite or complete.”
The somewhat bent alcoholic specimen measures 3 centim. from
head to tail, but would be at least 5 longer if it were straightened
out. The breadth of the back is 1°5 centim., of the foot °6. The
foot is long and narrow, pointed behind and truncate in front,
the corners not projecting.
No caruncle or nuchal papille are visible, but it is somewhat
difficult to reconcile the head-parts of the preserved specimen with
the description quoted above. According to a rough drawing
made from the living animal, it would seem that the mantle-edge
passes between the rhinophores and forms a sort of velum in front
of them; but in the preserved specimen it appears to le behind
them as in an ordinary Plewrophyllidia, and not to pass through
them at all.
The salient character of the genus is that there are no branchize
and no trace of a branchial cleft. Lateral lamelle are, however,
present. They are situated exclusively on the under edge of the
mantle, and not on the sides of the body. They extend from
the head to the tail, and are about 30 in number on each side.
They are irregular in size and shape. Some terminate before
they reach the mantle-edge, and some run from the mantle-edge
only halfway to the body. The genital papilla is 4 millim. and the
vent 1-2 centim. from the anterior end of the body. Cnidophores
are distinctly visible round the edge of the mantle.
The mouth is ventral, and forms a large transverse slit, with
slight indications of a T-shape. The jaws are yellow, narrow, and
united so as to form a shape like a boat. They bear no denticles,
but there are a few irregular coarse indentations of the edge, due
apparently to its being jagged by use. The radula consists of
about 30 rows, the formula for each of which is about 50+1-+ 50.
The central tooth consists of a squarish basal plate with a long
cusp, which bears about 10 denticles on each of its sides, The first
lateral is larger than the others and resembles the central tooth,
except that the denticles are only on the internal side. The
second and third laterals are also denticulate; the rest appear to
be simply hamate.
This form appears to be intermediate between Pleuroleura,
which has neither branchial clefts nor side lamelle, and Pleuro-
phyllidia, which has both. I have indicated its affinities by the
name Pleurophyllidiella.
BAOLIDIA MAJOR, N. sp. }
(Bergh, in Semper’s Reisen im Arch. Philipp., Malac. Untersuch.
vol, iii. pp. 778-780, 1880.)
1903. | FROM EAST AFRICA AND ZANZIBAR. 253
One specimen from Chuaka, E. coast of Zanzibar, under a stone
between tides.
The living animal was about 4 centimetres long. The body and
appendages were of a uniform greyish white, with spots of a dull
opaque white. The whole animal closely resembled a kind of
detachable sea-anemone which is very common at Chuaka, and
appears to be sometimes almost free-swimming.
The alcoholic specimen is 3 centim. long and 1 centim. broad
at the widest part, including the cerata. The foot is moderately
broad, and has fairly long tentacular expansions in front; but
its most remarkable character is the size and distinctness of the
anterior groove, which measures 2 millim. across. The upper lip
is separated into two parts bya deep cleft. The oval tentacles are
large and very thick. The rhinophores are shorter and studded
with minute knobs, which, in the preserved specimen at any rate,
appear not to be set in rings. The cerata are much flattened and
almost leaf-like, and the hepatic diverticula within them are
ramified. They bogin at the anterior end of the large peri-
cardial prominence, and are arranged in about 17 groups on each
side, each containing about 10 cerata. There are very distinct
gaps between the anterior groups, and a broad bare space runs
down the middle of the back, but towards the end of the body the
cerata are huddled together and continue until the extreme tip,
there being no tail. The outermost cerata of all the rows ave
smaller, and the inner considerably larger, but at the base of the
innermost are frequently quite small, some hardly larger than
tubercles. The genital orifice is below the first group of cerata,
and the lateral vent behind the second.
The specimen was only partially dissected. The jaws are very
large, colourless and transparent, with a perfectly smooth edge.
The radula consists of 32 pectinate teeth, very similar to those
of B. mebii (see Bergh, /. c. pl. Ixxix. fig. 16), with striations
under each denticle. They are, however, very much broader, the
widest measuring 2 millim., and the denticles are more irregular
in shape, being probably worn by use. There are about 150 of
them on the broader teeth. The three or four central denticles
are generally, but not always, smaller than the others.
This specimen is clearly referable to Bergh’s genus Beolidia,
and the difference between it and the type is mainly one of size,
B. mebii being only 8 millim. long. The similarity of habitat
makes one think that this may be merely a full-grown individual
of the same species ; and we know so little of the variations which
the radula and arrangement of cevata may present in AXolids at
the different periods of their growth, that I am not prepared to
reject this hypothesis. Still, the single specimen examined by
Bergh appears to have been sexually mature, and this being so
the two animals each present peculiarities amounting to specific
differences :—(1) In B. mebii the tentacles are said to be “ iibge-
plattet fingerformig” ; the cerata begin behind the rhinophores and
are set in rows: in &, major the tentacles are stout and round ;
254 SIR CHARLES ELIOT ON NUDIBRANCHS | Mar. 17,
the cerata begin further back and are set in groups. (2) In
B. mebu the reproductive orifice is under the third row of cerata,
and the vent between the sixth and seventh rows, almost dorsal :
in BL. major the reproductive orifice is under the first group of
cerata, and the lateral vent after the second. (3) In B. major
the basal part of the teeth is proportionally narrower than in
&. mebii, and the denticles are more irregular.
Var. ORNATA.
One specimen captured at Chuaka, May 1902, seemed to be a
typical Beolidia major, except for a somewhat more ornate
coloration. The ground-colour was of a yellowish-white with a
yellowish-brown pattern, consisting of a series of irregularly shaped
lozenges containing white spots, down the middle of the back as
in B. major. The oral tentacles were white with green stripes.
The cerata were white with yellow tips, below which was a bright
blue band.
CERBERILLA AFRICANA, 0. Sp.
(Bergh, Journ. Mus. Godef. Heft ii. 1873, & Heft vi. 1874;
Beitr. zur Kenntniss der Aeolidiaden, Theil 11.; Semper’s Reisen
im Arch. Philipp., Malac. Untersuch. 11. p. 879.)
One specimen from the reef Jembiani, Zanzibar, 3°5 centim. long
and 1:3 broad in life. The back was almost entirely covered with
cerata, so that the body-colour was hardly visible. Most of the
cerata were very dark green with a bright yellow ring, but the
innermost were white with bluish tips, with only a ring of dark
green. The oral tentacles were dark blue, with green bases. The
rhinophores had four bands of colour, which were, starting from
the base, greenish brown, white, blue, white. On the head were
two yellow lines, extending from the oral tentacles to the rhino-
phores, and the margins of the head and foot were also of a bright
light yellow.
The animal was stoutly built. The foot projected considerably
beyond the body on either side. Its anterior angles were expanded
into long tentacular processes. The head had also two lateral
expansions, from which projected at right angles the very large
and conspicuous oral tentacles. In life the rhinophores were
quite simple and fairly long. In the alcoholic specimen they were
contracted and somewhat wrinkled. The numerous and thick-set
cerata were disposed on peduncles. There was a bare triangular
patch behind the rhinophore, and a narrow bare space down the
middle of the back, but the cerata folded over the latter so that
neither it nor the transverse bare areas were visible. There were
about twenty transverse rows of cerata. The first row at the side
of the rhinophores consisted of about 10 cerata, much smaller than
the rest. After the third row was a distinct gap, and a smaller
gap after the fourth. After that the rows were so close together
that they could not be distinguished superficially. The innermost
cerata were larger than the others, and sometimes bifid: small
cerata extended almost to the end of the very short tail.
1903. ] FROM EAST AFRICA AND ZANZIBAR. 255
The jaws were large, with a smooth cutting-edge. The radula
consisted of a single series of 13 transparent yellowish teeth of the
shape usual in the genus, viz., pectiniform, with large irregular
denticles and small accessory denticles. It was sometimes hard
to decide whether the rather small denticles should be considered
main or accessory; but the average number of main denticles on
each tooth was 10, and the largest number (in one case only) 13.
The central nervous system was somewhat concentrated. The
specimen was only partly dissected.
A new species must, I think, be provisionally created for this
animal, though the discovery of intermediate forms may perhaps
render its retention unnecessary. In some ways it is itself a con-
necting-link between C. longicirrha and C. annulata, for the former
is said to have 7, and the latter 5 denticles on each side of its
teeth, whereas C. africana has 4, 5, 6, ov 7 indifferently. It can
hardly be C. annulata, for the difference in colour is too great,
and besides there is much less bare space on the back. Neither
can it be C. longicirrha, because (1) the coloration, though similar,
is still distinct ; (2) C. longictrrha has the back bare up to the 7th
row of cerata, and some of the cerata are very long, which is not
the case here; (3) the rhinophores are not perfoliate.
This last pomt is of some importance for the characterisation
of the genus. In the present animal the rhinophores were
undoubtedly quite simple in life, and in alcohol they are wrinkled,
though it is still possible to see that they are not really perfoliate.
In C. longicirrha, Bergh says the perfoliations are 14 or 15 in
number, and not deep. Of C. annulata he says that the rhino-
phores have 12-14 well-marked perfoliations, and that Garrett has
wrongly represented them as simple. But im Semper’s ‘ Reisen,’
xvii. he states that C. annulata var. affinis has simple rhinophores,
and gives as a generic character: ‘‘ Die Rhinophorien scheinen
nicht perfoliirt zu sein.” I have not access to part 1x. of his
‘ Beitriige zur _Kenntniss der Aeolidiaden,’ which perhaps explains
the matter; but it looks as if the rhinophor es are simple, but have
a tendency to simulate perfoliations when preserved.
PYrERAEOLIDIA SEMPERI.
(Bergh, Beitr. zw Kennt. der Aeolidiaden, iii. p. 22, and in
Semper’s Reisen, Malac. Untersuch. vol. i. p. 18 (1870); under
Flabellina.)
Four specimens, which seem probably referable to this species,
were dredged from 3 fathoms near Chuaka in July 1901. The body
is very long, narrow, and vermiform, the largest individual being
5°5 centim,. long and only 3 millim. broad. The ground-colour of
the body in the living animal is brown, with opaque markings of
very light green on the sides and back. The cerata are also dark
brown, with numerous thin lines of the same green. The top of
the head and the ends of the oral tentacles are opaque yellowish
white. The lower part of the tentacles brown, with three rings of
Proc. Zoot. Soc.—1903, Vout. I. No. XVII. Ai
256 SIR CHARLES ELIOT ON NUDIBRANCHS [Mar. 17,
vivid crimson-lake, The body of the rhinophores is brown and
inconspicuous, but the tips are whitish and have a crimson ring.
Although the colour of the animal when analysed is as described,
the general effect in most lights is that it is purple with a silvery
glaze.
The specimens were active in their movements, and in particular
the long oral tentacles were very mobile. The anterior margin of
the foot was produced into two deeply-grooved processes, and its
sides projected in two laminz along the whole length of the body.
The cerata, which were not at all caducous, were set on fan-like
projections of the dorsal margin, from 13 to 20 in number, on
each of which were from 10 to 20 cerata. The largest individual
appeared to have about 640 cerata in all. None of the cerata were
large, but they became more crowded and smaller towards the end
of the body, where they extended to the tip of the tail. In three
of the preserved specimens there was a very distinct raised border
on each side of the back, connecting the fan-like projections, and
almost resembling the mantle-edge of Dorids; but in the fourth
this remarkable character was not visible. The oral tentacles were
very large and long; the rhinophores, which stand between the
first clumps of cerata, were small and lamellated, except the tips,
which were narrow and smooth, In the individual dissected the
radula consisted of a single series of 18 yellowish teeth. The
separate teeth were much as represented in Bergh’s plate (J. c.
pl. i. figs. 5, 6, 7), but somewhat more regular in shape, having
nine denticles very symmetrically arranged on each side of a
central cusp, but not taking the form of indentations of the cusp
itself. The cutting-edge of the jaw bore a row of fine but very
distinct denticles, at the side of which were several rows of less
distinct accessory denticles. No spine or other armature was.
discovered in the reproductive system.
These specimens seem to clearly belong to Bergh’s genus Pteraeo-
lidia. There are some small differences (such as the shape of the
rhinophores and of the teeth and the lateral ridges) between them
and his description and plates of Pteraeolidia (Hlabellina) sempert,
but the ridges were absent in one specimen and the other characters
were slight. It is possible, however, that a comparison of the living
animals might show a specific difference.
ERCOLANIA ZANZIBARICA, 0. Sp.
(For the genus Hrcolania see Trinchese, ‘Un nuovo genere della
fam. degli Holididei,” Ann. del Mus. di Stor. Nat. di Genova, ii.
1872; id. ‘“‘ Aeolidee del Porto di Genova”; Bergh, Beitr. z. Kennt.
der Acolidiaden, v. pp. 9-18, Wien, 1878; Vayssiére, Recherches
sur Moll. Opisthobranches, Marseilles, 1888, pp. 121-128.)
Two specimens captured at Chuaka, East Coast of Zanzibar,
in February 1901. The living animal was rather more than
2 centimetres long, very fragile and delicate, transparent and of a
uniform bright green, the hepatic diverticula in the cerata not
1903. ] FROM EAST AFRICA AND ZANZIBAR, PATI
having been distinguished by any difference or intensity of colour.
It exactly resembled a piece of the racemose seaweed (Caulerpa)
on which it was found,
The length of the alcoholic specimens is 2 centim.; the extreme
breadth of the back with cerata 8 millim., and of the foot 2°3 millim.
The rhinophores are long and distinctly canaliculate. There are
no oral tentacles, but two lobes over the mouth. Behind the
rhinophores are two very distinct black eyes. The cerata are
club-shaped as in Galvina, of varying size, the largest inside. On
each side of the back are four clumps of about nine cerata each,
and there is a thick bunch on the tail, which, however, projects a
considerable distance behind the last cerata. Down the centre of
the back is a broad bare space, in the anterior portion of which
is the very large, elongated (not oval) pericardial prominence. In
front of this and fused with it is the vent, a large and conspicuous
tube. The foot is rounded in front.
I dissected one specimen, but was unable to obtain a clear view
of either the central nervous system or the reproductive organs.
The latter, as usual in this family, were extremely complicated,
both the prostate and albumen-gland appearing to be extensively
ramified. The verge was armed with a small spine. The hepatic
diverticula in the cerata, being colourless, were not easy to dis-
tinguish, but appear not to be much ramified and to resemble the
figure of those in Hreolania siottii in Trinchese, pl. ix. fig. 2.
The mouth-parts, buccal muscles, radula, &e. are of the usual
ascoglossan type. The teeth are not unlike those of Ercolania
viridis (v. Bergh, l. c. pl. xii. figs. 3 & 4), but the dorsal surface
is a simple curve and does not show any depression. The upper
portion of the radula contains 6 teeth, the lower 27, the last
members being arranged ina spiral like that depicted in Trinchese’s
plate of Z. siottii (pl. x a. fig. 1), from which it may be concluded
that the individual is young.
As the specimen presents all the characters of the genus Hrco-
lania, I describe it under that name, but I feel very doubtful if
the genus is valid. The only characters which differentiate it
from Stiliger, viz. that the rhinophores are canaliculate and the
pericardial prominence elongate and not oval, are surely very
slight. Vayssiére (J. c. p. 122) referred to the genus a species
(funerea) with entire rhinophores, which is probably in any case
a Stiliger.
The animal is not likely to be specifically identical with
L. viridis Bergh, for the coloration is not really the same, the size
is much larger, and the shape of the teeth somewhat different.
iW
258 MR. W. P, PYCRAFT ON THE 5h [ Mar. 17,
3. Contributions to the Osteology of Birds.
Part VI. Cuculiformes By W. P. Pycrart, F.Z.8., A.LS.
— [Received March 17, 1903. ]
(Plate XXII.’ & Text-figures 43-48.)
CoNnTENTS.
i. Introductory Remarks, p. 258. vii. The Pelvic Girdle, p. 280.
ii. The Skull of the Adult, p. 258. vill. The Pectoral Limb, p. 283.
iii. The Skull of the Nestling, p. 268. ix. The Pelvic Limb, p. 284.
iv. The Vertebral Column, p. 271. x. Summary, p. 286.
v. The Ribs, p. 274. xi. Key to the Osteology of the
vi. The Sternum and Shoulder-girdle, Cuculiformes, p. 288.
p. 275. xil. Explanation of the Plate, p. 291.
1. InrRopUCTORY REMARKS.
Although numerous contributions to the osteology of the
Cuculiformes have been made during the last quarter of a
century, the present paper claims to be the most comprehensive
survey of the group from this point of view which has yet been
made. Nevertheless, much remains to be done before our know-
ledge is anything like complete on this subject. How much, may
be gathered from the fact that out of 45 generally recognised
genera of the Family Cuculide, only 20 are represented in the
osteological collections of the British Museum, and this appears to
be the largest collection of the group extant. Of the 6 genera of
the Family Musophagide, only 2 are represented in our series.
1, THE SKULL OF THE ADULT.
The skull of the Cuculiformes recalls, on the one hand—through
the Cuculi—the skulls of certain Coraciiformes, e. g. Coracude
and Buccones; and on the other—through the Musophagi—the
skull of the aberrant Galliform Opisthocomus.
Nevertheless the Cuculiform skull may at all times be readily
distinguished, and by the following characters :—
The pterygoid is always free, and articulates with the palatine
—after the fusion of the hemipterygoid—by a more or less
obliquely transverse joint; the lachrymal is always present and
free; the antorbital plate (prefrontal) is large, and generally (if
not always) bears an os uncinatum; the vomer is vestigial or
wanting; the palate is indirectly desmognathous ; basipterygoid
processes are wanting ; and the postorbital processes are small and
inconspicuous, never extending down to the level of the quadrato-
jugal bar,
The skull of the Musophagi may always be readily distinguished
from that of the Cuculi by the fact that the mesial borders of
1 For Part V., see P. Z. S. 1902, vol. i. p. 277.
2 Hor explanation of the Plate, see p. 291.
P.Z.S.1903,vol. I. PL XXII.
H. Gronvold. del. Photogravure by Bale «Danielsson
OSTHEOLOGY OF THE CUCULIFORMES
1903.] - OSTEOLOGY OF THE CUCULIFORMES. 259
the palatines are cut away so as to expose the parasphenoidal
rostrum. &
The Occipital Region.
The plane of the occipital foramen slopes obliquely backwards ;
the orifice is either reniform or cordiform. In the obliquity of
this foramen the Cuculiform skull differs conspicuously from many
Coraciiform skulls wherein the foramen looks directly downwards,
its vertical axis lymg parallel with the long axis of the skull.
The supra-foraminal ridge is but ill-defined, but can be indistinctly
traced running downwards, on either side, into the processus ale
occipitalis.
The lambdoidal ridge in the Cuculiis generally well-defined and
bluntly conical in outline. In some genera, e. g. Crotophaga,
Centropus, Guira, Rhopodytes, Rhinococcyx, and Sauwrothera, rts
extremities bifurcate, sending forward an anterior band across
the inferior border of the squamosal to terminate in a pointed
squamosal prominence overhanging the quadrate; and a posterior
branch, which turns downwards and backwards to disappear on
the free edge of the lateral occipital. Where the lambdoidal
ridge is well- -developed, the cranial roof, lying immediately in
front, is more or less flattened, often, indeed, marked by deep
temporal fossee.
In many genera the lambdoidal ridge is much less distinct.
This occurs where the cranial roof is str ongly arched so as to rise
above the ridge with a strong curve, and Rae as fossee are
wanting or confined to the sides of the cranium, e.g. Coua,
Cacomantis, Chrysococeysx.
The supraoccipital region is marked by strong muscular depres-
sions, which in the case of Cowa, for example, take the form of
widely- separated oval scars, each surrounded by a swollen border ;
whilst in others, as in Budynama ys, these depressions are only
divided in the middle line by a narrow ridge, and are bounded
infero-laterally by slightly swollen bull produced by the lateral
occipitals.
The lateral occipital wing is produced downwards on either side
into a small or medium-sized processus ale exoccipitalis inferior
(paroccipital process).
In the Musophagi the lambdoidal ridge is fairly well-defined in
Schizorhis, but only very imperfectly so in T'uracus.
The ridge in Schizorhis is formed by the scooping-out, so to
speak, of the supraoccipital to afford depressions for the insertion
of the cervical muscles. On either side, and inferior to these
depressions, are two rather conspicuous bull lodging the posterior
and horizontal semicircular canals of the ear. The distal ends of
the lambdoidal ridge terminate before reaching the free edge of
the lateral occipital wing. The processus ale exoccipitalis inferior
is only feebly developed.
The lateral or exoccipital wings are not greatly developed in
the Cuculiformes.
260 MR. W. P. PYCRAFT ON THE | Mar. ilies
The Cranial Roof.—The cranial roof is never marked by supra-
orbital grooves.
The parietal region, in certain genera of Cuculi, is marked by
deep temporal depressions (temporal “ fossee ”), which may meet in
the middle line soas to form a faint sagittal crest, e.g. Khopodytes,
Piaya, Rhinococcyx, and Rhamphococcyx; ov they may be divided
by a low broad ridge, as in Geococcyx, Taccocoua, Saurothera,
Eudynamys, Guira, Centropus, and Crotophaga. In Coua,
Coccystes, Cuculus, and Cacomantis the temporal fossa is either
barely perceptible or confined entirely to the lateral aspect of the
cranium. Geococcyx, it should be remarked, presents an inter-
mediate stage in this character; the width of the ridge dividing
the right and left fossee being extreme, so much so that they
barely pass beyond the lateral aspect of the skull.
The interorbital region is moderately wide in all the Cuculi,
save in Sawrothera, but even in this genus it is not conspicuously
narrow. In the middle line the region may be marked with a
distinct furrow as in Cowa, or even by a low ridge as in Hudynamys.
The interorbital region is bounded anteriorly by the lachrymals ;
these do not develop outstanding supraorbital processes, and are
not conspicuous from this aspect of the skull. There is no trace
of the fusion of the frontal and nasal bones.
In the Musophagi the temporal fosse are not strongly marked,
and are confined to the lateral aspect of the skull. The parietal
region is more rounded than in the Cuculi; and in Zwracus is
marked by a median furrow, so that the form of the cerebral
hemispheres is distinctly indicated. In other respects this region
of the skull resembles that of the Cuculi.
The Base of the Skult.
The basitemporal plate, in the Cuculi, is flattened and triangular
in shape, and with the apex lying immediately beneath the
Eustachian apertures. Posteriorly it is bounded by a precondylar
fossa, which, though generally barely perceptible, in Hudynamys,
Centropus, and Geococcysx is fairly deep. The lateral angles of the
base are truncated, and, turning sharply downwards, cross the
paroccipital notch to form the lower portion of the rim of the
mouth of the tympaniccavity. In some genera, e. g. Coua, Guira,
these downwardly-directed angles are very prominent and simulate
mammillary processes.
The two sides of this triangular plate have fused with the
ossified connective tissue forming the anterior wall of the recessus
tympanicus anterior. Consequently the HKustachian grooves have
been converted into tubes opening directly above the apex of the
triangle.
The parasphenoidal rostrum bears slight vestiges of basi-
pterygoid processes in Hudynamys only among the Cuculi.
The rostrum is relatively long, and curves slightly upwards.
In the Musophagi the basitemporal plate, like that of the
1903. ] OSTEOLOGY OF THE CUCULIFORMES. 261
Cuculi, is triangular ; but it differs from the Cuculine plate in that
it is somewhat swollen, a feature which is especially well-marked
in Schizorhis. The two sides of the triangle are produced hack-
wards over the paroccipital notch to join the lateral occipital wing
of the tympanic cavity: thus they come to project beyond the
base of the triangle, though they are less conspicuous than in
some of the Cuculi. There is a barely perceptible precondylar
fossa.
The Lateral Aspect of the Cranium.
The tympanic cavity is oval or oblong in form and of moderate
size. It is bounded in front by the quadrate, behind by the
lateral occipital tympanic wing, below by the basitemporal plate,
and above by the squamosal prominence and by the head of the
quadrate.
Within the mouth of this cavity lie several smaller apertures.
The largest of these is that of the recessus tympanicus anterior.
Immediately below this is the mouth of the Eustachian tube.
The foramen ovale and foramen rotundum, three foramina of the
sinus petrosus, and the mouth of the posterior tympanic recess
open into the cavity by a common aperture—the fenestral recess—
which is very small, and lies mesiad of the articular surface for
the otic head of the quadrate. The mouth of the posterior
tympanic recess is completely concealed, and can only be made
out by cutting away its outer wall and passing a bristle through
from its cavity into the common aperture of the foramina in
question. In some birds, e. g. Malco, the posterior tympanic recess
communicates with the tympanic cavity by two apertures—one
caudad of the foramina ovale and rotundum, and lying within the
fenestra through which these are approached ; and the other external
to this fenestra, separated therefrom by a bony column, and lying
immediately beneath the articular surface for the otic head of the
quadrate. This external apertwre is wanting in the Cuculitormes.
Immediately above the head of the quadrate, and between the otic
and squamosal heads, will be found the aperture of the superior
tympanic recess, which is reduced in this group to extremely small
dimensions. The foramen of the 7th (facial nerve) opens also into
the tympanic cavity between the otic articular surface for the
quadrate and the rim of the anterior tympanic recess.
The squamosal and otic articular surfaces for the quadrate form
a dumbbell-shaped area immediately in front of the aperture of
the superior tympanic cavity. The actual articular surfaces are
formed by the expanded ends of the dumbbell only. Where the
pneumatic area is very large, as in the Striges and Falconiformes
for example, it breaks through the connecting portion of these two
articular surfaces so that they become quite isolated,
The Squamosal Prominence.—The size of this prominence
depends largely upon the development of the temporal fossa,
When this is deep the prominence is large; but when shallow the
prominence is quite inconspicuous. Its anterior angle is produced
262 MR. W. P. PYCRAFT ON THE [Mar. 17,
forwards into a more or less well-developed spine, the processus
zygomaticus squamost. The base of this angle serves for the
articulation of the squamosal head of the quadrate. This zygo-
matic process is especially well-developed in Cuculus, Crotophaga,
and Geococeyxz. It is small in Guira and Coua, for example,
and in the Musophagi. The processus articularis squamosi is
wanting.
The temporal fossxe are never very deep, but they may extend
inwards to within a short distance of the middle line. The
variations in the extent of these fosse have already been dealt
with (supra, p. 260).
The trigeminal foramen in the Cuculi lies immediately above
the mouth of the anterior tympanic recess, on a level with, and in
front of, the otic articular surface for the quadrate. In 7uracus
this foramen is divided by a median vertical bar of bone.
The orbito-nasal foramen (v') appears to be definite only in
Geococcyx. inother Cuculine and Musophagine the alisphenoidal
wall only partially ossifies, and consequently this and other
foramina around the optic foramen disappear in the dried skull.
The orbits are large, and roofed by moderately broad over-
hanging ledges formed by the frontals, and to a slight extent by
the lachrymals. The orbito-sphenoid does not ossify.
The interorbiial septum in the Cuculi is generally pierced by a
large fenestra, which attains its maximum in Dromococcyx and
Centropus. In © rotophaga, Coua, Piaya, Rhamphococcyx, Rhino-
coccyx, Khopodytes, and Zanclostomus, however, the fenestra is
greatly reduced.
In the Musophagi the fenestra is large in Twracus, small in
Schizorhis. I have not had an opportunity of examining other
genera.
The orbit is bounded in front by the antorbital plate (p. 263).
The lachrymal in the Cuculi is moderately large, with subequal
supraorbital and descending processes. The form of this bone
and its relation to the frontal and nasal bones vary considerably
however. In RLhamphococeyx calorhynchus only, apparently, is it
attached merely to the nasals. The orbital process looks entirely
forwards, instead of upwards: that is to say, it forms a shield
for the front of the eye instead of an overhanging ledge therefor ;
the descending process is slender. The frontals rest on the inner
angles of the orbital process of the lachrymal on either side, and
appear, in the adult skull, to be cut off from the nasals and pre-
maxillary process by a sharply-defined groove, which runs across
the skull from the hinder edge of the orbital process up the
lachrymal on the one side to that of the other, and simulates a
nasal hinge. Ahinococcyx curvirostris apparently closely resembles
Rhamphococeyx, but the lachrymal being completely fused with
the nasals and the frontal behind it, the precise relations of the
various regions cannot be made out.
In other Cuckoos, though exhibiting a distinct tendency to
form an anterior shield rather than a pent-house ledge, the
1903.] OSTEOLOGY OF THE CUCULIFORMES. 263
lachrymal extends backwards for some distance over the frontals.
In Hudynamys the frontals are deeply notched to receive the
orbital process of the lachrymal, which is more or less diamond-
shaped. In other genera the orbital process of the lachrymal
may be semilunar or rod-shaped. Where the antorbital process
is very large, the descending process of the lachrymal is always
more or less markedly degenerate. In Cuculus it has disappeared
altogether, whilst the orbital process has fused with the frontal
and nasal bones. In Zanelostomus javanicus it is reduced to a
mere vestige. The descending process is unusually large in
Crotophaga, one of the genera, it will be remembered, with a small
antorbital process.
Geococcyx has the largest lachrymal of all the Cuckoos. It is
nearly columnar in form, flattened antero-posteriorly, and deeply
grooved immediately above the quadrato-jugal bar.
The lachrymal in the Musophagi in its general shape closely
resembles that of the Cuculi: the orbital process is, however,
inclined more upwards. The descending process, judging from
what obtains in Schizorhis and Musophaga, is fairly strongly
developed. In Schizorhis the free end of the descending process
is twisted in itself, rests upon the quadrato-jugal bar, and articu-
lates by its postero-internal angle with the enormous ossiculum
lachrymo-palatinum.
The lachrymo-nasal fossa varies considerably in size. Among
the Cuculi, it may be described as large in Hudynamys, Centropus,
Guira, and Crotophaga. In Coua, Rhamphocoecyx, and Geo-
coccyx, for example, it is extremely reduced by the encroachment
of the lachrymal.
The Ethmoidal Region.—The mesethmoid, both in the Cuculi
and the Musophagi, terminates abruptly immediately in front of
the antorbital plate. The horizontal plate formed by the expan-
sion of its dorsal border is but feebly developed.
The antorbital plate (prefrontal), which, with the lachrymal,
bounds the orbit in front, is, in some Cuculi, e. g. Cacomantis,
Centropus, Coua, Coccystes, Cuculus, Hudynamys, Geococcyx,
Piaya, Taccocoua, and Sauwrothera, conspicuously large, quadr: ite
in form, and has its dorsal border pierced by two focaiine end
inner for the passage of the olfactory nerve, and an outer for the
orbito-nasal (v').
In other forms, e. g. Guira, Crotophaga, the antorbital plate,
though large, has fie exter nal lateral and inferior borders
deeply, excised. In such cases the orbito-nasal foramen described
above is represented by a notch. Attached to the inferior border
of this plate is a vestigial ossiculwm palatinum, which, however, is
generally wanting in dried skulls.
In the Musophagi the antorbital plate is greatly reduced,
forming but a triangular process projecting from the mesethmoid,
From its inferior border depends a well-developed ossieulum
lachrymo-palatinum. In Schizorhis this bone is relatively
enormous, projecting downwards to articulate by a strap-shaped
264 MR. W. P. PYCRAFT ON THE [Mar. 17,
limb with the outer border of the palatine. Its further relations
with the lachrymal will be discussed presently.
The nasal septum is completely ossified, and the external nares
are In consequence imperforate. The ectethmoidal region presents
several points for comment. By the ossification of the alinasal
wall, the form of the external nares is greatly changed in many of
the Cuculi. In consequence of this ossification, the space between
the premaxillary and descending processes of the nasal becomes
obliterated. Thus in Geococcyx the nostrils take the form of
an oval aperture lying just cephalad of the middle of the beak.
In Rhamphococcyx this aperture is still further reduced, and
consists only of an elongated slit lying at the base of the beak
above the tomium. In /hinococcyx the aperture is nearly circular
at the base of the beak, and bounded by a deep groove in front.
The alinasal ossification 1s, however, imperfect, inasmuch as a
small round hole is left in the angle between the premaxillary
and descending processes of the nasals. In TVaccocowa this
superior foramen and the aperture of the nostril have blended.
In Cowa the floor of the anterior region of the nasal fossa is
raised up into a sharp longitudinal ridge.
In the Musophagi the form of the external nostril differs con-
spicuously from that of the Cuculi. It is situated slightly distad,
or proximad, of a line passing through the middle of the beak, and
is reniform mm shape. In Zwracus it hes distad, in Schizorhis
proximad of the middle line. In both genera, just within the
mouth of the aperture, lies a well-developed concha vestibulum.
This is especially large in Schizorhis. Furthermore, the genus is
peculiar in that immediately distad of the concha the nasal septum
is pierced by a minute foramen.
The Cranial Cavity.—The metencephalic fossa is somewhat
shallower in the Musophagi than in the Cuculi. The internal
auditory meatus in the Cuculi is sharply defined, strongly con-
trasting in this respect with the Musophagi, in which it is but
ill-defined. The vagus foramen, which lies caudad and ventrad of
the meatus, is small. The orbito-nasal foramen (v') pierces the
outer superior rim of the fossa near the outer angle of the dorswm
sell.
The cerebellar fossa is relatively smaller in the Cuculi than in
the Musophagi, and in both it is relatively smaller than in many
other groups, ¢. g. Falconiformes. In the Cuculi the floccular
fossa is very shallow, and lodges a deep slit-like vertical
depression. In the Musophagi this fossa is reniform, deeper, and
lodges a deep and wide depression which gives the reniform
character to the whole.
The mesencephalic fossa is fairly sharply defined in both groups.
Among the Cuculi, this is especially noticeable in Geococcyx. Its
floor, in both groups now under consideration, is pierced by the
trigeminal foramen. The difference in position of this foramen is
worth noticing. In the Falconiformes it lies, together with the
orbito-nasal (v'), in a pit carved out of the superior border of
1903. ] OSTEOLOGY OF THE CUCULIFORMES. 265
the metencephalic fossa, midway between the sella turcica and the
floccular fossa, and is overhung by a sharp ledge formed by the
inferior border of the metencephalic fossa.
The pituitary fossa is large, deep, and tubular. The hinder
border of the fossa—the dorswm selle—forms a tumid lip.
Anteriorly the fossa is bounded by the pre-pituitary ridge, which
forms a broad triangular optic platform. This platform 1s carried
far forward into the interorbital septum, rising gently in its
course, and then turning abruptly backwards to terminate at the
pre-optic vidge, which may be traced into the tentorial ridge.
The form of the optic platform contrasts strongly with that in
some other types, ¢. g. Accipitres. The tentorial ridge in the
region of the pre-optic platform is but feebly developed. On the
parietal wall, however, it becomes tolerably distinct, especially in
Geococeyx and the Musophagi. In these, on its way to the median
falx it meets, near the crest of the epiotic, the sharply-defined
internal border of the mesencephalic fossa, and forms therewith
a prominent angular boss of bone, especially well marked in
Geococeye. The bony falx is not very strongly developed.
The oculo-motor (iii.) and abducens (vi.) nerves leave by a
common aperture, forming a deep groove across the posterior
angles of the optic platform.
The cerebral fosse lie quite in front of the cerebellar fossa, and
ave more flattened dorso-ventrally in the Cuculi than in the
Musophagi.
The olfactory fosse axe relatively feebly developed.
The Premacilla.
The external, as distinguished from the palatal, portion of the
upper jaw is made up mainly by the premaxilla. This region of
the jaw, in the Cuculiand Musophagi, varies considerably in form.
Normally, it may be described as about as long as the cranium.
In Crotophaga it is slightly longer than this, im Geococeyx and
Saurothera very much so. Typically, it may be described as
hemicrescentic in outline, depressed from above downwards, and
tapering from the base forwards. In Geococcyx and Saurothera
it is long and pointed; in Crotophaga surmounted by a sharp,
high keel; in the Musophagi more or less inflated.
The premaxillary portion of the jaw is slightly decurved at the
tip, and the palatal surface is level with the culmen. The nasal
processes of the premaxilla fuse completely with the nasals.
A nasal hinge oceurs in Z’uracus among the Musophagi, but is
wanting in the Cucull.
The Mawxillo-jugal Arch.
The maxilla is indistinguishably fused with the premaxilla.
The maxillo-palatine processes are expanded horizontally, and
meeting the ventral border of the nasal septum in the middle line,
form an indirectly desmognathous palate.
266 MR. W. P. PYCRAFT ON THE [ Mar. 17,
Among the Cuculi, the least specialised palate is probably to be
found in Cowa and Hudynamys. In Coua the maxillo-palatine pro-
cesses are widely separated and spongy in character; the consequent
palatal vacuity is filled by the nasal septum, which is also spongy
in character, and, fusing with the maxillo-palatine on either side,
forms an indirectly desmognathous palate. Hudynamys closely
resembles Cowa in this respect, but the bony tissue is less spongy,
and the fusion between the palatine processes and nasal septum is
more complete. Cuculus, Guira, and Centropus resemble Hudy-
namys. Further specialisation of the palate is seen in the still
further obliteration of the boundaries between the maxillo-palatines
and the nasal septum, and the tendency to shorten and broaden
the palatines. Forms like Gleococeyw and Rhinococcyx show how
these modifications have come about, whilst in Rhamphococcyx we
may see the maximum development of these peculiarities.
In the Musophagi the maxillo-palatines never appear to com-
pletely coalesce in the middle line, nor does the nasal septum
descend to the level of the ventral surface. The palate is never-
theless desmognathous, since this septum fuses with the dorsal
surfaces of the maxillo-palatines.
No separate elements can be distinguished in the quadrato-jugal
bar.
In the Cueuli the quadrato-jugal bar is almost or quite con-
tinuous with the tomium of the maxilla: in the Musophagi, on
the contrary, 1t rises considerably above the level at its distal end.
This difference is due to the fact that in the Cuculi the floor of
the maxillo-palatine process lies low, being only just raised above
the palatines at its postero-external angle, whilst in the Muso-
(=)
phagi this region is raised high above the palatines.
The Vomer, Palatines, and Pterygoids.
The vomer is absent in the Cuculi, vestigial in the Muso-
phagi. In the latter group it is spicular in form and fused with
the palatines, which it joins by means of a pair of very short
limbs. Anteriorly it touches the nasal septum.
The palatines, in the Cuculi, vary in shape. In the less
specialised forms, such as Hudynamys, Cowa, Gwira, they are
moderately long, and exposed slightly outwards. Anteriorly, from
the level of the forward face of the antorbital plate to the point of
fusion with the maxillo-palatines, the shaft of each is rod-like;
cephalad of this point the bone spreads out into a moderately
broad blade, the postero-external angles of which are rounded off,
whilst the mesial border is produced ventrad to form a more or
less pronounced keel. In OCwculws there is but little difference
in the width between the anterior and posterior moieties, and the
shaft is nearly straight.
In the heavy-billed forms like Rhamphococcyx and Rhinococcyx
the palatines are relatively shorter, and have the maxillo-palatine
extremity laminate instead of rod-like. The skull of Seythrops, I
regret to say, is not contained in the National Collection.
1903. ] OSTEOLOGY OF THE CUCULIFORMES. 267
In all the Cuculi, the palatines meet one another in the middle
line immediately beneath the parasphenoidal rostrum, which they
asp laterally through their fusion with the hemipterygoids.
In the Musophagi the palatines are relatively longer than in
the Cuculi, from which they also differ in that they taper, instead
of broaden, rapidly as they approach the pterygoids. Furthermore,
they do not meet below the parasphenoidal rostrum, but, on the
contrary, are separated one from another by the whole width of this
rostrum (Pl. XXII. fig. 2). The dorsal surface of the pterygoid
end sends up a long and deep, incurved keel, the antero-internal
angles of which fuse with the vestigial vomer.
The pterygoids, in the Cuculi, are moderately long and straight,
and in some, e. g. Crotophaga, Rhinococeyx, Gwira, Taccocoua,
have the dorsal border raised up into a high, sharp crest, the
palatine end of which embraces the parasphenoidal rostrum, whilst
in Eudynamys, Coua, Geococeyx, and Piaya, tor example, this
crest is absent. The palato-pterygoid articulation is in the form
of an obliquely transverse hinge-joint, permitting only a lateral
motion. Basipterygoid facets are absent.
The hemipterygoid element of the pterygoid is conspicuous only
in Taccocoua and Geococcyx. Herein it forms a continuation of
the dorsal crest of the shaft of the pterygoid, and rests on the
palatine at a slight angle. Secondary fusion between the hemi-
pterygoid and the main shaft of the pterygoid would reproduce
exactly the conditions of the palato-pterygoid articulation which
obtain in the Bucconide and Momotide.
In the Musophagi the pterygoids are relatively short, somewhat
twisted, rods, bearing vestiges of basipterygoid facets. They
articulate with the palatines as in the Cuculi.
The Mandibles.
The mandible in the Cuculi has a short, blunt angular process,
and a moderately long internal angular process.
In Eudynamys, Cuculus, and Guira the vami are pierced by a
long lateral vacuity, which is partly closed by a long and slender
coronoid. In Coua the coronoid terminates midway across this
vacuity; whilst in Zaccocowa and Centropus this vacuity is quite
open, the coronoid forming its ventral border. In Piaya, Rhino-
coccyx, and Rhamphococcyx the vamal vacuity is absent. In
Geocoecyx it is partly closed anteriorly by the hinder end of the
splenial.
The mandible of Cuculus possesses one conspicuous feature in
the presence of a triangular flange of bone formed by the deflection
of the superior border of the ramus at the point corresponding,
in the living bird, to the gape, in the skeleton to the region
immediately behind the lachrymo-nasal fossa. The flange,
especially conspicuous in Cuculus canorus, is also fairly distinct m
Cacomantis, and is traceable in Piaya and Rhamphococeyx. The
internal angular process is well marked in all the Cuckoos, but is
especially so in Coua and Centropus.
268 MR, W. P. PYCRAFT ON THE [ Mar. 17,
In the Musophagi the angular is sharply truncated posteriorly,
and the internal angular process is short and blunt.
In Schizorhis the ramal vacuity is closed posteriorly, in Twuracus
it is closed completely, by the coronoid.
The Hyoid.
Unfortunately the hyoid bones have been preserved only in one
or two of the skeletons of this group in the Museum Collection.
Years ago, when these skeletons were made, according to the
prevailing custom, the hyoid bones were not regarded as of value.
In Seythrops the basihyal is slender and rod-shaped ; the Ist
‘basibranchial triangular, with concave sides, the 2nd of medium
length, styliform and tapering. The ceratobranchial and epi-
branchial are subequal in length and offer no points for special
comment.
In the Musophagi the basihyal bones are reduced to mere
vestiges. The lst and 2nd basibranchials are short and fused.
The ceratobranchials are only slightly longer than the epi-
branchials.
ii. THe SKULL OF THE NESTLING.
The National Collection of nestling skulls of this group is
extremely limited, containing only one skull of Cuculus canorus
and one of Greococcyx californianus. A series of skulls of nestlings
and of immature individuals of various species of Cuckoos, as
well as of Plantain-eaters, would be a useful addition to the
Collection. At present the latter contains no skeletons of nestling
Plantain-eaters and few skeletons of adults.
a. Cartilage-bones.
The basioccipital is somewhat linguiform in shape, and widens
gradually from behind forwards. It is bounded on either side by
the lateral occipitals, in front by the basitemporal plate. Its
posterior border forms the greater portion of the occipital condyle.
The exoceipital, or lateral occipital, viewed externally, takes the
form of a broad horizontal plate bounding the basioccipital on
either side, and extending backwards and outwards, expands into
a fan-shaped plate, the superior half of which is wedged in
between the supraoccipital, parietal, and squamosal, whilst the
ventral, or downwardly directed, moiety forms the posterior
boundary of the tympanic cavity. The lateral segment of the
foramen magnum is formed by the horizontal plate of the lateral
occipital, which also, by the way, forms the extreme outer angle
of the occipital condyle. The fact that the exoccipital comes into
contact with the parietal is noteworthy: a similar relationship
obtains also in Dromeus among the Paleognathe, and will
doubtless be found among several other Neognathine forms. The
exoccipital is only just visible, from the cranial cavity being
overlaid by the opisthotic.
1903. ] OSTEOLOGY OF THE CUCULIFORMES. 269
The supraoccipital is extremely short antero-posteriorly, and
is deeply cleft in the middle of its superior border. Its external
lateral border fuses with the lateral occipital; within the cranial
cavity it is bounded by the epi- and opisthotic. The small size
of the supraoccipital recalls the skull of the Owls, but this of
course is but a coincidence. The character will doubtless be
found to obtain in the Kingfishers, Bucconide, and Capitonide,
which have many characters in common with the Cuckoos,
The provtie does not appear externally, Within the cranium
it occupies considerable space, forming the floor of the mesen-
cephalic fossa, as well as a considerable portion of the lateral walls
of the basin-like metencephalic fossa. It entirely excludes the
squamosal from the cranial cavity.
The epiotie is only partially ossified, and in such a way that its
boundaries cannot be made out.
The opisthotic has fused completely with the prootic.
The basisphenoid is not visible externally, being underlaid by
the basitemporal plate. Concerning its internal boundaries,
nothing satisfactory can be gathered from the two skulls in the
National Collection, the younger being damaged, whilst in the
more adult skull it has fused with the neighbouring bones.
The alisphenoid, in the younger of the two skulls, is not yet
completely ossified. As a result, between the external ventral
angle and the squamosal there is a wide gap, which extends
inwards below the inferior alisphenoid border, dividing its outer
moiety from the basisphenoid. This gap, when viewed from
without, is seen to be filled up by the prodtic. Its supero-
external angle is produced outwards to form the postorbital
process.
The orbito-sphenoid is not yet ossified in these skulls.
The presphenoid has fused with the basisphenoid.
The mesethmoid, in the two skulls now under consideration,
is yet incompletely ossified, forming but a linguiform plate; the
interorbital septum formed by the backward extension of the mes-
ethmoid having been represented only by cartilage. ‘The anterior
border of the linguiform plate is sharply truncated so as not to
extend beyond the level of the free end of the parasphenoidal
rostrum below and the anterior extremities of the frontals
above, ‘This truncation occurs at the cranio-facial fissure, which
has cut the mesethmoid into two parts: the one forming the
linguiform plate just described, which ultimately forms the
interorbital septum; the other, the septwm nasi, which in these
skulls is yet cartilaginous.
The cranio-facial fissure appears to be peculiar to the neo-
gnathine (Carinate) skull; but traces thereof are appa rently to be
met with in the Paleognathe (Ratitz), inasmuch as, in the skulls
of nestlings of Dromeus and Rhea in the Museum Collection, the
ossification of the mesethmoid commences, as in the Neognathe,
by the formation of a more or less linguiform plate, and this has
its superior border deeply excised, at a point exactly corresponding
270 MR. W. P. PYCRAFT ON THE [Mar. 17,
to the vertical face of the posterior division of the neognathine
mesethmoid, that is, exactly in front of the antorbital plate of the
adult. To make the fissure complete, the incision would have to be
continued downwards so as to completely bisect the plate, whilst
the rostrum would have to terminate at the free edge of the
posterior segment of the plate. In the adult skulls the incipient
fissure remains, placing the right and left olfactory chambers, in
the dried skull, in communication. The formation of a complete
eranio-facial fissure is correlated with, and perhaps consequent on,
the reduction in the length of the parasphenoidal rostrum, which
in all the Paleognathe is of great length. In a skull of a half-
grown Casuarius salvadorii it is, however, relatively much shorter
than in an adult C. australis, and much overhung by the mes-
ethmoid. Whether this peculiarity obtains also in the adult of
this species I am unable to say.
The olfactory cavities do not extend backwards in the Cuculi
or Musophagi, so as to lie on either side of the mesethmoid, as in
the Tubinares for example; the antorbital plate arising in the plane
of the cranio-facial fissure.
The guadrate does not differ materially from that of the adult.
The articulare is still distinct.
b. The Membrane-bones.
The parietal is oblong in shape, its anterior and posterior
borders sinuously curved, its mesial border straight, and its
external lateral border being slightly convex. It is bounded in
front by the parietal, behind by the supraoccipital, and laterally
by the squamosal.
The frontal has its hinder border simuously curved, and
throughout the greater part of its length applied to the parietal.
Its postero-external is closely applied to, and ultimately fuses
with, the supero-anterior angle of the squamosal and the post-
orbital process of the alisphenoid. In the supraorbital region it
turns downwards and inwards to form a broad overlapping plate
articulating with the alisphenoid. From the mid-orbital region
onward it becomes band-shaped, ultimately being produced into
an outwardly directed and blunt angle underlying the nasal.
The squamosal is roughly quadrate in form, and has the superior
anterior angle produced into a linguiform process which overlaps
the parietal and alisphenoid. Its antero-ventral angle is produced
into a small squamosal prominence, the under surface of which
affords the articular surface for the squamosal head of the quadrate.
The postero-ventral angle is obliquely truncated, and forms the
anterior segment of the rim of the tympanic cavity. The supero-
posterior angle is produced into a shght point which is wedged
in between the parietal and lateral occipital. The hinder border
of the squamosal serves to cover in the recessws tympanicus
superior, which is formed by the absorption of the diploé of the
lateral occipital.
1903. ] OSTEOLOGY OF THE CUCULIFORMES. 271
Internally the squamosal appears to be visible only in Cuculus
canorus and Centropus, where it appears as a small triangular
plate wedged in between the prootic, parietal, and alisphenoid.
The nasal, judging from the scanty material at my disposal,
differs slightly in form in different genera, inasmuch as in Cuculus
and Centropus it is not sufficiently large caudad to cover the
horizontal plate of the mesethmoid. In Centropus the hinder
border is produced into a sharp angle, and is rounded in Cueulus
and Geococeyx. ‘The nostril is holorhinal.
The lachrymal offers no special features for comment.
The premavilla is fully ossified only in the skull of Geococeyx.
Its nasal processes are long and slender, and the median suture
dividing them extends fax beyond the level of the anterior border
of the nasal fossa, The maxillary and palatine processes lie close
together and parallel with one another.
The mawilla in Geococcyx is elongated and triangular in form,
and sends backwards from its postero-external angle a long
slender bar to join the jugal and quadrato-jugal. The body of
the maxilla is slightly spongy and has the postero-internal angle
produced backwards and inwards into a maxillo-palatine process.
The body of the maxilla lodges a barely perceptible antrum.
The quadrato-jugal is long and slender in Geococcyx; the re-
lations between the jugal and maxilla cannot be made out, the
skull having become disarticulated.
The vomer is absent.
The palatine does not differ appreciably from that of the adult.
The pterygoid is rod-shaped, and produced forwards into a sharp
triangular spine (Pl. XXII. fig. 10), which articulates with a cor-
responding facet in the mesial border of the palatine. Later this
triangular process becomes segmented off from the main shaft
to form the hemipterygoid, which ultimately fuses with the
palatine. At the present stage this segmentation is faintly indi-
cated by a shallow furrow on the outer surface of the shaft. In
the adult, where the fusion with the hemipterygoid is complete,
the palato-pterygoid articulation is formed by the approximation
of glenoid surfaces sloping obliquely backwards.
The dentary and splenial do not differ from those of the adult.
The coronoid, angulare, and supra-angulare ave all as yet
distinct.
iv. THE VERTEBRAL CoLuMN.
The vertebral column of the Cuculiformes is not marked by the
same constancy of character which is so conspicuous in the skull,
In many respects it recalls that of the Coraciiformes, but even
among the Cuculi relatively wide differences obtain.
All the presynsacral vertebre are heteroccelous, and all the
thoracic vertebra are free.
The cervicals of the smaller Cuculi recall those of Leptosoma.
The odontoid ligament of the atlas is ossified, The axis vertebra
is very short antero-posteriorly, and has the neural arches produced
Proc. Zoou. Soc.—1903, Vou. I, No. X VIII. 18
272 MR. W. P. PYCRAFT ON THE oi Mlere alee,
outwards and upwards into a pair of pointed hyperapophyses.
The neural arches of the vertebre, from the third to the fifth
inclusive, are very broad, and with or without spines. From the
third to the sixth or seventh vertebie large metapophyses occur,
those of the fourth, fifth, and sixth being especially large, pro-
jecting like buttresses from the base of the anterior zygapophyses.
The metapophyses of the third vertebra send back each a slender
bar of bone to join the hyperapophysis. A similar bar of bone
occurs also from the fourth to the seventh vertebre, but instead
of joining the hyperapophysis, fuses with the antero- later: al angle
of the shield-shaped expansion formed by the neural arch. The
succeeding cervicals do not afford any particularly well-marked
characters in so far as this region is concerned.
The anterior cervicals of Scythrops and Cuculus (1 to 7) differ
from those of the Cuckoos just described in that they are
relatively slightly longer antero-posteriorly, that the bony bar
from the metapophysis to the hyperapophysis occurs only in the
3rd vertebra, and in that the shield-lke expansion of the neural
arches is very feebly developed. Short neural spines occur in the
2nd, 3rd, and 4th vertebre.
The 2nd, 3rd, and 4th vertebre bear hypapophyses; from the
5th to the 10th these are replaced by catapophyses, which from
the 11th vertebra backwards to the last cervico-thoracie are
replaced again by hypapophyses. The cervico-thoracie vertebree
are peculiar in that they bear more or less well-developed catapo-
physes in addition to the median hypapophysis. The catapophyses
never coalesce to form a carotid canal.
The cervico-thoracic vertebre may be two or three in number.
The number of the true cervicals varies, being either 11 or 12.
The thoracic vertebrz are free. The Ist or Ist to 3rd, as in
Hudynamys and Coua for example, bear hypapophyses. Pneu-
matic foramina pierce the centra of the vertebra beneath the
transverse processes.
The neural spines of these vertebrae gradually increase in height
from before backwards, and are more or less quadrate in form.
Coua, however, appears to form an exception to this rule, the spine
of the Ist thoracic being almost obsolete, and thus agreeing with
the cervico-thoracic vertebre ; whilst the spines of the 2nd, 3rd,
and 4th vertebre are, relatively, as high as in other Cuckoos.
The last two cervico-thoracic (2nd and 3rd) have apparently
only recently become separated from the thoracic series, inasmuch
as the vertebral segments of their respective ribs have undergone
no reduction in length. The ribs of the third pair still retain
their uncinate processes.
Only one thoracic vertebra, in the Cuculi, enters into the
synsacrum.
The synsacrum includes from 10 to 13 vertebre. The smaller
number appears to have been due to the reduction, possibly by
excalation, of the lumbar or lumbo-sacral vertebrze, or of caudal
vertebree, as in Geococcyx.
1903. ] OSTEOLOGY OF THE CUCULIFORMES. 273
The most complete synsacrum is that of Coua, and is made up
as follows :—1 thoracic, 3 lumbar, 3 lumbo-sacral, 2 sacral, and
4 caudal, making 13 in all.
Rhopodytes seems to possess the most reduced synsacrum : the
number of thoracic, lumbar, and sacral is the same as in Couwa, but
the lumbo-sacrals are reduced to 1 and the caudal to 2: making
therefore a total of 9 vertebra as against 13 in Coua.
Crotophaga and Geocoecyx each have 11 vertebrae in this region.
In the former, the reduction is from the lumbo-sacrals, which are
only represented by a single vertebra; in the latter there are
2 lumbo-sacrals, but only 3 caudals.
In other genera, the number of vertebrie appears constantly to be
12, composed as follows :—1 thoracic, 4 lumbar, 1 lumbo-sacral,
2 sacral, and 4 caudal.
In no case can distinct sacral vertebre be made out. The
synsacrum of Rhamphococeyx 1s vemarkable in that the single lumbo-
sacral vertebra bears a pair of exceptionally strong ventri-lateral
processes. Vestiges of these appear in Seythrops, Coua, and
Taccocoua.
In Centropus the third and fourth pairs of ventri-lateral processes
are apparently undergoing coalescence. In some skeletons this
fusion is complete; in others traces, more or less marked, of the
original buttresses still remain.
The last two caudal vertebra: combine to form a well-marked
planum anale.
There are 6 or 7 post-synsacrals—free caudal vertebra. Of these
the 4th and 6th bear pointed intercentra, which, however, are
completely fused with thew respective centra.
The total number of vertebre reaches its highest in Coua
with 37.
The vertebral column of the Musophagi is scarcely distinguish-
able from that of the Cuculi.
The cervical vertebree closely recall those of Cuculus. They may
be distinguished therefrom, however, apart from their greater size,
in that the 3rd and 4th both send backwards a bar from the meta-
to the hyperapophysis, and in that the 7th and 8th send back a
similar bar from the same region to the middle of the neural arch.
There are only two cervico-thoracies. The vertebra corresponding
to the third cervico-thoracic of certain Cuckoos, e.g. Scythrops,
becomes in the Musophagi thoracic, being joined to the sternum by
a sternal rib. The thoracic vertebre differ from those of the
Cuckoos, in that the transverse processes send backwards from their
postero-external angles a long bony spike to overlap the transverse
process of the vertebra next behind it. These connecting-rods
are either wanting or very feebly developed in the Cuckoos, e. g.
Seythrops.
The cervical vertebr are 12 in number, the cervico-thoracic 2,
thoracic 6, the last being fused with the synsacrum.
The synsacrum contains 13 vertebre, and is made up as
follows: thoracic 1, lumbar 3, Iumbo-sacral 3, sacral 2, caudal 4,
tS
274 MR. W. P. PYCRAFT ON THE | Mar. 17,
The 1st lumbo-sacral shows vestiges of ventri-lateral processes.
Only the 2nd sacral bears ribs.
There are 8 free caudals, but the 7th is nearly fused with the
pygostyle. The total number of vertebre is 41.
I must here point out that the determination of the sacral
vertebre in the skeletons herein described is a purely arbitrary one ;
they cannot possibly be determined with certainty without ex-
amination of the sacral nerve-plexus, and this I] am unable, at the
present moment, tomake. Thus, it may be that the plexus will show
that in some cases what is here described as the last lambo-sacral
in reality is the first sacral. If this should be the case then, of
course, the vertebra herein described as the 2nd sacral would be
the Ist caudal.
v. THe Riss.
The cervical ribs, in the Cuculi, form a series of very broad pleur-
apophyseal lamellee enclosing the usual vertebrarterial canal. These
plates extend backwards rather: beyond the middle of the centrum.
Seen from the ventral surface, the rib is free for about half its
length. Certain of the vertebre in the middle of the cervical
chain have this lamella pierced by a large fenestra. In Guwira,
Coua, Centropus, the 5th, 6th, and 7th are so distinguished; in
Crotophaga the 6th, 7th, and 8th; in Scythrops the 7th and 8th;
in Taccocoua, the 4th, 5th, and 6th. Cuculus, Chrysococcyx, and
probably other smaller forms, differ from the forms just described
in that the anterior cervical ribs are comparatively long and
slender, not lamellate; whilst from the 7th vertebra backwards
the ribs are reduced to vestiges.
The cervico-thoracie ribs number two or three pairs. The third
pair always bear uncinates.
There are five pairs of thoracic ribs, though not more than four
are attached to the sternum. The fifth pair are overlapped by
the preacetabular ilium, and may be reduced to the merest vestiges,
e.g. Rhamphococcyx and Coua, or they may be complete, as in Sey-
throps and Cuculus. They vary somewhat in form. For instance,
in Coua, Centropus, and Guira the proximal end of the rib is
extremely broad, and the shaft after leaving the vertebra almost
at right angles, curves abruptly downwards. In other Cuckoos
the broadening of the shaft is not conspicuous, and the shaft
slopes gently downwards and backwards, so that the thorax 1s
conspicuously broader at the articulation with the sternal ribs
than above.
The uncinates are broad and strong, and show a tendency to
develop a sharply defined postero-inferior angle at the junction
with the shaft of the rib; this is especially noticeable in Coua.
The sternal segments of the 5th pair of thoracic ribs never reach
the sternum. They may persist as vestiges, even the thoracic
segment of the rib reaching the verge of disappearance, as in
Coua and Rhamphococcyx, or they may be of considerable size,
1903. | OSTEOLOGY OF THE CUCULIFORMES. 275
and extend more than halfway down the sternal segment of the
Ath rib as in Seythrops.
The sternal ribs of Coua reynaudi ave relatively longer than in
other Cuckoos, and this fact appears to be correlated with
degenerate powers of flight. As a result of the lengthened rib-
segments and a change in the angle formed by the corpus sterni
and the vertebral column, to be discussed presently, the form of
the thoracic cavity differs conspicuously from that of other Cuckoos
and recalls features characteristic of ‘“ Ratite.” These peculiar
features ave wanting in Cowa cerulea.
The cervical ribs of the Musophagi closely resemble those of
the Cuculi, taking the form of broad pleurapophyseal lamelle.
From the 5th to the 8th these lamelle are fenestrated as in the
Cuckoos, so that the lamella appears to be joined to the centrum
by a slender bar of bone, the bar extending to the level of the base
of the postzygapophysis. There are six pairs of thoracic ribs, five
of which articulate with the sternum. The 6th pair are long,
but have lost connection with their sternal segments, which
remain as small spicules anchylosed with the sternal segment of
the 5th pair. In the Cuckoos, it will be remembered, there
are only 5 pairs of thoracic ribs, the 5th pair of which, like the
6th in the Musophagi, fails to reach the sternum.
vi. THE STERNUM AND SHOULDER-GIRDLE.
The sternum of the Cuculiformes recalls that of the Coraci
rather than that of any other group.
The sternum of the Cuculi may be readily distinguished from
that of the Musophagi in that it is relatively conspicuously shorter,
being indeed in many cases as broad as long. Moreover it
appears to be undergoing yet further reduction.
The form of the posterior margin of the corpus sterni and the
relative development of the spina eaterna present considerable
variations.
With regard to the posterior margin of the sternum, this appears
to be entire in Cacomantis only. Two extremely thin oval areas
indicate, however, the presence originally of a pair of posterior
lateral processes, the space between which and the median border
of the metasternum has been filled up. ;
In Scythrops, Cuculus, Coccystes, Guira, Centropus, and Croto-
phaga the posterior border is interrupted by a single pair of
notches, the outside of the notch being bounded by the posterior
lateral process.
All the other genera appear to have a doubly notched sternum,
but the character of the notches varies considerably. In Piaya,
for example, the posterior lateral process arises directly behind the
articulation of the 4th rib, and extends outwards and backwards
to terminate in a spatulate extremity some distance cephalad of a
line passing across the free border of the metasternum, Between
this process and the metasternum is a straight processus intermedius,
276 MR. W. P. PYCRAFT ON THE [ Mar. ris
which, like the external process, fails to reach the level of the meta-
sternal border. In Rhinococcyx, Rhamphococeyx, and Taccocoua
the posterior lateral process arises, as in Piaya, immediately behind
the articulation for the 4th rib, but it forms a wider angle with
the long axis of the sternum, and carries with it the intermediate
process, so that it looks like a branch thereof. In Cowa and Geo-
coccyx the posterior lateral process leaves the sternal plate further
back than in the forms just described. In other respects these
two sterna are quite distinct.
In Coua cerulea the posterior lateral process is very broad, and
extends backwards to the level of the posterior border of the meta-
sternum, whilst theintermediate processisshortand normaily placed.
Coua reynaudi, with an obviously degenerate sternum, differs from
C. cerulea in that the posterior lateral process is slender, and the
intermediate process shows a tendency to fuse with the base of
the posterior lateral. In Geococcyx the posterior lateral process
is long and slender, but fails to reach the level of the free border
of the metasternum ; whilst the intermediate process lies nearer the
posterior lateral than the metasternum. Further, the sternum is
peculiar in the great depth of the fissure on either side of the
metasternum, giving this a long and narrow shape.
A conspicuous pointed spina externa is found in Guira, Cuculus,
Geococcyx, and Scythrops. It is vestigial in Centropus. A small
spina interna occurs in Taccocoua.
In Rhamphococcyx the spina externa and interna are both
present, and fused to form a moderately large spina communis. In
Piaya, Rhinococcyx, and Coua there is a vestigial spina communis.
In Cacomantis the two spines remain distinct, but are extremely
reduced. A
Both sping externa and interna are wanting in Crotophaga.
The carina sternt is relatively largest in Cuculus (text-fig. 43,
p. 277), Cacomantis, and Scythrops. Its free ventral border is
strongly arched, and its anterior border is deeply concave. The
antero-ventral angle of the keel affords a firm articulation for the
fureula. In Coua, Rhamphococcyx, Crotophaga, and Piaya, the
anterior border of the keel is very deeply emarginate, causing the
free border of the kee! to project forward in the form of a long
style. With the dorsal surface of the style the clavicle articulates
by means of a long hypocleideum. Apparently the length of the
hypocleideum is directly correlated with the retreat of the carina
caudad. In the Galli this is well seen, the climax being attained
in Opisthocomus.
The depth of the carina is extremely reduced in Coua reynaude
(text-fig. 44, p. 277), so much so indeed that the power of flight
in this species must be extremely limited. In other Cuckoos the
depth of the carina is nearly or quite equal to half the breadth of
the corpus sterni; in the species in question the greatest depth
of the keel is barely one-fourth the breadth of the sternal plate.
Correlated with the reduction in the keel and the consequent loss
of flight, is a marked change in the position of the sternum, which
1903. | OSTEOLOGY OF THE CUCULIFORMES. 277
appears to have been thrust forward by a considerable increase in
the length of the posterior sternal ribs. As a consequence, the
shaft of the coracoid and the long axis of the sternum form nearly
a right angle with the vertebral column, and the shaft of the
scapula lies parallel with the vertebral axis, instead of forming an
oblique angle therewith. This forward movement of the sternum,
furthermore, has brought the acrocoracoid upwards to the level
of the neural crests of the vertebree, whilst the distance between
the acetabulum and the free edge of the metasternum has been
enormously increased. Jn fact, the thorax of this bird approaches
very closely, in the relative position of the sternum, that of the
“* Ratitee.”
Text-fig. 43. Text-fig. 44.
Text-fig. 43.—Left side view of sternum and shoulder-girdle of Cuculus canorus, to
show the great size of the carina sterni and the form of the posterior border
of the sternum.
Text-fig. 44.—Left side view of sternum and shoulder-girdle of Coua reynaudi, to
show the degenerate carina sterni, which is actually more reduced than the
figure indicates. Note also the long hypocleideum and the doubly notched
corpus sterni.
Shortening of the sternum is most marked in Piaya, Zanclo-
stomus, and Taccocoua.
The sternum of the Musophagi is relatively longer and more
Coraciiform than that of the Cuculi, butisatthesametime thoroughly
Cuculine in character. In both Zwracus and Schizorhis, the only
genera represented in the National Collection, the hinder border
of the sternum is doubly notched, and the notches are of consider-
able extent, the outermost being the deepest. The posterior
lateral process is slender, and directed straight backwards. The
intermediate process lies midway between the posterior lateral
process and the metasternum. The outermost notch is about
twice the depth of the inner.
The corpus sterni of Turacus is relatively much shorter antero-
278 MR. W. P. PYCRAFT ON THE [ Mar. 17,
posteriorly than in Schizorhis. The sternal notches of Z'wracus,
it is to be noted, though preserving the same relative proportions
between themselves, are yet relatively shallower than in Schizo-
rhis with regard to their relation to the corpus sterni.
There is a well-developed spina externa, but no trace of a spina
interna. In Schizorhis the spina is flabellate and projects from a
rounded base. In Zuracus it is quadrate in form. In 7’. buffoni
it projects nearly as far forwards as the antero-ventral angle of
the carina, and is distinguishable from the anterior border of the
carina only through the medium of a notch. Were this notch
filled up, the spina would disappear and the anterior border of
the keel would present a vertical face projecting Hae beyond the
coracoid grooves, as in certain Coraciiform birds, e. g. Cyanops.
The .coracoid grooves overlap one another. The dorsal lip 1 1s
extensive. <A median notch oceupies the place of the spina
interna, and this is bounded on either side by a conspicuous
oblong glenoid surface, which articulates with a special facet on
the coracoid. In the Cuckoos the dorsal lip does not overhang
the ventral, the coracoid grooves do not overlap nor in some
cases even reach the middle line, and the oblong articular facet
on the dorsal lip for articulation of the coracoid is only slightly
developed. In the Musophagi the dorsal surface of the base a
the spina externa affords articulation for the right coracoid ;
the Cuckoos this is never the case.
In the Musophagi the articular surfaces for the ribs are fairly
widely spaced, less so in Z'uracus ; in the Cuckoos these surfaces
are crowded together.
The coracoid in the Cuculi is relatively long and slender, being
nearly or quite as long as the sternum. From the ventral aspect,
one of the most conspicuous features is the large procoracoid.
This forms a large oblong shelf projecting inwards and downwards
from the shaft, at about the level of a line drawn across the shaft
from behind the glenoid cavity. The acrocoracoid is large, and
not twisted on the shaft so as to conceal the foramen interosseum.
The processus lateralis is well developed and the foramen swpra-
coracoideum is absent.
The processus lateralis basalis may be either broad and quad-
rate, with its antero-external angle produced forwards into a
spine, as in Cuculus, Rhopodytes, Hudynamys, Taccocoua, Rham-
phococcyx, Scythrops, and Guira ; or narrow and directed outwards
and backwards, as in Centropus, Cowa, and Dromococcyx. In
Taccocoua, and to a less marked extent in Centropus, the impres-
sion for the sterno-coracoideus muscle, on the dorsal aspect of
the coracoid, is bounded in front by a sharp vertically directed
spine.
The scapula is long and narrow, and differs somewhat in shape
in the different genera; but the variations are unimportant, and
not sufficiently large to justify description here. The acromion is
short and stout, save in Cowa reynaudi, wherein it is reduced in
width to form a somewhat cylindrical process.
1903.] OSTEOLOGY OF THE CUCULIFORMES. 279
The furcula has long, round, slender limbs, gently arched.
There is a moderately long and styliform hypocleideum in Geo-
coccyx, Coua, Rhamphococey yx, and Piaya, for example. In Croto-
phaga it is spatul: ite. In Centropus, Cuculus, Hudynamys, and
Cacomantis it appears to be of a degenerate spatulate type;
whilst in Seythrops the hypocleideum appears to be wanting.
The nature of the combination of the elements forming the
inner wall of the foramen triossewm—the acromion of the scapula,
the procoracoid, and the free end of the clavicle—may prove,
when exhaustively worked out, to have some slight value from a
systematic standpoint. I propose, however, here to offer only a
few remarks, selecting a few genera as examples.
In Cuculus, Scythrops, Cowa, the whole free end of the furcula,
passes up cephalad of the acromion of the scapula and separates the
latter from the procoracoid. In Guwira, Geococcyx, Rhamphococcyx,
Rhinococeya, Rhopodytes, the acromion turns forwards so as ne: rly
to join the procoracoid, and forms an opposing surface to the free
end of the clavicle, which accordingly turns forwards to terminate
ina point wedged in between the acromion and procoracoid. Cro-
tophaga vesembles Guira in this respect, but the free end of the
clavicle is much broader and the articulation with the procoracoid
more developed. In Centropws the acromion and acrocoracoid
meet and embrace, as in a wedge, the styliform free end of the
slender furcula. The procoracoid, which is large, does not come
in contact with the scapula.
No two of the genera, however, exactly agree, but the differences
between them are too slight to be described in words.
The coracoid of the Musophagi, though closely resembling that
of the Cuculi, is yet readily distinguishable therefrom by the fact
that the procoracoid turns downwards and outwards to fuse with
the acrocoracoid, thus encircling the foramen triosseum with a
continuous bar of bone. The procoracoid is large and passes
insensibly backwards into the coracoid shaft, and there is a
supracoracoid foramen. The processus lateralis is well developed,
and directed outwards and upwards into a point. In the region
where the pro- and aecrocoracoid fuse there is an elongated facet
for the articulation of the fureula. The procoracoid, however,
appears to take the greater part of the share in furnishing this
surface. On the dorsal or internal aspect of the coracoid there is
developed a special articular surface for the dorsal lip of the
coracoid groove of the sternum.
The furcula has relatively shorter, broader, and more laterally
compressed limbs than in the Cuculi. A hypocleideum is wanting.
The dorsal extremity bears a special projecting facet for articu-
lation with the procoracoid.
The scapula resembles that of the Cuculi, but is relatively
slightly broader and has a large acromial process.
The pro- and acrocoracoid meet and fuse one with the other, thus
excluding the furcula from participation in the formation of the
foramen triosseum.
280 MR. W. P. PYORAFT ON THE [Mar. 17,
vil. THE Petyic GIRDLE.
The pelvic girdle of the Cuculiformes, though recallmg in
certain features that of the Coraciiformes, is yet quite distinct
therefrom. Within the group it presents a comparatively wide
range of form.
Among the Cuculi, the least specialised pelves ave those of the
smaller forms, such as Cacomantis (text-fig. 45) and Chrysococcyx.
In Cacomantis the preacetabular region of the ilia is sepa-
vated by a low swollen ridge formed by the centra of the lumbar
vertebre. The dorsal border is nearly straight; the ventral,
external, border is deeply emarginate ; the anterior border trun-
cated and rounded, curving outwards to form a hook-shaped
antero-ventral angle. The postacetabular region of the ilium has
Dorsal aspect of the pelvis of Cacomantis merulinus, X 5, showing the widely
separated preacetabular ilia, and the broad dorsal plane of the postacetabular
region.
a broad dorsal plane which abuts against the long and slender
transverse processes of the vertebre. These last, by the way, are
separated by a row of intertransverse sacral foramina, one on
either side of the column. The dorsal plane of the postacetabular
ilium is not produced outwards so as to overhang the ilio-ischiadic
fissure, and its postero-external border is markedly depressed,
not forming an upturned crest as in forms to be presently
described. That portion of the ischium which, by its upturned
growth, converts the obturator fissure into a foramen is very
narrow ; below and behind the foramen the ischium is produced
1903. | OSTEOLOGY OF THE CUCULIFORMES. 281
backwards into an obliquely sloping bar. The pubis is long,
slender, and much bowed and closely bound to the posterior half
of the ventral border of the ischium, beyond which it extends for
some considerable distance. The obturator foramen is bounded
posteriorly by a broad bar of bone descending from the ischium
and fusing with the pubis. The pectineal process is reduced to a
vestigial condition.
In the form of the pelvis, Cucwus, Coccystes, Scythrops, differ
but little, save in size, from Cacomantis merulinus. In Scythrops,
however, the obturator foramen is not cut off posteriorly by the
descent of a bony plate from the ischium. In all save ¢ ‘acomantis
and Cuculus the intertransverse sacral foramina are more or less
completely filled up.
In Rhopodytes and Hudynamys the mesial border of the extreme
anterior end of the preacetabular ilium rises to the level of the
neural ridge as in Geococcyx (text-fig. 46, p. 282). In Taccocowa this
elevated region extends somewhat further backwards and forms
a quadrate plate, the free border of which is lightly applied to the
neural crest. In Crotophaga, Rhamphococcyx, Coua, and Piaya
actual fusion takes place between this plate and the synsacrum. In
Centropus and Dromocoecyx the relationship between the fused
region of the ilium and the synsacrum has become still further
modified, so that the extremity of the ilium has acquired a T-shaped
form, the dorsal limb of the cross-piece bridging a widely open
canalis ilio-lumbalis. A similar canal is of course formed in the
other cases where the ilia meet the neural crest of the synsacrum,
but it is inconspicuous. )
Further modifications in the form of the pelvis are the
enormous lateral expansion of the dorsal plane and the short-
ening of the pubis. In forms like Crotophaga, Coua, or Rham-
phococcyx, the early stages of the first-mentioned modification
may be studied. Passing through forms like Centropus and
Taccocoua, we reach a climax in Geococcyx (text-fig. 46, p. 282).
Herein the width across the dorsal plane equals the length of the
synsacrum. This outward extension of the postacetabular ium
has resulted in the formation of a huge ledge passing far beyond
the level of the antitrochanter, and finally, at its free edge,
turning abruptly downwards and then suddenly upwards, back-
wards, and inwards, like a pair of wings, so that the surface of the
dorsal plane acquires a peculiar saddle-shaped appearance. Seen
from the side (text-fig. 47, p. 282) or from below (text-fig. 48,
p. 282), this overhanging ledge forms an enormous penthouse
above and slightly behind the ilio-ischiadic foramen. A similar
modification obtains elsewhere only among certain Rails. It
appears to be correlated with a terrestrial mode of life, these forms
flying but little.
The shortening of the pubis is most marked in Geococeys,
Piaya, Centropus, and Rhinococeyx, and to a less extent in Coua
and Crotophaga.
A large pectineal process occurs in Geococcyx, Cowa, Centropus,
282 MR. W. P. PYCRAFT ON THE [ Mar. 17,
Rhamphococeyx, Rhinococcyx, and Piaya; it is small in Rhopodytes
and Zaccocowa, and vestigial or wanting in other genera.
The fovea lwmbalis is very small, and the fovea ischiadica and
pudendalis are confluent. There is a well-marked iliac recess.
Text-fig. 46. Text-fig. 48.
Text-fig. 46.—Dorsal aspect of the pelvis of Geococcyx mexicanus, showing the
narrow preacetabular ilia, extremely broad postacetabular dorsal plane, and
large pectineal process.
Text-fig. 47.—Side view of same, to show the broad ledge-like plate of the post-
acetabular ilium, the short pubis, and large pectineal process.
Text-fig. 48.— Ventral aspect of same, showing the great overhanging ledge of the
dorsal plane. All figures of natural size.
The pelvis of the Musophagi resembles that of the more
specialised Cuculi. Though relatively wide, its breadth is due
rather to the great length of the synsacral transverse processes
than to the dorsal plane of the postacetabular ilium. The pre-
acetabular region of the ilium differs conspicuously from that of
the Cuculi, inasmuch as in this group the dorsal aspect is so much
cut away as to le far below the level of the synsacral crest, save
where it rises at the extreme antero-dorsal angle. In the Muso-
1903. ] OSTEOLOGY OF THE CUCULIFORMES. 283
phagi, the preilium rises upwards and sweeps inwards in the form
of a broad vertical blade with a gently rounded dorsal border,
which ultimately meets and fuses with the synsacral crest, termi-
nating in a truncated anterior border cephalad of the synsacral
fusion. The canalis ilio-lumbalis is wide and spacious, The
dorsal plane of the postacetabular ilium is moderately wide, and
tapers posteriorly into a blunt, slightly upturned point. Its free
outer border is somewhat thickened in Z'uwracus to form a broad
lip with sharply defined upper and lower edges; the inferior of
these edges, immediately behind the ilio- ischiadic for: amen, 1S pro-
duced into a blunt spine. In Sehizorhis the free border of the
dorsal plane is sharply defined, so that the blunt point caudad of
the ilio-ischiadic foramen is apparent on the dorsal aspect of the
pelvis, instead of below this level. The ilio-ischiadic foramen is
relatively much smaller in Z’wracus than in Schizorhis. The ischium
expands caudad into an extremely broad plate, especially so in
Turacus. The free hinder border of this plate is slightly convex.
In Turacus the obturator foramen is cut off from the fissure of
that name by a descending plate of bone from the ischium. This
is not the case in Sehizor his, the foramen and the fissure being
confluent. The pubis is very long, attached for some distance
poster iorly to the ventral border or the ischium. The pectineal
process is large in both the genera in question.
The fovea lumbalis is larger than in the Cuculi, and the fovea
ischiadica is distinct from ‘the fovea pudendalis, the latter being
cut up into separate compartments by the synsacral ventri-later: al
processes. The iliac recess is fairly spacious.
vill. THE Pecrorat Limp,
The character of the wing is very uniform throughout the
group. The humerus only is pneumatic. Most nearly resem-
bling that of the Trogons in its general character, the limb of
the Cuculiformes may be distinguished by the greater size of the
pectoral crest of the humerus, which forms a linguiform or even
triangular plate, and by the prominent collum trochlee, the
strongly bowed curve of metacarpal TIT. ., and the absence of a
backwar dly projecting spur on the upper 3 of metacarpal IT.
In the humerus the coraco-humeral groove (sulcus transversum)
is wanting. The crista superior is cently arched, rising from the
base of the tuberculum externus ee ter minating about the upper
3 of the shaft. In some cases, e. g. Crotophaga, “the pectoral crest
may be more or less eed ‘instead of rounded. There is a
very small ectepicondylar “tubercle. The pneumatic foramen is
small; the incisura capitis sharply defined. The humerus of the
Musophagi may be distinguished from that of the Cuculi by the
fact that in the former the proximal border of the crista superior
is long, low, and concave, the distal border short and strongly
arched,
In the Cuculi there is considerable variation in the relative
284 MR. W. P. PYCRAFT ON THE [ Mar. 17,
lengths of the segments of the wing. Thus, in Scythrops, the
brachium and antebrachium are of equal length, whilst the
manus is only a very little less in length than either of these
segments. The brachium is longer than the antebrachium, and
the latter is in turn longer than the hand in Cowa, Rhamphococcyx,
Rhinococcyx, Hudynamys, Guira, and Crotophaga. 'The relative
disproportion between the length of the brachium and ante-
brachium is especially marked in Dromococcyx and Piaya. The
brachium is longer than the antebrachium, but the latter and the
manus are equal in length in Centropusand Taccocoua. In Cuculus
and Cacomantis the brachium and manus are subequal and
shorter than the antebrachium. In Coccystes the antebrachium
and manus are subequal and longer than the brachium.
The variation in the length of the wing among the Cuculi
becomes still more forcibly illustrated when compared with the
hind limb, the difference apparently varying with the power of
flight.
Tn Cuculus, Coccyzus, and Cacomantis, for example, the wing
is considerably longer than the leg. In Chrysococcyx the two
limbs are subequal. In Geococcyx, Crotophaga, and Couwa the
wing is much shorter. Taking the combined length of the femur
and tibio-tarsus of each species as a standard of measurement, it
will be found that in Crotophaga the wing exceeds this length
only by the distance from the middle of Ph. I. D. II. to the tip
of the wing. In Geococcyx the wing is slightly less than the
length of the standard. Coww has the shortest wing of all, it
being about + shorter than the combined length of femur and
tibio-tarsus.
In the Musophagi the brachium is somewhat longer than the
antebrachium, but the latter is considerably longer than the
manus. The ulna, as in the Cuculi, bears prominent bosses for
the attachment of remiges.
As in the Cuculi, the collwm trochlee is very conspicuous, giving
the distal end of the humerus an obliquely truncated form. The
coraco-humeral groove is wanting. The proximal border of the
crista superior is very long and concave, the distal border short
and convex. The ectepicondylar tubercle is very small.
The Ph. of D. III. in the Cuculi sends outwards from its post-
axial border a prominent spur, this is wanting in the Musophagi.
ix. THe Petyrc Lines.
The pelvic limb in the Cuculiformes has, in common with the
Psittaci and many Coraciiformes, a zygodactyle pes. The hypo-
tarsus is complex, and the tibio-tarsus has an ossified extensor
bridge. In all save Geococcyx the limb is non-pneumatic, and in
this genus only the femur is pneumatic.
The pelvic limb of the Cuculiformes has the 2nd and 3rd
trochlee nearly equal in length, but the latter is conspicuously
larger in size. The third trochlea is placed comparatively high
1903.] OSTEOLOGY OF THE CUCULIFORMES. 285
up the shaft, and has the glenoid articular surface conspicuously
laterally compressed and twisted, so as to cross the long axis of
the shaft transversely. This limb may be readily distinguished
from that of the Psittaci, inasmuch as in the latter the trochles
of D. IT. stands out at right angles to the tarso-metatarsal shaft,
whilst the trochlea for D. IV. has rotated so that its articular
surface is turned to look directly backwards. Thus, the outer
condyle comes to lie next the shaft, and, furthermore, is produced
into a hook-shaped process of considerable size.
Amongst the Coraciiformes, probably the pelvie limb of Zepto-
soma most nearly resembles that of the Cuculiformes, but the
conspicuously higher position on the shaft of the trochlea of D.TV.,
which obtains in this last group, is sufficient to prevent confusion.
The fibular ridge is confined to the upper end of the shaft, and
increases in depth from above downwards.
The length of the fibula may vary considerably even in different
species of the same genus. Usually it is greatly reduced, but in
Centropus toulou, for example, it extends considerably beyond the
middle of the tibial shaft, whilst in C. madagascariensis it falls
considerably short of this.
In Crotophaga, Kudynamys, and Rhopodytes it barely extends
beyond the level of the tibial fibular ridge.
The cnemial crests are, as a rule, feebly developed, indeed only in
Geococcyx do they attain to any respectable size. In this species
the entocnemial crest is more or less quadrate in form; its outer
border is convex, its inferior concave. The ectocnemial crest is
triangular in form, and terminates in a small pointed process
immediately in front of the head of the fibula.
The shaft of the tibia is long, slender, and cylindrical, and bowed
slightly forwards. In Geococcyx the posterior lateral borders of
the internal mesotarsal condyle are produced backwards and
upwards to form a rather prominent spur.
The popliteal fossa of the femur is obsolete.
The pelvic limb of the Musophagi is less specialised in some
respects than that of the Cuculi. This is especially marked in the
character of the tarso-metatarsus. As in the Cuckoos, the hypo-
tarsus is complex, and the tarso-metatarsal shaft grooved in front ;
but the arrangement of the distal trochlez is of a less specialised
character, and this fact is especially marked in Schizorhis, the foot
of which in this respect closely resembles that of Zeptosoma. The
foot of Schizorhis, however, may readily be distinguished from
that of Zeptosoma inasmuch as in the latter the trochlea for D. IV.
bears a very strongly-developed outer lip, which is produced
inwards towards Me.I. In Sehizorhis this trochlea is almost
indistinguishable from that of an ordinary eleutherodactyle foot.
In Turacus, however, the form of the outer trochlea nearly
resembles that of the Cuculi, but is less markedly compressed,
and looks backwards and slightly inwards, instead of being twisted
so as to cross the shaft transversely. Moreover, it is not raised
high up on the shaft as in the Cuckoos.
286 MR. W. P. PYCRAFT ON THE [ Mar. 17,
x. SUMMARY.
The isolated position of the Cuculiformes among the Coracio-
morphe is as evident from a study of the osteology of the group
as from other points of view. Their nearest allies, judged from
an osteological standpoint, would appear to be the Coraciide
(Coraciine and Leptosomatine) and Bueconide on the one hand,
and—more remotely—the Opisthocomi on the other. Their
relationship to the Psittaci, which is generally agreed upon,
would, | think, never be suspected from a comparison of the
skeletal framework. This fact is probably to be explained by the
great amount of specialisation which the Parrots have undergone.
Concerning the association of the Cuculiformes with the
Bucconide, a few words of explanation are necessary. Although
this connection has several times been made by the older orni-
thologists, Fiirbringer appears to be the only modern systematist
who, on anatomical grounds, takes a similar view. Gadow (8)
regards the Bueconide as close allies of the Galbulide, placing the
two in the same family—Galbulide. Beddard, on the other hand,
widely separates these two forms, and I think rightly so. He
places the Bucconidee between the Pici and the Rhamphastide,
but remarks that this is a family which “is at present little
known, and whose aftinities are therefore doubtful. It is only
provisionally that I place them in the present position.”
The claims to relationship, of the group now under consideration,
to the Opisthocomi are, on osteological grounds, not at first sight
very strong. <A careful study, however, of the skulls of Cuculi
and Musophagi, and a comparison with Opisthocomus, will show
points of agreement which suggest affinity rather than convergence.
The Musophagi most nearly approach Opisthocomus. The pelvis
and sternum of Opisthocomus are Cuculine.
That the Cuculide and Musophagide are very near allies there
can be no doubt. Indeed, the relationship between these two is
the only relationship about which we can speak with any real
certainty.
According to the British Museum Catalogue of Birds, vol. xix.,
the Cuculide (suborder Cuculi) embraces 47 genera and includes
202 species grouped under 6 subfamilies. This scheme is
the work of Capt. Shelley. Beddard (1) recognises only 3 sub-
families, and these, I think, will be found to meet all demands.
Further, it is quite open to question whether even these should
not be regarded as groups @, 6, ¢, rather than subfamilies.
Osteologically, the Cuckoos vary far more widely in respect of
the pelvis and sternum than in any other character. The great
feature in the skull is the uniform plan of the palate. Trusting
to this alone, one might divide the Cuckoos into two groups—
Cowa in the one, and all the rest of the Cuckoos in the other:
yet the difference in the two palates is trivial. The only other
characters offered the systematist to choose from are the form
of the lachrymal and temporal fossz and the shape of the beak,
1903.] OSTEOLOGY OF THE CUCULIFORMES. 287
but the latter at any rate is the direct outcome of adaptation.
Probably the remarkably specialised condition of the pelvis also
is adaptive.
In the accompanying table it will be found that I have arranged
the Cuckoos on a plan formed of a compromise between that of
Beddard, based on the pterylography and voice-muscles, and one
based on the osteology and voice-muscles. The two will be found
to disagree in many points; and it is on this account that
I have published the results of my comparison, since it will, I
hope, stimulate further research by directing renewed attention
to the anatomy of the discordant forms.
A. Sternum with single pair of notches ; pectineal process of pelvis vestigial.
Muscle-formula AXY+, except Hudynamys which has the formula
ABXY-+.
. : Cuculus.
a. Syrinx tracheo-bronchial.
Muscle-formula AXY+, except eee si
Eudynamys which has ABXY+. | 77, ae as
Old World. Sey th pane ;
b. Syrinx bronchial. i
Muscle-formula ABXY +. { hated aa
New World.
B. Sternum with single pair of notches; pectineal process of pelvis large.
e. Syrinx bronchial. Cent
Muscle-formula ABXY-+. { CREEL
Old World. Dromococeyx.
C. Sternum with two pairs of notches ; pectineal process large.
d. Syrinx bronchial.
Muscle-formula ABXY-+.
ING MW/@AIGE sacoscop snsoco sone ecocconss Cra ENE
@OLARWrorlde ene tec eco cei OOUde
( Rhinococeyx.
e. Syrinx tracheo-bronchial. FR aaa
Muscle-formula ABXY-+, except Baenowae
Z ; ies ypodytes.
Piaya which has AXY+. Wancinstonius:
Old World. Taccocoua.
Piaya. (New World.)
D. Sternum with two pairs of notches ; pectineal process vestigial.
f. Syrinx tracheo-bronchial. Che ,
Muscle-formula AXY-+. irysococeya.
Old World. Cacomantis.
Mr. Beddard has shown, in tabular form, the points wherein
the skulls of the Cuculi and Musophagi differ one from another.
These are not many, and I cannot but think that they are really
of slight importance. The most striking are the differences
in the size of the antorbital plate, the position of the maxillary
end of the quadrato-jugal bar relatively to the tomium, and the
relation of the mesial borders of the proximal ends of the palatines.
The desmognathism of the palate seems to me to be precisely
similar in both forms. If the skull of Z’wracus, for example, be
compared with that of Cowa, it will be found that the chief
difference lies in the more spongy nasal septum of the latter,
Proc. Zoot. Soc.—1903, Vou. I. No. XIX. 19
288 . MR. W. P. PYCRAFT ON THE © [Mar. 17,
which descends so as to lie between and force apart the maxillo-
palatine, causing the palate to be indirectly desmognathous. In
Turacus the nasal septum fuses with the dorsal borders of the
maxillo-palatine, and does not descend between them, but in this
case also the palate is indirectly desmognathous. In both Coua
and Juracus, but for the septum, the palate would be schizo-
gnathous. But the maxillo-palatine region in Schizorhis, for
example, is practically identical with that of normal Cucull.
Coua has the least specialised palate among the Cuculi; Z’uracus
among the Musophagi. In the more specialised skull of both
groups indirect desmognathism appears to have taken place by the
descent of the ventral border of the septum nasi between the
maxillo-palatine processes; resulting in the formation of a flat
palatal surface, or one with a slight median groove.
The very striking differences in the form of the pelvic girdle
stand in strong contrast with the uniformity found in the skull.
How considerable these differences in the girdle are may be seen
in the figures on pp. 277, 280, and 282.
Finally, I may remark that a great deal yet remains to be done
before our knowledge of the Osteology of this group is complete.
Of the nestling skeleton of the Musophagi we know nothing, and
only a very little concerning the early stages of the Cuculi.
Only about one-third of the total number of genera of Cuculi
are represented in the Museum Collection of skeletons. These,
however, fortunately, include all the more important forms.
xi, Key TO THE OsrEOLOGY OF THE CUCULIFORMES.
A. SKULL.
Palate desmognathous, and without conspicuous anterior palatal vacuity ; nostrils
holorhinal, imperforate ; basipterygoid processes wanting; vomer vestigial or
wanting; pterygoid free; lachrymal free; antorbital plate large; the postorbital
process never extending down to the quadrato-jugal bar.
A. Mesial border of palatines not meeting in middle line, leaving parasphenoidal
rostrum exposed; with a large os uncinatum .................. MusorPuHaaGi.
B. Mesial border of palatines meeting in middle line below and concealing para-
ONS AGC SRO SADT, “|. ooo sca coosem ons cedooncasonsedoassoosseeoodssese CWCKN
Key to the Genera of Cuculide.
a. Beak long and pointed. :
a’. Lachrymal with massive descending process; nostrils linear near the middle
of the tomium ; temporal fossz confined to lateral aspect of skull.
Geococcyx.
2’. Lachrymal very small ; temporal fossee meeting one another to form a broad
GAYS GREE, Gos ous coblseboc Sone aupdansercsnser bed concobosr cannounae | | MMOMROMACTR
b. Beak with a high median keel; lachrymal large ............ Crotophaga.
c. Beak swollen, with an arched culmen.
ce’. Palatines fused in median line posteriorly .................. ° Seythrops.
d’. Palatines free posteriorly.
a’. Nostrils linear immediately above toinium ; postorbital process of lachry-
TIENEN Eat ReR nhs net uoniice dob poseennidabe sa HeaBAUres Hocecroncaua a: Rhamphococcyx.
b’’, Nostrils oval immediately above base of tomium, and bounded above or
confluent with a conspicuous fenestra; postorbital process of lachrymal
Sina yl a eee cters eae else cine teitssenencmmntereacer ems Rhinococeyx.
9
1903. | OSTEOLOGY OF THE CUCULIFORMES. 289
d. Beak not exceeding the cranium in length, of medium size, and with a more or
less strongly arched culmen.
e’. Descending process of lachrymal vestigial or wanting ; temporal f oss
meeting to form a sagittal crest; nostrils more or less markedly reni form,
being formed like those of Rhinocoecyx by the confluence of the narial
aperture with a superior fenestra.
oe’. Descending process of lachrymal in form of delicate incurved st yle.
a’, Sagittal crest reduced to a mere line.................. Rhopodytes.
BUC a Sagittal Chest) DLOR ioc sesuenaceeaceaesesesstac-1 == Zanclostomus.
d'’. Descending process of lachrymal reduced to a stump.
Piaya.
é’. Descending process of lachrymal wanting; antorbitai plate slightly
Taccocoua.
inflated nok CORO E a OCH CIO NE ATE CRO CER CE ECC EERIE:
f’. Descending process of lachrymal moderately strong, bent outwards and
backwards upon the supraorbital portion; temporal fosse meeting to form
a broad sagittal crest.
f’’. Horizontal process of lachrymal longer than descending ; pterygoid with
dorsal keel; interorbital fenestra large ............... Guira.
g'. Horizontal process of lachrymal not exceeding descending process, and
having free orbital border sharply truncated ; interorbital fenestra small.
Budynamys.
temporal fossa meeting to form a
g’. Descending process of lachrymal slender ;
Centropus.
broad sagittal crest ; interorbital fenestra very large ...
h’. Temporal fossze confined to lateral aspect of skull.
hk’, Lachrymal moderately large and not distinctly divided into supraorbital
and descending processes; antorbital plate spongy ; interorbital fenestra
large ; maxillo-palatines not meeting in middle line, but palate edged by
MASA SOPLUMN eee ee eee cence earoe sere e nese pscacs Coua.
i”, Lachrymal moderately large, with slender descending process ; nostrils
ericular, just above base of tomium Secrets Coccystes.
i’, Lachrymal vestigial; mandible with broad flange at gape.
CU Memporall fossa) PLCSCMU e easawaesscccteraies sea sestr feleccia= Cuculus.
: d’”’, Temporal fosse obsolete .............s0200ccresrssereeeee Cacomantis.
B. VERTEBRZ’.
All the presynsacral vertebra are free and heterocclous ; neural arch of atlas
broad; hypapophyses blade-shaped, not extending beyond 2nd thoracic vertebra.
Axis vertebra not conspicuously shortened antero-posteriorly, and not having
conspicuously upturned hyperapophyses.
A. Lateral borders of thoracic vertebra terminating in a sharp backwardly directed
oe Mvsopuaal.
RIDE Ghsoc cocoon: 096 CBB ASBROSHSERED ONRCES a0OC cad HONEEOOGO CoE
B. Lateral borders of thoracic vertebre not produced into spikes. CUCULI.
CG. PrecroraL GIRDLE AND STERNUM.
Coracoid with a well-developed procoracoid; clefts of the posterior notches of
sternum never extending as far forwards as the level of the articulation of the ribs ;
furcula large.
A. Procoracoid fused with acrocoracoid; furcula with a well-marked facet for
articulation with the fused pro- and acrocoracoids ; anterior border of carina
sterni continued forwards to level of spina externa ............. MUSoPHAGI.
acrocoracoid; furcula without articular facet for
¢ continued forwards as far as
Cucutt,
B. Procoracoid not fused with
acrocoracoid ; anterior border of carina sterna no
free end of spima externa ...........0e cece eee seeeee cee eee ne eee eee eee
1 The presynsacral vertebr of the Cuculiformes are scarcely to be distinguished
from those of the Coraciine and Leptosomine.
Wg
AO) MR. W. P. PYCRAFT ON THE [ Mar. 17,
Key to the Genera of Cuculidee.
a. Sternal plate with spina communis.
ae spe border with two pairs of notches.
’. Hypocleideum styliform, free end only overlapped by carina sterni.
Rhamphococcye.
6”. Hypocleideum styliform, overlapped by carina sterni. Hae Ts
ce’. Hypocleideum oval, overlapped by carina sterni ...... ee te
b’. Posterior border with single pair of notches ............... Coccystes.
6. Sternal plate with spina externa only.
ce’. Posterior border of sternal plate with two pairs of notches. :
OD ’, Hypocleideum underlapped by carina sterni ............ Zanclostomus.
’, Hypocleideum with free end only underlapped by carina. Geococeyw.
ie “Posten ior border of sternum with a single pair of notches.
f’”. Hypocleideum obsolete ; procoracoid small, affording articulation for
scapula. ;
a”, itt cula large and strongly arched ..................... Seythrops.
wr ”” Burcula slender and slightly arched. —
at, Posterior lateral process broad ...................... Hudynamys.
64. Posterior lateral process narrow .... 2 Guia.
g’. Hypocleideum obsolete ; procoracoid large, not ‘affording articulation for
scapula... WREST ONC Cntropus:
e. Sternal plate with | spina ‘interna ‘only. srodursvne avadagaooenasdased « /MKAROCONIE
tos
@. Sternal plate without spimeS <-: 2... s-2-2--29-2+ ee. eee een eens ee { poe nae
e. ae nal plate with spina externa and interna distinct.
. Posterior border of sternum with a single pair of notches.
Cuculus.
i’, Posterior border of sternum with two pairs ofnotches. Chrysococcys.
D. Prtyic GIRDLE.
A. Preacetabular ilium with strongly arched border meeting in the median line above
the syusacral spines to form a large canalis ilio-lumbalis... Musopuact.
B. Preacetabular ilia with dorsal border never meeting in median line above syn-
SAGCKALIS DING eericee me ee CO CeCe iste CO CoE Eee ee sold OnU CELTS
Key to the Genera of Cuculide.
a. Presynsacral neural arches broad and without neural spines; ilia widely
separated. : Cacomantis
. With large intertransverse sacral foramina.................. ; Cini ‘
b’. me intertransverse sacral foramina. :
. Length not exceeding 1 WHC? soossoosapadccasonenqasaccsace —CLURIROGRRETER.
Bi Length not less than 2} inches... vee. Seythrops.
b. Presy nsacral neural arches Theenibns a high ‘median ridge ; no intertransverse
, sacr al foramina.
ee process vestigial.
. Antero-dorsal angles of pre-ilium not meeting in mid-dorsal line; pubis.
very long....... mes Eudynamys.
’. Antero-dorsal angles of pre- -ilinm meeting | in middle Geer
line; pubis short ep Ms : a Guir oe Moke
d’. Pectineal process large.
e’’. Ischio-pubic fissure slit- shaped and very narrow.
a’’’, Antero-dorsal angles of pre-ilium produced into Gorn
narrow bars meeting WA, TANEOUS IOUT \s.cncossncoace0 000 d Dronscoeae
b’’”. Antero-dorsal angle of pre-ilia produced inwards to form a quadrate
plate embracing neural spines; postacetabular ilia produced into an
enormous saddle-shaped WHEW soa ccsecnocosssccsecasovsne CGHANADBOI AD
Sf’. Ischio-pubic fissure extremely reduced.
e’’’, Antero-dorsal angle of pr e-ilia produced into a quadrate plate with con-
cave superior border just reaching neural spine ..._ Taccocoua.
d’’’, Antero-dorsal angle of ilia pr oduced into rounded Roe
angles, not reaching the neural spines te :
’ Tschio-pubic fissure very wide, closed ae Rhamphococcyx.
1903.] OSTEOLOGY OF THE CUCULIFORMES. . 29
REFERENCES.
(1) Bepparp, F. E.—Classification of Birds. 1898.
(2) Firsrixcer, M.—Vergleich, Anat. (. Brustschulterapparates.
Theil v.: Vogel. 1902.
(3) Gapow, H.—Bronn’s Thier-Reich. Bd. vi. Vigel. Syst. Theil.
1893.
(4) Parker, W. K.—Trans. Linn. Soe., 2nd ser. Zool. vol. i., 1879.
xii; EXPLANATION OF PLATE XXII.
Fig. 1. Ventral aspect of the skull of Coua veynaudi, showing the most primitive
form of the indirectly desmognathus skull of the Cuculide. Herein the
maxillo-palatines do not meet in the middle line, but the palate is bridged
by the swollen nasal septum. Basipterygoid processes are wanting. There
is no vomer.
Fig. 2. Ventral aspect of the skull of Twracus buffoni, showing the indirectly desmo-
gnathous type of palate of the Musophagide. The maxillo-palatines do not
meet in the middle line, but the palate is bridged by the fusion of the ventral
border of the nasal septum with the dorsal aspect of the maxillo-palatines.
The vomer is wanting, but present in a vestigial condition in Schizorhis.
Note the appearance of the parasphenoid between the palatines.
3. Dorsal aspect of the skull of Coua reynaudi, showing the absence of a nasal
hinge, and the general character of the root of the skull.
Fig. 4. Dorsal aspect of the skull of Turacus leucotis, showing the nasal hinge and
the general conformation of the root of the skull.
. Dorsal aspect of the skull of Sawrothera vetula, showing the long upper jaw
and the large temporal fossz.
6. Lateral aspect of the skull of Schizorhis zonura, showing the form of the
narial apertures, the os weeinatum and the general form of the skull.
Fig. 7. Lateral aspect of the skull of Rhinococeya calorhynchus, to show the form
8
9
=
Ig
ou
of the narial aperture and the general form of the cranium.
. Lateral aspect of the skull of a nestling Centropus, showing the arrangement
of the bones. Outer view.
. Lateral aspect of the inner view of the same skull, showing the separate
arrangement of the bones.
Fig. 10. The distal end of the pterygoid of the nestling Centropus, to show the
hemipterygoid.
Explanation of Letters.
als.=alisphenoid. n.h.=nasal hinge.
a.p.=antorbital plate. op.=opisthotic.
a.p.v.=anterior palatine vacuity. 0.u.=0s8 uncinatum.
}.oc.=basioccipital. p-=parietal.
b.s.=basisphenoid. p.a.=palatine.
e.=occipital condyle. par. =parasphenoidal rostrum.
ep.o.=epiotic. p.0.p.=postorbital process.
ex.0.=exoccipital. pro.=prootic.
-f.=tloccular fossa. pt.=pterygoid.
v.=tfrontal. g-=quadrate.
h.pt.=hemipterygoid. s.c.=sagittal crest.
.=lachrymal. s.0.=supraoccipital.
me.=imeatus internus. sq.=squamosal.
mes.=mesethmoid. t.f.=temporal fossa.
na.=nasal.
292 MR. H. SCHERREN ON [ Apr. 21,
April 21, 1903.
Dr. Henry Woopwarp, F.R.S., Vice-President,
in the Chair.
The Secretary read the following report on the additions made
to the Society’s Menagerie in March 1903 :-—
The registered additions to the Society’s Menagerie during the
month of March were 67 in number. Of these 14 were acquired
by presentation, 43 were received on deposit, and 10 in exchange.
The total number of departures during the same period, by death
and removals, was 89.
The following papers were read :-—
1. Linnzeus and Hunter on Feather-Tracts.
By Henry Scuzrren, F.Z.S.
[Received March 12, 1903. }
(Text-figure 49.)
The credit of using the feather-tracts of birds as a means of
classification belongs undoubtedly to Nitzsch, whose results, edited
after his death, by Burmeister, were published at Halle in 1840
under the title ‘ Pterylographie’. An English edition, translated
by W.S. Dallas and edited by Dr. Sclater, was brought out by
the Ray Society in 1867. Pterylosis, or the distribution of these
feather-tracts, is, according to Prof. Newton, “of prime taxonomic
importance in Ornithology, though more in the investigation of
small than of large groups.” This also seems to have been the
opinion of Nitzsch himself, who, however, was not aware that
anything at all had been done even in noting the existence of
such tracts and of the featherless spaces which he called apteria.
In his Introduction he says :—
T may, therefore, flatter myself with the hope of awakening the interest
of naturalists by the announcement of my new results, and, by the enumera-
tion and detailed description of the feathered regions of the bodies of birds
to which I give the name of feather-tracts (pteryle, Federnfluren), of
proving that these, new and surprising as they may appear to many on the
first glance at my figures, really furnish equally significant and important
characters for the certain and natural discrimination of the families of
birds.
Professor Newton (‘ Dict. Birds,’ Introd. p. 63, note 1) says that
the only men before Nitzsch’s time who seem to have noticed
feather-tracts were the great John Hunter and the accurate
1903.] FEATHER-TRACTS IN BIRDS. 293
Macartney. The observations of the latter were published in 1819
(Rees’s Cyclopedia, article Feathers) :—
Although the common feathers cover the whole body, they do not
grow from every part of the skin; they are thickest upon the shoulders and
loins, along the underpart of the neck and breast, and do not exist upon
the lateral lines of the neck or breast, or about the umbilicus. This
arrangement, and their being directed downwards and backwards, allows
them to cover the body more neatly, and to remain unruffled during the
motions of the bird.
The observations of Hunter appeared first in Owen’s ‘ Catalogue
of the College of Surgeons’ (vol. iii. pt. ii. p. 311), dated 1836 :—
Although the feathers of birds appear to be an entire and uniform
covering, they do not arise equally from every part of the body, but only
from such parts of the skin as are least liable to be attected by the motion
of the contiguous parts, such as the motion of the limbs.
Hunter, however, seems to have done something more, and to
have discriminated the feather-tracts, for, on the page quoted
above, we read :—
To these groups or thickets of feathers I shall give particular names,
taken from their situation.
It is, of course, only a coincidence that Hunter used the terms
“thickets of feathers,” and that Nitzsch, in his Essay, ‘ Pterylo-
graphie Avium, Pars prior, 1833, chose a very similar name,
pteryle, which Prof. Newton has translated “ feather-forests.” It
may perhaps be of service to record the fact.
If the date of Hunter’s observations be taken at about 1785, in
which year he built his Museum, an earlier notice of the feather-
tracts exists by at least twenty years. This occurs in the treatise
by Linnzus, ‘ Fundamenta Ornithologica,’ presented at Upsala by
A. P. Beckmann, May 4, 1765 (Ameenitates Acad. vii.). Premising
that the feathers are arranged in the form of a quincunx, the
author proceeds :—_
Nuda vero cutis est (h. e. pennarum gwincwnce non perforata aut tecta)
utrinque ad colli latera, a capite interscapulium versus, et ab axillis per
latera pectoris ad inguina usque, atque per femora postica, remotis integu-
mentis, videnda.
This is as clear as language can make it. If, however, any
doubt could exist, this would be at once removed by reference to
the plate (see text-fig. 49, p. 294), where, in a schematic bird, the
contour-feathers are arranged in quincunces, and there the bare
space running from the neck to the femur is connected with that on
the opposite side by the naked patch in the interscapular region.
The naked spaces, as figured, would, if looked at from above, form
a quincunx ; and the text recalls a passage in the ‘Garden of
Cyrus,’ where Browne writes of the quincuncial arrangement in
the “feathery plantation about birds.” Although the primary
reference by Browne is to the papille in the feather-tracts, I suggest
(and in this I am glad to have the support of Mr. Southwell)
294 MR. OLDFIELD THOMAS ON [Apr. 21,
that the passage alluded to shows that Sir Thomas Browne was fully
aware of the distribution of the feathers in well-defined regions,
and that these differed in position and extent in different birds.
Moreover, I believe the passage in the ‘ Ameenitates’ shows
that Linneus knew of Browne’s work. Besides the reference to
the quincunx already quoted, there is yet another—“ Penne que
Text-fig. 49.
ey wey \
wh Mo
Topographical diagram ae feather-tracts and bare spaces in schematic bird.
(Reduced from ‘ Amcenitates Academice,’ vii. tab. i. fig. A.)
(preter alas et caudam) reliquum corpus servant, In guincuncem
digestee sunt”; and his phrase ‘‘ mirando nunquam satis artificio,
quod in Colymbi presertim corio, alutariorum arte preeparato .
est conspicuum,” 1s strangely reminiscent of the note of the famous
Norfolk naturalist : ‘“‘ Elegantly conspicuous on the inside of the
stripped skins of the dive-fowl.”
On some Mammals collected by Capt. H. N. Dunn,
R.A.M.C., in the Soudan. By O.pFietp THomas,
F.R.S.
[Received March 11, 1903.]
The National Museum owes to Capt. H. N. Dunn a collection
of Mammals, mostly small, made by him in the Egyptian Soudan
in the course of 1902. Among these no less than five prove to
need new names, thus showing how much still remains to be done
1903. ] MAMMALS FROM THE SOUDAN. 295
in studying the mammals of this interesting region; and it is
hoped that other officers will follow Capt. Dunn’s example.
The localities at which collections were made are mostly rather
to the southward of Khartoum, the majority of them being in
Kordofan, a province hitherto almost entirely unrepresented in
our collections.
Besides the earlier writings of Sundevall ‘ and Heuglin * on the
Mammals of this region, reference may be made to a paper by
Mr. de Winton on a collection from Shendy*, to the north of
Khartoum, to the same author’s work* in conjunction with the late
Dr. J. Anderson on the Mammals of Egypt generally, and to a
small paper’ of my own on Mr. Hawker’s collection from the
Fashoda region of the Nile.
1. RovsErrus stRAMINEUS Geoff.
4. $. Khartoum. 15 August, 1902.
2. HIPPOSIDERUS CAFFER Sund.
6
5. o. El Obeid, Kordofan. 22 October, 1902.
H. caffer had already been recorded from this region by
Temminck °,
3. MEGADERMA FRONS Geoff.
9. Wad Medina, Blue Nile. 21 September, 1902.
4, TAPHOZOUS PERFORATUS Geoff.
1,2,38. 6992. Khartoum. 28 June, 1902.
5. CANIS ANTHUS SOUDANICUS, subsp. n.
60. @. El Obeid. 2 October, 1902. (B.M. No. 3.2.8.8.)
Type.
119. Habessa Wells, W. Kordofan. 7 December, 1902.
The eastern representative of the Senegal C. anthus F. Cuv.
Closely allied to the typical form, but paler and with markedly
smaller teeth.
General characters as in true C. anthus. Colour rather paler,
a clear sandy buff. Hairs of back and of tail broadly tipped with
black. Skull, as may be gathered from the measurements below,
smaller and more delicate.
Dimensions of the type :—
sate and body 650 mm.; tail 230; hind foot (s. u.) 137; ear
Skull: basal length 136; zygomatic breadth 78; length of
1 “Om Professor J. Hedenborgs insamlingar af Diggdjur i Nordostra Africa,”
K. Vet.-Ak. Handl. 1842, p. 189.
2 Reise N.O.-Afrika, ii., 1877.
3 Nov. Zool. viii. p. 397 (1901).
4 Anderson & de Winton, ‘Mamm. Egypt,’ 1902.
5 Ann. Mag. N. H. (7) viil. p. 273 (1901).
6 Fide Anderson, ‘Mamm. Egypt,’ p. 102.
296 MR. OLDFIELD THOMAS ON [ Apr. 21,
nasals (diagonally) 49; interorbital breadth 26; breadth across
postorbital processes 37; breadth of brain-case 49°5; palate,
length 72°5; breadth between outer corners of p.* 44.
Teeth: length of p.* 9-6, of p.* (on outer edge) 14:2, of m.* 11,
of m.1 and m.”* combined 16°8; breadth of m.* 14, of m. 10.
(Below), length p., 8°7, of p., 10, of m., 17°3, m., 81.
Hab. and type as given above.
This is the Jackal figured by Cretzschmar’* from Riippell’s
specimen as C. anthus, but is clearly at least subspecifically
different from that animal. Mr. de Winton has recently shown ?
that none of the earlier names of Hemprich and Ehrenberg or
other authors apply to this form, and I therefore venture to bestow
one on it.
6. VULPES VULPES ZGYPTIACA Sonn.
40. 9. Khartoum. 4 September, 1902.
7. VULPES PALLIDA Cretzschm.
8. ¢ (young). Wad Medina, Blue Nile. 18 September, 1902.
8. IcroNyX FRENATA Sund.
118. ¢. Gebel Haraza, W. Kordofan. 6 December, 1902.
The rediscovery of this species is of interest, as there has always
been some doubt whether it was or was not the same as the more
northern J. lybica. It proves to be readily distinguishable by its
smaller size, by certain differences in its body pattern, and,
especially, by the absence of the black ends to the caudal hairs.
9. DipopDILLUs stiGMoNYyx Heugl.
5. 2. Khartoum. 19 August, 1902.
A topotype of the species. As already noted °, the specimen in
the Stuttgart Museum marked Meriones stigmonyx does not agree
with Heuglin’s description of this animal, and is more like that of
his If. dongolanus.
It is to be observed that the present Gerbille and the next one
are so extremely alike, that it is almost impossible to distinguish
them apart except by an examination of the soles and skulls.
There is, however, a darker median area (“‘ Scheitel und Riicken-
mitte satter gefirbt”) in the Dipodillus not present in the
Gerbillws, and this confirms my previous allocation of the name,
which in any case having once been made should be adhered to.
10. GERBILLUS AGAG, sp. n.
96. g. Agageh Wells, W. Kordofan. 17 November, 1902.
(B.M. No. 3.2.8.11.) Type.
A small species of true hairy-footed Gerbillus, with compara-
tively short tail.
1 Atl. Riipp. Reise Mamm. pl. 17 (1826).
2 Anderson & de Winton, Mamm. Egypt, p. 218 (1902).
3 Ann. Mag. N. H. (7) vii. p. 276 (1901).
1903. | MAMMALS FROM THE SOUDAN. 297
Size small, and feet short. General colow above soft sandy
buff, slightly lined on the back with the dark tips to the hairs, but
without any marked darker dorsalarea. Along the back the bases
of the hairs are plumbeous, but laterally, still within the sandy area,
the hairsare broadly ringed with white subterminally, though this
colour does not show on the surface. Under surface pure sharply
defined white as usual. Cheeks, a prominent patch above and
behind each eye, and another behind the ear white. Whole of
fore limb white, hind limb with a sandy line down its outer side,
the inner side and whole of feet snowy white; palms and soles
thickly hairy. Tail short for this group, pale sandy above,
darkening towards the pencilled end; white below.
Skull unfortunately broken in the single specimen. Molars
markedly smaller and lighter than in the common Soudanese
G. pygargus.
Dimensions of the type :—
Head and body 87 mm.; tail 100; hind foot (s. u.) 24; ear 11.
Length of upper molar series 3°7.
Hab. and type as given above.
This little Gerbille is distinguished from its neighbour
G. pygargus by its smaller size and shorter tail. Its close resem-
blance to Dipodillus stigmonyx has already been noted.
11. ARVICANTHIS DUNNI, sp. n.
103. ¢. Kaga Hills, W. Kordofan (about 120 miles W. of El
Obeid). 20 November, 1902.
“ Dug out of reddish sandy cultivation soil, from among the
natives’ crops of dukhan.”—H. 1. D.
A many-striped species of the A. barbarus group; allied to
A, zebra, but smaller, paler, and with the ight and dark stripes less
contrasted.
Size small, the smallest of the group. General pale ground-
colour buff, the lateral darker stripes brown instead of black.
Head coarsely grizzled buffy and brown. Central dorsal stripe
beginning on the crown, very narrow, blackish, but not so deep a
black as in A. zebra; outside this there are on each side five
uninterrupted buffy stripes, separated from each other by broad
brown bands, each of which is divided down the centre into two
by an interrupted band of light, an arrangement essentially as in the
other species. The light spaces are throughout clear buff, and the
dark lines brown, a clear buffy line passing along below the outer-
most dark line and edging the pure white of the belly. In 4. zebra
the outer lines at least are white, only those near the spine being
buffy. Eye-ring buffy. Ears dull ochraceous, without darker
marking. Arms and legs pale buffy, becoming white on the
fingers and toes. ‘Tail well-haired ; dull ochraceous above, with a
narrow and inconspicuous mesial line of black ; whitish below.
Skull conspicuously smaller than in 4. zebra and the other
species of the group, with rather larger bull ; incisors narrower,
but molars rather larger in proportion.
298 MR. OLDFIELD THOMAS ON [Apr. 21,
Dimensions of the type :—.
Head and body 90 mm.; tail 70; hind foot (s. u.) 23; ear 14.
Skull—ereatest length 28 ; basilar length 22 ; zygomatic breadth
13; nasals 10x 3:4; interorbital breadth 4:4; breadth of brain-
case 12; palate, length 12-2; diastema 6°7 ; palatal foramina 5°6 ;
length of upper molar series 4°9.
Type. Old male. B.M. No. 3.2.8.15. One specimen only.
A Fashoda striped rat presented by Mr. R. M. Hawker in
1901 being identified as Arvicanthis zebra, the present species
may be readily distinguished from it by its much smaller size
and more buffy coloration.
Capt. Dunn tells me that this rat was very common in the
cultivated fields of the natives, burrowing in their crops of dukhan.
It is a very handsome and distinct species, and I have much
pleasure in connecting his name with it.
12. AcCoMYS WITHERBYI de Wint.
73. 3. Katul Hills. 30 October.
13, LEGGADA TENELLA, sp. 0.
7. Old 2. Roseres, Blue Nile. 14 September, 1902. (B. M.
IN@s Bee) | Wgyae.
‘Found in burrows in cornfield ; had 7 young in the womb.”—
IND.
A very small species of the Z. minutoides group.
General colour of cheeks and sides a clear sandy or ochraceous
buff, with a distinct darker median dorsal area, commencing as a
narrow well-defined line on the nose, broadening on the crown,
and extending, though less sharply defined, all down the back,
and dying away on the rump. Under surface pure sharply
defined white. No lighter markings round eyes. Ears small,
evenly rounded, grey, their edges faintly whiter; a large and
prominent white patch behind and below their posterior bases.
Fore limbs wholly white; hind limbs with a narrow line of the
body-colour continued down on the hinder side to the heel, other-
wise white. Tail about as long as the body without the head,
brown above, inconspicuously lighter below.
Skull smaller and narrower than in the Cape Z. minutordes,
with square and well-defined supraorbital edges. Palatal foramina
ending level with the anterior fourth of m.’. Posterior palate
elongated, its hinder edge about equidistant from the last molars
and the front of the bulle.
Dimensions of the type :—
Head and body 50 mm.; tail 35; hind foot 11:5; ear 9.
Skull: greatest length 17:2; basilar length 14; zygomatic
breadth 9; nasals, length 6°3 ; interorbital breadth 3; brain-case, -
breadth 7:5; palate, length 9:3; palatal foramina 4-0; diastema
5:0; length of upper molar series 3:0.
Hab. and type as given above.
1903. | MAMMALS FROM THE SOUDAN. 299
This beautiful little species is readily characterised by its strong
sandy colour, the marked dorsal darkening, and elongated palate.
14, JACULUS GORDONI, sp. 2.
85. 9. Gebel Agageh, W. Kordofan. 12 November, 1902.
104. 105. oo. Kaga Hills, W. Kordofan. 20 November,
1902.
106, g¢. Gebel um Durragh, W. Kordofan. 25 November,
1902.
Allied to J. jaculus Linn., but larger and differently coloured,
and with longer ears.
Size rather greater and build stouter than in J. jaculus.
General colour above, as compared to the yellowish “ buff” of
J. jaculus, darker, and nearly approaching to “ vinaceous buff ”
of Ridgway. Laterally, the dark colour seems to pass rather
sooner into the pure white of the under surface. White markings
more extended than in J. jaculus, the cheek, supraorbital, and
postauricular white patches all large. White hip-stripe large,
weakened in colour by a faint buffy or brownish sprinkling. Fore
limbs wholly white. Hinder aspect of thighs like back. Fine hairs
of feet silvery white, the terminal half of the long digital haus
sandy. Tail of the usual pattern, its basal portion isabelline buffy
above; black subterminal band rather over an inch in length;
white tip 3-2 in.; longest hairs at end of tail 16-17 mm. in
length.
Skull shaped quite as in J. jaculus, but larger and heavier
throughout.
Dimensions of the type, measured in the flesh :—
Head and body 120 mm. ; tail 200 ; hind foot (s.u.) 63; ear 25.
Skull: greatest length in middle line 34; basilar length 2775;
zygomatic breadth 24; tympanic breadth 2477; length of nasals
on outer edge 12°5; interorbital breadth 12°8; interparietal
5-7 x87; palate, length 17°3; palatal foramina 47; diastema
9:6 ; length of upper molar series 5:1.
Hab. (of type). Kaga Hills, W. Kordofan. Also oceurring at
Omdurman.
Type. Old male. B.M. No. 3.2.8.16. Oviginal number 104.
This Jerboa, which I have named in memory of the famous
General Gordon, differs decidedly from the ordinary Egyptian
J. jaculus by its larger size and different colour. The Museum
had previously received a specimen of it from Omdurman,
collected by Mr. W. L. 8. Loat, but as that was young and without
skull, it could not be described.
The only other species which need be referred to is Dipus
microtis Reichenow ' from “Samar, in Nord-ostafrica.” That was
founded on a young specimen, but the description of its teeth
shows that it was sufticiently adult not to be the young of the
present form, than which it is very much smaller (‘ Lauflinge
1 Zool. Anz, x. p. 369 (1887).
300 ON MAMMALS FROM THE SOUDAN. [ Apr. 21,
35 mm.”), with shorter ears, and with the remarkable character
(if not due to accident) of having no white at the end of its tail.
15. Lepus aruroricus H. & EK.
6. g. Shendy. 1 September, 1902.
92.93.94.97. 65622. Agageh, Kordofan. November 1902.
39. S. Wad Medina, Blue Nile.
16. Procavia RuFIcErS H. & EH.
77. 6. Kaga Hills, Kordofan. 2 November, 1902.
78.79.80.83.88. Agageh Hills. November.
17. ORYX ALGAZEL DAMMAH Cretzschin.
¢. Kordofan.
Practically a topotype of the subspecies.
When the ‘Book of Antelopes’ was published, owing to the
predilection of the senior author for ‘‘ well-established” names,
the term lewcoryx was used for the species to which Lichtenstein
erroneously transferred that name, while the true leucoryx of the
Persian Gulf was called by Gray’s name beatriz. ‘To put matters
more in accordance with modern ideas on nomenclature, the
Scimitar Oryx should bear the name algazel Oken; but as that
name was founded on Cuvier’s figure of a Senegal specimen, and
it is practically certain that the Eastern Soudanese form is at
least. subspecifically distinct from the Western, a second name is
required, and this we find in the “ Antilope dammah” of Riippell
and Cretzschmar. A. dammah was erroneously identified by
Riippell, and following him by Sclater and myself, with the Beisa,
but by its locality (‘die grossen Steppen von Haraza ”) is clearly
shown to be the Scimitar Oryx. This is fortunate, as the name
dammah, being earlier than beisa, would have had to be used in
the latter’s place had Riippell’s identification been correct, but
now the Beisa is left with its familiar name unaltered.
Briefly put, the nomenclature 1s as follows :—
Scimitar Oryx.
ORYX ALGAZEL Oken.
O. leucoryax of authors generally, not of Pallas.
Western form :—
Cemas algazel Oken, Lehrb. Nat. ii. pt. 1. p. 741 (1816);
ex L’Algazelle, F. Cuv. H. N. Mamm. 1. pl. 376 (1819).
(Senegal. )
Eastern form :—
Antilope dammah Cretzschm. Atl. Riipp. Reise, Mamm.
p- 22 (footnote), 1826. (Haraza, Kordofan.)
White Oryx.
Oryx LEUCORYX Pall.
Antilope lewcoryx Pall. Spice. Zool. xii. p. 17 (1777).
Oryx beatriz Gray, P. Z.8. 1857, p. 157, and of authors
generally.
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POLYCLADS FROM THE STRAITS OF MALACCA
1903.] ON THE POLYCLADS OF THE ‘‘ SKEAT EXPEDITION.” 301
18. GAZELLA RUFICOLLIS H. & E.
61. ¢. Gebel Tueis, 90 miles E. of Omdurman. 9 October,
1902.
This fine Gazelle is a most acceptable addition to the National
Collection, in which the species is still badly represented.
19, GAZELLA RUFIFRONS Gray.
3$ 2. Agageh Wells, Kordofan. November 1902.
Capt. Dunn tells me that this Gazelle does not range beyond
about 50 miles to the northward of El Obeid.
3. On a Collection of Turbellaria Polycladida from the Straits
of Malacca. (Skeat Expedition, 1899-1900.) By F.F.
Laipiaw, B.A. Cantab., Assistant Lecturer and Demon-
strator in the Owens College.!
[ Received March 19, 1903. ]
> (Plate XXIII. & Text-figures 50-56.)
The collection described below was made entirely by Mr. Evans
on the shores of a small islet called Pulau Bidan, a few miles north
of Penang. It proves a most interesting one, and includes, so far
as I can discover, only one previously known species, Thysanozoon
auropunctatum Coll.
As in deseribing Mr. Gardiner’s collection from the Maldives
[4], I have to note the scarcity or absence of the Euryleptide,
represented doubtfully only by a much damaged fragment in
Mr. Evans’s series,
The only previous record of species from the shores of the
Malay Peninsula that I have been able to discover was made by
Collingwood, who described and figured the following species from
Singapore :—
Thysanozoon allmani. Hurylepia kelaartii.
Proceros hancockianus. | Elasmodes obtusum.
ss buskit. | Leptoplana aurantiaca.
Sphyngiceps lacteus.
Grouping these specimens with those of the Skeat Collection and
adding Pseudoceros bedfordii from Singapore, kindly given me by
Mr. Lanchester, we get the following list classified according to
Lang’s system [6] :—
Acotylea.
PLANOCERIDA.
* Planocera sp.
1.* WNotoplana evansit, sp. n.
1 Communicated by Dr. S. F. Harmer, F.Z.S.
2 For explanation of the Plate, see p. 318.
302 MR. F. F, LAIDLAW ON THE [ Apr. 21,
Acotylea (con.).
LEPTOPLANIDE.
2.* Semonia penangensis, sp. Nn.
3.* Leptoplana malayana, sp. n.
as obtuswm (Coll.).
CRYPTOCELIDIDA.
5.* Bergendalia anomala.
LATOCESTIDA.
6.* Latocestus argus.
Cotylea.
PSEUDOCERID&.
7. Thysanozoon allman.
Cee a auropunctatum.
9. Pseudoceros hancockianus (Coll.).
10. a6 buskit (Coll.).
Wil. - kelaartvi (Coll.).
1A. i bedfordit, sp. n.
a" ws collingwoodi, sp. n.
14.* Pseudoceros ? rubellus, sp. n.
DrpPostHIDz.
15.* Asthenoceros woodworthi, sp. n.
PROSTHIOSTOMID&.
16.* Prosthiostomum pallidum, sp. v.
if 3 aurantiacum (Coll.).
Species marked with an asterisk were collected by Mr. Evans.
In this communication I have not ventured to give an account
of the anatomy of any of the species, contained in the collection,
which can be regarded as being at all complete. Such an account
would have expanded the paper beyond limits reasonable in a
systematic description. I have attempted only to give such a
diagnosis of each species as shall render its future identification a
matter of tolerable certainty, and to call attention to any of the
more striking features which presented themselves.
Family PLANOCERIDA.
PuaNnoceRA sp. (Plate XXIII. fig. 1.)
A fragment of tissue of which I made microscopic sections
proves to belong to a species of this genus, but being only a
fragment I will not attempt to describe or name it, though it is
obviously new. The section passes through the penis and prostate,
and the spines lining the lumen of the former are of such a remark-
able character that I venture to figure one of them, in the hope
that a perfect example may be obtained ere long. The length of
each spine is roughly ‘08 mm., its breadth at the base ‘02 mm.
NoroPLana, gen. nov.
NovoPLANA EVANSII, sp.n. (Plate XXIII. fig. 2.)
A number of specimens belonging to this species were taken by
1903. ] POLYCLADS OF THE ‘‘ SKEAT EXPEDITION.” 303
Mr. Evans on the shores of the island of Pulau Bidan, where it is
evidently one of the commoner species.
The body of this worm is rather elongated, rounded in front
and with a pointed hinder extremity. The arrangement of its
eye-spots is shown in the accompanying figure (text-fig. 50).
Text-fig. 50.
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Eye-spots of Notoplana evansii.
Unfortunately I have no notes as to colour, but, judging from the
spirit-specimens, the creature is probably of a yellowish-grey
colour, with a small number of irregularly scattered black spots.
The gut-branches are numerous, and there is no anastomosis.
The dimensions of an average individual are as follows :—
Iya t Ae edbacoucon an ohenonemarn ac ghas do 25 mm.
BCG 2 retrace tite. sectors stelatstteioe cielo gee Laat,
Tentacles from ant. margin............ ens;
Mouth-opening ok ne weno. EP ak},
d. Aperture behind mouth ......... BD) 45
Oe - behind male ............ TED: 555
Genital Apparatus (see text-fig. 51, 304).
The most striking features of the male copulatory apparatus
ave the great length of the antral chamber and the chambered
prostate organ, the latter recalling the prostate of certain
Leptoplanas, e. g. Z. aleinoi. From the male aperture the antral
cavity (text-fig. 51, a.m.) extends forwards as far as the level of the
hinder end of the pharynx. It has very muscular walls and runs
sloping in an upward direction. Lang, in his diagram of the
genital apparatus of Hoploplana (Planocera insignis group [6}),
calls this chamber the penis-sheath (‘‘ Penisscheide), but this term
should, I think, be reserved for the sheath-like folds of the walls
of the antral chamber occurring in Cestoplana and in certain
Cotylean genera.
The lining of the walls of the antral chamber in the present
species consists of a flattened ciliated (?) epithelium ; the muscular
wall consists of a thick layer of interlacing circular fibres.
Proc. Zoou. Soc.—1893, Vor. I. No. XX. 20
304 MR. F. F, LAIDLAW ON THE [ Apr. 21,
The penis (p.) itself is small and projects into the antral chamber
at the upper anterior end of the latter.
The penis is a fleshy organ, composed of longitudinal and
circular fibres which exhibit numerous nuclei (see Plate XXIII.
fig. 2). It is armed with a chitinous stylet.
The base of the penis is entered by the short ductus ejaculatorius
(d.e.) which leads to the prostate region. It turns shortly after
leaving the penis and runs in a backward direction, so that a
Text-fig. 51.
Male organs of Notoplana evansii.
For explanation of lettering, see p. 318.
transverse section passing through the penis passed also through
the prostate (see Pl. XXIII. fig. 2 and also text-fig. 51). |
The prostate consists of a number, some nine in all, of small
chambers (pr.c.) which lie about the ductus ejaculatorius and
open into it at their distal ends. The ductus, elsewhere with
muscular walls, here is lined only with’a very flattened epithelium,
outside which lies a second layer of similar cells belonging to the
1903. ] POLYCLADS OF THE ‘‘SKEAT EXPEDITION.” 305
inner walls of the prostate chambers. Between each of these
chambers a fine double septum runs out from this inner wall (Plate
XXIII. fig. 2, s.). The prostate cells stain very feebly, and the
lumen of each chamber is well defined, and contains in most cases
a certain amount of secreted matter. Outside the chambers is a
fairly thick layer of circular muscle-fibres, and, beyond these, an
ill-defined layer of cells which appear to be glandular. Here
and there are faint indications of processes from these outer
glandular cells, piercing the muscle-layer of the prostate organ.
This muscle-layer is also traversed by processes from the septa of the
chambers. There is no longitudinal muscle-coat to the prostate.
Towards the level where the prostate-chambers open into the
ductus ejaculatorius, some of them open into each other, so that
the number of chambers seen in cross-section is reduced to four
or five.
On the proximal, hinder, side of the prostate, the ductus is
continued back into the vesicula seminalis (v.s.), which is long and
contorted. It is lined with a ciliated epithelium, and its walls are
composed of regularly arranged circular muscle-fibres. At its
extreme hinder end it turns sharply forward and ends blindly.
Just before it takes this turn forward, the two vasa deferentia
open into it, one on either side, after piercing through its
muscular wall.
There is a spacious antrum femininum surrounded by the large
shell-glands, the secretions of which it receives. The vagina leaves
the antrum dorsally, and runs forward for the first part of its
course through the shell-glands, the secretion of which it also
receives. It then turns first dorsalwards and then backwards,
acquiring in its course a fine wall of circular muscle-fibres. After
running back for a short distance, it receives the short common
duct from the two uteri. It then continues to run back as far as
the level of the antrum femininum, where it ends blindly.
It is evident that this species is closely allied to von Plehn’s
Plagiotata promiscua [7]; in fact, the terminal parts of the male
apparatus in the two species have almost identically the same
structure,
It differs from von Plehn’s speciesin having the pharynx of the
normal type, not elongated transversely. This character is of
sufticient importance to justify the placing of these species in two
separate genera. In addition Wotoplana is characterised by
differences in the female apparatus, and by the great length of
the antral chamber of the male organ.
The genus Votoplana may be defined, then, as follows :-—
A Planoceroid genus with styliform penis and without a bursa
copulatrix. The male antral chamber is very long; there is a
complicated prostatic organ consisting of several chambers lying
around the ductus ejaculatorius in front of the vesicula seminalis.
The latter is long and twisted. Body rather elongate, without
marginal eyes ; mouth-opening rather behind the middle ; pharynx
normal ; gut-branches numerous, without anastomosis.
20*
306 MR. F. F, LAIDLAW ON THE [ Apr. 21,
Family LepropLaNip&.
LEPTOPLANA MALAYANA, sp. n. (Plate XXIII. fig. 3.)
A number of specimens; some, preserved in formol, are of a
uniform greyish-brown,
Length, about......... 30 mm.
Kiyes, abouthnceese a. 8 ,, from the anterior end.
Mouth-opening, about 15 a i
3 aperture, about ...10 ,, from the hinder end.
2 i ws ey Le eetromepnialle:
The arrangement of the eye-spots is that usual in the genus:
there are two small clusters of ‘ tentacle-eyes” about 1°5 mm.
apart, and in front of these are a few scattered spots on either side
of the middle line.
This species is closely allied to Z. pacificola von Plehn [7].
Genital Apparatus (see text-fig. 52).
The vasa deferentia (v.d.) open into the proximal (anterior) end
of the vesicula seminalis (v.s.); the latter, of considerable length
and much twisted, is lined with ciliated epithelium, and has thick
Text-fig. 52.
Genital apparatus of Leptoplana malayana.
a,f., antrum femininum ; acc.ves., accessory vesicle; sh.gl., shell-glands ;
u.t., uterine duct; v.d., vasa deferentia. For other letters see p. 318.
muscular walls made up of circular fibres. At its distal end the
vesicula narrows into the ductus ejaculatorius (d.e.), which is
shorter, but, like the vesicula, much twisted. Its walls are like
those of the vesicula, but it is much narrower and has a less well-
developed muscle-layer. The penis is small, fleshy, and unarmed,
and projects into the large antral chamber, which has thick mus-
cular walls and extends vertically upwards almost to the level of
the dorsal body-wall muscles. (See Plate XXIII. fig. 3.)
The female opening lies close behind the male, and leads into a
moderately large antral chamber (a.f.) with muscular walls. Into
this the vagina opens from above. This latter organ isa long and
tortuous duct of varying diameter; the shell-glands (sh.gl.) open
into it through nearly its whole length. At its hinder end, near
1903. ] POLYCLADS OF THE “‘SKEAT EXPEDITION.” 307
the point where the uteri open into it, the muscular walls, which
are elsewhere feebly developed along its course, become thicker,
but not very markedly so (cf. LZ. californica von Plehn). The
two uteri (wé.) unite as they enter it from below, and behind them
there is a very small accessory vesicle (ace.ves.) with muscular
walls.
The genus Leptoplana, as at present constituted, contains a
considerable number of species, many of them only referred by
Lang with doubt to the genus. In the majority of the species,
the anatomy of which has been investigated by means of serial
sections, the genital apparatus presents the following characters :—
The penis is directed backwards. The vasa deferentia open
into a muscular vesicula seminalis ; from this the ductus ejacula-
torius runs backwards to open into a prostate organ, which may be
chambered ; leaving this, the duct runs back into the penis,
which may be armed with a chitinous stylet.
The vagina runs back from the antrum femininum, receives the
secretion of the shell-glands, and further back the two uteri open
into it ventrally, usually by a common duct. An accessory vesicle
is generally present. The most familiar species of the genus, Z.
tremellaris, differs from the majority of species in the structure of
its male apparatus sufficiently to permit us to put it on one side
to form of itself a section of the genus further characterised by
the possession of a ventral sucker between the genital openings.
Another species, Z. swbviridis von Plehn [8] (=Z. pardalis mihi
[4]), approaches Discocelis tigrina in the structure of its female
organs, and accordingly we may put this species also in a section
apart.
The remaining species I have attempted to classify below, so far
as is at present possible, according to the structure of the penis
and prostate. I trust that I will not be thought to have laid too
much stress on the structure of these organs in dealing with this or
other genera. It seems to me that, of all the characters that
present themselves for classifying this order of Turbellaria, these
are the most useful, and that they are as reliable as any of the
other characters employed for this purpose, such as structure of
the pharynx, number and arrangement of eye-spots, presence or
absence of tentacles, &e. &e.
I propose, then, to group the species in two sections, A, and B,
so far as our knowledge of their anatomy permits:
A. Penis provided with a stylet.
(a) Prostate complicated by radially arranged ‘“ Drusen-
kaniile.”
1. L. kukenthali v. Plehn. Spitzbergen [7].
(6) Prostate divided into chambers lying parallel to the
ductus ejaculatorius.
L. vitrea Lang. Mediterranean ret
wo bo
. L. aleinoi Schmidt. 6].
9
308 MR. F. F. LAIDLAW ON THE (Apr. 21,
(c) Prostate not chambered.
4. L. panamensis v. Plehn. Gulf of Panama [7].
5. L. californica v. Plehn. California, Chatham Is. [11]
6. LZ. nationalis v. Plehn. Ascension Is. [8].
The following also belong to section A, but their position in
the section cannot be determined without information as to the
prostate characters.
L. variabilis Giard. |
L. ellipsoides Vervill. New England [13].
L. virilis Verrill.
L. drebachensis Oersted. | Norway [6].
L. fallax (de Quatvefages) [6] should perhaps form a separate
section of the genus on account of the great_length of its stylet.
It occurs on the south side of the English Channel.
B. Penis unarmed.
a. Prostate chambered.
7. L. chierche von Plehn. Callao [7].
(3. Prostate not chambered.
8. L. pallida (de Quatrefages). Mediterranean [6].
9. sp. (unnamed). Maldives [4].
y. No definite prostate.
10. Z. pacificola von Plehn. Valparaiso [7].
ll. LZ. malayana, sp. n. Straits of Malacca.
Belonging to this section but of uncertain position in it is
L. angust Vervill [13] from New England. JL. lacteoalba Vervill
[14] is insufficiently characterised, but said to be like L. pallida.
SEMONIA PENANGENSIS, Sp. 0.
Two specimens from Pulau Bidan. No note available as to
colour. This interesting species agrees fairly closely with the
only other species of the genus at present known, viz., S. macu-
lata von Plehn, from Java [7]. It is, however, readily distinguished
from that species by its external characters, the arrangement of
the eye-spots being obviously very different and showing some
approximation to the condition found in Discocelis tigrina (see
text-fig. 53, p. 309). |
The dimensions of the larger specimen are as follows :—
Total length, about ...... 22 mm.
Bread thwaboutperrssere.cr. INS
Mouth-opening, about ... 9 ,, from the hind end.
Genital aperture ......... 1 ,, behind mouth-opening.
Brainy about mesessceerrreec 4 ,, from the anterior margin.
The marginal eyes extend back for about 8 mm. on either side.
The arrangement of the brain and tentacle-eyes is shown in
text-fig. 53, p. 309.
An important distinction between this species and Semonia
maculata is afforded by the fact that in the latter the testes and
1903. ] POLYCLADS OF THE ‘‘SKEAT EXPEDITION.” 309
ovaries lie on the dorsal side of the body (see von Plehn [7]),
whilst in the present species the position of the testes is normal.
z.e. on the ventral side.
In one or two respects this species approaches the closely allied
Discocelis tigrina, and is perhaps intermediate between that species
and S. maculata. But the latter species is also certainly closely
allied to Diseocelis, and I am rather surprised to find that von
Plehn has not instituted any comparison between the two genera.
S. penangensis differs from S. maculata, and approaches Discocelis
tigrina, not only in the features already mentioned, viz., the
arrangement of the eye-spots and the position of the testes, but
also in that the uteri unite to enter the vagina by a short common
duct, and in having a very blunt, almost square penis.
Text-fig. 53.
“ Brain-eyes ” of Semonia penangensis.
It differs from Discocelis, and resembles S. maculata, in being
without the characteristic large prostatic cells which occur in the
epithelium of the penis and of the antrum, and without the
remarkable paired structures which run forward from the accessory
vesicle of Discocelis tigrina. Further, like S. maculata, it possesses
a definite vesicula seminalis, which appears to be absent in
Discocelis.
Lastly, S. penangensis is without the curious glandular vesicles
found along the anterior ends of the vasa deferentia in S. maculata;
and the vagina is prolonged, beyond the point where it receives
the openings of the uteri, into a very small accessory vesicle—so
small, in fact, that it almost escaped observation. This may be
due to the fact that the only specimen available for section-cutting
was not quite mature; or more probably, since the uteri were full
of apparently ripe eggs, that this organ is undergoing degeneration.
It is obvious that the distinction “between Discocelis and Semonia
is a very slender one, but still, I think, sufficient to warrant the
retention of the latter as a valid genus. The most important
characters separating the two genera are the absence of a vesicula
seminalis in Discocelis, and the curious horseshoe shi ape of the
accessory vesicle in that genus.
310 MR, F. F. LAIDLAW ON THE [ Apr. 21,
Family CryPTocELIDID&.
BERGENDALIA ANOMALA, gen. et sp. nov. (Plate XXIII. figs. 4,
5, 6, a 8.)
A most remarkable and interesting form, probably allied to
the anomalous genera Cryptocelides and Polypostia, described by
Bergendal [1], and provisionally referred to the same family with
them.
The structure of the female terminal ducts is, so far as I know,
unique, and approached only by the species referred to Z’rigono-
porus and Polyporus (see von Plehn [11)}).
Only a single specimen was obtained. It is rather a large
form, with a total length of about 60 mm. and breadth about
27 mm. The mouth-opening is some 15 mm. from the hind end,
and the genital pores lie halfway between these two points.
The margin of the body is complete, surrounded by a continuous
rather dense row of eye-spots. There are none of these, apparently,
over the brain. Hine
Colour in the spirit-specimen uniformly greyish white (see
Pl. XXIII. fig. 8). The specimen appears to be in an early
stage of sexual maturity, since no trace of ovaries or testes can be
discovered.
The pharynx is large and much folded, the gut-branches are
numerous and anastomose freely.
The cells of the epidermis are elongated, especially on the dorsal
surface. True rhabdites are absent, but in place of them the
epidermal cells are crowded with pseudorhabdites which are of a
coarsely granular texture, faintly stained and columnar in shape.
In my sections (stained with Grenacher’s hematoxylin) certain
gland-cells lying within the muscle-layers of the body-wall,
especially on the ventral side, are deeply stained ; from these cells
run processes which pierce the muscle-layers and basement-
membrane, and make their way to the surface through the
epidermal cells. These deeply-lying gland-cellsand their processes
have rather a spongy appearance, due to their preservation not
being quite perfect.
Genital Organs (text-fig. 54, p. 311).
The penis (p.) is a small fleshy organ composed of nucleated
longitudinal fibres (Pl. XXIII. fig. 4). Its outer side is lined with
cells continuous with those lining the antrum masculinum, but
whereas the latter are ciliated, those covering the penis are non-
ciliated. The base of the penis is pierced by a duct running
nearly vertically upwards. Immediately after leaving the penis
this is joined by two small ducts—the vasa deferentia (v.d.)—
which run forward on either side of the middle line. In one of
them, at the level of the hinder end of the pharynx, there is a
slight dilatation containing spermatozoa.
The vesicula (pr.) lying above the penis is small. Its wall
1903.] POLYCLADS OF THE ‘‘ SKEAT EXPEDITION.” 311
consists of a very thin layer of circular muscle-fibres, lined with
a cubical epithelium, the nuclei of the cells of which lie close to
the lumen, and which has rather the appearance of an exhausted
secretory tissue. This organ I call the prostate (Pl. XXITT. fig. 4,
pro.). About halfway between the penis and female aperture lies
an organ which bears some resemblance to the penis already
described. This organ constitutes one of the most interesting
features of this anomalous species. It consists of a small pro-
trusible penis-like organ (p.?), about one-third the size of that
first described, lying ina small antrum. It is pierced by a short
duct, which runs into it from a small non-muscular vesicle which
lies immediately above it (p7.?). This small vesicle is of about
- the same size as the prostate. There is no communication, so far
as I can discover, between the duct connecting these structures
and the vasa deferentia (Pl. XXITT. fig. 5).
Text-fig. 54,
Genital apparatus of Bergendalia anomala.
ace., accessory part of vagina; p. ?, penis-like organ; pr. ?, prostate-like organ ;
s., spiral coils of vagina. For other letters, see p. 318.
I can suggest only two explanations of the presence of this
second penis-like organ and its accessories.
Firstly, that it may be regarded as a prostatic structure which
has lost its connection with the penis and developed an intro-
mittent terminal part of itsown. Such a state of affairs is found
in the Cotylean family Diposthiide ; but in that case both penis
and prostate open into a common antrum, and there is what
appears to be a prostate gland in connection with the penis in
Bergendalia, a fact which makes strongly against this view.
The second possible explanation is that the organs under dis-
cussion are the vestiges of asecond penis. The fact that they bear
a close resemblance to the functional penis tends to support this
view, as does also, I think, a comparison with Cryptocelides of
Bergendal [1].
This is the only other Polyclad described, so far as I know, in
which two penial organs lie behind each other on the middle line ;
but, according to Bergendal, they lie behind the female aperture,
and, further, both open intoa common antrum, whilst in some cases
312 MR. F, F, LAIDLAW ON THE al Apri2it,
there may be four or even six penes present. In another genus
presumably allied to this, viz. Polypostia Bergendal [1], the penes
lie in large numbers around the female aperture.
The female apparatus is no less remarkable than that of the male.
The vagina (va.) runs forward for some little distance from the
aperture, then turns upwards. As it does so, it is twisted into
a remarkable spiral coil, making some five complete turns. . It
then runs backwards, narrows considerably, and soon receives the
openings of the two uteri (wt.) on its ventral side. Beyond this
point it is continued back as a narrow accessory vesicle (acc.ves.)
about as far as the level of the female aperture, when it turns
sharply ventralwards and opens to the exterior by the antrum.
The only other Polyclads with which I am acquainted that
possess a secondary female opening are Zrigonoporus of Lang [6]
and Polyporus of von Plehn [11], but in both these cases the
second opening is quite distinct from the primary one (the true
antrum femininum).
The curious spiral twisting of the vagina in the present species
is, so far as I know, unparalleled in the order.
For the first part of its course, 7. ¢. whilst it is running forwards,
the vagina is lined with elongated columnar ciliated cells, the nuclei
of which lie near their bases. Outside this epithelium is a thin layer
of circular muscle-fibres, and beyond these, in my sections, can be
seen a great number of nuclei massed round the vagina, and
probably belonging to gland-cells. Where the vagina turns
dorsalwards and becomes spirally twisted it has narrowed slightly,
but its epithelium retains the characters already mentioned. The
muscle-sheath does not follow the individual folds of the spiral, but
forms a continuous covering for that part of the duct (Pl. XXIII.
fig. 6, m.s.). The rest of the terminal female ducts are precisely
similar in character to the first part of the vagina, only narrower.
There seem to be no special shell-glands present.
The genus may be defined as an Acotylean genus in which behind
the functional penis a small second penial organ occurs. The
accessory vesicle of the female apparatus opens to the exterior
through the antrum femininum. Body pointed at either end.
Marginal eye-spots present ; mouth subcentral.
Family LArocestipa.
LATOCESTUS ARGUS, Sp. n.
Four specimens were collected by Mr. Evans. One of these is
labelled ‘‘ Chocolate-brown above, slightly lighter below.”
The largest specimen has a total length of about 30 mm. and a
breadth of 5mm. The mouth is 2°5 mm. from the hinder end of
the body.
This species is proportionately longer than the other member of
the genus which I have had an opportunity of examining. Its
most striking feature is the presence of a crowded row of eye-
spots running completely round the margin of the body. The
1903. ] POLYCLADS OF THE ‘‘SKEAT EXPEDITION.” 313
arrangement of the eyes at the anterior end of the body is shown
in text-fig. 55 A and towards the hinder end in 59 B.
The only other elongate form exhibiting this character with
which I am acquainted is Cestoplana ? maldivensis mihi. This is
almost certainly a Latocestus.
Text-fig. 55.
Eye-spots of Latocestus argus.
I can detect no rhabdites in the epidermis. The parenchyma
is very dense, and nervous tissue abundant on the ventral side
just within the body-wall muscles.
The terminal parts of the reproductive apparatus are almost
exactly similar to those figured by von Plehn [7] for Latocestus
paciticus.
314 MR. F. F, LAIDLAW ON THE [Apr. 21,
As in that species, the testes as well as the ovaries lie on the
dorsal side of the body.
THYSANOZOON AUROPUNCTATUM Coll.
Thysanozoon auropunctatum Lang [6]; von Stummer-Traunfels
[13].
One specimen, unfortunately immature and in a poor state of
preservation, although it has retained its colour fairly well.
PSEUDOCEROS BEDFORDI, sp. n. (Plate XXIII. fig. 9.)
This strikingly handsome and large species was obtained by
Messrs. Lanchester and Bedford in Singapore Harbour. The
single specimen, kindly given me by Mr. Lanchester, was unfortun-
ately in fragments and fully one-half of the specimen missing, so
that a satisfactory examination was not possible. I believe, how-
ever, that it was provided with a pair of penes. The colouring is
very beautiful, and quite sutticiently marked to render the identifi-
cation of this species in the future a simple matter, thanks to the
careful drawing for which I am indebted to Miss Dust.
On the label accompanying the specimen is written “Singapore
Harbour, from tide-marks to 10 faths.” I venture to associate
with this species the name of my lamented friend the late
Mr. Bedford.
PSEUDOCEROS COLLINGWOODT, sp. nh.
Closely allied to Ps. cerebralis (Kelaart) and Ps. striatus
(Kelaart), but smaller than either. It has the same regular
convolutions at the margin of the body that are shown in
Collingwood’s figures, and the coloration is similar especially to
Ps. cerebralis.
ene thyalboubets...-- 30 mm.
Bread t hirmness cs.) 25
Mouth-opening,about 3 ,, from the anterior margin.
Sucker about ......... 8 ,, behind the mouth.
The specimen isimmature and the penis is unpaired. Colour—
dorsal surface mottled dark brown and brownish white. The
extreme margin is white, but the white rim is exceedingly thin ;
just inside this is an equally fine black line. Ventral surface pale
brownish white, becoming darker towards the margin, which is
edged with black and white just as on the dorsal surface.
PSEUDOCEROS ? RUBELLUS, Sp. n. (Plate XXITI. fig. 10.)
A number of specimens of this very small species were collected.
One of these was “found under a stone between tide-marks. It
was magenta-red in colour. Nov. 1899.” Though very small,
these specimens appear for the most part to be mature. They
bear a close resemblance to Pseudoceros kelaartii (Coll.) in colour,
but the latter species is much larger and has the eye-spots arranged
quite differently. Thearrangement in the present species is shown
1903. | POLYCLADS OF THE ‘“‘SKEAT EXPEDITION.” 315
in fig. 10 of Plate XXIII. The tentacles also in Collingwood’s
species are much more prominent. The penis is unpaired. The
body is nearly circular and the sucker median. ‘Total length
about 5 mm., breadth 4mm. The arrangement of the eye-spots
is very different from that normally found in Psewdoceros, but I
cannot find any other characters distinguishing if from that genus.
At least two other species of this genus are represented in the
collection. One of these is a very small individual of the
P. superbus-group (perhaps P. hancockianus); whilst several
specimens belong to a species coloured exactly as Proceros con-
cinnwm Coll., viz., they are blue, with orange margin and median
stripe. They are, however, very much smaller, about 8 mm. to
10 mm. in length, and of a different shape, not pointed at their
hinder end, and have small, folded tentacles, whereas Collingwood’s
species has pointed tentacles. Proceros concinnwm in fact is almost
certainly a Euryleptid, whilst these specimens aie undoubtedly
members of the genus Pseudoceros.
I prefer, however, not to name them at present.
Family Dirostu1p«.
ASTHENOCEROS, gen. nov.
ASTHENOCEROS WOODWORTHI, sp. n. (Plate XXIII. fig. 7.)
Two specimens, without notes as to colour; but, to judge from
these spirit-specimens, the species is of a reddish-brown colour,
darkest on the middle line.
Length, about .... 17 mm.
Bread tints scents Ta 35
Mouth-opening... 7 ,, from the anterior margin.
SUCKER i ivr tteaioce 1:5 ,, behind mouth.
The male and female apertures lie in the usual order between
the ‘ mouth” and sucker.
The pharynx is large and much folded, its opening being sub-
median. This feature will serve at a glance to distinguish this
species from any of the Pseudoceridze. In describing the type of
the family, Woodworth [16] makes no statement as to the pharynx,
but, to judge from the figure given, it 1s median or submedian as
in the present species,
The body is flat and almost circular; the anterior margin is
feebly folded, and there is a small group of eye-spots on either
side of the middle line, but there are no very definite tentacles,
The prostate body hes in front of the penis.
In this respect Asthenoceros differs from the type-genus Dipos-
thus. There are no brain-eyes.
Only one of the two specimens was cut into sections, trans-
versely ; the other specimen appears, from an examination of it
when cleared in oil of cloves, to be quite immature. The speci-
men serially cut presents certain peculiar features, which I am at
316 MR. F. F, LAIDLAW ON THE [ Apr. 21,
a loss to explain satisfactorily. In the first place, the testes and
ovaries are very immature, and apparently only the terminal parts
of the genital ducts are developed. On the other hand, there is,
immediately over the penis, a large mass of mature spermatozoa
lying in a chamber the character of which cannot be determined
from my specimens, since it appears in places to have indications of
a proper lining-epithelium of its own, and again in places seems to
be merely a gap in the parenchymatous tissue. There are very
faint indications of a duct running from this chamber down in
the direction of the penis, but this duct cannot be traced far.
Woodworth [16] suggests in connection with the specimens of
Diposthus described by him, that they were in a late stage of
sexual activity, and that in consequence of this the sexual organs
were in a reduced condition. Possibly in my specimens of
Asthenoceros a similar state of affairs occurs, but, as I have
already stated, testes and ovaries in a very immature state are
present. There is also a possibility that the ripe spermatozoa
may be derived from another individual by hypodermic injection ;
but I do not think this is the case—firstly, because the penis is
not armed with a stylet; and, secondly, because the spermatozoa
lie over the penis, and because, as already stated, there are traces
of a duct running towards the penis from the chamber in which
they lhe.
The terminal parts of the male apparatus histologically resemble
those of Diposthus. The prostate and penis are separated into
two distinct organs, “both of which are doubtless intromittent,”
both opening by a single gonopore. The prostate lies immediately
in front, the penis directly over the aperture. It is rather
feebly supplied with muscle-fibres, and the secretory cells le in
the middle of the organ, but there is no lumen apparent
(cf. Diposthus corallicola).
The penis ismuch more muscular, and on the outer side has a
very definite series of circular and longitudinal fibres. Nuclei in
it too are much more numerous (Pl. XXIII. fig. 7).
In connection with the female organs there are traces of two
pairs of uterine vesicles. The uteri are very small and difficult to
distinguish from the surrounding tissue, and the ducts running
to them are merely solid rods of cells of an embryonic appearance.
The uteri open into the hinder end of the vagina. The antrum
is deep, and widens at its upper end where it receives the secretion
of the shell-glands.
The body epithelium is very densely crowded with small
rhabdites, and on the dorsal side especially with pseudorhabdites.
Scattered through the parenchyma, more particularly in the region
of the sucker, are numbers of small rounded darkly-staining cells
the nature of which is doubtful. The ventral surface of the body
projects immediately behind the pharynx into a prominent median
ridge which carries the gonopores and the sucker ; behind the
sucker the ridge disappears.
I have given as complete an account as possible of the cha-
1903. | POLYCLADS OF THE ‘‘ SKEAT EXPEDITION.” 317
racters of the genital organs, but, owing to the condition of the
specimen, it is obviously far from being satisfactory, and in order
to deal fully with this interesting species more material is
necessary.
PROSTHIOSTOMUM PALLIDUM, sp. 0.
One specimen, from the sea-shore, Dec. 1899.
Total length, about ......... 20 mm.
Breadth, about ..............- 4
Anterior margin rounded. Arrangement of eye-spots shown
in text-fig. 56.
9
Text-fig. 56.
br. e.
Eye-spots of Prosthiostomum pallidum.
e, marginal; b7.e, brain eye-spots.
Closely allied to P. siphunculus of the Mediterranean by the
arrangement of the eye-spots, and agreeing with it apparently in
being, so far as the spirit-specimen shows, of an uninterrupted
dull grey-brown colour, it differs sutticiently in that the two rows
of brain-eyes diverge continuously from one another from before
backwards, whilst in the Mediterranean species these two rows
converge at their middle.
It is also readily distinguishable, I think, from the latter species
by its smaller size. The single specimen obtained by Mr. Evans is,
to judge from the state of the sexual apparatus, fully mature.
P. siphunculus, moreover, has the genital sexual organs relatively
much smaller, to judge from Lang’s figures (see Lang, ‘ Poly-
cladida,’ pl. 5. fig. 3).
P. pallidum is certainly very distinct from any species from
the Indian Ocean that I have had the opportunity of studying,
and also, I believe, from the species found in the Pacific.
I believ e that Leptoplana aurantiaca ot Collingwood is really a
Prosthiostomum. It has the shape and proportions of a member
of that genus, whilst its eye-spots have the characteristic Pros-
thiostomum arrangement. It may, of course, be identical with the
318 ON THE POLYCLADS OF THE “‘ SKEAT EXPEDITION.” [ Apr. 21,
species described above, but, judging from Collingwood’s figure,
in the arrangement of the eye-spots it is sufficiently distinct.
Lnterature.
1. Bercenpat, “ Kongl. Fysiogr. Sallskapet,” Lund Handlingar,
Ny Foljd, 1892-93, Bd. 4.
2. CoLyinewoop. ‘Trans. Linn. Soc. Lond. ser. 2, Zool. i. 1875,
p. 83, pls. 17-19.
3. GRAFF, von. Zeitschr. wiss. Zool. lv. 1893, p. 189.
4. Larpnaw. Fauna and Flora of the Maldives and Laccadives,
ilo [Dy Pars JOS Kaen YE
5 Mem. & Proc. Lit. & Phil. Soc. Manchester, xlvii. p. 1
(1903).
6. Lane. Naples Monogr. xi., 1884.
7, PLEHN, von. Jena. Zeitschr. xxx. p. 137, t. vili. & xiii.
8
)
Ergebn. Plankt.-Exp., Heft 1., 1896.
Semon’s Zoologische Forschungsreisen, Bd. v. pp. 329-
304, t. XXlil.
Abh. Senckenb. Gesell. xxiv. 11. pp. 145-146 (1897).
It —— Jena, Zertschr. xxxi. p. 90; t. v. (1898):
Zool. Jahrb. Syst. xii. pp. 448-452 (1899).
13. Srummur-TRAUNFELS, von. Zeitschr. wiss. Zool. lx. p. 688,
t. XXXV.—XXXVII1.
14. Verritt. Trans. Connect. Acad. vill. p. 459, pl. xliv.
a a 1x, pl O23.
16. WoopwortTH. Bull. Mus. Comp. Anat. xxxii. 4, pp. 63-67,
pl. i. (1897).
EXPLANATION OF PLATE XXIII.
Fig. 1. Spine from the lumen of the penis of Planocera sp., p. 302.
2. Section passing through the penis and prostate organ of Notoplana evansii,
. 302.
3. Teaneverss section through the penis of Leptoplana malayana, p. 306.
4. ‘Transverse section through the penis and prostate gland (?) of Bergendalia
anomala, p. 310.
5. Section through the rudimentary organ of penis-like structure of the same
species, p. 311.
6. Section through the spiral part of the vagina of the same, p. 312.
7. Section across the antrum masculinum of Asthenoceros woodworthi, p. 315.
These organs are directed obliquely forward, and consequently both appear in
some of the sections. The granular appearance of the prostate in this region
is rather exaggerated in the figure, but it becomes more marked towards its
extremity.
8. Bergendalia anomala, p. 306.
9. Pseudoceros bedfordi, fragment of anterior part of body, p. 314.
10. Pseudoceros ? rubellus, p. 314.
Explanation of Lettering of Plate and Text-figures.
a.m., antrum masculinum. pr.e., chamber of the prostate.
a.m.l., muscular walls of antrum mascu- pro., prostate.
linum. ppr., prostate-like organ.
d.e., ductus ejaculatorius. S., Septum between chamber of
d.pr., prostate duct. prostate.
ep., epidermis. st., stylet of penis.
m.pr., muscular walls of prostate. | t., tentacles.
m.v., muscular sheath of vagina. va., lumen of vagina.
p., penis. | v.d., vasa deterentia.
pp., vestigial penis ? v.8., vesicula seminalis.
pr., prostatic cells.
1903.] ON THE TRANSPOSITION OF MAMMALIAN TESTES, 319
4. On the Phylogenetic Cause of the Transposition of the
Testes in Mammalia: with Remarks on the Evolution
of the Diaphragm and the Metanephric Kidney. By
W. Wooptanp, University College, London.*
{Received March 3, 1903.)
(Text-figure 57.)
The subject of the descent of the testes in Mammalia, notwith-
standing its bionomic interest, has hitherto received but little atten-
tion from. the phylogenetic sti mdpoint; indeed, so far as I know,
its interpretation has not hitherto been attempted. ‘To exhibit
the general nature of the problem and the apparent difficulty
attending its solution, the following paragraph from Mr. Spencer's
‘Principles of Biology’ (vol. i. p. 573) may be quoted :—“ But
now let it be confessed that though all phenomena of organic
evolution must fall within the lines above indicated, there remain
many unsolved problems. Take as an instance the descent of the
testes in the Mammalia. Neither direct nor indirect equilibration
accounts for this. We cannot consider it an adaptive change,
since there seems no way in which the production of sperm-cells,
internally carried on in a bird, is made external by adjustment to
the changed requirements of mammalian life. Nor can we ascribe
it to survival of the fittest; for it is incredible that any mammal
was ever advantaged in the str uggle for life by this changed
position of these organs. Contrariwise, the removal of them
from a place of safety to a place of danger would seem to be
negatived by natural selection. Nor can we regard the trans-
position as a concomitant of re-equilibration; since it can hardly
be due to some change in the general physiological balance.” I
agree with Mr. Spencer that the descent of the testes can neither
have been a change adaptive in nature, nor a result of the
operation of natural selection; on the other hand, I must dis-
agree with the statement that the phenomenon cannot be
regarded “as a concomitant of re-equilibration,” since, as will be
shown, I attribute it to the direct action of the conditions of life
found in the Mammalia,
To ensure due appreciation of the significance of the following
statements, it will be as well to here indicate the nature of the
theory about to be advanced. Of possible causes effecting the
transposition of the testes, there are two categories : («) causes
which as regards their effects on the organism mediately
or immediately impart advantage, so supplying the requisite
material for natural selection ; at (6) causes which in their
effects on organisation bear no appreciable relation to either
advantage or disadvantage. Since those of the former category
are inconceivable in this connection, we are led to conclude
1 Communicated by Prof. E. A. Mincurn, F.Z,S.
Proc. Zoot. Soc.—1903, Vou. I. No. X XI, 21
320 MR. W. WOODLAND ON THE y [Avr 25
that the transposition is an inevitable concomitant of some other
constant feature of the animal’s existence, thus not having arisen
in relation to ulterior ends. Again, since the testes consist of
ordinary matter possessing inertia and mass, their translation
(involving rupture or distension of the mesorchium) implies
mechanical force. From both of the foregoing considerations,
and from the fact that no other efficient cause is imaginable,
it is probable that the displacement results from the reaction
on the part of the testes to the incident forces, which, it can
be shown, are generated by bodily activity under mammalian
conditions of life. The theory here advocated is to the effect
that the descent of the testes in the Mammalia has been pro-
duced by the action of mechanical strains causing rupture of the
mesorchial attachments, such strains being due to the inertia of
the organs reacting to the impulsiveness involved in the activity
of the animals composing the group.
Mechanical Aspects of Organisation.
Before entering upon a discussion as to the production of the
forces above mentioned and the manner in which they have
acted, it is well briefly to outline the entire argument from the
mechanical standpoint. In the first place, accelerations imparted
to the body as a whole generate strains (or stresses) In connection
with the attachment or other means of support of every com-
ponent viscus, and the degree of such accelerations (and therefore
of the strains and stresses) is obviously dependent upon the
reaction which occurs between the animal’s body and the medium
in or substratum on which it is supported, being directly pro-
portional to the product of the powers of resistance possessed by
the substances constituting the same. Secondly, it must be
pointed out that the accelerations to which we refer, and to
which alone we attribute any importance, are those involved in
the impulses communicated to the body during the actual con-
tinuance of locomotion, and which inevitably result from the
mode of action of the propelling agency, whatever its nature.
The ordinary non-impulsive accelerations involved in the starting
of an animal into motion from a state of rest, or its converse, do
not concern us. In considering, however, the arboreal habits of
e.g. Primates, the case is different, accelerations involved in
motion from rest here being decidedly impulsive in nature. Now,
among the Vertebrata, it is evident that fishes, aquatic amphibia,
and birds severally exist in media which, owing to their mobility,
negative the occurrence of accelerations of high degree, 2. e.
impulsiveness. Hence, as regards these accelerations, we have
to consider only terrestrial animals, which come into contact with
a substratum possessing sufficient power of resistance to afford a
reaction of marked intensity. Of these terrestrial animals the
mammalian group is at once distinguished from terrestrial
Amphibia and Reptilia, both by the high degree of activity which
1903.] TRANSPOSITION OF MAMMALIAN TESTES. 321
its members exhibit, and by the more perfect ad: aptation of the
body (relation of limbs to trunk, &e.) to the locomotion in which
the superior activity is mainly manifested. Consequently, in
mammals, the reactions which occur between the organism and
the substratum vastly exceed in magnitude those occurring in
lower groups, and we have in this distinction a factor which in
considering bodily conformation is worthy of all attention.
Since in mammals the organisation is subject to such considerable
forces, 1b follows that if stability of position be ca in con-
nection with organs possessing appreciable mass, ¢. e. if these
organs are to adjust themselves to those tr: ihepaaine influences
which arise out of impulsiveness, fixative structures must be °
developed in response to the demand. For, as remarked above,
a sudden acceleration imparted to the body is necessarily trans-
mitted to an attached organ through its attachment, which, if the
organ be of considerable mass, is thereby subjected to an intense
strain, possibly ending in disruption. Stability of position of an
organ may be necessitated either on mechanical or physiological
grounds: thus, apart from considerations of function, viscera of
large mass (stomach, liver, intestine) must conform, as regards
situation in the body-cavity, to the principle that, ceteris paribus,
the more anterior the centre of gravity of the body the greater
the facility of progression’; and again, from the functional
standpoint, the kidneys must in active animals be maintained
anteriorly in order to ensure proximity to a vigorous blood-supply.
It will be evident from this, that if the position of an organ be of
no economic moment—if there be no need of localisation ‘either on
account of mass or volume for preservation of bodily equilibrium,
or on account of nature of function for physiological efficiency,—
retentive structures will not tend to be formed, and the organ, if
of sufficient mass, will respond by transposition to any considerable
force incident upon it—a trait which in the course of generations
will be inherited.
I shall now proceed to examine more in detail the production of
the forces hitherto assumed to be capable of effecting the trans-
position of the testes—to enquire whether the intensity and
direction of these forces are such as are congruous with the
hypothesis advocated,
Mammalian Locomotion.
In view of the importance that is here attached to the character
of mammalian locomotion, it is needful to discuss the subject in
detail. Every unsupported mass descends to the earth with a
given acceleration, and the magnitude of the force required to
re-elevate the body to its initial position is directly proportional
1 Well shown by measurements of the position of the centre of gravity in Pisces,
and by the abdominal contours of active cursorial mammals as compared with those
of species more slothful in habit (compare e.g. a greyhound or race-horse with a
St. Bernard or cart-horse, or , more generally, herbivores with carnivores, though
the different capacities of the dige stive organs have here to be eausidsyred),
21+
322 MR. W. WOODLAND ON THE [ Apr. 21,
to the height through which upheaval is effected, and inversely
proportional to the period of time occupied by upheaval. This
statement in its application to terrestrial locomotion provides
the clue for the solution of the present problem. With the
exception of aquatic organisms, which exist in a medium so
similar in density to themselves that the influence of gravity is
not felt, all animals which fly, crawl, leap, or run are subject to
this primary condition of self-support imposed by the earth’s
attraction. In Aves the body is not entirely unsupported, the
resistance of the air to the large expanse of body yielding con-
siderable aid in this respect, and the small amount of self-
sustainment needed can be provided by the minor elevations
imparted by individual strokes of the wing. The reactions
between the wing and the air are of necessity small, owing to the
mobility of the latter substance, and hence, in spite of the great
activity of birds, no great accelerations are imparted to the body
—the locomotion is not impulsive. The conditions affecting
terrestrial locomotion are exceedingly unlike. Here, as before
remarked, the powers of resistance possessed by the substances of
organism and substratum both being of high degree, reaction
between the two is correspondingly great. In Reptilia and
(excepting the saltatory Anura referred to below) terrestrial
Amphibia, however, the forces involved in locomotion are not
conspicuous for their intensity, owing both to the fact that the
period of upheaval is prolonged (no sudden impact occurring
between the limbs and the earth) and the small amount of eleva-
tion effected by the action of the limbs, these two features
resulting from the conformation of the body and the general
inactivity of the animal. In Mammalia, on the other hand, the
period occupied by contact of the limbs with the earth is
extremely brief and the height of elevation considerable; and
hence, though the angulation of the lmbs tends to diminish
concussion, the organisation is subject to the intense strains and
stresses resulting from the enormous forces generated during
locomotion. Consider the gallop of a typical Ungulate or
Carnivore. The elongated trunk possessing two pairs of limbs,
each pair being in an opposite phase of motion compared with
the other, it follows that the two halves of the trunk will alter-
nately be upheaved through a considerable distance on contact
with the earth being made by their respective paurs of limbs, and
depressed in the interval which exists between successive contacts.
That is, the pendulous swing of each pair of extremities is accom-
panied by an elevatory impulse at the centre of the are each
describes, and these impulses effect the upheavals of the respective
halves of the body through the distances which they have fallen
in the time required for each pair of limbs to describe twice the
length of its appropriate path. Considering the mode of action
of either pair of limbs, and beginning at the horizontal stage of
the trunk’s position when the half of the body is being depressed
under the influence of gravity, this continues to descend until the
1903.] TRANSPOSITION OF MAMMALIAN TESTES. 323
limbs make contact with the earth, when, as already descriped,
the entire portion of the trunk is suddenly upheaved with con-
siderable force. If the horizontal velocity could be reckoned as
constant, the only accelerations which the trunk would acquire
would be in a vertical line; but since the horizontal velocity is a
quantity which, owing to the resistance of the air, is continually
decreasing, the upward acceleration suddenly imparted to the
trunk at each contact of the limbs with the earth has to be com-
pounded with a sudden forward acceleration in order to maintain
the pace, and hence the resultant acceleration which the half of
the body acquires at every upheaval is in an wpward and forward
direction. It is most essential to recognise this fact that the body
as a whole has periodically imparted to it large accelerations in
these two directions. As is obvious, the upward and horizontal
velocities due to these accelerations become respectively neutralised
by gravity and atmospheric resistance at the middle and end of
the interval which exists between successive contacts of the limbs
with the earth. From the instant when the maximum elevation
of the half of the trunk is attained, 7.e. when the upward velocity
is completely neutralised, until contact again occurs, the body
continues to acquire a downward acceleration due to gravity.
But on contact taking place, the large downward v elocity due to
this acceleration is suddenly destroyed, and as suddenly exchanged
for an equal upward velocity, in the brief instant of time occupied
by the impact of limbs and earth. Whence it is evident that the
degree of upward acceleration imparted to the half of the body
during the sudden loss and gain of downward and upward velocities
must be considerable. It is noteworthy that whereas each forward
acceleration, due to either pair of limbs, is imparted to the body
as a whole, each upward acceleration is solely imparted to that
portion of the trunk to which the pair of limbs causing the
elevation belongs; and hence the attachments of those organs
situated in the median portion of the trunk are evidently not so
subject to downward strains as in the case of organs placed more
terminally. Thus it is important to notice, with regard to what
follows, that the primitive pre-mammalian portion of the testes is
decidedly posterior (see diagram, p. 334), and that in consequence
the motion of the hind portion of the body is the factor of most
importance in the consideration of the present problem.
From the foregoing, it is sufticiently manifest that at each
elevation of the body in mammals a considerable force is imparted
to the organisation asa whole (and by necessary implication to its
constituent parts) inan upward and forward directior ras to
be proved. If the constituent parts of the body be cousidureah it
will be evident that such a force works differential effects among
them. For these organs, largely differing among themselves in
regard to mass, and the same force being incident on all, it follows
that the individual reactions are e different, and hence there exists
a tendency to segregation of those organs of greater mass fromm
those of less. Moreover, organs differ inter se in their relation
324 MR. W. WOCDLAND ON THE [Apr. 21,
to the rest of the body. If an organ be imbedded or otherwise
firmly aftixed to the main bulk, it will on account of its large area
of attachment and close apposition readily share both the eleva-
tions and depressions incurred in locomotion—the strains and
stresses are distributed in space and time, and hence their inten-
sity is lessened; if, on the other hand, a massive organ be merely
suspended from the main mass of the body, such a condition
subjects the attachment to the influence of severe strains periodi-
cally recurring, the direction of which approaches that of the
resultant acceleration of the trunk during its impulsive elevation.
The great severity of these strains on the attachment of an organ
thus disposed will be more fully realised if it be remembered
that, on the sudden elevation of the body, the organ possesses the
downward velocity due to the previous depression—a momentum
which in being destroyed throws additional strain on the means
of support. As will be seen, the preceding remarks apply in an
eminent degree to the testes.
The Genitalia of Mammalia.
The reproductive organs of the Mammalia are relatively small
bodies of great density and, in the case of the males, compactness,
and they primitively occupy a position similar to that found in
the lower Vertebrata. It is, however, characteristic of the majority
of the Mammalia that in the course of development the testes
forsake their primitive lodgment, and migrate posteriorly and
ventrally to the terminal periphery, where they protrude at the
surface of the body-wall. This protrusion constitutes the scrotum,
in the wide sense of the term, which may vary in character from
that of a pair of small slightly-elevated areas to that of a capacious
oval pedunculated sac. Apparently in man alone the section of
the ccelom contained within the scrotum becomes completely
separated off from the main cavity; in other mammals, com-
munication is retained by means of the inguinal canal, which,
however, is usually narrow, so negativing return of the testes to
the main body-cavity. This feature in the case of active
mammals possessing large testes is very important, since if the
unattached organs were permitted to return to the main cavity,
the forces to which they would be exposed during locomotion
would doubtless be exceedingly detrimental—a malinfluence which
both the narrowing of the inguinal canal and the possible tractive
function of the gubernaculum during development safeguard
against. In fact, contrary to the usual supposition, the internal
and not the external situation of the testes is the source of
danger. It will also be observed that in small animals which do
not possess narrow inguinal canals, there is no such dire necessity
for the restriction of the testes to the scrotal cavity, both on
account of the smaller size of the organs and the usually less
intense forces to which the testes are subjected. Thus, para-
doxical as it may seem, the increased protrusion of the testes
1903.] TRANSPOSITION OF MAMMALIAN TESTES. 325
{
beyond the surface of the body-wall, besides being a necessary
result of more massive organs and the incidence of more powerful
forces, is in itself a further means of preservation. In physical
character, the testes, as already mentioned, are definite bodies of
concentrated form, suspended dorsally in the body-cavity by the
thin mesorchial membrane. Also investigation shows that, with
the exception of the heart, the testes are the densest organs in
the vertebrate body, and hence, as they possess appreciable volume,
their mass is also considerable. Moreover, the testes (and ovaries)
are distinctly separated from the remainder of the body, this
separation (lending additional facility to transposition) arising
out of the fact that the gonads are from their very nature indi-
vidualised, reproduction essentially consisting of the separating
off of a portion of the organism; and hence the body primawily
serves as a mere carrier of these organs, which, unlike the
kidneys e.g., bear’ no relation to the economy of the animal.
From this it follows that the transposition of the testes is of no
concern to the rest of the organism, since the process can cause
no derangement of function in other portions of the body; and
hence the testes, differmg from all other organs in this respect,
do not, under ordinary conditions, require to be retained in their
pr imitive position by the special development of fixative structures.
Thus, as regards their definiteness and concentration of form,
their means of suspension, their superior density, and their
structural and physiological separateness from the rest of the
organism, the testes fully conform to the above-specified con-
ditions favourable to transposition. The transposition of the
testes occurring under conditions which permit the descent of
these organs alone (and the testes alone have descended), and a
cause capable of effecting this transposition solely existing in the
Mammalha (in which group descent has alone occurred), it is
probable that the latter phenomenon is the cause of the former. I
hold that, in the majority of the Mammalia, the testis attachment
has thr oughout the history of the race been constantly subjected
to severe “strains consequent on the character and conditions of
mammalian locomotion, and that on account of the resulting
disruption or distension of the mesorchium, the testis has migrated
in a postero-ventral line (7. e. in an opposite direction to the
forward and upward accelerations imparted), coming to lie at the
terminal periphery of the body-wall and forming the scrotal pro-
trusion. Just as when a man runs, a weight in his coat-pocket
will periodically “drag” and ultimately wear a hole in the lining
by constant distension, so the testis of mammals has responded to
like forces resulting in “ descent.”
I now proceed to consider the genital organs of the Mammalia
as a whole, the general conformation of which amply confirms
the foregoing conclusion, tending to show that the testes have,
in every case, reacted in a degree proportional to the forces
concerned. Indeed, the general correspondence between situation
of the testes and grade of impulsiveness displayed in the various
326 MR. W. WOODLAND ON THE Apr. 21
if )
orders and families of the Mammalia affords such conclusive
evidence as to the causal relation subsisting between the two
that a systematic review of this evidence is well called for. That
the correspondence is not absolute, however—that there exist
instances of the concurrence of scrotal testes with sluggish habits
—is without doubt mainly, if not wholly, to be explained by
phylogenetic considerations. For it must be remembered that
once the descent of the testes is inherited, the trait is a constant
one (unless acquired sluggishness is able in time to produce effect,
which is doubtful), and remains so in successive generations
whatever varied habits may be assumed, unless the bionomic
aspect so changes that natural selection cancels the ‘ plus varia-
tions,” so affording an ascendancy to reversionary factors. That
such has been the case in several instances will be illustrated
below.
In the Monotremes the activity (impulsiveness) is of a very
low degree. Ornithorhynchus is “aquatic in its habits, passing
most of its time in the water or close to the margin of lakes and
streams.” The ‘ body is rather long, compact, and almost every-
where of the same thickness. It rests on short, massive legs
......80 short that the animal in walking or running actually
drags its body along the ground.” Hchidna is “ usually found in
rocky districts, and more especially in the mountains...... and
is mainly of nocturnal habits.” It is described as ‘“ indolent.”
The testes in both instances are ‘abdominal in position through-
out life,” and afford the only example in mammals of a disposition
of these organs anterior to the kidneys. The low status of these
animals is well known.
Inthe Marsupials “ the testes are always contained in a scrotum,
which is suspended by a narrow pedicle to the abdomen in front
of the penis.” If it is permissible to assume that the ancestor of
the marsupials was kangaroo-like, the pre-penial position of the
testes may perhaps be attributed to the peculiar mode of loco-
motion characterising this animal. For the rapid locomotion of a
Kangaroo consists of a series of leaps, and such leaps would involve,
ag in ordinary mammals, a series of antero-dorsal tensions on the
suspensory membranes of the testes; but since the bodily depres-
sions are marked by a more prolonged termination, as compared
with those involved in the ordinary gallop, the testes would in
addition tend to be thrown: forward, and these two factors in
conjunction have possibly led to the peculiar position of the testes
found in marsupials. On no other theory than that of descent
from an ancestor characterised by pre-penial testes can the peculiar
genital conformation of the variously habited marsupials be corre-
lated with special bionomic conditions; and hence, on this ground
alone, it is possible, as implied above, that the ancestor of the
Marsupialia was of the type of the Macropodidee, the characteristic
saltatory progression originating the pre-penial situation of the
scrotum in the manner indicated.
The Edentata represent another ancient group. Their activity
1903.] TRANSPOSITION OF MAMMALIAN ‘TESTES. all
in general is small. The Bradypodide are arboreal inhabit,
and are characterised by ‘“ habitual sluggishness,” being ‘ most
inefticient walkers,” and in climbing, never leaping from bough
to bough (a form of activity involving more impulsiveness than
even the gallop, but probably not more than that involved in
saltation). Their testes are placed close to each other, lying on
the rectum between it and the bladder; 7. e. are retained in the
abdomen. The Myrmecophagide are also not noted for activity,
and their testes are disposed as in the Sloths. The Dasypodide
are similarly inactive, being ‘‘ harmless and inoffensive,” endeavour-
ing to escape by rapid burrowing. However, it is stated that
“they can run with great rapidity.” The testes are abdominal,
lying “above the brim of the pelvis.” The Manide are terrestrial
and burrowing in habit, but some members of the group can
climb trees. They are not very active. The testes lie in the
inguinal canal. The Orycteropodide (Aard-varks or African
Ant-eaters) are terrestrial and fossorial. Their testes are
“inguinal, but they appear to descend, at all events temporarily,
into a scrotum.” Their phylogenetic position is uncertain. In
the order Sirenia the testes, needless to say, are abdominal.
These animals are “slow and inactive in their movements, mild,
inoffensive,” browsing at the bottom of water.
The piscine locomotion of the Cetacea sufficiently accounts for
the abdominal position of their testes, which organs are placed in
the proximity of the kidneys. Even assuming the terrestrial
ancestor of the Cetacea to have been characterised by the exter-
nality of the testes (which is improbable considering their existing
localisation), such would inevitably have assumed an internal
position consequent on the serious risk of injury involved under
Cetacean conditions,
The Rodents are comparatively small animals. In habit they
are mostly terrestrial, but some are arboreal and some natatorial.
“The testes in the rutting- season form projections in the groins,
but (except in the Duplicident ita) do not completely leave the
cavity of the abdomen,” 72.e. the scrotal elevations are not
well-defined. The Duplicidentata comprise the Hares, Rabbits,
and the Picas or Tailless aie all extremely active animals,
the latter being described as “agile” and as living in crevices
among rocks. The Siilicidontats comprise the. remaining
Rodents, the principal families of which are the following :—
The Sciuridee “ vary between the two extremes presented by our
ordinary squirrels, the agile climbers, and the sluggish, clumsy
marmots, which live almost entirely underground.” Assuming the
sti stement to be correct that the condition of the testes is similar in
each of these two divisions (which is doubtful), it is evident that
the latter are specialised forms descended from active ancestors,
The locomotion of the Dipodide resembles that of the marsupial
Macropodidee and insectivoran Macroscelidee, ‘* The whole struc-
ture is adapted for jumping, and we find resemblances in their
structure on the one hand to the kangaroos, and on the other
328 MR. W. WCODLAND ON THE [Apr. 2],
hand to the jumping-shrews among the Insectivora.” These
animals, unlike the Macropodide, move their hind limbs alter-
nately in the process of walking. Unfortunately the writer has
been unable to discover detailed information with regard to the
testes in these groups. Other families are the clumsy, thick-set
Hystricide, the very active Castoride (Beavers), Myoxide (Dor-
mice), and Muride. During the rutting-season the testes of
Rodents migrate more posteriorly than at normal times; but this
merely indicates a slight displacement probably due to the periodic
enlargement of the organ, the cremaster muscle effecting return
by decreasing the capacity of the scrotal emergence. This can be
well seen in the Muride, where the slightest impact will cause a
dislodgment of the testis.
The order Insectivora comprises ‘‘small animals ...... of very
low type...... belonging to the oldest mammalian stocks......
which are generally terrestrial, although rarely of arboreal or aquatic
LDVOMS 5 bo bec the greater number are cursorial.” “ In the sub-
family Centetine, and Chrysochloris, the testes lie immediately
behind the kidneys, but in others more or less within the pelvis;
during the rutting-season they become greatly enlarged, forming
protrusions in the inguinal region.” The various families of this
group afford good illustrations of the correlation of impulsiveness
with degree of testis transposition. As stated above, in the
Centetince or ‘‘ Crawlers ”—the appellation denoting their mode of
locomotion—and Chrysockloris, which is active and fossorial,
the testes lie just posterior to the kidneys, 7. e. are abdominal.
In the companion subfamily, the Oryzorictine, the two genera
are represented by small animals—Macrogale being mouse-like
with a long tail, and Oryzorictes mole-like in form. The testes
are situated near the urethra. In the Hrimacemme or Hedgehogs
the testes are situated on the “‘ underside of the inguinal canal.”
Their ‘‘ movements are sluggish, their steps almost tottering, their
gait clumsy.” On the other hand, in the active Soricide (Shrews)
and saltatorial Macroscelide (Jumping-Shrews), the testes project
at the periphery of the perineum, and in the Solenodontide, the
feet of which are ‘‘formed for running,” the “testes are received
into perineal pouches.”
In the order Chiroptera, the members of which are so highly
specialised for flight, there is “no scrotum, and the testes are
either abdominal or inguinal. ” “ We find in the low organization
of their brain a proof of their inferior status ”—a fact otherwise
implied by the absence of the scrotum, unless a secondary reten-
tion has occurred owing to the danger of partial externality
involved in the conditions of flight and position of the hind limbs.
The order Ungulata is subdivided into the Ungulata vera and
the Subungulata, In the former suborder, needless to say, the
testes of the large majority reside in a well- ‘defined scrotum, cor-
responding to the eminent impulsiveness of the animals. In the
Nasicornia (Rhinoceroses), however, ce testes are inguinal, the
tunica vaginalis communicating freely with the body-cavity. As
1903.] TRANSPOSITION OF MAMMALIAN TESTES. 329
is well known, they are “huge, heavy, clumsy creatures, with
bent legs so short that the belly seems almost to drag on the
ground.” The Subungulata comprise the Hyracoidea and the
Proboscidea. The ancestry of the members of both these sub-
orders is as yet undecided, their status, decidedly low in the scale,
being obscure. The Hyracoidea are animals of about the size of a
rabbit or somewhat larger, and “resemble small marmots.” They
possess a “short, fat body” with “weak and short feet.” “In
most species there is a complete adaptation to a life among the
rocks,” by possession of curious clinging habits resembling those of
the Geckos. They are “agile in their sports, but rather lazy where
food is abundant.” Owen states that ‘‘the testes are abdominal,
below or beyond the kidneys.” The low status of the Proboscidea
is shown by several anatomical traits, such e. g.as the possession of
two ven cave and the structure of the hmbs. These animals
have been well described as “ peaceable colossi.” Their gait 1s
“ pretty slow, though the Colossus can run very fast w ee once
in full career, but this pace never lasts very lung and is always
maintained in a straight lime.” Owing to their huge size
Elephants never gallop. The most active members are the
“rejected males ”— “such extra activity thus beimg neutral so far
as the inheritance of any tendency to testis descent is concerned.
Elephants are vegetable feeders and are “ gregarious, generally
inoffensive and even timid, fond of shade and solitude and the
neighbourhood of water.” The testes are permanently abdominal
—a fact explicable by habits, though phylogeny alone can afford
a complete solution. Here, as possibly also in the case of the
Nasicornia, it may be pointed out that the huge size of these
animals itself implies inactive ancestors, for, according to one of
the conditions of growth enumerated by Spencer, great activity is
antagonistic to Increase by bulk, and the occurrence of the latter
negatives the past existence of the former.
The order Carnivora is subdivided into the Fissipedia and the
Pinnipedia. The members of the former group are all exceedingly
active animals of large size,and, together with the Ungulata, include
the swiftest of terrestrial siimals: Their testes, needless to say,
reside in a well-defined scrotum. The Pinnipedia afford an inter-
esting illustration of the secondary operation of natural selection.
In the Otariide the “‘ hind feet are turned forwards under the body,
and aid in supporting and moving the trunk as in ordinary
mammals...... They spend more time on shore, and range
inland to a greater distance than the true seals.” In the Otariide
the testes are “suspended in a distinct external scrotum”?. On
the other hand, in the Phocide, ‘the hind limbs are directed so
far backwards that they continue the horizontal direction of the
vertebral column ...... They move on land only with difficulty
by fixing themselves with their flippers in front and pulling up
their hinder parts, then drawing their bodies up into a curve and
1 Owen states that the scrotum is not distinct.
330 MR. W. WOODLAND ON THE [Apri2i,
throwing their front parts again forwards. They drag their belly
along the earth, show little suppleness, and soon become tired.”
There is no scrotum, the testes being abdominal and “ imbedded
in areolar tissue.” Thus, as before maintained, the embryonic
transposition of the testes tends to be checked if external conditions
become so changed as to render complete descent of these organs
disadvantageous to the animal, such being obviously the case in
the Phocidz, which have become further modified for an aquatic
existence than the Otariide. It may be objected that ordinary
mammals are subject, though in a less degree, to a like drawback
—that their testes are also exposed to considerable danger, and that
on the hypothesis descent ought to have been negatived by natural
selection, 2.e. by the development of ligaments or supporting
areolar tissue. Incidentally noting that the actual facts prove
that no such danger exists, it may be observed that the testes of
a typical mammal are very efficiently screened, not only laterally
by their position between the broad thighs of the hind limbs, but
also posteriorly by the tail; and they are well preserved from
contact with surrounding objects by the elevation of the body
upon its limbs, the case being otherwise in Phocide, Cetacea, and
lower animals.
The ancestral history and arboreal or other habits of the
Primates constitute sufficient warrant for the conspicuous exter-
nality of their testes. ‘That active arboreal habits involve
impulsiveness of the highest degree, is sufficiently manifest on
contemplating the movements of any of the ordinary monkeys,
more especially in the case of the Gibbons.
Thus in a review of the Mammalia, we encounter a considerable
mass of evidence testifying to the validity of the theory here
advocated, and more might be added. In the Monotremata,
Sirenia, Cetacea, most Edentata, Hyracoidea, Proboscidea, and
Phocidee, conditions prevail, or have prevailed, negativing the
descent of the testes; and these conditions have either consisted
of the absence of that type of terrestrial locomotion which has
been the sole cause of the transposition of these organs, or of
secondary factors which have either negatived the operation of
the primary agency, or effected a reversion of the pre-existing
effect of the same.
Since the transposition of the testes is mainly due to their mass,
and definiteness and concentration of form—their means of
suspension and physiological separateness from the body merely
constituting conditions to the transposition—it follows that the
small and diffuse ovaries of the Mammalia will not respond in
any degree to the forces incident on the body, the magnitude of
the strains on an attachment obviously being proportional to the
mass of the organ attached. But apart from the smallness of
mass possessed by the ovaries, there exists an important reason
for their retention at or near the primitive position i the body-
cavity. This reason is the necessity of the proximity of the ovary
to the oviducal aperture for the conservation of the ova, the ovary
1903. ] TRANSPOSITION OF MAMMALIAN TESTES. 331
and the oviduct usually possessing no anatomical connection.
How important this is may not only be inferred from @ priori
considerations, but also from the discovery of the many structural
devices adapted to this end. Thus there occurs the “ develop-
ment of special folds of the peritoneum, which practically ensure
the passage of the ova into the oviduct when they are extruded
from the ovaries. The oviduct, moreover, has a large and
fimbriated mouth, called in human anatomy the ‘morsus diaboli.’
This almost wraps round the ovary, and thus prevents the ova
from straying in the wrong direction. Moreover, the ovary itself
is often so arranged that it can easily be withdrawn into a pocket
of the peritoneum, from which the obvious exit is by the gaping
mouth of the oviduct. This disposition of the generative parts is
still further modified in a few animals, such as the Rat and the
Kinkajou. In these animals the mouth of the oviduct actually
opens into the interior of a closed chamber which contains the
ovary” (Beddard). Thus there exists ample reason for the
retentive ligaments usually associated with the mammalian ovary
(see Appendix).
The Genitalia and Conditions of Locomotion in Ichthyopsida
and Sauropsida.
If, as we have observed, the conditions of mammalian loco-
motion alone subject the organisation to those concussive influences
which have effected, among other changes, the transposition of
the testes; and if, as we have also seen, only the highest mani-
festations of terrestrial activity are capable of producing complete
testicular descent, then we may be certain that neither in the
relatively inactive terrestrial Reptilia, the active aerial Aves, nor
the aquatic Pisces will a ike phenomenon occur. In Reptilia,
though Ophidia and many Lacertilia are capable of brief spasms
of great activity, the total impulsiveness is very small and of low
degree. This is not only due to general passivity, but also to the
fact that “the body of a reptile is, as it were, slung between its
limbs like the body of an eighteenth century Ghanigt between its
four wheels,” with resulting i imperfection of the ‘relations of the
limbs to the trunk from the point of view of a terrestrial
creature” (Leddard). Apart from the absence of the causal
conditions, the depression of the trunk and consequent liability
of injury to the testes would in itself have negatived descent
(cf. Phocidee above).
In the Anura, the only terrestrial amphibia which concern us,
adoption of terrestrial habits, resulting in a higher degree of
impulsiveness, has as usual for its concomitants structural con-
centration and increased retentivity or transposition. Here the
testes have assumed a “full oval form, compact and undivided :
they are situated, as shown in the Frog, on the ventral side of
the anterior half of the kidneys” (Owen). That the testes
have slightly descended in correspondence with the saltatory
332 MR. W. WOODLAND ON THE [ Apr. 21,
progression, is shown on comparison of two such forms as Triton
and the Frog (see diagram below, p. 334). If, as we admit, anuran
progression involves on a small scale those impulse s to which we
attribute the descent of the mammalian testes’, why have not
these organs migrated more than they have in this instance? In
the first place, it may be contended that the total impulsiveness of
a frog, though quite appreciable, is yet of small value as compared
with that of a mammal of the same dimensions, say the Common
Mouse. Secondly, the physiological and strucbur: al relations
obtaining between the testes and the kidneys afford sufficient
grounds for the special retention of the former, and, moreover,
the firm attachment of the testis to the kidney is shielded from the
abruptness of incident strains by the slight mobility of the latter
organ. Finally, in addition to the above considerations, it is
probable that the descent of the testes would here be prohibited
in consequence of the depression of the hinder portion of the
trunk and resulting proximity to the external surface. However,
that such a cause for retention might be operative, it would
obviously necessitate transposition to an extent that enabled the
organs to experience adverse influences, and it is questionable as
to whether they have travelled so far.
In the typical Aves, as in Mammalia, the expenditure of energy
is very great, but in the former, as also in Pisces, aquatic
Amphibia, &ec., the bionomic conditions negative impulsiveness of
locomotion, and their organisation is not subject to the kinetic
influences which affect terrestrial animals.
One noteworthy subject is the large increase in volume of the
testes during the brief courting-season: thus Owen provides a
figure illustrating the periodic enlargement of these organs in the
Common Sparrow, showing that their size ranges from that of a
pin’s head to that of a fair-sized marble, half an inch in diameter;
and this same periodic increase of volume, though not so marked,
occurs in sundry orders of the Mammalia, e. g. in Rodents and
Insectivores. It is easily comprehensible that if an animal’s
activity, however impulsive, were largely restricted to the inter-
breeding periods, the testes, possessing small mass during that
time, would exhibit less tendency to displacement; but as yet it
is not possible to decide whether this factor has any significance.
Connected with a consideration of the position of the testes in
birds, is the important case involved by the habits of the
Ostriches and their allies. The considerable speed attained by
these birds, comparable indeed with that of the swiftest mammals,
must involve a certain amount of impulsiveness ; and such being
the case, why have not the testes responded by descent ? For the
situation of the testes in Ostriches is normally avian, being “‘ placed
above and a little external to the kidneys,” which latter structures
are “elongated, flattened, glandular masses...... lying deeply
seated, and extending from the posterior edge of the diaphragm
1 “The creat Pull-Frog may clear six feet at a leap, and repeat them so rapidly
as to escape a pursuer, unless chased at a great distance from the water” (Owen).
¥903.] TRANSPOSITION OF MAMMALIAN TESTES. 333
to the anterior extremity of the pelvic cavity ” (Macalister). It
is obvious that non-descent cannot be ascribed to the serious risk
of injury that externality would involve (for, unlike as in the
mamumals, the position of the proportionately thin legs and the
absence of a tail ensure no protection from external influences),
since such an explanation is negatived by the present position of
the testes. However, if the several characteristics of the mam-
malian gallop be called to mind, it is easy to afford an explanation
of this apparently anomalous instance of the non-transposition of
the testes. For, as I shall show, the bipedal progression of the
Ostrich does not involve impulsive upheavals of the body to any-
thing like the same extent as in the gallop of a mammal. The
great upward impulses associated with the upheavals of the mam-
malian body are in large part due to the elongation of the trunk,
for the depression of one half of the body is lareely accelerated by
the impetus imparted to it on the sudden elevation of the other
half. That is, the upheaval of one half of the body rotates the
whole length of the trunk about its centre of gravity, and so
causes the other half to descend with greater acceleration than
it would do under the sole influence of gravity. Hence the
marked intensity of the contacts of the limbs with the earth,
which have not only to destroy the downward velocity due
to gravity, and the additional downward velocity due to this
lever-like action of the trunk as a whole, but have also to
impart an upward velocity equal in amount to the two it has
neutralised. In the Ostrich, on the other hand, there is no elon-
gated trunk possessing two pairs of limbs, and hence there can be
none of that lever-action just described. Moreover, the individual
elevations of the body are small compared with those of mammals,
owing to the alternate action of the limbs (¢f. the Kangaroo, in
which the action of the limbs is not alternate, and in which the
elevations are consequently great). And lastly, the gliding
motion adopted by the Ostrich in running is yet another factor
tending to diminish the intensity of the involved concussions.
Tfence for these three reasons the individual impulsive accelera-
tions imparted to the body of the Ostrich are very small compared
with those which exist in the case of mammals; and therefore it
can easily be understood how it is that, despite the terrestrial
locomotion and the perpendicular limbs, the ‘testes have not
descended in this instance. It is also possible that the posterior
extension of the sternum and ribs in the struthious trunk,
leading to tight packing of the viscera, so serves as a support for
the testes (and other organs), but whether this is the case can
only be determined by referring to the facts.
In conclusion, it will not be out of place to here give a brief
summary of the entire argument concerning the cause of the
descent of the testes, since it will show how strong is the proba-
bility of the theory advanced. We have seen that in mammals
alone there exists a mechanical cause competent to effect trans-
position. If any doubt remains in the mind of the reader as to
334 MR. W. WOODLAND ON THE { Apr. 21;
the competency of this alleged cause, contemplation of the actual
locomotion of a horse or dog, or reference to the works of Marey,
Pettigrew, and others will soon dispel it. Further, I have
shown that the degree of transposition of the testes corresponds
Text-fig. 57.
CLUPEA
SCYLLIUM
TRITON
LACERTACCLUMBA
Diagram illustrating Testis Descent.
This diagram, based on a series of measurements, illustrates the positions
and lengths of the testes relatively to the length of the trunk proper (minus tail) in
Clupea, Scyllium, Triton, Rana, Lacerta (muralis), Columba, the common mouse,
and the cat, all measurements being reduced to one scale. The vertical sides of
the rectangle represent the length of the animal to which the other measurements
are proportional, and the dots the approximate positions of the centres of gravity of
the testes. Such a figure well exhibits the concentration and transposition corre-
sponding to higher grades of impulsiveness. To fully illustrate the relationship a
similar diagram drawn up from several hundred dissections is necessary, together
with the average weight of the testis and intensity of the reactions incurred in
locomotion, appended in the case of each animal.
1903. ] TRANSPOSITION OF MAMMALIAN TESTES. 335
to the intensity of the forces concerned, and have thus supplied
the inductive evidence required to verify the conclusion arrived
at on & priort grounds. It has also been pointed out that
transposition of the testes can, from the nature of the function
of these organs, the mode of emission of their genital products,
and the absence of adverse external influences, neither cause
derangements in the individual economy nor impair the fertility
of the race ; ; and that in consequence of a similar change of
position in the case of other organs inevitably entailing one or
other of these penalties, the testes alone have descended. More-
over, the superior density, the disposition of the attachment, and
the appropriateness both as regards volume and definiteness and
concentration of form for change of position, appertaining to the
testes, lend greater additional facilities to their transposition than
in the case of other organs not so characterised. Hence we can
not only show why the testes have descended, but also why they
alone have thus responded to the incident forces—other or eans,
for one or more of the reasons supplied, requiring to be m: aintained
in position by the special retentive structures developed to that
end. Combining with these facts the additional evidence derived
from a study of mammalian anatomy, showing that impulsive
locomotion is not alone responsible for the descent of the testes,
but that it has also been the prime cause of the evolution of the
diaphragm, the metanephric kidney, and other minor anatomical
features, we see still more clearly “ how strong is the probability
of the theory advanced.”
Other Illustrations of the Relationship between Visceral Con-
formation and Impulsiveness : the Evolution of the Diaphragm
and the Metanephric Kidney.
Although differmg from the reproductive elements of the body
as regards their economic relations to the rest of the organism,
the non-reproductive organs, possessing like material properties,
may be expected to portray similar structural characteristics in
relation to locomotor impulsiveness. A consideration of such will
not only prove of utility as affording indirect confirmation of
the foregoing, but will provide opportunity for tendering similar
explanations in the case of other anatomical features. The con-
stant interrelations of position obtaining between the various
organs in the vertebrate body are in every case easily accounted
for, either on mechanical or physiological grounds, and it is
needless to discuss every organ from these points of view; it will
suffice if we treat of two or three structural phenomena, the
interpretation of which is not quite so self-evident or well
known.
The complete descent of the testes and the presence of a fully-
developed diaphragm both being common characteristics of the
Mammalia, it is possible that the cause to which the former is
attributable may also suffice to explain the evolution of the latter,
Proc. Zoot. Soc.—1903, Vor, I. No. XXII. 22
336 MR. W. WOODLAND ON THE [ Apr. 21,
and enquiry confirms the supposition. Though, in the above
account of mammalian locomotion, most stress has been laid upon
the impulsive elevations of the body and resulting strains on the
attachments of organs, yet it must not on that account be inferred
that on descent of either half of the trunk the shock consequent
on contact of the limbs with the earth (equal in intensity to the
elevatory impulse) is negligible. (See above in case of Kangaroo
in which it is taken into account.) Such shocks have the effect
of causing those viscera which are closely adherent to the mass of
the body either to exert a considerable pressure on structures
anterior to them or to be dislodged from theix normal position ;
and if in either case such a result is to be avoided, organs of
support must be developed. In mammals, the lungs together with
the heart occupy the anterior portion of the ccelomic cavity, and
behind these are situated the liver, stomach, and intestines, these
together constituting a considerable mass. These massive organs,
unless prevented, would on each contact of the limbs with the
earth exert great pressure on the fragile compressible lungs
immediately anterior to them. Hence, in order to obviate
ensuing derangements, we find in all mammals, and in many other
terrestrial animals, a stout partition separating the cavity of the
liver and gut from the cavity of the lungs—a structure which can
only have been originated by natural selection. Evidence sup-
porting the conclusion that the diaphragm arose as an adaptation
to the forward pressure of the liver and gastric mass, is not only to
be found in the fact that the diaphragm is convex anteriorly, but
also in that the convexity is, cewteris paribus, proportional to the
impulsiveness of the animal’s activity and to the mass of the
liver and gut. In herbivores the mass of the gut is greater than
in carnivores; on the other hand, ‘‘ carnivorous (fat-eating) animals
generally possess a larger liver than herbivores” (Wredersheim),
so that these respective characters tend to defeat comparison *.
But in the Perissodactyla we find a fairly large liver (considerably
larger than in Artiodactyla), a long herbivoran gut, and a high
degree of impulsiveness, the combination of which on our hypo-
thesis should be correlated with a highly convex diaphragm—an
inference which proves correct. Owen remarks that “in the
perissodactyle Ungulates, in which the movable ribs are numerous
and continued to near the pelvis, the diaphragm is also extensive
and much arched towards the thorax ”—indicating that the con-
vexity of the diaphragm is markedly above the normal. And,
doubtless, other illustrations are adducible. Additional evidence
as to the primary function of the diaphragm is, moreover, afforded
by the case of the Struthiones (and allies) which have adopted
the mammalian mode of locomotion. In these birds there exists
a “ well-marked diaphragm forming a partition which divides the
thoracic cavity into two parts, one posterior and small containing
the lungs, and the other anterior and large containing the heart
1 The small capacity of the herbivoran thoracic cavity as compared with that of
the carnivora must be remembered in this connection.
1903.] TRANSPOSITION OF MAMMALIAN TESTES. 337
and liver. It is a fibrous membrane, concave forwards, with a
muscular attachment at either side to the ribs and intercostal
tissues, which it joins in about the middle of their course......
The pleural cavity is closed above and below by the fibrous dia-
phragm becoming blended with the first and last ribs. The
anterior thoracic cavity, which contains the pericardium-coated
heart in its upper part, entirely independent of the pleural
cavity, is divided into two by a dense fibrous membrane
which springs from two vertebral crura, much as the human
diaphragm, and extends above the line to join the sternum along
the border which articulates with the ribs, leaving the heart entirely
in front of it; its concavity is directed downwards and forwards,
and it is separated from the diaphragm proper by very large air-cells,
The liver is completely separated from the abdominal cavity by a
fibrous membrane, so that when the included viscera are removed,
it is not at all brought into view. The mesentery is very dense
and strong” (Garrod & Darwin). A comparison of the struthious
and mammalian diaphragms affords further confirmation. In the
Ostrich, owing to the anomalous position of the avian lungs (their
close application to the costal skeleton ensuring the intactness of
the air-cell connections), the liver has practically assumed their
role, as regards conformation, in relation to the heart and dia-
phragm. Hence the diaphragm is concave anteriorly, consequent
on the necessity for the retention of the liver. In the mammal,
on the other hand, the lungs are of large volume and freely sus-
pended in the ventral portion of the thorax, enveloping the heart.
If the liver were anterior to the diaphragm it would, as before
remarked, during locomotion periodically exert great pressure on
the lungs ; hence the diaphragm here becomes the forward sup-
port of the liver and other alimentary viscera. The envelopment
of the heart by the liver in the Ostrich possibly affords an illus-
tration of that alternative to the formation of a diaphragm above
referred to.
From these and other considerations, we can more precisely
interpret the anterior convexity of the diaphragm, which is due
to two causes: the forward pressure in the median line of the
large abdominal viscera, and the backward lateral extension of
the capacious lungs. Evidence as to the cooperation of this last
factor is afforded by such cases as that of the Manatee, in which,
on account of its subaqueous habits, unusually developed organs
of respiration are required, it also being essential that their ex-
tension should be dorsal in order to ensure the ventral situation
of the centre of gravity of the body. Here, as Mivart remarks,
the diaphragm is so oblique that the thorax ‘“ extends backwards
above the whole length of the abdominal viscera”; and a similar
conformation is found in Cetacea and perhaps other groups. In
development, the diaphragm is ‘formed from a couple of septa,
dorsal and ventral respectively, which arise independently, and
are for some time quite distinct from each other” (Jarshall) :
from which fact it follows that the part the diaphragm plays in
20%
338 MR. W. WOODLAND ON THE [ Apr. 21,
connection with respiration cannot have constituted its primitive
raison a étre.
Thus, observing the necessity for the development of the dia-
phragm, we find that its variations in disposition and contour
found in the Struthiones and Mammalian orders fully confirm
the conclusion deduced on a@ priori grounds.
Another organ which, though not peculiar to the Mammalia,
is yet a diagnostic feature of terrestrial vertebrates, is the meta-
nephric kidney, and that there is possibly a relationship between
terrestrial modes of lecomotion and the evolution of this organ
T will now endeavour to show.
It is well established that the metanephros is a development
of the definitive mesonephros which has lost its nephrostomata,
acquired a separate duct, and become more or less concentrated
in form. It has been pointed out above that concentration
of structure is an essential concomitant of that increase of
impulsiveness which results from the increased activity of the
animal under conditions which involve marked reactions between
the body and the medium or substratum. That this is so in the
case of the kidney, it is only necessary to compare the elongated
mesonephric bodies of Pisces and aquatic Urodeles with the same
organs of the terrestrial Amphibia and the metanephric bodies
ot Reptilia, and again to compare these latter with their repre-
sentatives in Mammalia, in which both concentration of structure
and impulsiveness of locomotion attain their maxima. With
regard to the loss of nephrostomata, the same relation holds.
In all Pisces, with the exception of certain Elasmobranchii, the
mesonephric bodies retain connection with the ccelom by means
of the nephrostomial tubules. Whether the absence of these
fragile structures in Elasmobranchs (their function with regard
to the celom perhaps being assumed by the abdominal pores) is
to be attributed to the fact that these fishes either are or are not
descended from the most active members of their class (and it
must be remembered that they are surface forms) is not certain,
but it is possible. In the aquatic Urodeles the nephrostomes are
present, but in the terrestrial Anura they have completely lost
their connection with the celom (which latter, as in Elasmo-
branchs, has had to discover another means of exit for its waste
products). The causal relation between disruption of the nephro-
stomata and the adoption of a terrestrial life is here clearly shown
by the ontogeny. In the development of the frog, the meso-
nephros is at first in communication with the coclom: by means of
the nephrostomata, but at the period of metamorphosis this con-
nection is severed, and the kidney, becoming more concentrated,
finally assumes the definite form of the adult : structure. Needless
to say, the metanephros of Reptilia, Aves, and Mammalia is
totally devoid of nephrostomes, that of the last not possessing
them at any stage of development. The division of the meso-
nephros into two portions, one coming into relation with the
testes, and the other—the definitive mesonephros—yretaining its
1903. ] TRANSPOSITION OF MAMMALIAN TESTES. 339
urinary function, is evidently a physiological phenomenon, though
the ultimate structural separation of the two parts must largely
be attributed to the influence of greater activity of the organism.
Also the secondary development of the definitive mesonephros is
indirectly due to increased activity, since the work of the excretion
resulting from the latter is largely increased. In fact, the whole
ontogeny of the kidney—the successive developments of the pro-,
meso-, and metanephros in space and time, the differences obtaining
between the pro- and mesonephric tubules, the disappearance of
the pronephros, &c., &e.—can be directly or indirectly attributed
to the increasing activity of the organism in the course of phylo-
geny, as a little thought will show.
With regard to the position of the kidney—a body of consider-
able mass, and in Amniota similarly disposed to the testes—no
relation between such and the impulsiveness of the animal being
traceable, it is evident that some other factor must be involved,
and enquiry shows that the localisation of the kidney is correlated
with the total activity of the animal. Excretion heing facilitated
by proximity to a vigorous plood-supply, we find that in active
animals the kidney tends to be situated anteriorly, i. e. nearer the
heart ; on the other hand, close connection with the vent to the
exterior is similarly advantageous, and in inactive animals, such
as Reptilia and Urodeles, we find the kidneys situated at the
posterior extremity of the ccelom.
The kidneys of mammals are retained in position by a circwm-
ambient development of areolar tissue, which usually contains
much fat; support is also contributed by the “tonicity” of the
muscles of the abdominal wall and by the terminal ribs.
Tn conclusion, I must acknowledge my indebtedness for details
vespecting the habits and anatomy of the mammalian orders to
the following authors and works: Flower and Lydekker, Owen,
Wiedersheim, Beddard and Gegenbaur, Lydekker’s ‘ Royal
Natural History,’ ‘Mammals’ by Vogt & Specht, Schmeil’s ‘ Text-
book of Zoology,’ and Allen’s & Lloyd’s editions of Jardine’s
‘ Naturalist’s Library.’
L also wish to thank Prof. Minchin, Mr. J. T. Cunningham, and
Mr. H. S. Shelton for kindly reading through the manuscript and
for making several suggestions and criticisms.
APPENDIX.
In respect to the non-descent of the ovary in the Mammalia, it
is as well to emphasise one factor ensuring its retention. Assum-
ing that the females of mammals are approximately as active as
the males, it is evident that if the mammalian ovum had retained
its yolk (7. e. if the mammalian ovary possessed the size and mass
of that of the Sauropsida and Monotremes), the retention of the
ovary within the body-cavity by the development of special liga-
ments, &c., would have been, if not impossible, yet extremely
hazardous. And such being the case, it would obviously benefit
the race if the potency of one of these two factors were decreased.
340 MR. R. I. POCOCK ON THE [ Apr. 21,
Now, under the conditions of mammalian life—conditions in which
survival of the fittest attains within its limited sphere of operation
a maximum degree of efficiency—it is obvious that diminution of
activity would be fatal, the speed of mammals being one of the
most important conditions to survival in the struggle for existence
(as is shown by the fact that this trait is so highly developed in
these animals) ; hence, a less degree of activity bemg prohibited,
any decrease in the mass of the ovary would be of service to the
organism in which it occurred. Natural selection may legitimately
be supposed to operate here, since, although it has been clearly
demonstrated that the higher the life of the organism the less
range of application does this principle possess, yet it doubtless
applies in the case of any feature which is of paramount importance,
and modifications concerned with the genital structures must
necessarily possess such importance. It is therefore possible, and
even probable, that the loss of yolk suffered by the mammalian
ovary and the alternative adoption of a placental mode of nutrition
both indirectly result from that same cause of impulsive locomotion
to which we have traced several other features of mammalian
structure.
It may also be worth while to add that many minor features of
mammalian anatomy, the significance of which is usually over-
looked, are only explicable on the assumption that they are related
to impulsive locomotion. Instances of these minor structures are :
the accumulation of fat at the base of the heart, the fatty cushion
surrounding the neck of the bladder, the fatty development about
the kidney already noticed, and the various ‘“ suspensory ligaments ”
and other “ fixative organs” referred to above, associated with the
stomach, liver, and other viscera of large mass.
d. On the Geographical Distribution of Spiders of the
Order Mygalomorphe. By R. I. Pococxr, F.Z.S.
[ Received March 17, 1903.]
(Text-figures 58-61.)
Parr [.
InrRODUCTORY REMARKS UPON THE PALHZONTOLOGY AND THE
MEANS OF DISPERSAL OF SPIDERS.
(a) Summary of the Paleontological History of Spiders, and
its bearing on the Phenomena of Distribution.
Owing to the enormous chances against the preservation of
fossil Spiders in sedimentary rocks, the paleontological history
of this Order is very imperfect. One or two types have been
diseovered in Carboniferous strata of Europe and North America
(Arthrolycosa and Protolycosa), and also a fairly large number of
specimens from amber and from gypsum and lacustrine deposits
of Oligocene and Miocene age in those countries. But absolutely
1903.] GEOGRAPHICAL DISTRIBUTION OF SPIDERS. 341
nothing is known of the forms that inhabited the world during
the enormous lapse of time represented by the Mesozoic strata,
and nothing except inferentially of the types that occupied the
southern countries of the world during Tertiary times.
Spiders of the type that lived in the Carboniferous period
succeeded in holding their own in Europe until the Oligocene, and
are represented at the present time by the genus Liphistius, which
is restricted to the Indo-Malayan area of the Oriental Region.
Apart from the genus Liphistius, all existing Spiders, including
the Mygalomor phe, belong to the group Opisthothele. There is no
evidence that this group existed in the Carboniferous period ; but
since most of the Oligocene and Miocene fossils belong to existing
families, or sometimes indeed to existing genera, It is permissible
to suppose that the Opisthothele originated some time during the
Mesozoie epoch, and may, in fact, be. coeval with the mammalia.
Whether any of these hypothetical Mesozoic forms survive to the
present day, it is quite impossible to say. All that paleontology
allows us to infer is that during the Tertiary period there was
a rich and varied spider -population spread over the Northern
hemisphere, contaiing forms that have undergone but little
metamorphosis since that date. The existence of ‘Mygalomor phe
at that time is attested by the discovery of one form referred to
Mygale in the gypsum-beds at Aix, and of another, Hoatypus,
in the Eocene strata at Garnet Bay in the Isle of Wight. But
since it is impossible to classify these forms with an approach
to certainty in any of the existing families, their only value
from the geographical standpoint is the evidence they supply
that the Mygalomorphe had come into being in Tertiary times, and
were living in the Northern hemisphere.
The imperfections in our knowledge above alluded to permit
only a provisional acceptance of the theories put forward in the
following pages to explain the distributional phenomena of the
Mygalomorphe. But all the available evidence, little enough
though it be, points to the conclusion that the Mygalomorphe
and the rest of the Opisthothele appeared first in the Northern
hemisphere, and spread thence over the southern countries of
the world,
(6) Means of Dispersal of Spiders, and the importance of the
Mygalomorphe from the Geographical standpoint.
It cannot be claimed that Spiders as a whole are a favourable
group to study from a geographical point of view; for, although
exclusiv ely terrestrial when adult, and, like other flightless animals,
dependent upon continuity of land-surfaces for migration, a great
many species are known to have the power, and the instinct to
put it in foree, of dispersing themselves over wide areas by
practising when young the habit of flight, using silk-threads as
aerial floats upon which they may be carried long distances before
the wind. This phenomenon is well known, and has given rise to
342 MR. R. I. POCOCK ON THE [ Apr. 21,
the belief in the existence of a ‘ gossamer’ spider which is supposed
to be the cause of the fine threads which fall from the air and
carpet the fields with silk at certain times of the year. It is now
known that the ‘gossamer’ spider is a mythical species, and that
species of the most diverse habits belonging to widely different
families are responsible for the floatmg threads. The habit is
practised alike, and, so far as is known, to an equal extent, by
snare-spinning forms belonging to the Argiopidee and Theridiide,
by hunting-spiders like the Lycoside and Attide, or by sedentary
species that lurk in flowers, like the Thomiside.
That this method of locomotion may considerably influence the
distribution of spiders may be inferred from the fact that cobwebs
thrown out in this way, and affording support to little spiders,
have been found at the tops of our highest buildings, and have
become entangled in the rigging of ships 200 miles from land.
There are reasons for thinking, however, that the habit 1s for
the most part restricted to phanerozoic diurnal species, namely,
those that hunt their prey or spin their webs in the open; and
that cryptozoic forms, that live in burrows or under stones or logs
of wood, and that are for the most part nocturnal, do not indulge
in it’.
Clearly, therefore, these eryptozoic groups, in which the restric-
tions to dispersal are presumably the same as in other terrestrial
animals which can neither fly nor swim to any distance, have more
value for the establishment of geographical areas than those species
with powers of dispersal analogous to flight.
Owing to the relatively large size and great weight of the newly
hatched | young of the Mygalomory phee, coupled with the reduction
in the number of spinning- appendages and the greater simplicity
of the silk-glands, it seems probable that aer ial sailing is not
practised to any great extent by the members of this: suborder 2.
Especially true will this be of the Aviculartide, a family which
contains the largest spiders known of this or any other epoch,
with newly-born young rivalling or excelling in size the adults of
many species of the Arachnomorphe.
Consideration of these facts, coupled with the impossibility of
dealing in detail, in one paper, with the distribution of all the
genera of the Aranez, has led to the selection of the Mygalomorphee
as the fittest group to illustrate the geographical distribution of
Spiders in general.
1 Simon states that the Spider-fauna of the Sandwich Islands is composed wholly
of species of the former category, with the exception of some few forms which appear
to owe their presence in that Archipelago to human ageney (‘Fauna Hawaiiensis,’
sranee; 1902).
2 The young of the only known British representative of this group, namely
Atypus, one of the smallest types of Mygalomorphe, have been seen to scatter over
small areas by this method of tray elling (EB. BENE Tr. Ent. Soc. 1885).
In this connection it is instructive to remark that Atypus has a wider distribution
than any other known genus of the suborder, ranging from Ireland and Algeria to
Japan and over the Hastern (? the Western) ‘States of North America, that is to say
ene the Northern hemisphere from the eastern to the western shores of the
tlantic.
1903.] GEOGRAPHICAL DISTRIBUTION OF SPIDERS. 343
Parr II.
DISTRIBUTION OF THE FAMILIES, SUBFAMILIES, AND GENERA OF
MycGatomorpPu a, and the evidence thus supplied as to their
Original Habitat and the Lines of Migration followed in
Dispersal.
Family DipLurip#.
The Dipluride are the most widely distributed of all groups of
Mygalomorphe, being found practically all over the world to the
south of about the 40th parallel of north latitude. The numerous
groups, however, into which the genera fall present some features
in their geographical range of considerable interest.
1. Subfamily Dreturin2.—The genus Drachythele’ is met with
in the Mediterranean Region, Central Asia, and the Southern
States of North America. Nearly allied to it are Hapalothele from
Madagascar; Brachytheliscus from Natal; dAname, Txamatus,
Chenistonia, and Dekana from Australia and Tasmania; /ufius
from Central and South America; Z’rissothele from Chili; Lycinus
from the Argentine. South America is also the home of Z7rechona,
Diplura, Uruchus, Harmonicon, and Melodeus.
2. Subfamily Macrornerin2.—This subfamily is divisible into
four groups.
(a) The Macrothele :—JMJacrothele occurs in Spain, China,
Burma, Singapore, and Java. Nearly related are Phyxioschema
from ‘Transcaspia, Stenygrocercus trom New Caledonia and
Queensland, and Porrhothele from New Zealand. /schnothele
is represented by species from India, Madagascar, 8. & W. Africa,
and Central and South America; and Hvagrus is known from
S. Africa and Central America, whence it extends into the
Southern States of North America (Idaho).
(b) The Hexathele contain two genera—Hewxathele from New
Zealand, and Scotinecus from Chili.
(c) The Atraces contain the genera dAtrax and Hadronyche, from
Eastern Australia.
(dz) The Masteriz, comprising Accola and Masteria, which are
probably identical, occur in Venezuela, the Philippine Islands, and
Upolu. :
The presence of Drachythele in the Mediterranean and Sonoran
areas, and of nearly allied forms in South Africa, Madagascar, and
all over South America, suggests immigration from the north
into these countries of the Southern hemisphere. On the other
hand, the entire absence of related types from the area lying
between and including India and Austro-Malaysia, and the
reappearance in Australia of genera closely allied to Brachythele
1 This genus has also been recorded from S. Africa, Madagascar, 5. America, and
Australia. It is probable, however, that the species referred to it belong to one or
other of the allied genera from these areas.
[ Apr. 21,
MR. R. I. POCOCK ON THE
344
ay Iouay ‘sooDdap ‘aLdagsnyy ‘wjayjJ0uon AT
pace pos St SS ES) rE
‘opiimnjdyuy JO eIouNs suUT[EYJOIOV_Y, 94} Jo UOTyNqIAASIG: TworyderSoey oy} oyvaysnTI 04 dey
"BG “SH-4xo],
1903. ] GEOGRAPHICAL DISTRIBUTION OF SPIDERS. 345
as well as to the South-American and Afro-Mascarene forms, points
equally forcibly to the peopling of Australia from either one or
the other, or perhaps both, of the southern continents just
mentioned,
The remaining genera of Diplurine, namely, 7’rechona, Diplura,
Harmonicon, Melodeus, and Uruchus, all of which are more
specialised types than Brachythele and its allies, probably arose
within their present area of distribution.
A great contrast to the distribution of the Diplurine is presented
by that of the Macrotheline.
The occurrence of Macrothele in Spain, China, Burma and Java;
of Porrhothele, which is scarcely separable from it generically, in
New Zealand ; of Phyxioschema in the Transcaspian area; and of
its near ally, Stenygrocercus, in Queensland and New Caledonia,
suggests a southward migration of these types from the northern
provinces of the Old World into Australia and New Zealand by
way of China and Indo-Malaysia. Similarly /schnothele, a more
specialised type than Jacrothele, perhaps descended from the north
by way of India into Madagascar, South Africa, and crossed
thence into South America, where with Hvagrus, which is also
represented in South Africa, it is the only representative of this
group of Dipluride. The entire absence from the Sonoran Region
of forms related to Maecrothele, Hvagrus, and Ischnothele, is opposed
to the supposition that the two last-mentioned genera had a
northern origin in America.
The Masteriz appear to be degenerate forms of the Macrothele.
They are the smallest of all known Mygalomorphe, and are
essentially eryptozoic or lucifugous, living in caverns or under old
decaying vegetation in the dark, damp forests. Hence we can
only pretend to a partial knowledge of then distribution, and it
would be rash to draw deductions from the fact of their having
been discovered hitherto only in Venezuela, the Philippines, and
Upolu.
The two genera of Atraces, Atrax and Hadronyche, confined to
Australia, appear to be Macrothelinz specialised for a fossorial
life, with which is correlated certain features imparting to them a
superficial similarity to the Ctenizide.
The Hexathele, resembling the Macrothele except in the
retention of an additional pair r of spinning-maimille, undoubtedly
a primitive feature, are confined, so far as is known, to New
Zealand and Chili. There seems no reason to doubt that they
passed from one of these countries to the other by a southern
Jand-connection.
Family PARATROPID®
This family, specialised both in structure and habits, is repre-
sented by three genera, Paratropis, Anisaspis, and Anisaspoides,
confined to the Neotropical Region. Its affinities are doubtful,
but some primitive genus of Dipluride allied to Brachythele must
346 MR. R. I. POCOCK ON THE [Apr. 21,
probably be looked to for its ancestry. There seems no reason to
doubt that it originated in the area it now occupies.
Families Arypip#, BRACHYBOTHRIIDH, MECICOBOTHRIIDA.
The genera of Atypide, two in number, scarcely pass south of
the Equator. Atypus, occurring in the Mediterranean Region,
and spreading northwards into Central Europe, beyond the 50th
parallel of north latitude, is the most northern type of the Mygalo-
morphe. It is also met with in Japan, Burma and Java, and in
North America, where it ranges to the east of the Mississippi
from Wisconsin (45th parallel of latitude) to Florida. Calommata
is more restricted and more southern in range. It is confined to
the Old World, and has been recorded from Japan, Siam, Burma,
Sumatra, Java, and the Camaroon area of tropical West Africa.
The Brachybothriide contain the genera Acattyma from Japan,
and Brachybothriwm and Atypoides from North America, the latter
from California, the former from British Columbia, North Carolina,
Texas, &e. ;
The Mecicobothriidee comprise two genera—Hexura from the
two north-western States of North America (Washington and
Oregon); and MMecicobothrium from the Argentine.
Atypus seems to be a genus which, like Puchylomerus and others,
extended in Tertiary times sufticiently far to the north to avail
itself of the land-connection that then existed across the area
now covered by the Behring Sea. Its disappearance from the
countries to the north of its present distributional area must be
assigned to the refrigeration of this region of the globe with the
advent of glacial conditions.
The distribution of Calommata in the eastern part of the
Oriental Region and in Tropical West Africa has many parallels
amongst the mammalia, and points to the former extension of the
genus across an intervening forest-clad tract, and its survival in
districts where the conditions remained favourable to its existence.
The explanation given of the distribution of Atypus applies
equally to the Brachybothriide, except that Atypoides has no
representative in the Old World. Srachybothrium extends in
Western America northwards to Queen Charlotte Island, that is
to say to the latitude of the Aleutian Islands and the Alaska
Peninsula. Hence its similarity to, perhaps identity with, the
Japanese Acattyma is no matter for surprise.
The apparently discontinuous distribution of the Mecico-
bothriide is very remarkable; but since the only known example
of Mecicobothriwm measures only 6 mm. in total length, it is
highly probable that the existence of the genus in South America
to the north of the Argentine has been overlooked. Considering
the close structural similarity between Heauwra and Mecicobothrium,
and the admitted relationship between these two genera and the
Brachybothriide and Atypide, both northern groups, it seems
d47
GEOGRAPHICAL DISTRIBUTION OF SPIDERS
1903.]
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348 MR. R. I. POCOCK ON THE [ Apr. 21,
probable that the Mecicobothriide first appeared in North America
and penetrated thence into South America in the latter half of
the Tertiary epoch.
The three families just discussed form an ascending series in
specialisation, starting with Hexwra, which leads from the
Dipluride, passing through the Brachybothriide, thence to Atypus,
and culminating with Calommata, the most specialised of all.
Assuming, moreover, that dAcattyma and brachybothrium are
identical, it is noticeable that of the six known genera, three are
eculiar to the Western (American) hemisphere, two are common
to both the Eastern and Western, and one only peculiar to the
Eastern, namely Calommata. In other words, the most primitive
genera are exclusively American, one that occupies an intermediate
position (Acattyma) extends only as far as Japan, and the two
genera of the most specialised Atypide extend from east to west
throughout the Hastern hemisphere, and one of them, Atypus, to
the extreme east of the Western hemisphere.
Since we should expect to find the primitive types of a group
persisting in the area of its origin and the specialised forms in
districts remote therefrom, the modifications being brought about
by the varying conditions of existence incident to wandering,
North America may be looked upon as the probable home of this
section of the Mygalomorphe.
If this be the explanation of their distribution, it presents
analogies with that of the Camelidee amongst the mammalia, which
originated in the Sonoran area of America, and thence extended
into South America and into Central Asia by way of Alaska and
Kamchatka.
Family CyRTAUCHENIIDA.
The genera of Cyrtaucheniide are exceedingly numerous, and
rival the Dipluride in extent of distribution ; but, owing to the
complex and puzzling nature of their relationships, it is not easy
to deduce any satisfactory conclusions from their distribution.
They are referable to the following sections, which are arranged
as nearly as possible according to their lines of descent, starting
with the most primitive, z.e. the Nemesiz, which have departed
least from the Diplurid type.
1. Nemesie.—The genus Wemesia itself is abundant in the
Mediterranean Region, and is met with also in China; Anemesia
oceurs in Afghanistan, Vemesiellus in 8. India, and Scalidognathus
in Ceylon. Hermacha, Hermachastes, Lepthereus, Pionothele, and
Spiroctenus inhabit 8. Africa; Genysa, Madagascar; Arbanitis,
Australia, Tasmania, and New Zealand; Cantuaria, New Zealand ;
Rhachias, Carteria, and Hermacha (the last also in 8. Africa) in
South and Central America.
2. Cyrtauchenii, differing from Nemesixe in the procurvature
of the fovea.—Atmetochilus and Damarchus occur in Burma and
Sumatra; Cyrtauchenius and Amblyocarenum in the Mediter-
ranean, the latter also in California ; Pelmatorycter in KH. and 8.
1903.] GEOGRAPHICAL DISTRIBUTION OF SPIDERS. 349
Africa; Mierobatesia in tropical W. Africa; Stictogaster, Dessia,
and Homostola in 8. Africa; Stenoterommata, Pselligmus, Neo-
cteniza, Eucteniza, and Enrico in South and Central America ;
Aptostichus and Actinowia in California ; and Myrmeciaphila in
Virginia and Carolina.
Bn Aganippe, differing from the preceding in the specialised
arrangement of the eyes.—dAganippe, Blakistonia, Anidiops,
Idiosoma—all confined to Australia.
4, Aporoptychi, differing from the Cyrtauchenii in the
enlargement of the labium and maxillee.—Aporoptychus and Lolo-
stromus occur in tropical West Africa and South America ;
Phrissecia, Pheoclita, Celidotopus, Phenothele, and Rhytidicolus
in 8. America.
Of the above mentioned groups, that of the Nemesiz is the
only one that is represented at the present time in India, Mada-
gascar, and New Zealand—a fact in keeping with its primitive
status, and suggestive of an earlier migration into the Southern
hemispheres. “Singularly enough, the group is unrepresented in
North America. This and the fur ther facts that South Africa is
the richest of the regions in number of genera, and that one of its
genera, Hermacha, occurs also in Brazil, make it almost impossible
to doubt that the group entered South America from South Africa.
Again, since the group is also apparently absent from the whole
of the area lying between India and Australia, we must look to
South Africa, Madagascar, or South America as the source whence
it entered Australia ; and since the Mascarene genus Genysa is
said to be nearly allied to the Australian Arbanitis, Madagascar
and §. Africa were perhaps the feeders to the Australian area.
Since the Cyrtauchenii are specialised allies of the Nemesiz,
and therefore later developed forms, it is interesting to note their
apparent absence from Madagascar, India, Australia, and New
Zealand, which indicates a later southern migration from the
north. The following hypothesis seems to explain the facts of their
distribution. In early Tertiary times the group was continuously
distributed throughout Europe, Asia, and North America. In
Eastern Asia it descended a short distance into Indo-Malaya after
the severance of Australia, and penetrated Africa after the forma
tion of the Mozambique Channel. Similarly from the Sonoran
Region it passed into Central and South America, after the
Pliocene union of the latter with North America. Whether any
of the South-American fauna was derived from Africa or vice
versa, there is no reliable evidence to show.
If the Aganippe entered Australia from South-eastern Asia,
it is strange that no related forms have been discovered in Austro-
Malaysia. They may have come from South America or South
Africa; but I am disposed to think that they have had an origin
independent of the Cyrtauchenii in Australia from the Nemesiz,
from which they differ practically only in the procurvature of the
fovea—a feature which is known to have arisen more than once
390 MR. R. I. POCOCK ON THE [ Apr. 21,
within the limits of the Mygalomorphe. This conclusion as to
their descent is supported by the fact that the specialised arrange-
ment of the eyes is foreshadowed in Dyareyops, one of the
Nemesize, and by the presence of a single row of ungual pectinations
in the Aganippz and all the Australian Nemesize, two such rows
being characteristic at least of the southern forms of the Cyrt-
auchenil.
The Aporoptychi are specialised Cyrtauchenii. Their distri-
bution suggests that they originated in South America or South
Africa, and crossed from the one continent to the other when the
two were united. This conclusion is strongly supported by the
occurrence of two of the genera in both of the areas in question.
Family Crenizip&.
The genera of the subfamily Ctenizine are almost exclusively
northern. Cteniza, Hpycephalus, and Cyrtocarenwm are restricted
to the Mediterranean; Sterrhochrotus occurs in Turkestan ;
Latouchia 11 China and the Himalayas; Pachylomerus in Spain,
Algeria, Japan, N., Central, and 8. America, and the West Indies ;
Bothriocyrtwm in California; Conothele ranges from Burma to the
Solomon Islands. The only genus which is exclusively southern
is Stasimopus, which is confined to 8. Africa.
The structural features of this group suggest that it is a
specialised offshoot from some northern forms of the Cyrtau-
cheniide. This conclusion is borne out by the more restricted
distribution of the genera, which, in virtue of their later appearance
in the Northern hemisphere, have had less time to distribute
themselves over the southern continents. The only genus which
has entered the Ethiopian Region, namely Stasimopus, appears to
have done so not earlier than the Pliocene, since it is seemingly not
represented in Madagascar. Conothele, too, seems to have
migrated southwards by way of Burma to the Solomon Islands,
after the separation of Australia; Pachylomerus, the only form
which enters the Neotropical Region, appears almost certainly to
be a northern immigrant from the Sonoran region,
The Idiopine section, apparently a specialised offshoot of the
Ctenizine, has, on the contrary, a more southern distribution
than the typical Ctenizine. The genus /diops (Acanthodon) itself
has representatives in Central Asia, Syria, Arabia, India and
Burma, Tropical and South Africa, and 8. America ; Heligmomerus
occurs in India, Ceylon, and Tropical Africa; Gorgyrella in
S. Africa; while the aberrant Pseudidiops is confined to the
forests of South America.
The presence of the genus /diops so far north as Central Asia
and Syria, and in India, Burma, Tropical and Southern Africa,
and Brazil, attesting as it does considerable powers of adaptation
to varied climatic and other physical conditions, justifies the
supposition that the genus never formed part of the Sonoran
fauna of America. Otherwise it would be difficult to account for
GEOGRAPHICAL DISTRIBUTION OF SPIDERS.
1903.]
‘mph ‘purdompT "DUIzwUagd
Wiiia SS NMI,
‘apuhyy pur apizruay oy} Jo uoynqiaystq peorydeasooy oy} oywaysnqr 03 depy
NH WII li ;
‘09 ‘SY-9X0],
1893, Vou. I, No. XXIII.
23
Proc. Zoon. Soc.
352 MR. R. I. POCOCK ON THE [ Apr. 21,
its absence at the present time from this region, as well as from
the West Indies and Central America. These considerations
justify the view that South America acquired this element in its
fauna, not from North America but from Africa. But by
whatever route Zdiops reached South America, it is probable that
Pseudidiops, a specialised offshoot of that genus, arose within the
Neotropical Region.
Family HaLonoprocrip#.
The three known genera of this family, a specialised offshoot of
the Ctenizide, namely, Halonoproctus from China, Chorizops and
Cyclocosmia from the Sonoran Region, also constitute a northern
group, which has not migrated into the southern continents.
Family AcTINOPODIDA.
Of the two genera of this family, Hriodon (Missulena) is restricted
to Australia, and Actinopus to the Neotropical Region.
The descent of this family is doubtful, but its structure and
distribution suggest its origin in the Southern hemisphere from
the Cyrtaucheniidee of the group Aporoptychi, which exist in
South America and South Africa. Its present distribution may be
accounted for on the hypothesis of the migration of the ancestral
form either from Africa to Australia and thence to South America,
or the other way about.
Family Micip#.
The Migidee are essentially a southern group. Two or three
genera (Myrtale and Thyropeus) occur in Madagascar; Moggridgea
in Sokotra, Nyasaland, Natal, and Cape Colony ; Heteromigas in
Tasmania; J/igas, nearly related to Moggridgea, in New Zealand ;
and an undescribed genus, close to Heteromigas, in Chili.
The recurvature of the thoracic fovea in this family suggests
its descent from Cyrtaucheniide of the section Nemesie, an
ancient and very widely distributed group, with representatives
in South Africa, Madagascar, South America, Australia, and
New Zealand, the countries to which the Migid are now restricted.
The distribution of the latter attests not only a southern origin,
perhaps in the Afro-Mascarene area, but also migration thence
to Australia and New Zealand, and from Australia to South
America. s
Family BARYCHELIDA.
The genera of this family fall into three sections, which may be
arranged in order of specialisation as follows :—
a. Barycheli: Leptopelma, Mediterranean Region ; A trophothele,
Sokotra; Hubrachythele,Somaliland; Cyphonisia, W. Africa;
Pisenor, hi, Africa; Brachionopus, Cape Colony ; Tigidia,
1903.] GEOGRAPHICAL DISTRIBUTION OF SPIDERS. 353
Mauritius ; Sasonichus, 8. India; Sipalolasma and Plagio-
bothrus, Ceylon; Hneyocrypta, Singapore to Australia ;
Barychelus, New Caledonia; Jdioctis, Upolu; fdiommata
and TZrittame, Australia; Psalistops, Stothis, Huthycalus,
Epipedesis, Cosmopelma, Trichopelma, ? Acanthogonatus,
Homeoplacis, Idiophthalma, Stropheus, Cyrtogranmommna,
Neotropical Region.
b. Diplothele : Diplothele, India and Ceylon; Sorsythula,
Madagascar.
c. Sasones: Sason, Seychelles, Maldives, Ceylon, India, and
Celebes; ? Rianws, Pinang.
The existence of a primitive type, Leptopelma, in the Medi-
terranean area, and the entire absence of the group from the
Sonoran Region and from China, suggest its origin in the western
part of the Old World. Moreover, the presence of genera in
Sokotra, Mauritius, Madagascar, and all over the Ethiopian
Region attests a southern migration at a very early date,
Similarly, the extension of the group over the Oriental Region,
from India to Australia, suggests a perhaps contemporaneous move-
ment in a south-easterly direction over the area in question, after
the isolation of New Zealand. Again, the absence of genera of this
family from the Sonoran Region, coupled with the relationship
between the Mediterranean genus Leptopelma and many of the
Neotropical types on the one hand, and between the remainder of
the latter and the Tropical African genera on the other, points to
a transatlantic connection between Africa and Europe and South
America. At the same time, the possibility of a migration into
South America from Australia across the area of the Pacific must
be borne in mind,
In the absence of evidence to the contrary, the genus Sason,
specialised both in structure and habits, may be regarded as
having arisen from a primitive type in the area it at present
occupies. That this must have taken place in very early times, before
the severance of the Seychelles from India and Ceylon, w ould be
an unavoidable conclusion, were we sure of the distinctness of the
Seychellesian from the Ceylonese species. But the occurrence
of one of the Ceylonese species in the Maldives proves, I think,
artificial introduction into that Archipelago; and the same expla-
nation may apply to the presence of the genus in the Seychelles,
One other point of interest remains, and that is the unques-
tionably close relationship that obtains between the Indian and
Ceylonese genus Diplothele and the Mascarene Forsythula, the latter
being a more specialised type. This is almost the only undoubted
case of similarity between the faunas of Madagascar and India
that the Mygalomorphe supply.
Family AvICULARIIDS.
The distribution of this family is most instructive.
The heterogeneous group of genera associated together as
9%
ae
3504 MR. R. I, POCOCK ON THE [ Apr. 21,
Aviculariine is represented in the Mediterranean area by the
primitive genus [schnocolus, which ranges from Spain and Algeria
to Syria, and perhaps also occurs in Burma; by Chetopelma and
Cratorrhagus, which ave perhaps identical, found in the eastern
countries of this sea; in India by the genera Phlogiodes, Plesio-
phrictus, and Heterophrictus; and in tropical West Africa by
Scodra, Heteroscodra, Selenogyrus, Heterothele, and Solenothele.
In America the group is represented by a wealth of genera
ranging from the Southern States of the Union (Sonoran),
throughout the West Indies and over the whole of 8. America to
Chili and the Argentine. It is necessary to mention only a few
of the principal genera, such as Aphonopelma from the Sonoran,
Brachypelma, Cyrtopholis, Hapalopus, Acanthoscurria, Lasiodora,
Pamphobeteus, Avicularia, Tapinauchenius, Citharoscelus, Metrio-
pelma, Theraphosia, &e. &e.
The Selenocosmiine range from India and Ceylon over the
Indo- and Austro-Malayan Islands and Australia; the principal
genera being Pecilotheria, confined to India and Ceylon; Chilo-
brachys, occurring in India, Ceylon, and Burma; Lyrognathus in
Assam; Selenocosmia, extending from the Himalayas into
Australia; and Selenotypus and Selenotholus, which are confined
to Australia.
The Thrigmopceine, represented by Thrigmopeus and Haplo-
clastus, are confined to 8. India.
The Ornithoctonine—Cyriopagopus, Melopweus, Ornithoctonus,
Phormingochilus, Citharognathus—extend from Burma to the
Moluccas.
The Eumenophorine, containing the genera Humenophorus from
Sierra Leone, Hysterocrates from the Congo, &c., Phoneyusa and
Pelinobius from the Congo and Masailand, Batesiella from the
Camavroons, Citharischius and Anoploscelus from British East Africa,
Monocentropus from Sokotra and 8. Arabia, and Hneyocrates from
Madagascar, are thus restricted to Madagascar and the northern
portions of the Ethiopian Region, since they do not appear to
pass south of the Congo or the Zambesi.
The Harpactirine, on the contrary, are confined to South and
East Africa, ranging from Masailand to Cape Colony, and crossing
the area of the Eumenophorine north of the Zambesi. The
principal genera are Pterinochilus, Hucratoscelus, Ceratogyrus,
Celogeniwm, Harpactirella, and Harpactira.
Of the above-mentioned subfamilies the least specialised is that
of the Aviculariine, which contains genera like Heterothele and
Mitothele, serving to link in a measure the Aviculariide with the
Macrotheline Dipluride. It is, moreover, the most widely
distributed and most northerly in its range of all the sub-
families, and far the richest and most diversified in its genera.
Certain facts point to the conclusion that the Selenocosmiins
originated in South-eastern Asia. The two most primitive
genera of the subfamily, Phlogiellus and Selenocosmia, occur at the
‘DUD doubhrey T, *DULLOJIOYPULO *DULUSOIOUII AS PORTE BLUE & DULY Cour wn ‘DUD NII
‘mprimjnoapy ayy} JO Woryqrayst] jeorydersoay oy} oyeaysnq{[r OF deyq
355
GEOGRAPHICAL DISTRIBUTION OF SPIDERS.
1903.]
"19 ‘SU-9X0,
356 MR. R. I. POCOCK ON THE [ Apr. 21,
present time in this area, the former in Java and the Nicobars,
the latter all over the Indo- and Austro-Malayan Islands and
Australia. Selenocosmia, which must be regarded as a direct
descendant of Phlogiellus, must itself be looked upon as the anaestor
of the Malayan Coremiocnemis, the Assamese Lyrognathus, the
Burmese, Indian, and Ceylonese Chilobrachys, and of the two
Australian genera Selenotypus and Selenotholus, all of which, in
the absence of evidence to the contrary, may be held to have
originated within the areas of their present distribution. The
affinities of the aberrant genus, Pacilotheria, which is specialised
both in structure and habits, are more doubtful; but there is no
evidence against the hypothesis that it has been evolved in India
itself.
The presence of Pwcilotheria and Chilobrachys in Ceylon and
India attests their occupation of the latter area before the severance
of Ceylon; and the extension of Selenocosmia into Australia but
not into New Zealand, similarly attests a southern migration into
the former country before its separation from South-eastern
Asia, but after the isolation of New Zealand. It seems probable,
indeed, that this southern immigration into Australia synchronised
with that of the ancestors of the Australian marsupial mammals,
and in this connection it is significant to note that the latter are
believed to have originated in South-eastern Asia* and to have
entered Australia in Eocene times. The great difference, however,
in distribution between the Marsupials and Selenocosmiine may
be explained by the survival of the latter, and the extinction of
the former, in the area of their birth.
The Thrigmopeeine, which are confined, so far as is known, to
India, appear to have been developed from the Aviculariine stock
ata late date, namely after the depression of the connecting land
with Ceylon; and the Ornithoctoninz, which range from Siam
and Burma to the Moluceas, seem similarly to have put in an
appearance after the separation of Australia from the continent
to the north of it.
The Eumenophorine must have entered or been developed in the
Ethiopian Region at an early date, antecedent to the separation of
South Arabia and Sokotra from what is now Somaliland, and
before the formation of the Mozambique Channel divided Mada-
gascar from Hast Africa.
The Harpactivine, on the contrary, which range from Somali-
land to Cape Colony but are unknown in Madagascar, seem to
have originated independently within the Ethiopian Region
subsequent to the Miocene period. The least specialised of the
genera (Harpactirella and Pterinochilus) have not departed far
from the type of structure found in the Aviculariine.
There remains the South-American fauna to be accounted for.
Owing to the general similarity that obtains between the —
genera of this area and those of other parts of the world, the
! Lydekker, ‘ Geogr, Hist, Mammals,’ p, 5 (1896).
1903. ] GEOGRAPHICAL DISPRIBUTION OF SPIDERS. 3507
differences in extremest cases being merely accorded subfamily
rank and in others only generic rank, it is hard to believe that
the ancestors of the existing fauna entered South America from
North America in the early part of the Secondary Period, that is
to say in pre-Jurassic times, and that they have been isolated since
that date until the close of the Miocene. Again, if it be supposed
that they passed southwards from the north after the union of
the two Americas with the close of the Miocene, the conclusion is
inevitable either that the wealth of genera now existing in South
America has been evolved since that date, or that there has been
such a wholesale destruction of genera in the north as to leave
but one genus (Aphonopelma) in the Southern States of N, America
at the present day’.
The occurrence of this genus in the area in question may be
equally well attributed to migration of its ancestors northwards
into the Sonoran Region after the union of North and South
America at the close of the Miocene.
Again, since no members of the Aviculariine occur at the
present time in Australia or Austro-Malaysia, It is needless to
look to this area as the source of the South-American fauna of
this subfamily.
Africa, therefore, and the Mediterranean area alone remain as
the centres whence the incursion could have taken place. In
support of the view that preference in this connection should be
given to Tropical Africa may be urged the following facts. The
Aviculariine of this region are confined to the forest-region of
West Africa, which is roughly conterminous with the basin of the
Congo and includes also the forest-covered district to the north of
the Gulf of Guinea, and are unknown in South Africa. Secondly,
the West-African genus Scodra is apparently the nearest living
ally of the Brazilian genus Avicularia; and Heterothele and
Solenothele, from the same region as Scodra, are equally nearly
related, especially the former, to the Patagonian J/itothele.
Perhaps also some significance must be attached to the cir-
cumstance that the stridulating-organ which attains such a
state of perfection in the Kumenophorine is represented in an
imperfect and unspecialised state in many of the South-American
genera of Aviculariine.
Summary of the preceding pages.
In very early Tertiary times, or, perhaps, still earlier, the primitive
Macrotheline Dipluride arose in Eastern Asia and spread thence
in four directions :—(1) South-eastwards into Australia and New
1 Three other genera, namely Rhechostica, Tapinauchenius, and Avicularia, have
been recorded from the Southern States of North America; the first from Texas, the
second from ‘Texas also and * Indian Territory,’ and the third trom California. ‘The
generic affinities of the first are doubtful; it may indeed be reterable to Aphono-
pelma. The records of the second and third have little value, since the specimens
determined were immature females.
308 MR. R. I. POCOCK ON THE [ Apr. 21,
Zealand, where the most primitive type of all, Hexathele, still sur-
vives amd whence the ancestor of its near ally, Scotinecus, crossed
to South America, (2) South-westwards into India, Madagascar,
and Tropical Africa (fschnothele and Hvagrus), whence these
genera migrated into South America, passing thence into North
America in later Tertiary times. (3) North-westwards into the
Mediterranean Region (Phyxioschema and Macrothele). (4) North-
eastwards into North America, to give rise to the Mecicobothriide.
On the other hand, the genera of Diplurine, which are more
specialised than the Macrotheline, and therefore of later origin,
were represented in Tertiary times by genera (Brachythele) both
in the Sonoran and Mediterranean Regions. From the latter,
perhaps in the Oligocene era, they descended into Africa and
Madagascar, but do not appear to have entered the Oriental
Region at all. From Africa and Madagascar they probably
entered South America, and perhaps Australia also, though they
may equally well have passed from South America to Australia.
In later Tertiary times also it is probable that there was a
commingling of Sonoran and Neotropical forms due to southward
and northward migration.
The Mecicobothriide, which arose in the Sonoran Region from
a form, like Hexura, allied to Heaathele, themselves gave rise in
early Tertiary times to the Brachybothriide, which still exist
there and have succeeded in reaching Japan (Acattyma), and to
the Atypidee as well, which also crossed into Eastern Asia, and
thence extended westwards as far as Ireland and Algeria, and
southwards into Burma and Java. In South-eastern Asia from a
primitive Atypoid genus origimated Calommata, which probably in
the Pliocene extended right away from the Oriental Region into
West Africa, the existing species being known only from the
latter area and from Indo-Malaysia and Japan.
In later Tertiary times the Sonoran Mecicobothriide moved
southwards into South America, where the existing genus
Mecicobothrium met the genus Scotinecus, the ancestors of which,
according to my hypothesis, reached the same country by the
southern route from New Zealand.
The Cyrtaucheniide of the primitive group Nemesize at an
early date entered India, Africa, and Madagascar from the north,
or equally likely originated in Africa itself and spread thence
into the Mediterranean Region, into Madagascar and India, into
South America and into the Australian Region, as is attested
by the closeness of the similarity between the South-African,
South-American, and Australian species, and the absence of the
group from the eastern parts of the Oriental and from the Sonoran
Regions. Within Australia itself they seem to have given rise
to the group Aganippe.
The explanation given above of the distribution of the
Diplurine Dipluride applies in almost every particular to that of
the Cyrtaucheniidee of the Cyrtauchenii group, except that the
latter were later in the southward movement into Africa, reaching
1903. ] GEOGRAPHICAL DISTRIBUTION OF SPIDERS. 359
that region after the separation of Madagascar, and except that
they have left isolated genera in Indo-Malaya. The Aporoptychi
arose in Africa from the Cyrtauchenii before the severance of that
country from South America.
The Actinopodide were perhaps evolved from an Aporoptychus-
like form in South America, and crossed thence to Australia.
The Migidee probably originated in the Ethiopian Region before
the isolation of Sokotra and Madagascar. The similarity between
the South-African and Novo-Zelandian, the Tasmanian and Pata-
gonian genera suggests that the migration of the family from its
hypothetical home took an eastward direction.
The Ctenizide, a specialised northern offshoot from the
Cyrtauchenu, crossed during the Tertiary period from Asia to
North America, and occur at the present time in the Senoran and
Mediterranean Regions. It entered South-eastern Asia after the
separation of Australia; and Africa after the formation of the
Mozambique Channel, but before the sinking of the connecting
land with South America. In late Tertiary times the union of
North and South America admitted the Sonoran and Mediter-
ranean genus Pachylomerus ito the Neotropical area, where it
mingled with the Idiopine element that had come from Africa.
The Halonoproctidze must similarly have crossed from America
to Asia, or vice versa, during the Tertiary Period, since the
existing genera persist at the present time in the Sonoran Region
and China.
It is impossible to determine the original home of the Bary-
chelide. Possibly the family originated at an early date in
Africa, and became distributed all over the area they now inhabit
from that centre. Possibly it arose in the northern parts of
the Old World, and at an early date extended southwards into
the Ethiopian, Oriental, and Australian Regions. However that
may be, there seems no reason to think that the group entered
the Neotropical Region from the Sonoran,
The Aviculariide arose in the northern portion of the Old
World, whence emigrants passed into the Oriental and Ethiopian
Regions. From one primitive type in India sprang the Thrig-
mopeeinze ; from another in Indo-Malaysia the Ornithoctonine ;
and from a third the Selenocosmiine, which distributed themselves
from India, Ceylon, and the Philippine Islands into Australia,
perhaps in Eocene times. In the Ethiopian Region, from a
primitive type arose the Eumenophorine, at a sutticiently early
date to reach Sokotra, 8. Arabia, and Madagasear. Later on, after
the severance of Madagascar, arose the Harpactirine. The genera
of these specialised subfamilies probably supplanted to a large
extent both in the Ethiopian and Oriental Regions the Avicu-
lariine, but the latter attained an enormous development in
South America, which they seem to have reached from Africa or
possibly from Europe. With the Pliocene union of North and
South America certain forms spread northwards from South
America into the Sonoran Region.
360 MR. R. I, POCOCK ON THE [ Apr. 21,
Parr III.
GEOGRAPHICAL REGIONS AND THEIR PRINCIPAL GENERA.
Mediterranean Region.
Dipluridee.
Diplurine.—Brachythele: Greece, Cyprus, Central Asia.
Macrotheline.— Wacrothele: Spain, China, LooChoo. Phyxio-
schema: Transcaspia.
Brachybothriidee.— A cattyma: Japan.
Atypide.—Atypus: South and Central Europe, Algeria, Japan.
Calommata: Japan.
Cyrtaucheniidee.—Wemesia: 8. Europe, N. Africa, China. <Ane-
mesia: Afghanistan. Cyrtauchenius : Southern countries
of Mediterranean. <Amblyocarenum: 8S. Europe, N.
Africa.
Ctenizide.—Pachylomerus: Spain, Algeria, Japan. Jdiops :
Syria. Cteniza: France, Italy. dpycephalus: Sicily,
Sardinia. Cyrtocarenum: Eastern countries of Medi-
terranean. Sterrhochrotus : Turkestan. Latouchia :
Loo Choo Islands, China.
Halonoproctidee.— Halonoproctws : China.
Barychelidee.—Leptopelma: 8. Italy, N. Africa.
Aviculariidee.
Aviculariinse.—Tschnocolus: 8. Europe, N. Africa. Crator-
rhagus and Chetopelma: Eastern area of Mediterranean.
Of the above-mentioned genera, 22 in number, those belonging to
the Aviculariide and Barychelide are peculiar ; also Sterrhochrotus,
Cyrtocarenum, Zpycephalus, and Cteniza amongst the Ctenizide ;
Cyrtauchenius, Anemesia, and Nemesia amongst the Cyrtau-
chentide; Phyxioschema amongst the Dipluridz; and perhaps
Acattyma amongst the Brachybothriide. Of the others, Brachy-
thele and Amblyocarenum are also Sonoran ; J/acrothele and
Latouchia also Oriental; Atypus, Orientaland Sonoran; Calommata,
Oriental and Ethiopian; Pachylomerus, Sonoran and Neotropical ;
Idiops, Oriental, Ethiopian, and Neotropical.
Thus of the 22 genera enumerated, more than half, namely 12,
are peculiar. Of the rest, 4 are Sonoran and 5 Oriental, the latter,
with exception of /diops, occurring in the Indo-Chinese area ;
2 Ethiopian and 2 Neotropical. With the Australian Region the
Mediterranean has no genera In common, although Porrhothele
and Stenygrocercus of the former are nearly related to MJacrothele
and Phyxtoschema of the latter. These data make it impossible
to attach the Mediterranean area to either of the others with
which it has genera in common. Hence it must be regarded as a
Region apart. The only genus which extends into the Holarctic
is Atypus.
It may be defined as the Region lying north of the Ethiopian
and Oviental Regions, and south of the great mountain-chains of
urope, north of which the Mygalomorphe, excepting Atypus,
do not pass.
1903. ] GEOGRAPHICAL DISTRIBUTION OF SPIDERS. 361
Ethiopian Region.
(Africa south of the Sahara, South Arabia (Sokotra), and
Madagascar.)
Dipluride.
Diplurine.—apalothele : Madagascar.
Brachytheliscus: 8. Afvica.
Macrotheline.—Hvagrus: 8. Africa. Ischnothele: Mada-
gascar, 8. & W. Africa.
Atypide.—Calommata: W. Africa,
Cyrtaucheniidee.—H ermacha, Hermachastes, Lepthereus, Pi onothele,
and Spiroctenus: S. Africa. Genysa : Madagascar.
Pelmatorycter: 8. Africa. Microbatesia: W. Africa.
Bessia, Stictogaster, and Homostola: 8. Africa. Aporo-
ptychus and Ancylotrypa : W. Africa.
Ctenizide.—Stasimopus: S. Africa. diops : E., W., & 8. Africa.
Heligmomerus: E. & 8. Africa. Gorgyrella: 8. Africa.
Migidee.— Moggridgea : 8. Africa, Sokotra. Myrtale and Thyro-
peus : Madagascar’.
Barychelicee. Atrophothele: Sokotra. Eubrachythele, Pisenor,and
Pisenorodes: EK. Africa. ‘yphonisia: W. Africa.
Brachionopus: $%. Africa. Lorsythula : Madagascar.
Tiyidia: Mauritius. Sason: Seychelles.
Aviculariide.
Aviculariine.—Scodra, Heteroscodra, and Selenogyrus : We
Africa. Heterothele: W. & E. Africa. Solenothele :
W. Africa.
Eumenophorinse.—Hysterocrates : W. Africa. Loxomphalia :
BE, Africa. Phoneyusa and Pelinobius: W. & K. Africa
(probably identical). Citharischius: Ki. Africa, Batest-
ella: W. Africa. Anoploscelus: KE. Africa, MJono-
centropus: Sokotra, 8. Arabia. Lneyocrates: Mada-
gascar,
Harpactirine.—Pterinochilus: E. & S. Africa, Huecrato-
scelus: E. Africa. Ceratogyrus, Calogeniwm, Harpac-
tirella, and Iurpactira: 8. Africa.
Perhaps the same.
Of the 54 above-mentioned genera, all are peculiar to the Region,
with exception of Mvagrus, Tschnothele, Idiops, and Hermacha,
which also occur in South America; Sason, Ldiops, Ischnothele,
and Calommata in the Oriental Region; possibly also with the
exception of Ancylotrypa and Moggridgea, if the former is identical
with the South-American Bolostromus, and the latter with the
New-Zealand JMigas.
Apart from its distinctive genera, the Region is essentially
characterised by the two subfamilies of Aviculariide, the Kumeno-
phorine and Harpactirine, which are peculiar.
The fauna of Madagascar does not seem to warrant the
ascription of more than subregional importance to this island,
Apart from the occurrence of Zschnothele in India and Mada-
gascar, and the relationship between Diplothele and Forsythula,
362 MR. R. I. POCOCK ON THE | Apr. 21,
Madagascar shows no similarity to the Oriental Region, with which
Simon united it. (Trouessait, La Géogr. Zoolos. pp. 208-211,
1890.)
The African portion of the Ethiopian Region is divisible into
two well-marked subregions, namely, the West-African or forest
area of the Congo, which extends from the shores of the Gulf of
Guinea to Uganda, and a South and Hast-African area. The
former is essentially characterised by the presence of the Avicu-
lariine, Kumenophorine, and Barychelide; the latter by the
Har pactiri ine, Cyrtaucheniide, and Migide. In East Africa,
north of the Zambesi, the two regions cross.
Oriental Region.
Dipluride.
Macrothelinee.— Macrothele : Burma, Malacca, Java. Ischno-
thele: India.
Atypidx.—Atypus: Burma, Java. Calommata: Burma, Siam,
Java, Sumatra.
Cyrtauchentida
Damarchus: Burma, Malacca. <Atmetochilus: Tenas-
serim.
Ctenizidee.—TJdiops : India, Burma. Heligmomerus: Ceylon, 8.
India. Latouchia: Himalayas. Conothele: Burma to
Solomon Islands.
Barychelidee.—Diplothele : India, Ceylon. Sasonichus: India
(Travancore). Plagiobothrus and Sipalolasma: Ceylon.
Sason: India, Ceylon, Saleyer. Hncyocrypta: Malacca,
Borneo, to Queensland.
Aviculariide.
Aviculariinse.— Phlogiodes and Heterophrictus: India. Plesio-
phrictus: India, Ceylon, ? Burma. Acolischnus ?
Burma.
Thrigmopeeinse.-—Thrigmopeus and Haploclastus : Western
and Southern India.
Selenocosmiinee.—Pecilotheria: India, Ceylon. Chilobrachys :
India, Ceylon, Burma. Selenocosmia : Himalayas,
mama, Java, Sumatra, Borneo, New Guinea. Phlogi-
ellus: Java, Nicobars. Lyrognathus: India (Assam).
Coremiocnemis: Malacca. ~% Orphnecus: Philippine
Islands.
Ornithoctoninze.—Ornithoctonus: Burma. Cyriopagopus :
Burma, Malacea. JMelopeus: Burma, Siam. Citharo-
gnathus : Borneo. Phormingochilus: Borneo, Celebes,
Moluccas.
36 genera. Characteristic of this Region, and entirely confined
1 New name for the species from Tenasserim, described by Thorell as Ischnocolus
brevipes (Thorell, Ann. Mus. Genova, xxxvii. p. 170, 1897; also Pocock, ‘ Fauna of
British India: Arachnida,’ p. 183), which at least differs from the typical species of
Ischnocolus from the Mediterranean in having the tarsus of the palp short in the male,
1903.) GEOGRAPHICAL DISTRIBUTION OF SPIDERS. 363
to it, are the two subfamilies of Aviculariide, Thrigmopeine and
Ornithoctonine, while the third subfamily, Selenocosmiine, is
common to it and the Australian Region. Moreover, all the genera
of Aviculariine, of Cyrtaucheniide and Barychelide are peculiar,
with the exception of Hncyoerypta, belonging to the last-mentioned,
which extends to Australia. Conothele amongst the Ctenizide is
also peculiar, if the Solomon Islands be included. For the rest,
Macrothele and Atypus occur also in the Mediterranean Region ;
Calommatain Japan and West Africa ; Tschnothele in Madagascar,
Africa, and South America; Zdiops in Africa and South America ;
Heligmomerus in Africa ; Latouchia in China; and Sason in the
Seychelles.
The apparent richness of India, Ceylon, and Burma, as compared
with the eastern portion of the Region, may be merely due to
more extensive collecting in the former countries ; or it may be
attributable to failure to penetrate far into the latter before the
break-up of the area into islands. Four genera only are known
to cross Wallace’s line, namely, Selenocosmia, Phormingochilus,
Encyocrypta, and Sason ; but, apart from these genera, there is no
evidence to show that this channel constitutes the divisional line
between the Oriental and Australian Regions,
Australian Region.
(Australia and its adjacent Islands.)
Dipluridee.
Macrotheline.—Stenygrocercus : N. Caledonia, Queensland.
Porrhothele and Hexathele: New Zealand. Atrax and
Hadronyche: Australia.
Diplurine.—Aname, Ixamatus, Chenistonia, and Dekana :
Australia, Tasmania.
Cyrfaucheniide.— Cantuaria: New Zealand. <Arbanitis: New
Zealand, Australia. Dyarcyops, Aganippe, Llakistonia,
Anidiops, and Idiosoma: Australia.
Migidee.—Heteromigas: Tasmania. Migas: New Zealand.
Actinopodide.—Hriodon : Australia.
Barychelide.—/diommata: Australia. Hneyoerypta: Australia,
New Caledonia. Trittame: Australia. Barychelus:
New Caledonia. Jdioctis: Upolu.
Aviculariidee.
Selenocosmiinse.— Selenocosmia, Solenotholus, and Selenotypus :
Australia.
Of the above-mentioned 27 genera, all are peculiar to the
Region with the exception of Hneyoerypta and Selenocosmia, which
also occur in the Oriental Region, and possibly of J/igas, which
may prove indistinguishable from the South-African JMoggridgea.
The poverty of New Zealand in genera, and the distinctness of
two of the three forms that do occur there, entitle it to subregional
rank, but no more.
364 MR. R. I. POCOCK ON TIE [Apr. 21,
The Austro-Malayan Islands, which were referred by Wallace
to the Australian Region, belong, according to their Arachnid
fauna, to the Oriental Region—Wallace’s line being practically
non-existent.
Sonoran Region.
Dipluride.
Diplurine.—Brachythele : California, Texas, &e.
Macrothelinee.—Hvagrus : Idaho.
Mecicobothriide.—Hewura : Oregon and Washington State.
Brachybothriidee.—Lrachybothrium : From Texas and Virginia to
British Columbia. <Atypoides: California.
Atypidee.—<Atypus: Eastern States, from Wisconsin to Florida.
Cyrtauchentidee.—Amblyocurenwm, Aptostichus, and Actinoxia:
California. M/yrmeciaphila: Carolina, Virginia.
Ctenizidee.—Pachylomerus : Carolina, Alabama, &c., Mexico.
Bothriocyrtum : California, Utah, Texas, Mexico.
Halonoproctidee.—Cyclocosmia : California, Alabama. Chorizops :
Mexico.
Aviculariudee.
Aviculariinee.—4d phonopelma : Texas, Arizona, California.
And perhaps others.
Out of the above-given list of 15 genera no fewer than 10 are
peculiar to the Region, all, that is to say, with the exception of
Atypus, Brachythele, Amblyocarenum, and Pachylomerus, which
are also present in the Mediterranean Region, and Hvagrus,
which is met with in the Neotropical and Ethiopian Regions.
This Region, including the Mexican plateau and the Southern
States of the Union, extends northwards to the northern limits
of the Mygalomorphe, or at least to those of the Scorpiones and
Solifugee *.
Neotropical Region.
(Central America south of the Mexican plateau, West Indies,
Bahamas, and South America.)
The principal genera are :—
Dipluridee.
Diplurine.—Diplura: 8. America. Uruchus: Ecuador.
Trechona, Melodeus, and Harmonicon: Brazil. Fufius:
Central and South America. Trissothele: Chil. Lyci-
nus: Argentine.
Macrotheline.—IJschnothele: Central and South America,
W. Indies. Hvagrus: Central America.
Paratropide.—Paratropis : Ecuador, Amazons. <Anisaspoides :
Amazons. Anisaspis: W. Indies.
Mecicobothriidee.— MWecicobothrium: Argentine.
1 Compare also in this connection the distribution on the coast of Limulus poly-
phemus, trom Cape Cod to the Gulf of Mexico.
1903.] GEOGRAPHICAL DISTRIBUTION OF SPIDERS. 365
Cyrtaucheniidee.—Stenoterommata: Brazil, Chili. Pselligmus:
Brazil. Hutychides: Central America, Antilles. Aporo-
ptychus and Bolostromus: 8. America. Rhytidicolus :
Venezuela. Phrissecia: Ecuador. Pheochta and
Celidotopus : Venezuela. Hermacha: Brazil, Chili.
Carteria: Chili. Rhachias: Brazil. Neocteniza: Central
and South America. ucteniza and Hnrico: Central
America.
Actinopodide.—<Actinopus: Central America to the Argentine.
Ctenizidee.—Pachylomerus: Centraland South America, W. Indies.
Idiops and Pseudidiops : 8. America.
Migidee.—(Undescribed genus): Chili.
Barychelidee.—Psalistops: Venezuela. Stothis: Venezuela, W.
Indies. Huthycelus and Hpipedesis: Venezuela, Cosmo-—
pelma: Brazil. Trichopelma: St. Domingo, Brazil.
Idiophthalma: Brazil. Cyrtogrammomma: Demerara.
Homeoplacis: Brazil.
Aviculariide.
Aviculariine.—Mitothele : Patagonia. Thalerommata and
Tmesiphantes: Brazil. Dryptopelma: Ecuador. Ma-
gula: Braal. Adranochelia and Cheetorrhombus :
Venezuela. Stichoplastus: Venezuela, Trinidad. Hapa-
lopus: Central and South America. Jetriopelmea :
Central America to Argentine. Cyrlopholis: W. Indies,
Bahamas, Central America. Citharacanthus and brachy-
pelma: Central America. Homaomma: Southern Brazil.
Paraphysa and Phryxotrichus: Chili. Citharoscelus :
Chili, Argentine, Brazil. Zasiodora: 8. Brazil. Pampho-
beteus: Ecuador, Colombia. Phormictopus: WHayti.
Acanthoscurria: W. Indies to Argentine. Sericopelma :
Panama, Cayenne. phebopus: ‘Amazons. Avieularia:
Brazil, W. Indies, Central America. Tapinauchenius :
Central and South America. Psalmopeus: 8S. America.
This list of 69 genera, although incomplete, shows the Neo-
tropical Region to be far the richest of all in point of numbers,
Moreover, all are peculiar to the Region except Pachylomerus,
which occurs also in the Sonoran and Mediterranean Regions ;
Idiops, which is also met with in Africa, India, and the Mediter-
ranean Region; and Hermacha, Aporopt yohus, possibly Bolostromus,
Hvagrus, and Ischnothele, which have been recorded from Africa,
the last-named also ean Madagascar and India. So too ane
family Paratropidee is confined to the Region ; the Mecicobothriidee
exist elsewhere only in the Sonoran Region ; ; and the Actinopodidze
only in Australia,
366 MR. R. I, POCOCK ON THE fApr. 21,
Part lV.
DisTRIBUTION OF SOME OF THE FAMILIES OF ARACHNOMORPHE
THAT WERE REPRESENTED IN THE OLIGOCENE PERIOD.
The only fossil Spiders, with one or two exceptions, of which
the generic determination may be trusted, are those that have
been found in the amber-beds of Oligocene age. Those that have
been referred to genera not now known to exist have no special
importance from a geographical standpoint; but of those that
have been assigned to existing genera, the following may be taken
as a fairly complete list :—Segestria and Dysdera (Dysderide) ;
Eresus (Eireside) ; Amaurobius (Dictynide); Aranea, Zilla, Tetra-
gnatha, and Nephila (Argiopide) ; Archea (Archeidee) ; Tegenaria
and Agelena (Agelenide); Drassus (Drasside); Clubtona, Any-
phena, and Sparassus (Clubionide) ; Thomisus and Philodromus
(Thomisidee).
Although since the Oligocene these Spiders have had the same
time for dispersal, they nevertheless differ greatly in their distri-
bution. Dysdera, for example, is apparently indigenous only in the
Mediterranean and central portions of Europe and Asia’; Hresws is
found in South and Central Europe, Central Asia, China, and Africa
to the north and south of the Sahara; Amaurobius all over the
world with the exception of India, tropical Africa, and Madagascar ;
Aranea and Tetragnatha are cosmopolitan; Wephila is mainly
restricted to the tropics and the Southern hemisphere, though in
Eastern Asia it extends as far north as Japan, and in North
America enters the Southern States of the Union; Archea is
known only from Madagascar; Tegenaria is indigenous apparently
only in Europe and North America; <Agelena in Hurope,
South Africa, India, and Burma; Anyphena in North, Central,
and South America (Andes), Japan, India (in the mountains),
and Central Europe.
The Hersiliidee were represented in the Oligocene of Hurope.
The four existing genera are distributed as follows :—AHersiha
ranges from the southern area of the Mediterranean as far as
Malaysia in the Oriental Region, and over Africa and Madagascar.
Tts very near ally Murricia is confined to India; Herszliola is
known only from the Mediterranean Region and 8. Africa. Zama,
next to Hersiliola the most primitive of all the genera, occurs in
the Mediterranean, Oriental, Australian, and Neotropical Regions.
The distribution of the group offers no difficulties to the hypo-
thesis of a southern migration from EKurope. The absence of the
genus Zama from North and Central America strongly suggests its
existence in South America to be attributable to migration from
Australia.
Two genera of Hresidz have been recorded from the European
1 Specimens that have been recorded from the Southern hemisphere are mostly,
probably in all cases, referable to imported European species.
1903.] GEOGRAPHICAL DISTRIBUTION OF SPIDERS. 367
Oligocene. At the present time the family is represented by
Adonea and Dorceus in the southern area of the Mediterranean ;
by Dresserus and Seothyra in Tropical and South Africa; by
Stegodyphus in the Mediterranean Region, India, Ceylon, and
Burma in the Oriental Region, and East and South Africa in the
Ethiopian Region; by resus, which extends across Europe and
Asia from England to China, being especially abundant in the
Mediterranean Region and also occurs in South Africa. The
absence of this group from Madagascar points to a late (probably
Pliocene) incursion from the nor th into Afri ‘ica, which, in conjunc-
tion with its failure to reach North America and Australia,
also explains its absence from South America,
The only existing genus of Urocteidee, Uractea, was represented
in the European Oligocene. At the present time it is found in
the Mediterranean Region, China, Japan, India, and South Africa.
The apparent absence of the genus from Madagascar indicates a
late movement into South Africa.
The Palpimanide are also alleged to date back to the Oligocene.
The most primitive member of the family, Huttonia, constituting
the Huttoniine, is now restricted to New Zealand. The more
specialised Stenochilinze are confined to the Oriental Region,
where they range from Bombay to the Philippines. The Palpi-
manine, the most specialised of the three, to which the Oligocene
genus belongs, are represented by two groups, the Chedimez and
the Palpimanee. The latter, containing the single genus Palpi-
manus, occurs in the Mediterranean Region, Western India, and
East and South Africa; the former is represented by Boagrus in
the Malay Peninsula, Steriphopus in Ceylon, Sarascelis in the
Malay Peninsula and tropical West Africa, Diaphorocellus in
South Africa, and by Otiothops and Aniscedus in South America.
The survival of the primitive type, Huttonia, in New Zealand
assigns great antiquity to this group. The Stenochiline and
Palpimanine were perhaps evolved in South-eastern Asia and
spread thence in a westerly direction, the more specialised Palpi-
manine reaching Kurope in the Oligocene and passing into Africa
and thence to South America. The entire absence of Palpimaninze
from North America and Australia, and the near relationship
that obtains between the genera from South America and Tropical
Africa, almost compels the belief that the former country received
this element of its fauna from the latter.
The Anyphenide are represented in the Oligocene beds of
Europe and North America. At the present time, one genus
only, Anyphena, occurs in the temperate area of the Northern
hemisphere of the Old World; the same genus also occurs in
North America; while the remainder of the genera, sixteen or so
in number, extend into South America, principally along the
Andean chain as far south as Tierra del Fuego. These facts
suggest that the group had its origin in North America, perhaps
in Eocene times, crossed thence into Asia and Europe before or
during the Oligocene, and, when North and South America were
Proc. Zoou. Soc.—1903, Vou. I. No. XXIV, 24
368 ON THE GEOGRAPHICAL DISTRIBUTION OF SPIDERS. [ Apr. 21.
united at the close of the Miocene, descended into the latter area.
If so, it presents a striking parallel to that of the Camel family,
which was represented by many genera, now extinct, in North
America during the Tertiaries, from the Lower Oligocene upwards,
entered the Old World, where it is now represented by the Camels,
and South America, where the Llamas still exist and extend
principally along the mountains far to the south into Patagonia.
Here again, as in other cases, the exact parallelism between
the Mammalia cited and the Spiders fails apparently only on
account of the survival of the latter and the extinction of the
former in the area of their origin.
To pursue this subject further, and take all the families and
genera of Spiders into consideration, would require a special
volume. Enough, however, has been said to show that the
suggestions put forward to explain the distribution of other groups
of animals apply also to the distribution of Spiders.
Norre.—During the passage of these pages through the press,
Simon has issued two papers (Bull. Mus. d’Hist. Nat. 1902,
pp. 595-598; Ann. Soc, Ent. Belg. xlvii. pp. 21-23, 1903)
containing descriptions of the following new genera of Mygalo-
morphee :-—
Dipluride.—Hntypesa, allied to the Australian Jeamatus:
Madagascar. é
Cyrtaucheniide.— Diadocyrtus, allied to the Ceylonese Scalido-
gnathus; Genysochera, near Genysa: Madagascar.
Ctenizidee,— Hebestatis, near Pachylomerus: California,
Migide.—Calathotarsus, near the New-Zealand Migas: Chill.
Pacilomigas, near Moggridgea: S. Africa.
Barychelidee.—(Barycheli) Zophoryctes, allied to the Australian
Trittame ; (Diplothele) Cestotrema, Acropholius: Mada-
gascar.
The chief feature of interest is the six new genera from Mada-
gascar, two of which are related to Australian forms, one to a
Ceylonese genus, one to a previously known Mascarene genus; the
remaining two belonging to the Diplothele, hitherto represented
by two genera, one from Madagascar, the other from Ceylon and
India. Reference may also be made to the new Chilian genus of
Migidee, which I have already mentioned (p. 365).
Ablabes
dorie, 93.
hermine, 101.
semicarinatus, 101.
Acalyptophis
peronit, 95.
Acanthephyra
pellucida, 76, 77.
purpurea, 77.
Acanthedon, 350.
Acanthogonatus, 353.
Acanthoscurria, 354, 365.
Acartia
clausti, 119, 125.
Acattyma, 346, 348, 358,
360.
Accola, 3438.
Acetes, 53.
Achalinus
bracconieri, 88.
rufescens, 88.
spinalis, 88, 100.
Achtheres
sandre, 109.
Acolischnus, gen. nov.,
362.
Acomys
witherbyi, 298,
Acrochordus
Javanicus, 85.
Acropholius, 368.
Actinopus, 352, 365.
Actinopyga, 193.
Actinoxia, 849, 364.
Adonea, 367.
Adranochelia, 360.
Egisthus
aculeatus, 124.
atlanticus, 124.
longirastris, 124.
mucronatus, 124,
ZEpycephalus, 340, 360,
Aganippe, 349, 565.
Agelena, 366.
INDEX.
Aipysurus
annulatus, 95, 101,
Alces
bedfordieg, 147, 148,
150
machlis, 133-151.
Alestes
baremose, 6, 7.
Suchsii, 23.
intermedius, 22, 28.
kingsleyie, 23.
longipinnis, 22.
mucrolepidotus, 7, 28.
opisthotenia, 22, 28.
thollont, 22.
Alma
budgetti, 222.
emint, 222.
millsont, 222.
nilotica, 222.
stuhimanni, 221, 222.
Alvania, 219.
Alycveus
conformis, 198.
diplochilus, 198.
perakensis, 198.
thieroti, 197.
Amaurobius, 366.
Amblycephalus
moellendorfii, 97.
Amblyocarenum,
360, 364.
Ampbilius
longirostris, 25, 28.
Anabas
kingsleye, 27.
maculatus, 27.
pleurostigma, 27, 29.
Aname, 343, 363.
Anchicaligus
nautili, 108.
Ancistrodon
acutus, 98.
blomhoffii, 98, 102.
348,
Ancistrodon
intermedius, 101.
Ancylotrypa, 361.
Anemesia, 348, 360.
Anidiops, 349, 363.
Anisedus, 367.
Anisaspis, 345, 364.
Anisaspoides, 345, 364.
Anoplopterus
longirostris, 25.
Anoploscelus, 354, 361.
Anthropopithecus
troglodytes, 191.
Antilope
dammah, 300.
leucoryx, 300.
Anyphena, 366, 367.
Aphidium
Jicus, 106.
Aphonopelma, 354, 357,
364.
Aporoptychus, 349, 359,
361, 365.
Aptostichus, 349, 364.
Arachnactis
albida, 117, 118, 123.
Aranea, 366.
Arbanitis, 348, 349, 363.
Archaea, 366.
Argyroneta, 158.
Arthrolycosa, 340.
Artotrogus, 104.
Arvicanthis
barbarus, 297.
dunni, 297.
zebra, 297, 298.
Aspidosiphon, 37, 38, 104.
Aspidura,
copii, D4.
trachyprocta, 94.
Asthenoceros, gen. nov.,
315.
woodworthi, 302, 315,
318.
370
Atmetochilus, 348, 362.
Atrax, 8438, 345, 863.
Atrophothele, 352, 361.
Atypoides, 346, 364.
Atypus, 342, 346, 348,
360, 363, 364.
Auchenoglanis
ballayi, 25.
Augraptilus
zetesios, 125.
Avicularia, 354, 357, 365.
Beolidia
major, 252, 253, 254.
— ornata, 254.
mbit, 203, 254.
Bagrus
bayad, 6.
Balistes
biaculeatus, 182, 184.
bines, 185.
Barbus
batesti, 25, 29.
howkert, 24.
bynnt, 25.
camptacanthus, 25.
guirali, 25.
kessleri, 25.
potamogalis, 25.
progenys, 24, 29.
teniurus, 24, 28.
tanensis, 25.
Barilius
kingsleye, 25.
ubangensis, 25.
Barychelus, 353, 363.
Bassettia
congri, 108.
Batesiella, 354, 361.
Bergendalia, gen. nov.,
310.
anomala, 502, 310, 311,
318.
Beroé
cucwmis, 117, 118.
Bessia, 349, 361.
Bettonia
lagariensis, 213, 214.
Blakistonia, 349, 363.
Boagrus, 367.
Bolostromus, 349, 361,
365.
Bomolochus
onost, 108.
zeugopter?, 108.
Bothriocyrtum, 350, 364.
Brachionopus, 352, 361.
Brachybothrium, 346,
348, 364.
Brachypelma, 354, 365,
INDEX.
Brachythele, 343,
358, 360, 364.
Brachytheliscus, 343, 361.
Bryconzethiops
microstoma, 22.
Buccones, 258.
Bufo
vulgaris, 89, 90.
Bungarus
candidus, 96.
fasciatus, 96.
multicincta, 96.
Cacomantis, 259, 260,
267, 275, 276, 279,
280, 281, 284, 287,
289, 290.
merulinus, 280.
Calamaria
berezowski, 93.
pavimentata, 93, 101.
pfeffert, 93.
septentrionalis, 93.
Calanus
finmarchicus, 118, 128.
Calathotarsus, 368.
Caligus
labracis, 108.
pacificus, 108.
Callionymus
maculatus, 109.
Calliteuthis
sp., 77.
Callophis
macclellandii, 97.
Calommata, 9846, 348,
358, 360, 361, 362, 363.
Calyptopis, 129, 130,
131.
Canis
anthus, 295, 296.
— soudanicus, 295.
Cantuaria, 348, 363.
Carteria, 348, 365.
Casuarius
australis, 270.
salvadorit, 270.
uniappendiculatus, 1.
Cecrops
acanthie vulgaris, 108.
Celidotopus, 349, 365.
Cemas
algazel, 300.
Centropus, 259, 260, 262,
263, 266, 267, 271,
273, 274, 275, 276,
278, 279, 281, 284,
287, 289, 290, 291.
madagascariensis, 285.
toulou, 289.
a
345, |
Cephalophus
equatorialis, 6.
harveyt, 220.
egnifer, 226.
gohnstoni, 226.
weynst, 226.
Ceratella
fusca, 113, 114, 115,
116
16.
minima, 118, 114, 115,
116.
procumbens, 113, 114.
spinosa, 118, 114.
Ceratogyrus, 354, 361.
Ceratophyllidia, gen.
nov., 250.
africana, 250.
Cerberilla
africana, 254.
annulata, 25d.
—, var. affinis, 255.
longicirrha, 250.
Cercocebus, 12, 18, 20.
aterrimus, 191.
Cercopithecus, 12, 15, 20.
stairsi, 12.
Cereanthus
loydii, 117.
Cestoplana
maldivensis, 313.
Cestotrema, 368.
Cheetopelma, 354, 360.
Chetopterus, 170, 171,
172, 173.
Cheetorrhombus, 3865.
Charopinus
dubius, 109.
Chenistonia, 343, 363.
Chilium
chlorurus, 109.
Chilobrachys, 354, 356,
362. :
Chiridius
armatus, 126.
tenwispinus, 125.
Chitina
ertcopsis, 113, 114.
Chlamydosaurus
kingi, 191.
~ Chondracanthus
blackeri, 109.
ninnit, LOY.
ornatus, 109.
Chondrocarpus, gen. nov.,
sp., 105.
reticulosus, 105, 106.
Chorizops, 552, 364.
Chrysobagrus
longipinnis, 25.
Chrysochloris, 328.
Chrysococeyx, 259, 274,
280, 287, 290.
Chrysopelea
ornata, DA.
Ciralias
uranoscopus, 109.
Citharacanthus, 365.
Citharinus
sp., 6.
Citharischius, 354, 361.
Citharognathus, 354, 362.
Citharoscelus, 354, 360.
Clarias
lazera, 7.
liberiensis, 25.
Clavella
cluthe, 108.
Cloeosiphon, 38.
Clubiona, 366.
Clupea, 334.
Cobus
defassa, 6.'
leucotis, 9).
thomasi, 4, 6.
Coceygus, 287.
Coceystes, 260, 263, 275,
281, 284, 287, 289;
290.
Coelogenium, 354, 361.
Colobus, 12, 16, 17.
sp., 9.
guereza, 17.
vellerosus, 16.
Coluber
climacophorus, 92, 100.
conspicillatus, 100,
davidi, 92.
dione, 91, 100.
mandarinus, 91.
melanurus, 92.
mellendor ffit, 92.
phyllophis, 92.
porphyraceus, 91.
quadrivirgatus, 100.
radiatus, 92, 100.
rufodorsatus, 91.
schmackeri, 101.
schrenckii, 92, 100.
teniurus, 92.
Columba, 334.
Conchecia
borealis, 121.
hyalophyllum, 120,121.
maxima, 118, 120, 121.
porrecta, 120, 121.
Concheecilla
daphnoides, 120, 122.
lacerta, 122.
Conothele, 350, 362, 363.
Coracias
caudatus, 9.
INDEX.
Coregonus, 109.
Coremiocnemis, 356, 362.
Corvinella
affinis, 7.
corvina, 4.
Coryphodon, 201.
Corythornis
cyanostygma, 8.
Cosmetornis
vevillarius, 6.
Cosmopelma, 353, 365.
Coua, 259, 260, 262, 268, |
964, 266, 267, 272, |
Tis molaaloueto, |
281, 286, 287, 288,
289, 290.
cerulea, 275, 276.
reynaudi, 275, 276,
Di 278s 201.
Crateropus, 7.
Cratorrhagus, 354, 360.
Criodrilus, 221.
262, 263, 265, 267,
274, 275, 276, 279,
281, 283, 284, 285,
287, 288, 290.
Cryptocelides, 310, 311.
Cryptorhina
afra, 8.
Cteniza, 350, 360.
Ctenolabrus
rupestris, 108.
Cuculus, 260, 262, 263,
266, 267, 271, 272,
273, 274, 275, 276,
278, 279, 281, 284,
287, 289, 290:
canorus, 268, 277.
Cucumaria
planci, 192,
Cyanops, 278.
Cyclocaris
Ffaroensis, 127.
guilelmi, 126.
tahitensis, 126, 127.
Cyclocosmia, 352, 364.
Cyclophorus
borneensis, 195.
malayanus, 195.
saturnus, 195.
tuba, 195.
Cyclopterus
luimpus, 109.
Cyclostoma
anostoma, 197.
sectilabrum, 197.
Cynocephalus, 18, 19, 20,
21.
mormon, 12, 18.
porearius, 18,
260, |
371
Cynocephalus
sphinx, 18.
Cynognathus, 177, 178.
Cynopithecus, 12, 18, 19,
20, 21.
niger, 18, 19, 20.
Cyphonisia, 352, 361.
Cyriopagopus, 354, 362.
Cyrtauchenius, 348, 560.
Cyrtocarenum, 350, 360.
Cyrtogrammomma, 393,
365.
Cyrtopholis, 554, 365.
Cyrtopia, 131.
Damatiscus
sp., 9.
Damarchus, 348, 862.
Dehitella
atrorubens, 113.
Dekana, 343, 363.
Dendrophis
pictus, 92.
Desis, 158.
Diadoeyrtus, 368,
Dicrurus
assiinilis, 6.
Dinematura
musteli levis, 108.
Dinemelia
dinemelia, 9.
Dinodon
Japonicus, 100.
rufozonatus, 89, 100.
senicarinaius, LOU.
septentrionalis, 90.
tessellatus, 100.
Diocus, 104.
Diplommatina
laidlawi, 198, 199.
skeati, 198, 199.
Diplothele, 353, 362.
Diplura, 343, 345, 364.
Dipodillus
stigmonyx, 296, 297.
Diposthus
corallicola, 3\6.
Dipsadomorphus
kraepelini, 84, 94.
multi-maculatus, 94.
Dipus
microtis, 299.
Discocelis
tigrina, 307, 308, 309.
Distichodus
niloticus, 6.
notospilus, 23,
Distira
brugmansii, 96.
cyanocincia, 96, 101.
372
Distira
orientalis, 95.
ornata, 95, 101.
stokesit, 95.
subcineta, 95, 101.
viperina, 96.
Ditropis
caverne, 195, 199.
Dolomedes, 152, 153, 154,
158.
Dorceus, 367.
Doridicola, 104.
Dossenus, 153, 155.
marginatus, 102, 159d,
168.
Dranees, 153.
Drassus, 366.
Drauces, 153.
Dresserus, 367.
Dromeus, 268, 269.
Dromococcys, 278, 281,
284, 287, 290.
Dryptopelma, 365.
Dyarcyops, 390, 363.
Dysdera, 365.
Hehidna, 328.
Hehinodoris, 251.
Elaphis
virgatus, 92.
Hlasmodes
obtusum, B01.
Eminoscolex, 214.
Encyocrates, 354, 361.
Encyocrypta, 353, 362,
363.
Knhydris
hardwiekii, 96.
Enna, 153, 164.
Knoploteuthis
diadema, 77.
Hnrico, 349.
Enterocola, 106.
Entypesa, 368.
Hoatypus, 341.
Nolidicola, 104.
Hphebopus, 365.
Epipedesis, 353, 365.
Hpiplatys
infrafasciatus, 27.
sex fasciatus, 27.
Equus
burchelli, 2.
caballus, 2.
przewalskii, 200.
Brcolania
Junerea, 257.
siottii, 257.
viridis, 257.
eanzibarica, 256.
Bresus, 366, 367.
INDEX.
Hreasilus
biuncinatus, 107.
centrarchidarum, 107.
Eriodon, 352, 363.
Hrythropygia
ruficauda, 5.
Kubrachythele, 552, 361.
Eucalanus
attenuatus, 119, 123.
crassus, 129.
elongatus, 123.
Hucanthus
marchesettii, 107.
Eucheta
norvegicd, 119, 123.
Euchirella
carinata, 126.
Eueratoscelus, 354, 361.
Eucteniza, 349, 365.
Endactylina
acanthii, 108.
carcharie glauci, 108.
musteli levis, 108.
similis, 108.
squatina angelt, 108.
Eudriloides, 219.
Eudrilus, 212, 213, 214.
| Hudynamys, 259, 260,
2630 260qu 207 ees
278, 279, 281, 284,
285, 287, 289, 290.
Eumenophorus, 354.
Euphausia, 129, 130.
EHurylepta
kelaartiz, 301.
Euthemisto
sp., 127.
bispinosa, 127.
compressa, 127, 129.
libellula, 128.
Euthyeelus, 353, 368.
Hutropius
niloticus, 6.
Eutychides, 365.
Evagrus, 345, 345, 361,
364, 365.
Falco, 261.
Felis, 334.
Forsythula, 353, 361.
Fufius, 348, 364.
Furcilia, 131.
Gaetanus
armiger, 125.
major, 125.
Gaidius, 125.
pungens, 125.
Galeus
vulgaris, 108.
Galvyina, 257.
+ Gastrosteus
aculeatus, 107.
Gazella
cuviert, 191.
ruficollis, 301.
rufifrons, 301.
Gecko
subpalmatus, 91.
Genysa, 348, 349,
368.
Genysochera, 368.
Geococcyx, 260, 262, 263,
264, 265, 266, 267,
Pil, PARA Bless, 276;
278, 279, 281, 282,
284, 285, 287, 288,
290.
californianus, 268.
mexicanus, 282.
Georissa
monterosatiana, 199.
Gerbillus
agag, 296.
pygargus, 297.
Gobius, 109.
buccatus, 109.
minutus, 109.
Goldfingia, 38.
macintoshi, 37, 40.
Gomphognathus, 177,
179, 180.
kennemeyeri, 177, 180,
Gorgyrella, 350, 361.
Guira, 259, 262, 263, 266,
361,
207, 274, 275, 276,
278, 279, 284, 287,
289, 290.
Hadronyche, 343, 345,
363.
Heemobaphes
ambiguus, 109.
Halocyphria
globosa, 120, 122.
Halonoproctus, 352, 360.
Hapalopus, 354, 365.
Hapalotbele, 343, 361.
Haplochilus
elegans, 27.
infrafasciatus, 27.
sexfasciatus, 27.
Haploclastus, 354.
Harmonicon, 3438, 345,
364.
Harpactira, 354, 361.
Harpactirella, 354, 356,
361.
Hebestatis, 568,
Heligmomerus, 350, 361,
362, 363.
Heliodrilus, 218, 219.
Hemibungarus
japonicus, 101.
Hermaeba, 348, 349, 361,
365.
Hermachastes, 348, 361.
Hersilia, 566.
Hersiliola, 366.
Hesydrus, 153, 164.
bucculentus, 165.
estabanensis, 165.
habilis, 152, 164, 165,
168.
jJullient, 153, 164.
palustris, 152, 164, 168,
168.
Heterocheta
grimaldii, 124.
longicornis, 124.
major, 124.
papilligera, 124.
zetesios, 124.
Heteromigas, 352, 363.
Heterophrietus. 354, 362.
Heteroscodra, 354, 361.
Heterothele, 354, 357,
361.
Hexathele, 843, 558, 563.
Hexura, 346, 348, 358,
364.
Hipparion, 202.
Hippidium, 202.
Hipposiderus
caffer, 295.
Hirundo
rustica, 5.
Histioteuthis, 77.
TLomalopsis
buccata, 94.
Homcomma, 365.
Homeoplacis, 353, 365.
Homostola, 349, 361.
Huttonia, 367.
Hydroeyon
forskalii, 6.
Hydrophis
cyanocinetus, 96.
diadema, 99.
fasciatus, 95.
gracilis, 95.
melanocephalus,
101.
obscurus, 99.
Hydrus
major, 96.
platurus, 95, 101.
Hylobates
hoolock, 187.
Hyperiodrilus, 218, 219.
Hyperopisus, 8.
Hypbantornis |
cucullatus, 6.
95,
INDEX.
Hypsirhina
bennetti, 94.
chinensis, 94.
enhydris, 94.
plumbea, 94.
Hysterocrates, 354, 361.
Tanthina, 129.
Ichthyosaurus, 179.
Ictonyx
Srenata, 296.
lybica, 296.
Tdioctis, 353, 363.
Tdiommata, 353, 363.
Tdiopbthalma, 353, 565.
Idiops, 350, 352, 360,
361, 862, 363, 565.
Idiosoma, 349, 3638.
Trrisor
erythrorhynchus, 5.
Ischnocolus, 860.
brevipes, 362.
Ischnothele, 343, 3845,
358, 361, 362, 363,
364, 565.
Tsichthys
henryi, 22.
Tsmailia, 104.
Ixamatus, 343, 363, 368.
Jaculus
gordoni, 299.
jaculus, 299.
Kréyeria
(Lonchidium) ga/ii
vulgaris, 108.
Labdacus, 166.
monastoides, 166.
Labeo
annectens, 28, 28.
hosei, 6.
parvus, 24,
Labrus
maculatus, 108.
mixtus, 108.
turdus, 108.
Lacerta, 334.
Lachesis
flavoviridis, 102.
gramineus, 99.
jerdonii, 99.
luteus, 102.
mucrosquamatus, 99,
102.
okinavensis, 102.
Lagochilus
kobelti, 194, 195, 199.
rollei, 198.
|
373
Lagochilus
tounsendi, 195.
Lagothrix, 16.
humboldti, 16.
Lamprocolius
purpureus, 6.
Laniarius
barbarus, 6.
erythrogaster, 9.
Lanius
collurio, 9.
excubitorius, 4, 6.
Lasiodora, 354, 365.
Lates
niloticus, 6, 7.
Latocestus
argus, 802, 512, 313.
pacificus, 313.
Latouchia, 350, 3860, 362,
363.
Leggada
minutoides, 298.
tenella, 298.
Lepthereus, 348, 561.
Leptopelma, 352, 3093,
360.
Leptoplana
alcinoi, 303, 307.
angusta, 308.
aurantiaca, 301.
californica, 307, 308.
chierche, 308.
drwhachensis, 308.
ellipsoides, 308.
Jallax, 308.
kukenthali, 307.
lactcoalba, 808.
malayana, 302, 3806,
308, 318.
nationalis, 308.
obtusum, 302.
pacificola, 306, 3808.
pallida, 308.
panamensis, 308.
pardalis, 307.
subviridis, 307.
tremellaris, 307.
variabilis, 308.
virilis, 3808.
vitrea, 307.
Leptopoma
asptrans, 194.
Leptosoma, 271, 285.
Lepus
ethiopicus, 8C0,
Lernxa
abyssicola. 109.
lumpr, 109.
minuta, 109.
Lernwepoda
extumescens, 109.
374
Lernanthropus
micropterygis, 108.
polynemi, 108.
tetradactylus, 108.
trifoliatus. 108.
tylosurt, 108.
Limulus
polyphemus, 304.
Liphistius, 341.
Lophogaster, 130.
Loxomphalia, 361.
Lucieutia
grandis, 124.
magna, 124.
Lucifer, 53.
Lutra
capensis, 191.
Lybiodrilus, 218.
Lycinus, 343, 364.
Lycocyprinus
sexfasciatus, 27.
Lycodon
aulicus, 88.
fasciatus, 88.
subcinctus, 88.
(Ophites) adbofuseus,
89.
Lyrognathus, 354, 396,
362.
Lystrosaurus, 179.
Macacus, 12, 15, 20.
Macrogale, 328.
Macronyx, 5.
croceus, 7.
Macrothele, 348, 345,
358, 360, 362, 363.
Magulla, 365.
Malapterurus, 8.
Marcusenius
brachyhistius, 22.
sphecodes, 22.
Mastacembelus
congicus, 28.
eryptacanthus, 28.
liberiensis, 28
loennhergi, 28.
marchit, 28.
sclateri, 28, 29.
Masteria, 343.
Mastigopus, 53, 56, 62,
63, 64, 65, 66, 67, 68,
69, 70.
Mecicobothrium, 346,
308.
Megaderma
frons, 295.
Melittophagus
bullockoides, 7.
pusillus, 8.
INDEX.
Melodeus, 343, 845, 364.
Melopeeus, 354, 362.
Meriones
dongolanus, 296.
stigmonyx, 296.
Merops
albicollis, 6.
nubicus, 9.
Metridia
longa, 119, 123.
Metriopelma, 354, 365.
Microbatesia, 349, 361.
Microcephalophis
melanocephalus, 9d.
Micropteryx
dumerilt, 108.
Microsynodontis,
nov., 26.
batesti, 26, 29.
Migas, 352, 363, 368.
Missulena, 352.
Mitothele, 354, 357, 365.
Mogeridgea, 352, 361,
363, 368.
Monocentropus, 354, 361.
Morinyrops, 8.
Mormyrus, 8.
Motella
tricirrata, 107.
Murricia, 366.
Mus, 334.
Musophaga, 263.
Myegale, 341.
Myrmeciaphila, 349, 564.
Myrmecobius
Jasciatus, 200.
Myrtale, 352, 361.
gen.
Naia
bungarus, 97.
tripudians, 97.
Nasalis, 12, 13, 14, 15,
16, 18, 20.
larvatus, 12.
Nebalia, 130.
Nemesia, 348, 360.
Nemesiellus, 348, 362.
Neocteniza, 349, 365.
Neothereutes, 166.
darwini, 166.
Neotragus
batesi, 193.
pygmaeus, 193.
Nephila, 366.
Notoplana, gen. nov., 302.
evansit, 801, 302, 303,
304.
Nyctiphanes
norvegicd,
130, 132.
119) 129;
|
Okapia
Johnston, 3.
Onchnesoma, 38.
Onos
cimbrius, 108.
mustelus, 108.
Ophiocephalus
obseurus, 27.
Opisthocomus, 258, 276,
286.
Opisthoporus
penangensis, 196.
Opisthostoma
annandalei, 198.
laidlawi, 198.
Opisthotropis
andersonit, 87.
Ornithoetonus, 354, 362.
Ornithorhynchus, 326.
Ornithoscatoides
migra, 50.
Orphnecus, 362.
Oryx
algazel, 300.
— daminzh, 300.
beatriz, 300.
leucoryx, 300.
Oryzorictes, 328.
Otiothops, 367.
Ovis
arcal, 103.
arkal, 102.
vigner, 102, 103.
— arkal, 103.
— blanfordi, 103.
—- cyeloceros, 102.
Pachylomerus, 346, 350,
309, 360, 364, 365,
368.
Pagodina
(Nemesis) charcharie
glauci, 108.
Paleohatteria, 179.
Palzornis
docilis, 9.
Palpimanus, 367.
Paimphobeteus,
369.
Pandarus
carcharti glaucus, 108.
musteli levis, 108.
spinacis acanthie, 108.
unicolor, 108.
Pantodon
buchholzi, 21.
Papio, 12.
Paraconchecia
oblonga, 120, 121.
variabilis, 121.
304,
Paradisea
apoda, 192.
Paradossenus, gen. noy.,
153, 155.
nigricans, 152, 155,
Paraphysa, 365,
Parathemisto
abyssorum, 126.
oblivia, 119, 126, 129.
Paratropis, 345, 564.
Pareiasaurus, 179.
Pareudrilus
sp., 220.
papillata, 216, 218,
219.
stagnalis, 216, 217,
5)
Parra
africana, 8.
Parus
leucopterus, 7.
Pelinobius, 354, 361.
Pelmatochromis
batesti, 28.
subocellatus, 28.
Pelmatorycter, 348, 361.
Peraderma
bellotti, 109.
peterst, 109.
Petalidinm
sp., 78.
foliacewm, 54, 55, 56,
69, 78.
obesum, 56.
Petrocephalus
simus, 22.
Phacocheerus, 202.
Pheenothele, 349.
Pheoclita, 849, 365.
Phascolion, 37, 38, 39.
Phascolosoma
abnorniis, 37.
cylindratum, 37.
elongatum, 37.
lobostomum, 37.
sanderi, 37, 38.
teres, 29-41.
vulgare, 37.
Philichthys
doderleini, 108.
fialole, 108.
Philodromus, 366.
Phiogiellus, 554, 356,
362.
Phiegiodes, 354, 362.
Phoneyusa, 354, 561.
Phormictopus, 3605.
Phormingochilus,
362, 363.
Phoronis, 35, 36.
354,
Proc. Zoo. Soc.—19038, Vou. I. No. XXV,
\
INDEX.
Phrissecia, 349, 365.
Phrynarachne
ceylonica, 50.
decipiens, 51.
fatalis, 50.
papulata, 50.
peeliana, 50.
rothschildi, 48, 50, 51.
tuberosa, 50.
Phryxotrichus, 365.
Phyllidia, 251.
Phyllidiopsis, 251,
papilligera, 251.
Phyllocheetupterus, 170,
173
aciculigerus, 170, 171,
172; 176.
claparedi, 171, 172,
173, L75, 176:
eliotz, V7, 172; 173;
175, 176.
fallax, 171.
gardineri, 171, 172,
1738.
gracilis, 171, 175.
major, 171, 172.
pictus, 171, 172, 174,
175, 176.
socialis, 171,
175.
Phymosoma, 33, 34, 35,
36, 38.
varians, 34, 40.
Phyxioschzema, 343, 345,
358, 360.
Piaya, 260, 262, 263, 267,
276, 277, 279, 281, 282,
284, 287, 289, 290.
Pionothele, 348, 361.
Pirata, 158.
Pisenor, 352, 361.
Plagiobothrus, 353, 362.
Plagiotata
promiscua, 305.
Planocera
sp., 801, 302, 318.
insignis, 303.
Platurus
colubrinus, 96, 101.
laticaudaius, 96, 101.
muellert, 96.
schistorhynchus, 101.
Platyctenus, 116.
Plesiophrictus, 354, 362.
Pleuroleura, 252.
Pleuromma
abdominale, 119, 128.
robustum, 123.
Pleuropbyllidiella,
gen. nov., Jol, 252.
horatii, 251.
172,
Ploceus
nigerrimus, 6.
Podon
intermedius, 118, 120.
Pecilia
sexfasciata, 27.
Peecilomigas, 368.
Pecilotheria, 354, 356,
Pceocephalus
meyert, d.
Polyodontophis
collaris, 85.
Polyporus, 310.
Polypostia, 310, 312.
Polypterus, 3, 6, 10, 11,
senegalus, 10.
Polytoreutus, 212, 219.
Porrhothele, 348, 345,
360, 363.
Potamon
africanum, 41, 42, 43,
44, 46.
aubryi, 42, 46.
consobrinus, 44.
decazei, 42.
emarginatum, 42.
Sfloweri, 42.
latidactylum, 42, 48,
44, 46.
pelt, 42, 46,
(Potamonautes)
canum, 47.
(—) latidactylum, 41-
47.
afri-
Prionops
plumatus, 5.
Procavia, 202.
ruficeps, 300.
Proceros
buskii, 801.
concinnumn, 315.
hancockianus, 301.
Procolophon, 179.
Prosthiostomuim
aurantiacum, 302.
pallidum, 302, 317.
siphunculus, 817.
| Protolycosa, 340.
Protopterus, 6, L1.
Psalistops, 353, 365.
Psalmopceus, 365.
Psammodynastes
pulverulentus, 94.
Pselligmus, 349, 365.
Pseudaetideus
armatus, 125.
Pseudidiops, 330, 852,
365.
Pseudoceros
bedfordiz, 301, 3802,
314, 318.
25
376
Pseudoceros
bushii, B02.
cerebralis, 314.
collingwoodii, 302, 314.
hancockianus, 802, 315.
kelaartii, 802, 314.
rubellus, 302, 314, 318.
striatus, 314.
Pseudopareas
vagus, 98.
Pseudoxenodon
dorsalis, 87.
macrops, 87.
Pteraeolidia
semperi, 255, 256.
Pterinochilus, 354, 356,
361.
Pterocyclos
blandi, 196.
subalatus, 195, 199.
Pterygioteuthis
margaritifera,
Ptychognathus, 179.
Pupina
aureola, 197.
lowi, 197.
Pyromelana
. flammiceps, 6.
Python
molurus, 85.
reticulatus, 85.
Tide
Raia
circularis, 109.
radiata, 108.
Rana, 334.
occipitalis, 6.
Ranatra, 158.
Ranzania, 170.
Rhachias, 3848, 365.
Rhamphococcyx, 260,
£62, 263, 264, 266,
267, 273, 274, 287,
289, 290.
calorhynchus, 262, 291.
Rhaphaulus
ascendens, 196, 199.
blunfordi, 197.
lorraint, 197.
perakensis, 197.
—, var. jalorensis, 197, |
199.
Rhea, 269.
Rhechostica, 357.
Rhinoceros
simus, 222, 223, 224.
Rhinococeyx, 259, 260,
262, 264, 266, 267,
DG 209, 26) 282)
284, 287, 288, 290.
INDEX, |
Rhinoeoecyx
curvirostris, 262.
Rhinopithecus
bieti, 224, 225.
brelichi, 224.
roxellane, 224, 225.
Rhiostoma
sp., 196.
houset, 196.
jalorensis, 196, 199.
Jousscaume?, 196.
Rhopodytes, 259, 260,
By, Ve, Vis, Bae,
OFT DisPay DIS, Pei,
289, 290.
Rhynchza
capensis, 5.
Rhytidicolus, 349, 365.
Rianus, 353.
Rousettus
stramineus, 295.
Salmo, 108.
Saltuinus, 154. »
Sarascelis, 367.
‘Sason, 353, 3861, 362,
363.
Sasonichus, 353, 362.
Saurothera, 259, 260,
263, 265, 288.
vetula, 291.
Scalidognathus, 348, 362,
368.
Schistoloma
anostoma, 197.
sectilabrum, 197.
Schizorhis, 259, 261, 262,
263, 264, 268, 277,
278, 283, 285, 288.
zonura, 291.
Sciacaris, 53.
Scina
borealis, 128.
Scodra, 854, 357, 361,
Scopelus
benotti, 109.
Scoptelus
notatus, 8.
Seopus
umbretta, 4.
Scotineecus, 343, 358.
Scyllium, 334.
Scythrops, 266. 268, 272,
278, 274, 275, 276, =78,
279, 281, 284, 287, 288,
290.
Segestria, 366.
Selenocosmia, 856, 362,
363.
Selenogyrus, 354, 361.
Selenops, 158.
Selenothelus, 354, 356.
Selenotypus, 354, 363.
Semnopithecus, 14, 15,
16, 17, 18, 20, 21.
entellus, 14, 15, 16.
Jemoralis, 14, 15.
hypoleucus, 14, 16.
maurus, 14.
priamus, 14, 15, 16.
rubicundus, 14, 15.
Semonia
maculata, 308, 309.
penangensis, 302, 308,
309.
Senoculus, 166.
albidus, 152, 167, 168.
canaliculatus, 167.
darwint, 166, 168.
gracilis, 166, 167.
tricolor, 166, 167.
maronicus, 166, 168.
monastoides, 166, 167.
pallidus, 166, 167.
parallelus, 152, 166,
167, 168.
plumosus, 166, 167.
prolatus, 166, 167.
purpureus, 166, 168.
rubromaculatus, 166.
ruficapillus, 166, 167.
Seothyra, 367.
Sergestes
ancylops, 65, 66, 71.
arcticus, 57, 59, 60, 61,
62, 65, 68, 70, 71,
78.
armatus, 63, 64, 71.
atlanticus, 52, 58, 59,
60, 61, 65, 66, 71,
loA®)
brachyorrhos, 56.
challengeri, 52, 53, 61,
Tes, 1d, To, Tis 7shs
corniculum, 65, 66, 70,
lewis
cornutus, 67.
diapontius, 62, 63, bd,
67, 68, 71.
dissimilis, 70, 71.
dorsispinalis, 62, 71.
edwardsti, 56, 64, 60,
70, 71, 72.
Sermerinkit, 67, 71.
‘frisii, 59.
incertus, 65.
inous, 69.
intermedius, 56, 71.
japonicus, 5d, 57, 58,
Tile
Junceus, 66.
Sergestes
kroyeri, 58, 61, 71, 78.
leviventralis, 68, 71.
laterodentatus, 62, 71.
longicollus. 67, 68.
lonyirostris, 66, 71.
longispinus, 66, 67.
mediterraneus, 70, 71.
mollis, 57.
nasidentatus, 62, 71.
obesus, 56.
oculatus, 64, 65, 71.
ovatoculus, 65.
parvidens, 65.
penerinkii, 63, 65, 67,
Gite
precollus, 68, 71.
prehensilis, 56, 58, 61,
Til Task
profundus, 55, 69, 71,
78.
rinkii, 64, 65, 67, 71.
robustus, O7, 48, 61.
sanguineus, 56.
semiarmis, 68, 71.
similis, 59, 60, 62, 71,
78.
spiniventralis, 68, 69,
71.
tenuiremis, 66, 67, 68,
72.
utringuedens, 70, 71.
ventridentatus, 70, 71.
vigilax, 65, 68, 69, 72.
Sericopelina, 365,
Simotes
chinensis, 93.
cyclurus, 92.
formosanus, 92.
purpurascens, 92.
vaillanti, 93.
violaceus, 92.
Sipalolasina, 3535, 362.
Solenothele, 854, 3057,
361.
Solenotholus, 863.
Spaniopholis
souliet, 93.
Sparassus, 366.
Sphenodon, 179.
Sphyngiceps
lacteus, B01.
Spiocheetopterus, 170.
Spiroctenus, 348, 361.
Splanchnotropus, 104.
brevipes, 104.
gracilis, 104,
Spreo
superbus, 9.
Squalus
acanthias, 108.
INDEX.
Stasimopus, 350, 361.
Stegodyphus, 367.
Stenoterominata,
365.
Stenygrocercus, 345, 345,
360, 363.
Steriphopus, 367.
Sterrhochrotus, 350, 860.
Stichoplastus, 365.
Stictogaster, 549, 361.
Stothis, 3538, 3865.
Stromateus
Jialola, 108.
Stropheeus, 303.
Struthio
asiaticus, 209.
camelus, 204, 205, 206,
207, 208, 209.
chersonensis, 209.
karatheodoris, 203, 204,
205, 206, 208, 209.
Stuhlmannia, 212, 213,
218, 219, 220.
gracilis, 213.
michaelseni, 210, 211.
349,
variabilis, 210, 211,
212, 213.
Synodontis
nigritus, 6.
obesus, 26.
Syntrechulea, 153.
Taccocoua, 260, 263, 264,
267, 273, 274, 276, 277,
278, 281], 282, 284, 287,
289, 290.
Tama, 366.
Tamandua
tetradactyla, 191.
Taphozous
perforatus, 295.
Tapinauchenius, 354, 357,
565.
Tapinophis
latouchii, 87.
Tegenaria, 366,
Telephonus, 7.
senegalus, 6,
Telepsavus, 170, 173.
Temora
longicornis, 119, 123.
Terpsiphone
perspicillata, 6.
Tetragnathus, 366.
Thalassius, 152, 158.
Thalerommata, 365.
Thanatidius, 153, 156.
dubius, 156.
spinipes, 52,
168.
156,
377
Thanatidius
tenius, 156.
undulatus, 156,
Thaumasia, 152, 153.
annulipes, 152,
168.
binotata, 154.
marginella, 153.
seupularis, 154.
scoparia, 154.
sentlis, 158.
velow, 152, 154, 168.
Thelphusa
apa 41, 47.
Theraphosia, 554.
Thomisus, 366.
Thrigmopeeus, 354, 362.
Thyropeeus, 352, 361.
Thysanoessa
longicaudata, 119, 128,
129, 130, 131.
tenera, 128.
Thysanopoda
longicaudata, 128.
microphthalma, 119,
132.
Thysanozoon
allinani, 301, 802.
auropunctatum, 801,
302, 314.
Tigidia, 352, 361.
Tilapia
lata, 28.
nilotica, 6.
zibli, 7.
Tinus, 153.
Tmesiphantes, 365,
Vracheliastes
gigas, 109.
Tragelaphus
scriptus, 6.
Trechalea, 153, 157, 164.
amazonica, 152, 159,
160, 163, 168.
connera, 152, 159, 162,
168.
ellacombei,
161, 168.
extensa, 152, 159, 162,
168.
hahilis, 165.
154,
152, 160,
keyserlingi, 152, 159,
1638, 168.
longitarsis, 152, 157,
158, 159, 160; 163,
164, 168.
macconnelli, 152, 159,
162, 168.
urinator, 152, 158, 159,
161, 162, 163, 168.
Trechoua, 343, 343, 364.
378
Triacanthus
angustifrons, 182,
biaculeatus, 180, 181,
182, 183, 184.
blocht, 180, 181, 182.
brachysoma, 184.
brevirostris, 180, 181,
183, 184, 185.
indicus, 180, 181, 184.
longirosiris, 182.
macrurus, 180, 188.
nieuhofi, 180, 181, 184.
oxycephalus, 180, 181,
183.
rhodopterus, 183.
russellit, 188.
strigilifer, 180,
182.
Trichopelma, 353, 365.
Triclaria, 157.
connexa, 162.
extensa, 162.
hubilis, 165.
Trigonoporus, 310, 312.
Trionyx
triunguis, 6.
Trirachodon, 177, 179.
kannemeyeri, 177, 178,
180.
Trirhinopholis
styant, 88.
Trissothele, 343, 364.
Triton, 332, 334.
Trittame, 353, 863, 368.
Tropidonotus
annularis, 86.
balteatus, 87.
181,
INDEX.
Tropidonotus
chrysargus, 86.
craspedogaster, 87.
nuchalis, 86.
percarinatus, 87.
piscator, 86.
pryeri, 100.
stolatus, 86.
subminiatus, 86.
swinkonis, 86.
tigrinis, 86, 99, 100.
vibakari, 86, 99.
Tropidostoma
dunni, 177.
Tryphana
boecks, 127.
malmi, 127.
nordenskioldi, 127.
Turacus, 259, 26'), 262,
265, 268, 277,
, 283, 285,
288.
buffoni, 278,
leucotis, 291.
Tylosoma, 38.
Tylosurus
impertalis, 108.
Typhlops
braminus, 85.
lineatus, 89.
291.
Unyoria, 219.
Urococcyx, 287.
Uroctea, 367.
Urolestes
equatorialis, 7.
Uruchus, 3438, 345, 364.
THE END,
Velella, 129.
Ventriculina, gen. nov.,
106.
crosslandi, 106, 107.
Vidua
hypocherina, 5.
principalis, 4.
Vinago
waalia, 9.
Vipera
ammodytes, 185.
— meridionalis,
186.
— typica, 185, 186.
raddit, 186.
Vulpes
pallida, 296.
vulpes egyptiaca, 296.
185,
Xenocharax
spilurus, 23.
Zamenis
korros, 90, 100.
mucosus, 90, 100.
spinalis, 91.
Zanclostomus, 262, 263,
277, 287, 288, 290.
Javanicus, 263.
Zaocys
dhumnades, 90, 100.
Zeugopterus
punctatus, 108.
Zilla, 366.
Zophoryctes, 368.
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Frttows pay an Admission Fee of £5, and an annual Contri-
bution of £3, due on the Ist of January, and payable in advance,
or a Composition of £30 in lieu thereof; the whole payment,
including the Admission Fee, being £35.
No person can become a Frttow until his Admission Fee and
First Annual Subscription have been paid, or the annual payments
have been compounded for.
Frttows elected after the 30th of September are not liable for
the Subscriptions for the year in which they are elected.
PRIVILEGES OF FELLOWS.
Frttows have Personal Admission to the Gardens with Two
Companions daily, upon signing their names in the book at the
entrance gate.
Ferttows receive a Book of Saturday and a Book of Sunday Orders
every year. These Orders admit two persons to the Gardens on each
Saturday and two on each Sunday in the year. But the Saturday
3
Orders are not available if the Frtrow shall have used his privilege
of personally introducing two companions on the same day.
Fertiows also receive every year Twenty Free Tickets (Green),
each yalid for the admission of one adult any day of the week,
including Sunday. Children’s Tickets (Buff) can be had in lieu of
Green Tickets in the proportion of two Children’s Tickets to one
Adult's. These Tickets, if not made use of in the year of issue, are
available for following years.
Fettows, if they wish it, can exchange the Book of Saturday
Orders for Twenty Green Tickets available for any day. The Book
of Sunday Orders can also be exchanged for a similar packet of
Twenty Tickets. These books must, however, be returned entire,
and the exchange can only be made during the year of their issue.
The annual supply of Tickets will be sent to each Frttow on the
1st of January in every year, on his fillmg up a form of Standing
Order stating in what way they should be made up, and to what
address they should be sent. Forms for this purpose are supplied
on application.
The Wire of a Frttow can exercise all these privileges in his
absence.
Ferttows have the privilege of receiving the Socicty’s Publications
on payment of the additional Subscription of One Guinea every
year. This Subscription is due upon the Ist of January and must
be paid before the day of the Anniversary Meeting, after which
the privilege lapses. Frttows are likewise entitled to purchase the
Transactions and other Publications of the Society at 25 per cent.
less than the price charged to the public. A further reduction of
25 per cent. is also made upon all purchases of Publications issued
prior to 1871, if above the value of Five pounds.
Frttows also have the privilege of subscribing to the Annual
Volume of the Zoological Record for a sum of £1, payable on the
1st July in each year, but this privilege is forfeited unless the
subscription be paid before the 1st of December following.
They may also obtain a TransrerasLe Ivory Ticker admitting
Two Persons, available throughout the whole period of Fellowship,
4
on payment of Ten Pounds in one sum. A second similar ticket
may be obtained on payment of a further sum of Twenty Pounds.
Any Fritow who intends to be absent from the United Kingdom
during the space of one year or more may, upon giving to the
Secretary notice in writing, have his name placed upon the
“« dormant list,” and will be thereupon exempt from the payment of
his annual contribution during such absence.
Any Frttow, having paid all fees due to the Society, is at liberty to
withdraw his name upon giving notice in writing to the Secretary.
Ladies or Gentlemen wishing to become Fellows of the Society
are requested to communicate with the undersigned.
P. CHALMERS MITCHELL, M.A., D.8c.,
Secretary.
3 Hanover Square, London, W.,
August, 1993.
MEETINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON
FOR
SCIENTIFIC BUSINESS.
(AT 3 HANOVER SQUARE, W.)
Session 1903-1904.
1903.
Turspsy, Novemper 3and17 | ‘Tuxspay, December 1
1904.
Turspay, JANUARY 19 Turspay, Aprin ..19
x Frprvuary 2 and 16 a May ....< Sandy
re IMA CHa saul aaa anit) FAME OED Clee. 7
The Chair will be taken at half-past Hight o'clock in the Evening —
precisely.
LIST OF THE PUBLICATIONS
OF THE
ZOOLOGICAL SOCIETY OF LONDON.
Tn scientific publications of the Zoological Society of London
are of two kinds—“ Proceedings,” published in an octavo
form, and ‘‘ Transactions,” in quarto.
According to the present arrangements, the ‘‘ Proceedings”
contain not only notices of all business transacted at the scien-
tific meetings, but also all the papers read at such meetings
and recommended to be published in the ‘ Proceedings” by
the Committee of Publication. A large number of coloured
plates and engravings are attached to each annual volume of
the “‘ Proceedings,” to illustrate the new or otherwise remark-
able species of animals describedin them. Amongst such
illustrations, figures of the new or rare species acquired in a
living state for the Society’s Gardens are often given.
The “ Proceedings” for each year are issued in four parts,
on the first of the months of June, August, October, and
April, the part published in April completing the volume
for the last half of the preceding year. From January 1901
they have been issued as two half-yearly volumes.
The ‘‘ Transactions” contain such of the more important
commuuications made to the scientific meetings of the Society
as, on account of the nature of the plates required to illustrate
them, are better adapted for publication in the quarto form.
They are issued at irregular intervals.
Fellows and Corresponding Members, upon payment of
a Subscription of One Guinea before the day of the Anni-
versary Meeting in each year, are entitled to receive the
Society’s Publications for the year. They are likewise
entitled to purchase the Publications of the Society at 25 per
cent. less than the price charged for them to the Public. A
further reduction of 25 per cent. is made upon purchases of
Publications issued prior to 1871, if they exceed the value of
five pounds.
Fellows also have the privilege of subscribing to the
Annual Volume of the Zoological Record for a sum of £1
(which includes delivery in the United Kingdom only),
payable on the Ist July in each year; but this privilege
is forfeited unless the subscription be paid before the Ist of
December following.
The following is a complete list of the publications of
the Society already issued. They may be obtained at
the Society’s Office (8 Hanover Square, W.), at Messrs.
Longmans’, the Society’s publishers (Paternoster Row, E.C.),
or through any bookseller.
[ August, 1908. ]
TRANSACTIONS OF THE ZOOLOGICAL SOCIETY OF LONDON,
4to. 15 vols. and Index. Eee
Vol I., containing 59 Plates.... (1833-35) .... £318 6.
x ETL TL ee MESS aly) (aus Mae KONO
5 Lane WRN Aries aes CECE wn 8 8 8.
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po eae Gi) | CRORE RUT ie es a
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aE aaa * (1869-72) 1 A. Oe.
SRVALNTC St, fo gone WOOD) a6 O 8 8.
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BRP OX aie) CORN, (IST TONNe ee OF Olea
Index: vViolshil Ney hie emana (1833-79) Or 6.
Vol. XI., containing 97 Plates.. (1880-85) .... 912 0,
ci oxen Mea \(sschooy ite re some
sy UL, yO) SU EEC UNS Gy
ORY, nar dae. | (18B6SGR NNN Ba 5a cone
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im oxeyale 2) 88g MO CCUIWIS), & 8 0.
Pov ilies Pt. 1 ss ag 6 ope etter USB Gg IL BG.
Cron 5 DB A ACETONE) 48 OI B,
PROCEEDINGS OF THE COMMITTEE OF
CORRESPONDENCE OF THE ZOOLOGICAL
LONDON. 8vyo. 2 vols. (Letterpress only).
Part I.
II.
”
1830-31.
1832.
1 vol. 8vo.
bh}
Price to
Fellows.
Price to the
Public.
ocoocoocoecocococeoqoa
SCIENCE AND
SOCIETY OF
Price to the
Public.
6s.f
6s.
PROCEEDINGS OF THE ZOOLOGICAL SOCIETY OF LONDON.
8vo. 15 vols. (Letterpress only) and Index. (First Series.)
Price to Price to the Price to Price to the
Fellows. Public. Fellows. Public.
Part I. 1833.1 vol. 8vo. 4s. 6d. .. 6s.f | Part IX. 1841.1 vol. 8vo. 4s. 6d. .. 6s.
oy) WO aKeees %p 4s. 6d. .. 6s. ap X. 1842. i 4s. 6d. .. 6s.
JOD, IIs 4s. 6d. .. 6s. i) XI, 1848. on 4s. 6d. .. 68.F
oy) Jue JUSBKGy Hp 4s. 6d. .. 6s. oh eaSIOE Ikul 3 4s, 6d. .. 6s.
30 V. 1837. of ds. 6d. .. 6s. » AIIL. 1845. s 4s. 6d. .. 6s.
» VI. 1838. % 4s. 6d. .. 6s. » AIV. 1846. op 4s. 6d. .. 63.+
oy AUT, 1St38 5 4s. 6d. .. 6s.+ oy OW USELY/, + 4s. Gd. .. 63.
» VIIL 1840. 66 4s. Gd. .. 6s. | Index 1830-1847. > 4s, Gd. .. 6s.
8vo. 13 vols. and Index. (Second Series.)
Letterpress only. With Plates coloured.
Price to Price to the Price to Price to the
Fellows. Publie. Fellows. Public.
Part XVI. 1848. 1 vol. 8vo. 4s. 6d. 65.0 ie £1.08) Or
2 XVII. 1849. 4) As, 6d. 6S. ae 1 0 8 Bead bens Zune (Gh P
} XVIII. 1850. as 4s, 6d. GSio)) dee LS eG 118 Of
. XIX. 1851, op 4s, 6d, So) Hee een 015 9 We Cap
i XX, 1852. 45 4s, 6d. G Susan aay, 015 9 We ih Ober
us XXI. 1853. os As, 6d. SA erase 018 O 1 ASO:
, XXII. 1854, 0 4s, 6d. (OXIA g ihe 019 6G to @ OF
“ XXIII. 1855. . 4s. 6d. CSE ne aes 286 SO,
. XXIV. 1856. 96 As. 6d. GIS ee eat Sea 10 8 Te Che
‘ XXYV. 1857. 5) 4s, 6d. COR ie er lt @ 8 IY eee Giz
“5 XX VI. 1858. 50 As, 6d. Gsei Gas eae 1 i & 2 2 OF
» XXVIII. 1859. As, 6d. CCP a Oa ta 111 6 ZnO
» AXVIII. 1860. , As, 6d. (SPAT ee eae ea mein I bea) (5) 2) oun On
Index 1848-1860, i As. 6d. 6s.
if Ont of print.
PROCEEDINGS OF THE SCIENTIFIC MEETINGS OF THE
ZOOLOGICAL SOCIETY OF LONDON. 8vo. 40 vols. and 4 Indices.
Letterpress only. , With Plates uncoloured. With Plates coloured.
Price to Price to the Price to Price to the Price to Price to the
Fellows. Public. Fellows. * Public, Fellows. Public,
MSGi As Odio cis (OSs alee ate 9s. Dparaa | eee eres W Oosy OO sins. ADaat
EP 5 ACA A Aas (o:tenincrice 9s, Brace LOSE crohns 308, Od. 4... 458.f
MSGS Aa yGd.© 2h .00' 08%. eta Qs. Meleler Loonies semua Od amretie nn cos:
IWstyh oI GEA Sawa OR Goa A toh 5010.0 ty aor reo eyo aouc. CLL
UeGbn ae 465 Gd. ane OSs ems: Stal O21: aie eae spihesiis pong aun
866). 48) (6d. 5,06 Gata. 29s! AIO Ea EMIS ROMO DT De. Grane Come
MSG Tiara ce ois ors ole clade a eee 9s, Mei AS Ae ots Lone OGL eps:
MEGS Resrcetasnpaiceie BAM ROT cr crac Qs, Mets EBS ele eieg. OOSe OL aD:
SCOP eters fi 2 Ne Oo ae enters 9s, BLO sy eee el a BB BE Boon Slap
SAO ea sire ake ecovavereie aiatene ree tee 9s, A Bee cia veteis 338, 9d. .... 408.
Index, T8611 870405, «ceri ements As, Gd..... 68;
IRSA Bemetaebidn or sxaloksbarets os 8 ees Lee. eas d ODai Ole. Ace OR
BS easvarcronre teeters Suacatakeke er stetoretelose Aa hol OAS ee BB Mh gous Ctaar
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TCV Let dhs ite cen ee Bwintelaie ates 9s, mg Te a eee 36s. coe a SRST
NS7ZOMe ae <,- Rt teh Pa Vatarn bi ciote 9s, Be era’ Pee 868: Cloke ene hoes
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MST OWES wiatetets CL rR PC VOHO ERROR RE 9s. Sey OSI a i 36s. saree Gage
MESO eet taucasteleteteh a tea Savevsttvatetorcte:s 9s, LeSaeaariryn OOS dc wrete: (4085
rec Tepietes gia ABs Obes sone. 6s.
ae I RIS Reet eee eeeibicnete aoe wee l2s: ae OOS: Roc beth,
1 Milica sie diners Ay Tebcvaharteenae tals 9s. ls: OSE biijemephose
ISEB Y's GR ohn o oro ORO Une CeC oe an 9s. els eee eve care 36s. 48s.
MOS Arr horelaieh cveraisiniavele.eve cicero otaleset OSs ial sia eee 36s. sivhis SOs
SBS raacten nate piaiatevarastonue matetole 9s. LO Sa rote eieOOSs weeoe:
MS BG ier ncisyes sitar ecrevereemmene 9s. 12s. 36s. . 48s,
NSS Teresi eische ache nachos eerie 9s. Bsc aeeoegnn LAAs ameter 36s AOC oh
1888 .... F ‘ : eistorin ee SDSS Oe 36s seen 4Ogs
TORO errs e kcal ete Stans evchenerers SUDO PUKE elas Cae ae 36s 5 48s,
E90 Rees. neo BOG OO Re 9s. HlZs. so dedermoos nic 48s.
Index, 1881-1890 ........ moon. hs 6d. . Ne OSs
OM rel exaievesaacie acu naksteerevcietes Adoccuoeee snaieie oie ateroNShoterens 36s. aieieay 4OSe
Oe avs veteval oterescuatste oheetetete MESA aR arches; a <elle.\ei'e. eie''e Biosansrersreterons 36s. Sono Cheky
NSO S ee Ae, aie aieeiecheiaterecne Pr ealsiole-savlerns AO CODED Te 36s. Scace 485:
SOAS TASS Wcietersiele/cvero mer TO rales caus og bits seletay OSs A Otten
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WOOT Vee se BAS ee Ore CRATE POTTS MME MOERTSTT ai'claeitie \iMiaiatetatebstec ave . 36s. Aghia ssh.
MOS ee raya cts Sio\cel etal aioe sods NCTC RrCIONSTOMCRCR STS: gavoter che \iosfo: tte cyvsileveveraiere te 36s. e(siensecAons
tet ae eS See nae FORO EO OUG.6 UCU OOO RGR ee oan bic 36s. Seon bet,
OOO ere ert cpe arte Rela CHO RCL TRI COTM e ie peeves wiatbeoletee 36s. wae hos:
Index, 1891=1900) ..)...c.cceauie ne Asi Odessa OS
* No perfect copies in stock t Out of print.
PROCEEDINGS or tor GENERAL MEETINGS ror SCIENTIFIC
BUSINESS or tute ZOOLOGICAL SOCIETY OF LONDON.
8vo. 4 vols.
Price to Price to the
Fellows. Public.
TAIOUA lb Leas oitonon os 15-05. ca. ckAMERTEOaAas sons ose clits 5 CER Blots 5/5) 24s,
. SAILS slofai os sie niaia eRe eos @oneee es Pred ce ica Ee MSSM rsteeeecas 24s,
OOD eee il. Gk aes p che heIPEeronteaviel’s 4 5 n10/ o\teictetelnye, evel wiercrenepaetens liebe 2l8-
9 0 10 re Reh 5.5 ays CRANE ILO Ma os 5. DAS.
TOO Se COT 2 oN, SRO, GME Train NIU 185), eiee 24s,
LISTS OF THE ANIMALS IN THE SOCIETY’S GARDENS.
List of the Vertebrated Animals now or lately Living in the Gardens
of the Zoological Society of London. (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.) 8vo. 1902. Price 6s.
These publications may be obtained at the Socrnry’s Orrick
(3 Hanover Square, W.), at Messrs. Lonamans’ (Paternoster Row,
E.C.), or through any bookseller.
THE ZOOLOGICAL RECORD.
—— 0593, 0o ——_
HE object of the Zoonoarcat 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 to form a repertory that will retain its
value for the Student in future years.
The ‘ Zoological Record’ is published for the Society by Messrs.
Gurney and Jackson at the price of 30s. per volume. But all
Members of the Zoological Society of London have the privilege
of receiving it, including the cost of delivery (within the United
Kingdom), at a subscription price of 20s. per annum. This Sub-
scription 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 sets. The
thirty-seven Volumes to the end of the nineteenth century, and the
Index-Volume 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.
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,
when filled up and signed, should bo sent to the Society’s office for
registration ; it will then be sent to the Agents named therein.
Learned Socicties and Institutions and members of the former
Zoological Record Association are permitted to subscribe to the
‘Record’ on the same conditions as are accorded to Members of
the Zoological Society.
Commencing with Vol. xxx1x., the divisions of the ‘ Zoological
Record’ may be obtained separately, as shown in the subjoined
List.
6b
SEPARATE DIVISIONS OF THE ZOOLOGICAL RECORD.
At present each Volume of the Zootoe1can Recorp consists of
20 separately paged Divisions. Commencing with Vol. xxxtx., for
1902, these Divisions may be obtained separately, in paper covers,
stitched and lettered.
The following are the Divisions and their prices, viz. :—
§
. List of abbreviations and journals
2
HGienerall Subjectem.)o))) sic) Merinn tom eho Werte nn
. Mammalia tte 2
ce OAWOS) 1) Rencraineeie oye 9 Tatee 6
. Reptilia and Batrachia. . 2
Pisces 2
. Tunicata 1
. Mollusca 4
. Brachiopoda .. 1
. Bryozoa 1
. Crustacea 2
. Arachnida ZW Vis aioe Cte aie aie aR ge 2
TBS, iGreen Me MERC doe oo Gon eas | dk
2
3
3
iL
2
2
2
pt
SOMOTIAMNRWON HS
et
bo
yeu
14. Insecta ..
15. Echinoderma
16. Vermes ..
17. Coelenterata ..
18. Spongie
19. Protozoa 3 ROSE ane! aig ae
20, Index of new names of genera and subgenera
SeSOCOROBRDORSOADT®PCSDDCA SOARS
On receipt of the price any Division will be forwarded as soon as
ready. The first part is expected to be ready about May 1908.
These separate Divisions can be obtained from the Zoological
Society, 8 Hanover Square, London. Post-Office Orders should
Le payable to “The Zoological Society,” and may be crossed
“Drummond’s.” Also from the following Agents, viz.: Friedlander
& Sohn, 11 Carlstrasse, Berlin; Gurney & Jackson, 1 Paternoster
Row, London; Librairie A. Hermann, 6 rue Sorbonne, Paris.
P. CHALMERS MITCHELL, M.A., D.Sc.,
Secretary.
August, 1903.
ZooLogicaL Sociery oF Lonpon,
3 Hanover Square, W.
LIST OF VOLUMES or tue ‘ZOOLOGICAL RECORD,’
The Record of Zoological Literature, 1864-1868. Vols. 1.-v.
Edited by Auserr C. L. G. Giwruer, M.A., M.D., Ph.D., F.Z.5., &e.
Price 10s. each Volume net.
The Record of Zoological Literature, 1869. Volume Sixth.
Edited by Axsert C. L. G, Gtnrner, M.A., M.D., Ph.D., F.B.S.,
F.Z.8., &c. Tondon, 1870. Price 30s.
The Zoological Record for 1870-1872, Vols. vir.-1x. Edited
by Atrrep Newron, M.A., F.R.S., F.L.S., V.P.Z.8., &e. Price 10s.
each Volume net.
The Zoological Record for 1873-1883, Vols. x.-xx. Edited by
Epwarp Carpwett Ryz, F.Z.8., M.E.8. Price 10s. each Volume net.
The Zoological Record for 1884, 1885, Vols. xx1.,xxm. Edited
by F. Jerrrey Bert, M.A. Price 10s. each Volume net.
The Zoological Record for 1886-1890, Vols. xxi1.—xxvit.
Edited by Franx KE. Brpparp, M.A., F.Z.8. Price 10s. each
Volume net.
The Zoological Record for 1891-1896, Vols. xxvi1.—xxxm1.
Edited by D. Suarp, M.A., F.R.S., F.Z.S., &. 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. yon
Lendenfeld. Edited (for the Zoological Society of London) by Davin
Suarp, M.A.,, F.R.S., F.Z.8S., &. 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. Boulenger, W. A,
Herdman, E. R. Sykes, E. A. Smith, G. C. Crick, A. W. Brown,
D. Sharp, F. A. Bather, Florence Buchanan, R. T. Giinther, and
R. von Lendenfeld. Edited (for the Zoological Society of London) by
Davin Suarp, M.A., F.R.S., F.Z.8., &c. 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. 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 R. von Lendenfeld.
Hidited (for the Zoological Society of London) by Davin Suarp, M.A.,
F.R.S., F.Z.8., &. 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 Suazrp,
M.A., F.R.S., F.Z.8., &. London, 1901. Price 30s.
The Zoological Record, Volume the Thirty-cighth; being
Records of Zoological Literature relating chiefly to the year 1901.
By J. A. Thomson, Rt. Lydekker, R. Bowdler Sharpe, G. A. Boulenger,
Alice L. Embleton, EH. 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., &c. 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
included in the ‘Nomenclator zoologicus’ of 8. H. Scudder. Com-
piled (for the Zoological Society of London) by Cuartes OwEn
Warernouse and edited by Davip Suarp, Kditor 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’” (88 vols.). Price £15 net; or without ‘Index Zoo-
logicus’ (37 vols.), Price £14 10s. net.
These publications may be obtained at the Socrnry’s OFFIcE
(3 Hanover Square, W.), of Messrs. GurNEY anp Jackson (Pater-
noster Row, H.C.), or through any bookseller.
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.
D)
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 oe Leopoldino-Carolinian Academy of Naturalists,
alle.
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 Wiirtemberg, 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 Athenzeum Club, London.
The British Museum of Natural History, London.
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 Ilinois State Laboratory of Natural History, Ilinois.
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.
ROO
The Publications (except in special cases) are sent out direct as
soon as they are issued. It is requested that they may be ac-
knowledged by the return of the form of receipt sent with them,
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 Savane, Lonpon, W.,
August, 1903.
PROCEEDINGS
OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS.
OF THE
ZOOLOGICAL SOCIETY
OF LONDON.
1903, vol. I.
PART I.
CONTAINING PAPERS READ IN
JANUARY ann FEBR BAR
JUNE 1903.
PRINTED FOR THE SOCIETY,
SOLD AT THEIR HOUSE IN HANOVER SQUARE.
LONDON :
MESSRS. LONGMANS, GREEN, AND CO,,
PATERNOSTER-RO
[Price Twelve ‘Shillings.]
LIST OF CONTENTS.
1003 Ver 1
Part I.
January 20, 1903.
Page
The Secretary. Report on the Additions to the Society's Menagerie in December 1902 -. 1
Mr. P. L. Sclater. On the Zebra-and-Pony hybrid living in the Society’s Menagerie .... 1
My. J. 8S. Budgett, M.A.,F.Z.S. Account of his recent J ourney to Uganda in search of the
Okapi and Polypterus ......++..-2..0.08 ore afar staltasatelegetchaiey's «ire (alevele a eiete e ceeh eneaee 2
1. Note on the Spiracles of Polypterus. By J.S. Bupaurt, M.A., F.Z.8. ........0008 Peete! i
2. On the Brains of Nasalis /arvatus and of some other Old World Primates. By Frank
EL Bepparp, FURS. Ge... eee ee cece ee cece ee ee eee ee te cece eens Pie fy NP Dav kietels 12
3. On the Fishes collected by Mr. G. L. Bates in Southern Cameroon. By G. A.
(BOULENGER, PEER Savile Se o (ealetes lV) iste cic/cic' +(e) «le clelsivieler elle «ec eyeliner 21
4, On the Anatomy of the Gephyrean Phascolosoma teres, n. sp. By W. K. Hurton,
M.A., M.B., Senior Demonstrator of Anatomy in the University of Soe!
(Plates (VE V Ue ei rh teretc eee Rei eisi soc 'm » ie) 0 \ein'e 5: re enenenenS g aete 29
5. On Potamon (Potamonautes) latidactylum, anew Freshwater Crab from Upper Guinea.
By Dr..dG: pr Man, of Terseke: Hollandse (Plate UX:)/2)0% sin. swale esters cee eee 4]
6. On a new “ Bird’s-dung” Spider from Ceylon. By R. I. Pococx, F.Z.8., and the
voetion. Ni Ce RopHSCHInD,bAt JHEH Seah 27S. \(bIAtecXs)\ ccs): se lelete clans «ate elaelageele . 48
7. On the Orustaceans of the Genera Petalidium and Sergestes from the ‘ Challenger,’ with
an Account of Luminous Organs in Sergestes challengeri, n. sp. By Dr. H. J. Hansen
(Copenhagen). (Plates XT. G XU.) oie ces ewes seams cae cies sues Sete oat allen e
February 3, 1903.
The Secretary. Report on the Additions to the Society's Menagerie in January 1903.... 79
- 1, Notes on the Hair-Slope of four Typical Mammals. By Water Kipp, ZS. oie dcceoe is he
2. A Prodromus of the Snakes hitherto Gecneded from China, Jape and the Loo Choo
Islands; with some Notes. ey, Captain F, Watt, Indian Medical Service ........ » 84
Contents continued on page 3 of Wrapper.
ContENTS (continued).
Deas February 3, 1903 (continued).
' 8. Note on the Wild Sheep of the Kopet-Dagh, By R, Lypexxer......... IY ek 102
4. On new Parasitic Copepoda from Zanzibar and East Africa, collected by Mr. Cyril
Crossland, B.A., B.Sc. By Staff-Surgeon P. W. Bassert-Saitn, R.N., F.Z.S. ...... 104
5. On the Original Home of the Tiger. By Col. C. EB. Srewart, O.B.,0.M.G.,C.LE, .. 109
6. On the Mode of Copulation of the Indian Elephant. By H. Stapr, Conservator of
SURES NEAVERVO, DUEMS 0.0’ c'e'een ares 'cluldie ¥'e ca cnse'ss au.es see ee us os ca tijeets weds 11]
7. On the Ceelenterata collected by Mr. C. Crossland in Zanzibar.—I. Ceratella minima,
n. sp. By Sypyey J. Hickson, M.A., F.RS., F.Z.S., Beyer Professor of Zoology
in the Owens College, Manchester. (Plate XIII.) .......... ele eluveich qretaid’ ele "sala iataigiata 115
8. Contributions to our Knowledge of the Plankton of the Faeroe Channel.—No, VIII.
By iG. Hurserr Hownmr B.A:) PHD EZ.S. oe secs clon ren sane cciss cumelesen clans 117
9. On the Present Condition and Habits of the Elk in Norway. By H. J. Exwes, F.R.S.. 133 |
February 17, 1903.
Mr. R. E. Holding, Exhibition of, and remarks upon, some Skulls of Mammals showing
abnormal dentition..........+ MMMIADTR Elan imelca se «sce 60 00ceun b¥icen een ean 151
1. On some new Species of Spiders belonging to the Families Pisauwride and Senoculide;
with Oharacters of a new Genus. By Freperick Pickarp-Camsrines, B.A,, F.Z.S.
CE Tabes 2c LV ce ee nme reels vinle te oo cs os ce nescence dccsietipaccinusviw ele cvs mmeUE
bo
On the Marine Fauna of Zanzibar and British East Africa, from Collections made by
Cyril Crossland in the Years 1901 and 1902.—Polychxta. Part I. By Cyr Orosstanp,
B.A., B.Se, (Plates XVI. & ee leSsdtis v Gulviege ook «ec 0:0 eau «aie hie en 169
8. On the Axis, Atlas, and Bioatlas in the lg Theriodonts. By R. Broom, M.D.,
B.Se,, C.M.Z.8. (Plate XVIII.).. Wied c.cseinsoie.wicle's eetsituatetertle =/ere ciate als aie isan nan
4, A Revision of the Fishes of the Genus Triacanthus, By O. Tare Ruaan, B.A,........ 180
or
. On the Geographical Variations of the Sand-Viper, Vipera ammodytes, By G. A.
BovureEncnr, F.R.S., V.P.Z.S. .
6. Notes on the Habits of the Hoolock. By Gro. Canpier, M.B.Cantab, ....0+++0++.0+ 187
LIST OF PLATES.
1903; Oe
PA Tylon:
Plate Page
I. 1. Alestes intermedius. 2. Alestes opisthotenia. 3. Ammphilius \
UQIUGUMOSURIS selene tie «4. 0)< (=:0'e oficiales ipod hte taielabele | e(e: ==) siete erate
II. 1. Labeo annectens. 2. Barbus teniurus ....0.eec.ssseeeee \ 21
Il. 1. Barbus progenys. 2. Barbus hatesit .. 11.00... ..eeee cee
LV.) -Microsynodontis Odtesit 0.0. oc. cvicces o> == 0+-svemsiance ou ee |
V. 1. Anabas pleurostigma. 2. Mastacembelus sclatert ......-... )
VALS
VII. | Anatomy and Histology of Phascolosoma teres ..........++.- 29
VIII.
IX. Figs. 1-6. Potamon (Potamonautes) latidactylum. Figs, {-9.
Potamon (Potamonautes) africanum «2-.eceecesereveee 4)
X. Phrynarachne rothschildi ........ SBE COU SO OOTOAe ra ecco - 48
XI. Fig. 1. Petalidium foliaceum. 2. Petalidiumjun. 3. Sergestes |
profundus. 4. 8. prehensilis. 5. S. kroyert. 6. 8. Be 52
XID. Big. 1. Sergestes arcticus. 2. S. challengert 0.02.22 .eus.--
RT Cer atella Went Wee clots eece 62 Soke. eles naelekae Dee eter 113
XIV. Spiders of the Families Pisauride and Senoculide .......... } 151
XV. Spiders of the Family Pisauride ......-.--.2+...02.0ceeeee
XVI. Figs. 1, 2, 3, 4,7, 8. Phyllochetopterus elioti. Wigs. 5, 6, 9.
JEL GURUE Sodio Coc oho genase AEE OSA adivooo sce 169
XVII. Phyllochetopterus €lt0tt 12.2060. c ee cece eset cence ceee
XVIII. Axis, Atlas, and Proatlas of Gomphognathus and Trirachodon. 197
NOTICE.
The ‘ Proceedings’ for the year are issued in four parts, forming two yolumes
as follows :—
VOL. I.
Part I. containing papers read in January and February, in June.
II. re » March and April, in August.
VOL. Il.
Part I. containing papers read in May and June, in October.
Ts n ie », November and December, in April,
PROCEEDINGS
OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
ZOOLOGICAL SOCIETY |
OF LONDON. 3
1908, vol. I.
PART IT.
CONTAINING PAPERS READ IN
MARCH anv APRIL.
nsomtian Insziy,,,,
a T5 72 ah
SEP 28 1903
National Muse A
AUGUST 1903.
PRINTED FOR THE SOCIETY,
SOLD AT THEIR HOUSE IN HANOVER SQUARE,
- LONDON:
MESSRS. LONGMANS, GREEN, AND CO.,,
PATERNOSTER-ROW.
LIST OF CONTENTS.
1903.—Vot. Tat:
Part II,
March 3, 1903.
The Secretary. Report on the Additions to the Society's Menagerie in February 1903..-.
The Secretary. Extract from a, letter from Major C. Delmé Radcliffe concerning skins of
an Monkey,and, an Otter. fromyu manda eteands 0+ viasinielsine ciein hele: selleia) aia tieteie aie c opetee <
Mr. F. B. Beddard, F.R.S. Exhibition of a mounted skin of the Gea: Bird of Beans
Mr. J. L. Bonhote, F.Z.8. Exhibition of, and remarks upon, a phone of Elephants
showing considerable growth of hair, and note upon the Sanskrit name of the Tiger..
Prof. F. Jeffrey Bell, F.Z.8. Exhibition of, and remarks upon, a Holothurian of the Genus
ee Sie cero seyeintelivlereyere clekc v= cevenctatete a
eo eoo ee ee cee see ee oe ee ee eo ee eeee re FTF OS oO
1. On a new Species of Pigmy Antelope of the Genus Neotragus from the Cameroons
District, W. Africa. By W. E. pp Wivtoy, F.Z.S. (Plate XIX.) -...+.........--
2. On the Land Operculate Mollusca collected during the “ Skeat Expedition” to the Malay
Peninsula in 1899-1900. By EH. R. Syxus, F.Z.S. (Plate XX.) ............. Saternte
3. The Significance of the Callosities on the Limbs of the Equide. By R. Lypzxxnr, F.Z8.
4, Note on some Remains of Struthio karatheodoris Maj. of the Island of Samos. By Rupour
Martin, of Basel University .-..00...0cee0-e 50 %
5. On a new Genus and two new Species of Harthworms of the Family Hudrilide, with
some Notes upon other African Oligocheta. By Franz E, Bepparp, M.A, F.R.S.,
TRASH His SRD EL CE RCCRT ERO Oe Boia 86 eae ead pabie
@eecv cece ee ce eoceeecsecerece @oe
March 17, 1908.
Prof. Newton, F.R.S. Exhibition of, and remarks upon, plotogmphs 0 of the White
Rhinoceros taken by Mr. OC. R. Saunders, C.M.G., in Zululand... seers
Mr. Oldfield Thomas, F.R.S. Exhibition of a skin and Beant of anew species of
Monkey, Rhinoptthecus br elichi, (Plate XXI.)
eves oce ee se eo so ee oo eer ese sees se ce
Page ub
191
199
203
210
Contents continued on page 3 of Wrapper.
ConTENTs (continued).
March 17, 1903 (continued).
NTE RCH Seal CL ETIGETCR IOP TTIC Mb w'e \iui's, oe sn aire bib W OUe whe/s. dw alate able ep vceew-s
Page
Mr, Oldfield Thomas, F.R.S. Exhibition of specimens and re of a new species of
ea Cephalophus ignifer.. wei eh Utara WEOe oeaetiy' 6 G''elale ie: sige ate\ahctamet aeole aeeretets 225
1, Observations and it ont on Japanese Long-tailed Fowls{ By J. T. Constnanam,
M.A., F.ZS. . eee tee ee ere rennet Peewee ee ee FOO eee ewe e ree eeereeeewn ere ewenn ee 237
2. On some Nudibranchs from Hast Africa and Zanzibar.—Part IT. ae Sir C. Error,
K.C.M.G., H.M. Commissioner for the East Africa Protectorate, F.Z.S 250
8. Contributions to the Osteology of Birds.—Part VI. Cucwliform2s. By W. P. Pycnrarr,
MEATS) (EIRCG Bc LL.) og eons soldimipiee inc a's oShea ne ealmetWbre ile aly’aat in iiale 258
April 21, 1903.
The Secretary. Report on the Additions to the Society's Menagerie in March 1903.... 292
1, Linnus and Hunter on Feather-Tracts. By Henry Scnerren, F.Z.S, «+. .000s 292
2. On some Mammals collected by Capt. H. N. Dunn, R.A.M.C., in the Soudan. By
OxupFietp Tuomas, F.R.S., F.Z.8. -..... aere awe SA SEAR ge ie ie ROPE fend Ch tr 294
3. On a Oollection of Turbellaria Polycladida from the Straits of Malacca, (Skeat
Expedition, 1899-1900.) By F. F. Larpiaw, B.A.Cantab., Assistant Lecturer and
Demonstrator in the Owens College. (Plate XXIII.) ...... 0... cece cece ereenees 301
4. On the Phylogenetic Cause of the Transposition of the Testes in Mammalia: with
Remarks on the Evolution of the Diaphragm and the Metanephric Kidney. By W.
Wooptanp, University College, London..........:seeee severe eeeeees NEP a prneiia3e 319
5, On the Geographical Distribution of Spiders of the Order Mygalomorphe. By R. I.
BOGOGRaTH: Aisa” erajelless/ eles cisieteseieie Baila, « '6 ciate el Anis eiakerhiahe eae Oma oie earn falas 340
Index os ccsscessee REE e tae cd, va were ema mete ae oo 6 ky Sle guna Wino «) iphOeleeN iy Be area cae 369
Titlepage .......... ae 1c Ua WE rs vee Gah wean ee
RPM Cir On MAC CN CBRGy 4 «J c'sine's edad + .0's oeteeg.ci's Sled pnlamiaeiia pe «sain Reh eee ii
ERAEREL QICREUOILLH «1 t)/ SECROIA slaieials cla e/a des wia'dsle seh ca oe se cbiggmalawisieme va stevdvenennis iii
* Erratum ......e00. 000: RPT Ee oo ose oanaiaheis oo. dle sibyacerel GR CentOS a" eine ce abaieinete) 6, 8 viii
Mipibeticdl Tnstiog Conthibutors ......06....ecenesce sect camy ips dipeiesasee see}. ix
UNIS EAP PES EUR MPEER Sf clbl cio la'usa(s's,cie's v's ejsle'sva\a ie elvis waye's «uit dts wiQnaaivelsc siaitlr sible Fa ORM xv
PARADOR RRES org Sw els ceed sececnyscnveteec¥iguean Sapien eerajanee sg bs cene xvi
LIST OF PLATES. or
1903.—VOL. I.
PAC oalek
Plate Page
DTK NGotr agus Oates.) steele cie's\ete ein sieve iene neste) seis i in afeke ate tee ee 192
XX. Mollusca from the Malay Peninsula ..-...--...+0. pee 194
PNONGL BPI 2WOPIENECUS ONCLICHY amr els toreleretn tleyategatorer= alesis tekelotelop kate 224
XXII. Osteology of the Cuculiformes ........00 0 ee tees cece ee ee ee 258
XXIII. Polyclads from the Straits of Malacca .....-.....-0-0-..00 301
NOTICE.
The ‘ Proceedings’ for the year are issued in four parts, forming two volumes,
as follows:—
VOL. I.
Part 1. containing en read in January and February, in June.
aly, a HA ». March and April, in August.
VOL. II.
Part I. containing papers read in May and June, in October.
1th # = » November and December, in April,
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