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PROCEEDINGS 


OF THE 
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS 


OF THE 


ZOOLOGICAL SOCIETY 
OF LONDON. 


1913, pp. 339-1104, 


witH 64 PuLatres and 128 Texvr-FIGtres. 


| : 230453 
PRINTED FOR THE SOCIETY, 
SOLD AT ITS HOUSE IN REGENT’S PARK. 
LONDON: . 
MESSRS. LONGMANS, GREEN, AND Co.,, 
PATERNOSTER ROW, 


Gy JL Sy 


OF THE 


COUNCIL AND? OF FECERS 


OF THE 


LOOLOGICAL 


SOCIETY 


OF LONDON. 


Ie, 


COUNCIL. 


His Grace Tue DuxKe or Beprorp, K.G., F.R.S., President. 


Sir JoHN Rosé 
IR CIE IN EGID 
F.R.S., Vice-President. 

Ricuarp H. Burne, Esq., M.A. 


Surg.-Genl. Str R. HAvetock | 


Cuarugs, G.C.V.O., M.D. 
THe Rv. Hon. tree Haru 
Cromer, P.C., G.C.B., 


BRADFORD, | 
DE Scea 


| 
| 
| 
| 


OF | 


Srrk WALTER Roper LAWRENCE, 
Bt., G.C.L.E. 

Sir Epmunp G. Loprr, Bt., 
Vice- President. 

Prof. Ernest W. MacBribs, 
MeAGy (DSc, oHARsS: /ace- 


President. 


| Prof. Epwarp A. MuUINcHIN, 


GOW, KOR, BIAS, | 


Vice-President. 


F. G. Dawrrey Drewirr, Esq., | 


M.A., M.D. 
CHARLES DRUMMOND, 
Treasurer. 
Sir Epwarp Duranp, Bt., C.B. 
FREDERICK GILLerr, Esa. 
THE LorpD GLENCONNER. 
F. Du Canr Gopman, 
D.C.L., F.R.S. 


EKsq., 


Ksq., 


| 
| 


M.A., F.R.S., Vice-President. 


| P. Cuatmers MircHe.y, Ese., 


M.A., D.Sc., LL.D., F.B.S., 


Secretary. 


| W. R. Ocitvir-Grant, Esq. 


ApRIAN D. W. Potiock, Ese. 

OLpFIELD THomas, Esgq., F.R.S. 

AntHOoNY H. WINGFIELD, 
Ksaq. 

Henry WoopwArp, Ksq., LL.D., 
F.R.S., Vice-President. 


PRINCIPAL OFFICERS. 
P, Cuatmers MircHetn, M.A., D.Se., LL.D., F.RS., 


Secretary. 


Frank E, Bepparp, M.A., D.Sc., F.R.S., Prosector. 

R. I. Pococn, F.R.S., F.L.8., Curator of Mammals and 
Resident Superintendent of the Gardens. 

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

Epwarp G. BouLencer, Curator of Reptiles. 

Prof. H. Maxweiu Lerroy, Honorary Curator of Insects. 

Henry G,. Purmmer, F.R.S., M.R.C.S., Pathologist. 

Henry G. J. Peavor, Librarian and Clerk of Publications. 


JoHn Barrow. Accountant. 
W. 4H. Cots, Chief Clerk. 


LIST OF CONTENTS. 


1913, pp. 339-1104. 


EXHIBITIONS AND NOTICES. 


The Secrerary. Report on Additions to the Society’s 
Menagerie during the month of March 1913 ......... 


Mr. H. J. Etwes, F.R.S., F.Z.S. Exhibition of the Head 
of an Asiatic Wapiti. (Text-figs. 83 & 84.) ............ 


Mr. R. H. Burne, M.A.,F.Z.S. Exhibition of malformed 
Antlers of an Axis Deer (Cervus aitis) ............-.-::: 


Mr. E. G. Bousenerr, F.Z.8., Curator of Reptiles. Exhi- 
bition of living specimens of the  Leaf-Insect 
(CMO GPO OUCOHIO)) Acco dab ccaees Bobs sscouedgankucéodeLee 


Mr. E. G. Boutencer. Exhibition of a living melanistic 
specimen of the Green Lizard (Lacerta viridis) ...... 


Dr. R. Broom, C.M.Z.S. A new Species of Golden Mole. . 


Dr. R. Broom. Exhibition of living specimens of the 
female and young of the South African Lizard, 
EUUEROUS Chua EmOUCOIGE (Vl IUD, OSIIN))) Gaadoncedoossosasrose 


The Secrerary. Report on Additions to the Society’s 
Menagerie during the month of April 1913 ............ 


The Rev. T. R. R. Sraesine, M.A., F.R.S., F.Z.S. Notice 
of Prof. F. KE. Schulze’s ‘ Nomenclator Animalium’... 


The LrprariaAn. List of dates of publication of the early 
parts of the Society’s ‘ Transactions’ ...............0.068. 


546 


lv 


“Page 
Mr. R. E. Hotpine. Exhibition of Antlers, Skulls, and 
Photographs illustrating Variations in the growth of 
the Antlers of Deer. (‘Text-fig. SAS) tector cohen eee 815 
Ir. D. Sera-Smira, F.Z.S., Curator of Birds. , Exhibition 
of the Egg and Young of the Mikado Pheasant 
(COMO DOCSIS TOUBEUD). 95 3ccdacd00e gaa506 do00 00 0bce scene oonconaae 818 
The Secrerary. Report on Additions to the Society’s 
Menagerie during the months of May, June, July, 
August, and September OMe tear ake cto ean eee 1090 
The Secretary. Exhibition of the Alimentary Tract of a 
young Elliot's Pheasant (Phasianus elliott) ............ 1094 


Mi. R. I. Pocock, F.R.S., F.L.S., F.Z.8., Curator of Mam- 
mals. Exhibition of a female and three young speci- 
mens of the Woolly Opossum (Philander laniger) ... 1094 


Mr. R. I. Pocock. Exhibition of two youne Degus 
(Cctodonidegus) ern raa. eee ceneee sat Secure dasaysecete ek 1095 


Mr. D. Seru-Suira. Exhibition of a spirit specimen of a 
young Toucanet (Selenidera maculirostris) showing 
the well-developed serrated Heel-pads. (Text-fig.191.). 1095 


Mr. C. Tare Reaan, M.A., F.Z.8. Exhibition of a speci- 
men of a Deep-sea Angler-Fish, Melanocetus johnsonii 


Giinnbly,. 39255 ats sie coe oe ee ere tT ee 1096 
My. E.G. Boutencer, Exhibition of a young specimen of 

the Matamata Terrapin (Chelys fimbriata) ......4..2.. 1097 
Mr. D. Sera-Smirn, Exhibition of a hybrid Black=winged 

Peafowl (Text-fig. 192), a hybrid Pheasant, and @ 

hybrid Jungle- Bowlin eer ne tie tte eee 1098 
Mr. R. I. Pocock, Exhibition of Armadillos to show the 

Dorsal Glands. (Ci sseotai NEB HOG)).  coeocsousbscocoode 1099 
The Secrerary. Report on Additions to the Society’s 

Menagerie during the month of October 1913 ......... 1108 
Prof. H. Maxwent Lerroy, M.A., F.Z.S, Remarks open 

the New Insect Efonee Sonu spbudacuna dacHAosEsaossedeoaes 1104 


Mr. R. H. Burne. Exhibition of specimens of Foetal 
Skeletons prepared by the Beale-Schultze method ... 1104 


28. 


29. 


30. 


31. 


34, 


35, 


PAPERS, 


. New Species of Rhopalocera from Costa Rica. By 


WAMLTAM SCHAUS. EIZ.S5) (Rist WeiiVi.)y-2-5-.25-2. 


. A Collection of Fishes made by Professor Francisco 


Fuentes at Easter Island. By C. Tare Reean, M.A., 
Le Acer jee {CLEA EBS UD: RRR RB iene alsin Ah Tiber Heap 


A Revision of the Fishes of the Genus Auhlia. By C. 
Tate Reean, M.A., F.Z.S. (Text-figs. 68 & 69.) 


The Affinities of the Antarctic Wolf (Canis antarcticus). 
By R. I, Pococn, F.R,S., F.L,S,, F.Z.8,, Curator 
OH Wilerairmelis,  (UNeexhies) (Oi) nonpAbpasubeooecoanbee se 


On the Patella in the Phalacrocoracidee. By Dr. R. W. 
EU PEED IE CoV eZ anes (dca [UIXGIs) Wa sa vdaseee sake seeecits- 


Experiments on the Metamorphosis of the Mexican 
Axolotl (Amblystoma tagrinwm), conducted in the 
Society’s Gardens. By E. G. Bourencer, F.Z.S., 
Curator of Reptiles, (Text-figs. 75 & 76,) ...,........ 


. Contributions to the Anatomy of the Ophidia. By 


Surgeon JosepH C. THompson, U.S.N, (Text-figs. 
TES Ths) eecene anh 2 voller oie sah oes POV OP a a aa 


The Polyzoa of Waterworks, By Srpney F. HARMeEr, 
SGD), Une dd4isy | (dallsaMUDCUL, we LpQUE Le) eee nade 


The Marine Fauna of British Kast Africa and Zanzibar, 
from Collections made by Cyril Crossland, M,A,, 
B,Sc., F.Z.8., in the Years 1901-1902. Bryozoa— 
Cheilostomata. By ArtHur Wm, Waters, F,[.5., 
F.G.S, (Pls. LXTV.-LXXIII. and Text-figs. 79-82.) 


Notes on Albinism in the Common Reedbuek (Cervicapya 
arundinum) and on the Habits and Geographical Dis- 
tribution of Sharpe’s Steenbuck (Raphiceros sharpei), 
By Major J, SrevEnson-Haminton, C.M.ZS, ....0010 


Page 


399 


368 


374 


382 


393 


403 


414 


426 


d37 


36. 


37. 


39. 


40. 


4]. 


43. 


44, 


45. 


A6. 


vi 


Contributions to the Anatomy and Systematic Arrange- 
ment of the Cestoidea.—X. On Two Species of Tape- 
worms from Genetta dongolana. By Frank KE. 
Bepparp, M.A., D.Sc., F.R.S., F.Z.S., Prosector to 
themsociety.sa(ext—tlesn8o—04,)\erne re eeseeeeE ee eeraee 


Pacific Salmon: An Attempt to evolve something of 
their History from an examination of their Scales. 


By Jonn ApAmM Mitne. (Text-figs. 95-118.) ......... 


. Notes on Peripatoides woodwardi Bouvier. By Karu- 


TARE NAD DOIN pcos eae elas eee teeta 


Field-Observations on the Enemies of Butterflies in 
(Cleydlarats 1837 di, Chie IMirwaoin, WING, INIIUSE S54 s¢ocnenae 


On the South-African Pseudosuchian Huparkeria and 
Allied Genera. By R. Broom, M.D., D.Sc., C.M.Z.S. 
(TIE) DDO. Viral BD.O.D. Gaia oe Shean peabaiaanas po5cocgace codon 


On some Cases of Blindness in Marine Fishes. By 
SG yea Deel Bo C1 ba ON Nseries oe Precine ebh A era ec 


2. Notes on Turacin and the Turacin-Bearers. By 


Sir ARpHpR JE CHrurcH KUCAVCOL SROs) aaeeeereeee 


Observations on the Anatomy of the Shoe-bill (Bale- 
niceps rex) and allied Birds. By P. CHALMERS 
Mircueni, M.A., D.Sc., LL.D., F.R.S., Secretary to 
the Society. (Pls. LXXX.-LXXXIII. and Text- 
LS Sel O13.) Cee crctets wetioceeceo ah et see eee EERE ae 


The Classification and Phylogeny of the Calcareous 
Sponges, with a Reference List of all the described 
Species, systematically arranged. By Arruur Denby, 
D.Se., F.R.S., F.Z.8., and R. W. Harotp Row, B.Sc., 
HES. -(@ext=fig. 133.) 02s a.en foi ngaie Wo teenie ees 


The Transvaal Race of the Cape, or Khama, Hartebeest. 
By R. Lyprxxker, F.R.S., F.Z.8. (Text-fig. 135.) ... 


On a Collection of Mammals from the Inner Hebrides. 
By G. E. H. Barrerr-Hamitton, F.Z.S., and Marrin 
A. C. Hinton. (Pl. LXXXIV. and Text-figs. 136- 
BGP) sec ca We gndsce oi dokets scot BEE eee eee 


Page 


D49 


572 


611 


613 


619 


634 


639 


644 


704 


818 


47. 


48. 


49, 


5]. 


BI 
O1 


vil 


Some Miocene Cirripedes of the Genera Hevelasma 
and Scalpellum from New Zealand. By Tuomas H. 
Wiruers, F.G.S. (Pls, LXXXV. & LXXXVI. and 
Memes a OAL) Clea saecijcmacscieen vide wesctoed valnceees 


Observations on Osteomalacia in the Zoological Col- 
lections of Manchester and Cleveland. By T. Wrneate 
Topp, M.B., F.R.C.S. (Els. LXXXVIT.-LXXXIX.) 


Contributions to the Anatomy and Systematic Arrange- 
ment of the Cestoidea.—XI. On a new Tapeworm 
from Wdienemus. By FRANK EK, Brpparp, M.A., 
D.Se., #.R.S., F.Z.8., Prosector to the Society. (Text- 
Heese oe MEA ME ae ac esas Sarai As tse else anaes Sela 


. The Brain and Brain-Case of a Fossil Ungulate of the 


Genus Anoplotherium. By R. W. Paumer, M Sc. 
(Manchester), Research Fellow in Zoology, University 
College, Reading. (Text-figs. 150-157.) ..,..,......... 


The Fossil Crinoils referred to Hypocrinus Beyrich. 
bya 105 Avo letemenaic. | WIS, IDES @ye Sita Nasi N/A iS 
(PIX and Mext=fiess VOSNG602) 5 eet eee se cnc cces- 


- On Freshwater Decapod Crustacea (Families Potamo- 


nide and Palemonide) collected in Madagascar by 
the Hon. Paul A. Methuen. By W. T. Catman, 
D.Sc., F.Z.8. (Pls. XCI. & XCII. and Text-fig. 161.) 


A New Trematode of the Genus Lechriorchis from the 
Dark Green Snake (Zamenis gemonensis). By Marre 
Y. Lesour, M.Sce., Assistant Lecturer and Demon- 
strator in Zoology, Leeds University. (Pl, XCIII.) , 


. Cirripedes from the Cenomanign Chalk Mar] of Cam- 


bridge, By Tsomas H, Wiruers, F.G.S. (Pls, 


. Batrachiderpeton lineatum Hancock & Atthey, a Coal- 


Measure Stegocephalian. By D, M, 8. Warzson, 
M.Sce., Lecturer on Vertebrate Paleontology in Uni- 
versity College, London, (Pls. XCVI, & XCVII, 
aide Dextre saat OL MOMs) caccecnutsasesecse meer niseieriens ; 


. On two New Actinians from the Coast of British 


Columbia. By J, Puayrarr McMourricn, C.M,Z8, 
as ce ONY LETS) “oopane oooenbupvonunseonenee on popapne pevevegvenes 


Paga 


840 


895 


878 


894 


914 


933 


937 


949 


963 


Vill 


Page 
57. Sponges in Waterworks. By W.N. Parker, Ph.D., 

F.Z.S., Professor of Zoology, University College, 

Wand iity A fugetcith..d Jaded Riehoretnareacecnee sean eae 973 
58, A Revision of the Cyprinodont Fishes of the Subfamily 

Peciliine. By C. Tare Recan, M.A., F.Z.8. (Pls. 

NC X= Cian Mext-figsilGS= ids) eeeceneeer sesso: Sa 
D9. On a Collection of Batrachians and Reptiles made by 

Dr. H. G. F. Spurrell, F.Z.8.,in the Choco, Colombia. 

By G. A. Boutencer, F.R.S., F.Z.8. (Pls. CII.= 

CWWaltisicvand Wext- tose (4 lis) i cepecee sees cere eck 1019 
60. The Peroneal Muscles in Birds. By P. CHALMERS 

Wilewornsine, = Wl oNGy IDES, IbIELIDL, GIRS, | Arsen 

Secretary to the Society. (Text-figs. 179-190.) ...... 1039 
61. On the External Characters and Biology of Bryde’s 

Whale (Balenoptera brydei), a new Rorqual from 

the Coast of South Africa. By Oran OLsEN, 

Zoological Laboratory, Christiania University. (Pls. 

CTE FOTN) so cis Sebi pa es Werder gael wae dene) ok eee 1073 
Alphabetical Mist) of Contributors) 2. «ssa. sseeseseeteneees 1X 
iNew. Generic WBC Siem. o.jshesr scrarccken tle cstroc Seeieee score ops xvi 
TroGlebe Git SONGTTUAUINS INEWINGS — dacooonpoonscodaonodonodoodobodeodoedse X1X 


indexvotqlilustrabvonssicats ccecee tes coe ie eeioe ees xlii 


ALPHABETICAL LIST 


OF THE 


CONTRIBUTORS, 


With References to the several Articles contributed by each. 


(1913, pp. 339-1104.) 


Page 
Barrett-Hamizton, G, E. H., F.Z.S., and Hryton, Martin 
A.C. 


On a Collection of Mammals from the Inner Hebrides. 
(Pb TOO ein) eesti USJONGISh) bcoeeseoncoseceonacas 821 


Batuer, Francis A., M.A., D.Sce., F.R.S., F.Z.8. 


The Fossil Crinoids referred to Hypocrinus Beyrich. 
CeEEXCr andi Dextre OS — 100s) esacacncaealaeiecdseete ace 894 


BEDDARD, FRANK E., M.A., D.Sc., F.R.S., F.Z.8., Prosector 
to the Society. 


Contributions to the Anatomy and Systematic Arrange- 
ment of the Cestoidea. 


X. On Two Species of Tape-worms from Genetta dongo- 
aiipoiig. (QUEERS, (35 8))) pee copqopgpabbecsoo se odenconconnogd: 549 


XI. On a New Tapeworm from @dicnemus. (Text- 
fos yA eA On ea Sind Sate inher Bann len ccce Salada cra dcte <a 86] 


x 


Page 
Bovutencer, Epwarp G., F.Z.8., Curator of Reptiles. 
Experiments on the Metamorphosis of the Mexican 
Axolotl (Amblystoma tigrinum), conducted in the Society’s 
Gardens: an (Next-fisss 1 160s) menses neh n tees tanner 403 
Exhibition of living specimens of the Leaf-Insect 
(CECH OUTD, CELIO OINTEDN). "achboebennaoagoosennscensebaebocesadee cc: 546 
Exhibition of a living melanistic specimen of the Green 
JINR (OL RIGIG, CHPUCHIS)) aoa oabepocpbadaedeadoooosoonbsacdoon oso: 546 
Exhibition of a young specimen of the Matamata 
UVeroenyjarin (Claas jUMULPOGTD) ccocascooossps0q0n0cacadbaen00900350 1097 
Bounrencer, Grorce A., F.R.S., F.Z.S. 
On a Collection of Batrachians and Reptiles made 
by Dr. H. G. F. Spurrell, F.Z.8., in the Choco, Colombia. 
(Pls. CII.-CVIII. and Text-figs. 174-178.) .,....,..,... 1019 
Broom, Rospert, M.D., D.Sc., C.M.Z.8. 
A new Species of Golden Mole..............-..:cscssseceoes 546 
Exhibition of living specimens of the female and young 
of the South African Lizard, Zonurus giganteus. (PI. 
NBD. BES) anne ere Ferien thn Vals AAMT ae Aad ond ohbonasbaa une 548 
On the South-African Pseudosuchian Huparkeria and 
Allied Genera. (Pls. LX X VX XO) eee ee 619 
BULLEN, G. E. 
On some Cases of Blindness in Marine Fishes _......... 634 


Burne, Rrenarp H., M.A., F.Z.S. 


Exhibition of malformed Antlers of an Axis Deer 


(CCE CD TIS) Danna eHn AN PENEERB HORM E an aco chin Gia jnbquanantandoc 545 


xi 


Page 

Exhibition of specimens of Foetal Skeletons prepared 
by the Beale-Schultze method ...............sseeeeeeeeeseeees 1104 

CaLMAN, Witii1aM T., D.Sc., F.Z.8. 

On Freshwater Decapod Crustacea (Families Pota- 
monidz and Paleemonidz) collected in Madagascar by the 
Hon. Paul A. Methuen. (Pls. XCI. & XCII. and Text- 
in, UG pesbdaddudeschsnedescodabosh aad satpocooceoesr suceecnecseatnc 914 

Cuurcsa, Sir Artuur H., K.C.V.O., F.R.S. 
Notes on Turacin and the Turacin-Bearers............... 639 
Drenpy, ArtHuR, D.Sc., F.R.S., F.Z.8., and Row, R. W. 
Haro, D.Sc., F.L.S. : 

The Classification and Phylogeny of the Calcareous 
Sponges, with a Reference List of all the described Species, 
systematically arranged. (Text-fig. 133.) .................. 704 

Ewes, Henry J., F.R.S., F.Z.8. 

Exhibition of the Head of an Asiatic Wapiti. (Text- 

HUGS HSA I0L) OAn) erence dnecgec madsen acecneede ce stewigwcieae aetna 542 
Fryer, J. C. F., M.A., F.E.S. 

Field-Observations on the Enemies of Butterflies in 

(CES TINOT ONS Sommer bred dee TOsOe acc Ine BOGE erick Ctra Tir LOCC Cees ORES 613 
Happon, Miss KATHLEEN, 
Notes on Peripatoides woodwardi Bouvier ............. 611 


Hamitron, G. EK. H. Barrerr-. See Barrerr-HAmItroy, 
Gals EH. 


> 


Hamiuron, J. SrevENsSON-. See SrrveNsoN-HAmILtoN, J. 


X1L 
Harmer, Sipney, F., Se.D., F.R.S., F.Z.S. 
The Polyzoa of Waterworks. (Pls. LXII. & LXITI.). 426 


Hinton, Martin A. C., and Barrerr-Hamitron, G. E. H., 


E.Z.S. 
On a Collection of Mammals from the Inner Hebrides. 
CEI IDXOXGXeVE and Mext=neseek3 Ota. a) ee eEee ee nae etary 821 
Ho.prine, R. EK. 
Exhibition of Antlers, Skulls, and Photographs illus- 
trating Variations in the growth of the Antlers of Deer. 
(Dext=fig. Moa) cise. dates oan, seinen oes cig aber enlace 815 


Lesour, Miss Marte V., M.Sc, 


A New Trematode of the Genus Lechriorchis from the 
Dark Green Snake (Zamenis gemonensis), (Pl, XCIII.). 933 


Lerroy, Prof. H. Maxwett, M.A., F.Z,8., Hon, Curator of Insects. 


Remarks upon the New Insect House..,....,,,........... 1104 


LYDEKKER, RicHarp, F,R$,, F.Z.8, 


The Transvaal Race of the Cape, or Khama, Harte- 
beest;  Bext-fig, 135.) :cscsd.. dads scutes tae eee eee eee 818 


McMoraicu, J. Prayrarr, C,M.Z.S, 


On two New Actinians from the Coast of British 
Columbia, «(PE XC VILL icc eee Cee eee ens 963 


Mine, Joun ADAM. 


Pacific Salmon: An Attempt to evolve something of 
their History from an examination of their Scales. (Text- 
eS eo) a Speen SME MRP ago Hrs Sudgaaconatcadon 572 


xiil 
’ Page 
Mireneuy, P. Caatmers, M.A., D.Sc., LL.D., F.R.S., F.Z.S., 
Secretary to the Society. 


Report on Additions to the Society’s Menagerie during 
Taeemomulmotevlanche MOMS ssrausetases ecisstccwetcrecse seas 542 


Observations on the Anatomy of the Shoe - bill 
(Baleniceps rex) and allied Birds. (Pls. LXXX.- 
LXXXIII. and Text-figs. 119-132.) ..................-.e 644 


Report on Additions to the Society’s Menagerie during 
UME MAL OLIN ON PADIEM MON ed therserasye s stisicistensints otras clerieletate ajtaietasier 813 


The Peroneal Muscles in Birds. (Text-figs. 179-190.) 1039 


Report on Additions made to the Society's Menagerie 
during the months of May, June, July, August, and 
Be pbemaly err USM AIG ae ee. ee SUI athe de Rlaateened aa torars 1090 


Exhibition of the Alimentary Tract of a young Elliot’s 
Bheasaintg (hasvaneuscelliote)i-saeses anes esse eee eee ee 1094 


Report on Additions to the Society’s Menagerie during 
iln® sonore Or Crewe orere OMS? ceccacogssacnpaco soedaeaccsesessecac 1103 
OLSEN, ORJAN. 


On the External Characters and Biology of Bryde’s 
Whale (Balenoptera brydei), a new Rorqual from the 
coast of South Africa. (Pls. CLX.-CXIIT.) 


Pater, R. W., M.Se.(Manch.), 
The Brain and Brain-Case of a Fossil Ungulate of the 
Genus Anoplotherium. (Text-figs, 150-157.) ve... eee. 878 


ParkKER, Prof. W. Newton, Ph.D., I'.Z.S. 


Sponges in Waterworks 


Pravor, Henry G. J., Librarian to the Society. 


List of dates of publication of the early parts of the 
Society’s ‘ Transactions’ 


X1V 


Page 
Pocock, Reeinatp I., F.B.S., F.L.S., F.Z.S., Curator of 
Mammals. 
The Affinities of the Antarctic Wolf (Canis antarc- 
dics). (Rext=figs MOS ( ANE. ca ja. sch epee eee eee neer te. eecr ance 382 
Exhibition of a female and three young specimens of 
the Woolly Opossum (Philander laniger) — ..........ece cece 1094 
Exhibition of two young Degus (Octodon degus) ...... 1095 
Exhibition of Armadillos to show the Dorsal Glands. 
(Mextetigs NOS S195) is iiaieeeencortuuatine scone cceren eee cre: 1099 


Reean, C. Tats, M.A., F.Z.S8. 


A Collection of Fishes made by Professor Francisco 
Fuentes at Easter Island. (Pls. LV.-LX.) ............... 368 


A Revision of the Fishes of the Genus. Kuhlia. 
(Aes ces tata lela (0) )) e Rada cing MogeBaonaaaesdscoddonadessaeasGoon 374 


A Revision of the Cyprinodont Fishes of the Sub- 
family Peeciliine. (Pls. XCIX.—CI. and Text-figs. 
Ios Nfe)8) Waser ota nardndnodeadanccesac SPE SID oe Og 


Exhibition of a specimen of the Deep-sea Angler-Fish, 
MUNG POCA MS NOMISO MID EMD tin phsq0bbs0a0006000s60a0e0be0G0D00. 1096 


Row, R. W. Harotp, B.Sc., F.L.S., and Drenpy, ARTHUR, 
ID S55 ISIS) NYAS 


The Classification and Phylogeny of the Calcareous 
Sponges, with a Reference List of all the described 
Species, systematically arranged. (Text-fig. 133.)......... 704 


ScHaus, WiLuiAM, F.Z.S8, 


New Species of Rhopalocera from Costa Rica. (Plates 


1,1 I gs ae Mrs 8 Ae A 339 


XV 
Sura-Suiru, Davip, F.Z.S., Curator of Birds. 


Exhibition of the Egg and Young of the Mikado 


Pheasant (Calophasis mikado) ........1sesereeeeete nsec een ees 


Exhibition of a spirit specimen of a young Toucanet 
(Selenidera maculirostris) to show the well-developed 
Semrated Heel pads.) (Mext-ton NON) oor. secaenentectmecee 


Exhibition of a hybrid Black-winged Peafowl] (Text- 
fig. 192), a hybrid Pheasant, and a hybrid Jungle Fowl . 


Suuretpr, Roperr W., M.D., C.M.Z.S. 


On the Patella in the Phalacrocoracide. (PL. Xe 
Sairn, D, Seru-. See Sera-Smiru, D. 


STEBBING, The Rev. T. R. R., M.A., F.RB.S., F.Z.S. 


Notice of Prof. F. E. Schulze’s ‘ Nomenclator Ant- 
malium ’ 


Pee eee ewe ee ea eee eee e eres ress sees eseesseseessssaessestsssose 


SLEVENSON-Hamiuron, Major J., C.M.Z.S. 


Notes on Albinism in the Common Reedbuek (Cervi- 
capra arundinum) and on the Habits and Geographical 
Distribution of Sharpe’s Steenbuck (Raphiceros sharpei) . 


THompson, Surgeon Josepu C., U.S.N. 


Contributions to the Anatomy of the Ophidia. (Text- 
LIS SHRM OD Sa) BAI. Menthe toeenemade sah aNen saute uctauncuedaa 


Topp, T. Wrineate, M.B., F.R.C.S. 


Observations on Osteomalacia in the Zoological Col- 
lections of Manchester and Cleveland. (Pls. LX XXVIT-— 
TE XSNOXSIENG nae 


ee ee i ee oe ry 


Page 


818 


1095 


1098 


393 


814 


537 


Al4 


XV1 
Waters, Artuur W., F.L.S., F.G.S. 


The Marine Fauna of British East Africa and Zanzibar, 
from Collections made by Cyril Crossland, M.A., B.Sc., 
F.Z.S., in the Years 1901-1902. Bryozoa—Cheilostomata. 
(Pls. LXTV.-LX XIII. and Text-figs. 79-82.) ............ 458 


Watson, D. M.8., M.Sc. 


Batrachiderpeton lineatum Hancock & Atthey, a Coal- 
Measure Stegocephalian. (Pls. XCVI. & XCVIT. and 
Pext-figs. [62-16 0.) cease rae eer ctr ree etedste screenees 949 


WITHERS, THOMAS H., E:G:S: 


Some Miocene Cirripedes of the Genera Hexelasma 
and Scalpellum from New Zealand. (Pls. LXXXV. & 
IDXOOXOVS andi Vext-figss 139d 140.) ree eset eeeeeeeee 840 


Cirripedes from the Cenomanian Chalk Marl of Cam- 
lover Fegen, (Ue CONAN 05 ACORVG)) So gecn ondoadssocsocsonacosntoc 937 


XVil 


NEW GENERIC TERMS 


PROPOSED IN THE PRESENT VOLUME (pp. 339-1104). 


Page | Page 
Ascute (Spongiz) .............:. 729 Pamphoria (Pisces) ............ 1003 
Brachyrhaphis (Pisces) ...... 997 Pamphorichthys (Pisces) 10038 
Browniella (Reptilia)............ 627 Peciliopsis (Pisces) ............ 966 
Corrachia (Insecta) ............ 351 Priapella (Pisces) ............... 992 
Diplopylidium (Vermes) ...... 569 Priapichthys (Pisces) ......... 991 
Epinosis (Insecta) ..............- 366 Pseudopecilia (Pisces) ...... 995 
Kugonodzum (Vermes)..... ... we Styriodes (Insecta) ............ 361 
Girellops (Pisces) ............... 369 Syeute (Spongia) ............... 763 
Hebius (Reptilia) ............... 424 Teichonopsis (Spongiz) ...... 761 
Kuarrhaphis (Spongiz) .. ...... 780 Titanolepas (Crustacea) ....... 943 
Leptorhaphis (Pisces) ......... 998 Uteopsis (Spongiz) ............ 766 
Leucopsila (Spongiz) ......... 7795 Zeugmatolepas (Crustacea)... 938 


Proc. Zoo. Soc.—1913. 


tainty 
Jy) 


TT tne CAM: 
: ahh! i a 4 ac ‘ i b Lets 


INDEX 


OF 


SCIENTIFIC NAMES, 


Acanthistius 
cinctus, 868, 369. 
Suscus, 868, 373. 
Acanthopacelus 
bifurcus, 1006. 
melanzonus, 1007, 
1008. 
reticulatus, 1008. 
Acauthorbynchus, 


1065. 
Achradoecrinus, 896- 
899. 
Achramorpha, 758, 766, 
798. 


glactalis, 765. 
grandinis, 765. 
nivalis, 76d. 
schulzei, 765. 
truncata, 765. 
Acineta, 428. 
Acleropogon 
piceus, 618. 
Acmepteron 
cinerascens, 356, 357. 
poasind, 896, 367. 
Acridotheres 
melanosternus, 1091. 
Acropeecilia, 1014. 
tridens, 1018. 
Actias 
selene, 618. 
Actinia 
clavus, 969. 
cylindrica, 966, 972. 
Actitis 
hypoleucus, 1054. 
Adeona 
Joliacea fascialis, 532. 
Adeonella, 492, 519, 
528. 
contorta, 504, 529. 
platalea, 463, 501, 514, 
529-532, 537. 
polystomella, 530, 531. 


| Adeonella * 


(Laminopora) 
contorta, 529. 
Adeonellopsis, 528. 
crosslandi, 463, 531, 
532, 537. 
distoma, 531, 532. 
imperforata, O31. 
subsulcata, 531. 
/Kchmophorus 
major, 6O4. 


| Agialitis 


nivosa, 861. 
AKigotheles, 1065. 
nove-hollandie, 1062. 
/Eluredus 
melanocephalus, 1065. 
Aiyuorea, 970. 
forskalii, 967. 
Actea 
anguina, 462-464, 532. 
recta, 463, 464. 


truncata, 462, 465, 
5a 
Aithopyga 


seherie@, 1091. 
Aétosaurus, 629, 630. 
ferratus, 624. 


x 
galericulata, 1047. 
sponsa, 1047, 1048. 
Afaro, 979. 
acutiventralis, 981. 
amazonum, 981, 990. 
cultratus, 981. 
Agalychnis 
calcarifer, 1023, 
1038. 
spurrelli, 1024, 1025, 
1038. 
Alea 
torda, 1056. 
Alcedo 
astaticus, 1060. 


Alcyoncellum, 787. 


Aleedo 
bengalensis, 1060, 
ispeda, 1060. 
gelatinosum, 746, 

787. 

Aleyonella 
articulata, 441. 
fungosa, 450. 

Aleyonidium 
duplex, 488. 

Alopoglossus 
coptt, 1038. 
fuliginosa, 1034. 

Alticus 
striatus, 372. 
variolosus, 373. 

Alytes 
obstetricans, 1025, 

Amabilia, 571. 

Amazilia 
felicie, 1063. 

Amblystoma 
tigrinum, 403-413. 

Ameiva 
bridgesti, 1032. 
festiva, 1033. 


| Ampheristocrinus, 911. 
| Amphisbeena 


fuliginosa, 1094. 
Aimphiute, 754, 798. 
paulini, 755. 
Amphoridium, 787. 
viridis, 787. 
Amphoriscus, 781, 798, 
804. 
buceichii, 782, 790. 
chrysalis, 782. 
cyathiscus, 782, 799. 
cylindrus, 782. 
elongatus, 782. 
flamma, 752. 
gregorti, 782. 
kryptoraphis, 782. 


xx 


Amphoriscus 
murmanensis, 733. 
oblatus, 711, 782. 
ovipurus, 782. 
poculum, 752. 
semoni, 782. 
testiparus, 782. 
thompson, 753. 
urna, 782. 

Amphorula, 787. 
solida, 787. 

Anadia 
oceliata, 1033. 
vittata, 1033, 1038. 

Anamixilla, 758, 786, 

798. 
torresi, 766. 

Anampses 
ceruleopunctatus, 372. 
pulcher, 371-373. 

Anaschisma, 957. 

Anastomus, 688, 690. 

Anchisaurus, 629, 630. 

Ancylus, 427. 

Anodonta, 428. 

Anolis 
breviceps, 1031, 1038. 
Jasciatus, 1030. 
godmani, 1031. 
macrolepis, 1032. 
maculiventris, 1030. 
notopholis, 1032. 
princeps, 1031. 

Anoplotherium, 879- 

882, 886-888, 890- 
893. 

Antedon, 903, 904. 

Anthropoides 
virgo, 1053. 

Anthropopithecus 
troglodytes, 1090, 

Aphaneramma, 956. 

Aphelocoma 
sordida, 1091. 

Aphroceras, 768, 776, 

798. 
aleicornis, 777, 789. 
asconoides, 729. 
cespitosa, 777. 
cataphracta, 

777. 
cliarensis, 777. 
corticata, 777. 
elongata, 777. 
ramosa, TO4, 
sericatum, 'T56. 
syconoides, 764. 

Apodemus 
hybridensis, 835-837. 
sylvaticus sylvaticus, 


835, 837. 


alu; 


Appias, 616. 

sp-, 615, 618. 

albina, 614, 618. 

paulina, 614, 617, 618. 
Aprosmictus 

cyanopygius, 1008. 
Aptenodytes, 1044. 

pennantii, 394. 
Apteryx, 1043. 
Aquila 

verreauxt, 1O49, 
Arachnechthra 

zeylonica, 1066. 
Aramides 

chiricote, 1052, 1104. 

ypecaha, 1052. 
Aramus 

scolopaceus, 1053. 
Archegosaurus, 956. 
Archispirostreptus 

nitidus, 1093. 


Archonias 
dismorphites, 356. 
intermedia, 3595, 

367. 


nigrescens, 396. 

Arcoscalpellum, 850, 
853. 

Ardea, 656, 690, 691, 
695, 694, 696-699, 
703. 

egretta, 1045. 
herodias, 1045. 
— lessoni, 1045, 
1091. 
purpurea, 1045, 
Ardetta, 674. 
minuta, 1045. 
Arginia 
sp., 614, 618. 
Argus 
giganteus, 1051. 
Artamus 
Suscus, 613, 615. 
leucogaster, 1066. 
Arthropoma 
cecillii, 462, 508. 
Artynas, 787. 
compressus, 787. 
villosum, 746. 
Artynella, 787. 
compressa, 787. 
Artynes, 787. 
compressa, 787. 
Artynium, 787. 
compressum, 787. 
Artynophyllum, 787. 
compressum, 787. 
Asarcia 
variabilis, 1055. 


Ascaltis, 717, 787. 


INDEX OF SCIENTIFIC NAMES. 


Asealtis 
botryoides solanderii, 
720. 
canariensis, 
787. 
cerebrum, G24. 
— decipiens, 725, 
compacta, "724. 
darwinii, 725. 
— caroli, (24. 
gegenbauri, 725. 
— charybdea, 724. 
goethet, 725. 
lamarckii, 726. 
— agassizti, 724. 
solanderti, "725. 
Aseandra, 717, 
787. 
angulata, 721. 
hotrys, T21. 
complicata ameboides, 
720. 
contorta, 721. 
conulata, 726. 
corduta, 721, 787. 
densa, 721. 
echinoides, 722. 
falcata, "17, 720, 722, 
788, 790. 
hermesti, "722. 
licherkiihnit. 723. 
nirabilis, 761. 
nitida, 723. 
panis, 723. 
pinus, 723. 
reticulatum, 728. 
— reticulata, 723. 
sertularia, 723. 
tenuis, 723. 
variabilis, 723. 
— arachnoides, 721. 
— cervicornis, 721. 
— confervicola, 721. 
— hispidissima, 722. 
Ascetta, 717, 788. 
blanca, 724. 
— phillipina, 726. 
celathrus, 725. 
coriacea himantia, 
726. 
flewilis, 725. 
macleayt, 726. 
primordialis, 726, 730, 
788. 
— dictyoides, 725. 
— loculosa, 726. 
— poterium, 726. 
— protogenes, 726. 
procumbens, 726. 
sagittaria, 727. 
sceptrum, 727. 


724, 


Ascetta 
spongiosa, 727. 
vesicula, 727. 
Ascilla, 7388. 
coriacea, 725. 
gracilis, 725, 788. 
— convallaria, 725. 
— japonica, 726. 
Ascometra, 788. 
primordiale, 788. 
Ascortis, 788. 
clarkit, 721. 
corallorhiza, 721. 
Sragilis, 722. 
horrida, 722, 788. 
lacunosa, 72%. 
Asculmis, 758. 
armata, 721, 788. 
Ascuris, 788. 
arrecifé, 788. 
Ascute, gen. nov., 719, 
29% 798; 799! 
asconoides, 729. 
uteoides, 710, 718, 
G29. 
Ascyssa, 717, 719, 767, 
798, 799. 
acufera, 729. 
troglodytes, 729. 
Asellus, 426, 451, 4384, 
437. 
aquaticus, 427. 
Asio 
otus, 1061. 
Aspatha 
gularis, 1060. 
Aspidelectra, 461. 
Aspidostoma 
giganteum, 487. 
Asterias 
rubens, 1104. 
Astrosclera, 788. 
willeyana, 788. 
Ate 
canace, 359, 367. 
Athene, 1062. 
noctua, 1061. 
Atheris, 1034. 
Atractus 
iridescens, 1035. 
multicinctus, 1035. 
Aulacorhamphus 
prasinus, 1063. 
Auloplegma, 7&8. 
haeckeli, 721. 
loculosum, 738. 
Aulorhiza 
intestinalis, 726, 788. 


Baeria, 768, 780, 798, 803. 


INDEX OF SCIENTIFIC NAMES. 


Baeria 
ochotensis, 779. 
Baleniceps, 645-703. 
rex, 644, 650, 701-708, 
1039, 1045, 1092. 
Balxnoptera 
acutorostrata, 1076, 
1077, 1085, 1086. 


borealis, 1073, 1074, 
1076-1030, 1083, 
1085-1087, 1089, 
1090. 

brydei, 1074-1977, 


1079, 1080, 1082- 
1087, 1089, 1090. 
musculus, 1076, LO89. 
physalus, 1076, 1077, 

L085. 

Balanus, 842, 846. 
callistoderma, 847. 
corolliforneis, 847. 
evermanni, S47. 
hirsutus, 847. 
hockianus, 847, 848. 
psittacus, 847. 

Balearica 
chrysepelargus, 1053. 

Basiliscus 
americanus, 1032. 
galeritus, 1032, 

Bathus 
hoitentotta, 1093. 

Bathystethus 
cultratum, 373, 374. 
orientale, 379. 

Batrachiderpeton, 949, 

955, 956, 958-961. 
lineatum, 950, 951, 
953, 954, 962. 

Batrachosuchus, 956. 

Beania 
hirtissima, 467. 
hyadesi, 487. 
intermedia, 467. 
magellunica, 

530. 
mirehilis, 462, 467. 
quadricornuta, 467. 
spinigera, 462, 467. 

Belodon, 623, 629. 

Belonesox, 979. 


467, 


belizanus, 977, 990, 
991. 
Bematiscus 
transvaalensis, 547, 
548. 
trevelyani, 547. 
villosus, 546-548. 
Bicellaria, 476. 
chuakensis, 462, 467, 


5384. 


XX1 


Bicellaria 
ciliata, 468, 470. 
Bicidium, 970. 
equoree, 967-969, 972. 
clavus, 969. 
parasiticum, 969, 972. 
Bifaxaria 
abyssicola, 528. 
corrugata, 527. 
denticulata, 527. 
levis, 528. 
minuta, 528. 
papillata, 528. 
reticulata, 527, 528. 
submucronata, 527. 
vagans, 463, 527, 528, 
582, 533. 
Biflustra 
armata, 486. 
Bithynia, 437. 
tentaculata, 427. 
Bithynis 
gaudichaudii, 930. 
hildebrandti, 926, 928, 
930. 
Biziura, 393. 
lobata, 396. 
Boa 
occidentalis, 1094. 
Bolla 
cylindus, 359. 
machuca, 359, 367. 
sodalis, 359, 367. 
Boobon 
fuliginosus, 1092. 
Botaurus, 652, 699. 
stellarts, 1045. 
Bothriceps, 956, 957. 
Bothrodytes, 424. 
Bothrops 
acrochordus, 1037, 
Boulengerina, 374. 
Brachypternus 
auranticus, 1063. 
Brachyrhaphis, gen. nov., 
997, 980. 
rhabdophora, 990, 997, 
998. 
Bradypus 
tridactylus, 1094. 
Brettia, 484. 
australis, 465. 
longa, 465, 
tropica, 462, 465, 532. 
Browniella, gen. noy., 
627. 
africana, 627, 628, 633. 
Bubalis 
caama, 818, 819. 
— selhornet, 820, 821. 
— typica, 821. 


Xxll 


Bubalis 
cokei, 819. 
Bubo, 151. 
lacteus, 1061. 
maculosus, 1061. 
maximus, 668, 1061. 
Bucorvus, 670, LO61. 
Buffonella, 505. 
Bufo 
hematiticus, 1022. 
hypomelas, 1022, 1038. 
marinus, 1022. 
typhonius, 1022, 
Bugula, 459, 470, 522. 
avicularis, 479. 
bicornis, 476. 
caliculata, 469, 479. 
capensis, 471, 472. 
dentata, 472. 
gracilis, 472. 
mirabilis, 472. 
murrayanda, 479. 
neritina, 472. 
— minima, 462, 471. 
— ramosa, 472. 
— rubra, 472. 
— tenuata 472. 
reticulata, 472. 
— unicornis, 472. 
robusta, 462, 471, 534. 
versicolor, 472. 
Bugulopsis 
peachit, 473, 474, 584. 
Buthus 
gibbosus, 542. 
Butleria 
faula, 360. 
lalage, 360. 
lethea, 360, 367. 
lysis, 360, 367. 
Butorides 
cyanurus, 650. 


Caberea, 474, 475, 481. 
crassimarginata, 480. 
darwinit, 477, 
ellisit, 534. 
retiformis, 479, 480. 

Caberiella, 475. 

Cacatua 
sulphurea, 1058, 

Cecilia 


intermedia, 1020, 1034. 


isthmica, 1020. 
nigricans, 1022, 


pachynema, 1020, 1022, 


palmeri, 1021. 

tentaculata, 1020. 
Caica, 1059. 

melanocephala, 1058, 


Caiman 
selerops, 1030. 
Calantica, 937, 938, 942, 
947. 
(Titanolepas) — ¢uber- 
culata, 943, 948, 
Calcispongia, 789. 
compressa, 789. 
Calodromas, 1051. 
elegans, 1050. 
Calophasis 
elliott, 1052, 1098. 
mikado, 818. 1094. 
Caloporella, 484. 
Calopsittacus 
nove-hollandie, 
1059. 
Calpidium, 483. 
Calwellia, 465. 
Campylorhynehus 
unicolor, LOG. 
Cancroma, 645, 649, 652, 
674, 689-691, 693- 
696, 698-700, 703. 
cochlearia, 699. 
zeledoni. 1045, 1091. 
Canda, 473, 474. 
arachnoides, 480. 
Jfossilis, 480. 
retiformas, 
481, 534. 
tenuis, 480. 
Canis 
antarcticus, 382-893. 
anthus, 384, 386, 393. 
aureus, 384. 
azare, 392. 
CUNCTLLOTUS, 
390, 333. 
familiaris, 384, 
Fulvipes, 390, 
gracilis, 390. 
Jubatus, 383, 384. 
latrans, 382-393. 


462, 479, 


lupaster, 386, 9890, 
392. 


lupus, 384, 390, 393. 
mesomelas. B84. 
mnicrotis. 390. 
occidentalis, 884, 390. 


1058, | 


382-384, | 


pallipes, 386, 390, 392, 


393. 
parvidens, 390. 
rudis, 390. 
sclatert, 390, 391, 1108. 
urostictus, 390. 
vetulus, 383, 384, 393. 
(Nyctereutes)  pro- 
cyonoides, 391. 
Capitosaurus, 956. 


Caprimulgus, 1065, 1072. | 


INDEX OF SCIENTIFIC NAMES. | 


Caprimulgus 
europeus, 1062. 
Caranx. 
cheilio, 370, 
guard, 3710. 
Cariacus 
leucurus, 817. 
Cariama 
burmeistert, 1094. 
cristata, 1054. 
Carphibis 
spinicollis, 654, 1047. 
Carpophaga 
rufiguia, 1056. 
Carystus 
commodus, 366. 
subrufescens, 865, 367. 
Castor 
canadensis, LOL. 
Casuarius 
keysseri, 542. 
scluteri, 1091. 
uniappendiculatus, 
1043. 
Catantica 
(Scillelepas), 947, 948. 
Catargynnis 
dryadina, 342, 367. 
Catenaria, 465. 
diaphana, 462, 
485, 532. 
lafontii, 462, 481, 482, 
489. 
parasitica, 489. 
Catenicella 
contei, 485. 
continua, 485. 
cornuta, 483, 484. 
delicatula, 488. 
elegans, 484. 
fusca, 483. 
internodia, 484, 485. 
perforata, 453, 
seplentrionalis, 48d. 
taurina, 483. 
umbonata, 433. 
Jathartes, 661, 1069. 
aura, 655, 1048. 
Cathrina, 717. 
Catopsilia, 616. 
crocale, 615. 
pyranthe; 614, 615, 
Cellaria, 461, 493. 
australis, 495, 496. 
bicornis, 495. 
contiqua, 497. 
dennanti, 496. 497. 
divaricata, 495. 
dubia, 42h. 
Jistulosa, 490, 496. 
gracilis, 490, 495-497, 


482, 


Cellaria 


gracilis tessellata, 492, | 


495, 497, 535. 
hirsuta, 496. 
Jullieni, 496. 
magnifica, 497. 
malvinensis, 495-497. 
oculata, 490. 
rigida, 495, 496. 
salicornia, 494. 
salicornioides, 497. 
tenella, 490. 
tenuirostris, 495, 497. 
variabilis, 495, 496. 
wandeli, 496. 
wasinensts, 462, 495— 

497, 533. 

Cellarinella, 528. 
Cellepora, 461, 529. 
albirostris, 522. 
— typica, 522. 
avicularis, 5V8. 
cidaris, 521. 
ciliata, 509. 
columnaris, 521. 
conica, 508. 
coronopus, 508-510. 
eatonensis, DO8. 
evexd, 508. 
hyalina, 509. 
incrassata, 509. 
megasoma, 508. 
pumicosa, 509, 510. 
ramulosa, 509, 510. 
redoutii, 508. 
rota, 510, 
sardonica, 509. 
simonensis, O08. 
spongites, 509. 
Cellularia, 461. 493. 
diplodidymoides, 492. 
spatulata, 478. 
tubucellaria, 494. 
Centropomus 
rupestris, 375. 
Ceraterpeton, 960, 962. 


galvani, 958, 959, 962. | 


reticulatum, 98d, 959, 
961. 

Cercopithecus 

randti, 542. 
Cereopsis 

nove-hollandie, 1047. 
Cerianthus 

membranaceus, 966. 
Cervicapra 

arundinum, 537. 
Cervulus 

muntjac, 817. 
Cervus 

axis, 040, 817. 


INDEX OF SCIENTIFIC NAMES. 


Cervus 
canadensis, 816. 
canadensis siberica, 545. 
dama, 817. 
elaphus, 543, 545, 816. 
eustephanus, D4). 
unicolor, 817. 
Ceryle 
alcyon, 1060. 
americana, 1060. 
anda, 1060. 
maxima, 1060. 
Ceyx 
rufidoris, 1060. 
Charadrius 
pluvialis, 1059. 
Charis 
chrysus, 352. 
subressellata, 351, 367. 
turrialbensis, BOK 
367. 
Chasmorbynchus 
nudicollis, \065. 
Chauna, 1042, 1043. 


chavaria, 1040, 1041, | 
1047. 
cristata, 1092. 
Cheilio 


inermis, 371. 
Cheilopora, 515, 
Chelys 

Sinbriata, 1097. 
Chilopora, 515. 
Chionis 

alba, 1055. 
Chironomus, 

459. 
Chizerhis, 641. 
Chlidonia 


433, 45C- 


cordiert, 462, 489, 490, | 


492, 583. 
Chloéphaga 
tornata, 1047. 
Choanoteenia, 564. 
coronata, 861. 
Chorizopora 
brongniarti. 507. 
Chrysaora, 969. 
Chrysochloris, 546, 
villosa, 547. 
Chrysopelea 
ornata, 420. 
Chrysotis, 1059. 
Cicada, 618. 


Ciconia, 645, 679, 688, | 


699. 
nigra, 1046. 
Cinnamopteryx 
rubiginosa, 1094. 


_ Cinosternum 


spurrelli, 1030, 1058. 


xxi 


Circus 
qouldi, 1049. 
hudsonius, 1104, 
Citharoscelus 
koekti, 1098. 
Cittotzenia, 569. 
Cittura 
cyanotis, 1060. 
sanghirensis, 1060. 
Clathrina, 730, 789. 
cavata, 724. 
clathrus, 789. 


daminoclathrata, 727. 
latitubulata, 787, 
738. 


osculum, 726. 

reticulum, 718. 

sulphurea, 725, 727. 

tripodifera, 706, 717, 

728. 

— gravida, 724. 

ventricosa, 724. 
Clava, 505, 522. 
Claviporella 

pulchra, 483. 
Clystolynthus, 789. 

vesicula, 789. 
Cnesterodon, 980, 

carnegie, 1000. 

decemmaculatius, 1000, 

1001. 

scalpridens, 1008. 
Cobalopsis 

latonia, 363, 367. 
Cobalus 

gabina, 362. 

lateranus, 362, 867, 

laureolus, 362, 367. 

nigrans, 862, 367. 

pindar, 362, 567. 
Codiacrinus, 896. 
Coendu 

mexicanus, 1108. 
Coenocystis 

richardsoni, 897. 
Ceenostomella, 789. 

caminus, 789. 
Ccenostomium, 789. 

crambessa, 789. 
Ccenostomus, 789. 

primigenius, 789. 
Coereba 

cyanea, 1065. 
Colaptes 

meaxicanoides, 1068. 
Colius 

capensis, 1063. 
Coluber 

helene, 424. 
Columba 

livia, 1056. 


XX1V 


Columbula 

picut, 1056. 
Colymbus 

septentrionalis, 394. 
Compsognathus, 629. 
Conger 

vulgaris, 635. 
Connochates 

albojubatus, 1090. 
Conurus 

jendayt, 1058, 
Copsychus 

saularis, 614. 
Coracias 

garrula, 1060. 

indica, 1060. 
Cordylophora, 427, 

431, 

Corrachia, 


gen. nov., 


leucoplaga, 351, 
367. 

Corynoporella, 465. 
Corythzola, 641. 
Corythaix 

albicristata, 629. 

persd, 1057. 
Corythophanes 

cristatus, 1032. 
Costaticella 

lineata, 483. 
Costicelia, 483. 
Cotugnia, 570, 571. 
Coturnix 

coromandelica, 1052. 
Cracticus 

destructor, 1065. 
Craspedozoum, 475. 
Creadion 

carunculatus, 1065. 
Crenothrix, 431. 

hiihniana, 430. 

polyspora, 430. 
Crex, 670. 
Cribricella, 488. 
Cribrilina, 529. 

figularis, 474, 476. 

radiata, 462, 501. 
Crisia, 473, 493. 

pilosa, 477. 
Cristatella, 431, 976. 
Crossoptilon 

manichuricum, 1052. 
Crotalus 

confluentus, 422. 
Cryptocrinus, 895-897. 
Cuculus 

canorus, 1057. 
Cumulipora 
angulata, 502. 


INDEX OF SCIENTIFIC NAMES. 


Cupularia, 481. 
owent, 508, 
Cursorius, 651. 
Cyanea 
arctica, 969. 
Cyanocorax 
luxuosus, L065. 
Cyanops 
flavifrons, 1068. 
Cyanorhamphus 
alpinus, 1058. 
Cyathiscus, 789. 
actinia, 777, 789. 


Cyclocorus 
lineatus, 419, 425. 
Cycloszemia 
subcerulea, 358, 
367. 


Cyclotosaurus, 956. 
Cydonocrinus, 913. 

parvulus, 894, 911. 
Cygnus 

nigricollis, 1047. 

olor, 1047. 
Cylindrecium, 464. 
Cymbirhynebus 

macrorhynchus, 1065, 
Cyneelurus 

Jubatus, 1093. 
Cynictis 

penicillata, 559. 
Cynocephalus 

anubis, 856, 859. 
Cyphosus 

cinerascens, 378. 
Cypselus 

apus, 1062. 


Dacelo 
gigantea, 1060. 
Danais, 619. 
aglea, 617. 
limniace, 616, 617. 
septentrionis, 615- 
618. 
Daption 
capensis, 1044, 
Dasylophus 
superciliosus, 643. 
Dasyprocta 
colombiana, 814. 
variegatus, 1094. 
Dasypus see Euphractus. 
Davainea, 570, 876, 
877. 
Decapterus 
sancte-helene, 378. 
Dendrobenia, 479. 


Dendrobates 


aurotenia, 1029, 1038. 

paraensis, 1028, 1029. 

tinctorius, 1026-1029, 
1038. 

— chocoensis, 
1028, 1038. 

—coctei, 1026, 1027, 
10388. 

— cocteawi, 1027. 

— daudini, 1027. 

trivittatus, 1029. 


1026, 


Dendrocopus, 1065, 


major, 1063. 


Dendrocygna 


Julva, 1047. 


Dendrograptus 


serpens, 471. 


Dendrohyrax 


dorsalis, 1104. 


Dendya, 706, 712, 716, 


717, 719, 730, 736, 

737, 742, 798-802. 
prolifera, 728. 
tripodifera, 710, 728. 


Dermatreton, 789. 


chartaceum, 789. 
hodgsont, 760. 


Diachoris 


spinigera, 467. 


Diastopora, 461. 
Diceum 


hirundinacewm, 1092. 


Diceratosaurus, 962. 


punctolineatus, 959. 


Dicrurus, 1066. 


cerulescens, 614. 
leucopygialis, 614. 


Didelphys 


azare, 1094. 


Dinetopia, 465. 
Diplocaulus, 960-962. 
Diplodidymia, 492. 


conplicata, 459, 462, 
489-491, 529, 534. 


Diploglossus 


monotropis, 1032. 


Diplophallus, 569, 570. 
Diploposthe, 569, 570. 


sui-generis, 570. 


Diplopylidium, gen. nov., 


559, 567-571, 872- 
874. 

genett@, 559-561, 563- 
566, 


Diplostomum, 635. 
Dipsadoboa 


unicolor, 10938. 


Dipsadomorphus 


blandingii, 1093. 


INDEX OF SCIENTIFIC NAMES. 


Dipsas 
JSorstenti, 1092. 


Dipylidium, 564, 57], | 
872. 


caninum, 564- 


599, 870. 


551, 


dongolense, 549, 550. | 


553-559, 561, 567. 
echinorhynchoides, 558, 
559. 
genette, 558, 559. 
gervaisit, Ho8. 
monoophorum, 5A9. 
monticellit, 5d9. 


pasqualii, 554, 558, 
BIS), 

trtnchesti, 554, 558, 
559, Siow 


triseriale, 559. 
eschokkei, 552, 559. 
Discopora, 461. 


Dissura, 681, 688, 693, | 


703. 
episcopus, 1046. 
Ditaxipora. 485. 
Djedda, 789. 
vtolacea, 753, 789. 


Dolichotis 
magellanicus, 1091. 

Dreissensia 
polymorpha, 427, 
439. 


Dromeus, 692. 
nove-hollandie, 1043. 
Dromas, 651. 
Drymobius 
dendrophis, 1034. 
Dryonastes 
cerulatus, 814. 
ruficollis, 814. 
Dules 
caudovittatus, 379. 
Suscus, 375, 377. 
leuciscus, 378. 
maculatus, 378. 
malo, 378. 
marginatus, 378. 
— boninensis, 378. 
mato, 378. j 
teniurus, 381. 
Dunstervillia, 789. 
corcyrensis, 746. 
elegans, 745, 789. 
Jormosa, 746. 
lanzerote, TA5. 
schmidtii, 746, 748, 
Dyssycarium, 790. 
egedii, 790. 
Dyssyconella, 790. 
caminus, 769, 
pumila, 790. 


Dyssyeum, 790. 
Jistulosum, 790. 
periminum, 773. 

Dyssycus, 790. 
primigenius, 790. 


Ebnerella, 790. 
buccichti, 790. 
gregortt, 782. 
kitkenthali, 753. 
lanceolata, 752. 
nitida, 752. 
schulzei, 765. 

Echelatus 
lucina, 358, 367. 

Echidna, 1072. 

Echinoencrinites, 895, 

896. 

Echinorhynehus, 427, 

Eclectus, 685. 
pectoralis, \Q58. 
roratus, 1058. 

Edwardsia 
kerquelensis, 969. 

Egernia 
depressa, 1092. 

Eilhardia, 768, 798. 
schulzet, 780, 781. 

Elaps 
corallinus, 1021, 1036. 
microps, 1036, 1038. 
mipartitus, 1035, 1036. 
rosenbergri, 1021, 1034, 

1036. 

Hnantia 
lua, 307. 

— costaricensis, 307, 
367. 

Enodes 
erythrophrys, 1065. 

Entomyza 
cyanotus, 1065. 

Enyalioides 
heterolepis, 1082. 

Ephydatia 
fluviatilis, 427, 436. 

Epimachus, 651. 

Epinosis, gen. nov., 366. | 
angularis, 367. 

EKresia 
cwla, 344, 567. 
pacilina, 344. 
sticta, 344, 367. 

Krithizon 
dorsatus, 542. 

Eryops, 956, 957, 961. 

Erythrosuchus, 623, 6380. 

Eschara 
hexagonalis, 529, 


platalea, 529. 


Escharina, 505. 
Kscharinella 
cecilleana, 529. 
Escharoides, 461. 
oceiusa, 463, 519- 
521. 
Estrelda 
phenicotis, 1065. 
Eubagis 
hecubu, 347, 367. 
vicaria, 247. 
HKueratea 
chelata, 462, 466. 
cordieri, 492. 
Tudocimus 
longtrostris, 650, 
Eudyptes 
chrysocome, 394. 
Eugonodeum, gen. nov., 
872, 874-877. 


edicnemi, 862-866, 
- 868, 870, 871, 873, 
877. 
Eunectes 


murinus, 814. 
Euparkeria, 620, 624,625, 
629, 630, 632. 
capensis, 619, 627, 628, 
632, 633. 
Hupetoieta 
poasina, 343, 367. 


| Euphractus 


sexcinctus, 1102, 1103. 

villosus, 1099. 

villosus, 1099, 
Euplectella, 787. 
Euplea, 617-619. 

core, 615, 616. 

coreta, 6195. 

kollart, 615, 616. 
Eupodotis 

kori, 564. 
Euptychia 

agnata, 842, 367. 

antonoé, 342. 

confusa, 342. 

drymo, 341, 367. 

juani, 342, 

labe, 342. 
Eurygona 

cataleuca, 349. 

leucon, 3849, 367. 

leucophryna, 849, 

367. 

matuta, 849, 867. 

mystica, 348, 367. 
EKurylemus 

cchromelas, 1065. 
Eurypyga, 645, 651, 

699. 
helias, 1054, 1092. 


XXV1 


Eurystomus 
orientalis, 1060. 
Eutimalphes 
indicans, Y70. 
Evotomys, Nb 
alstoni, 522, 827, 829, 
&30, 839. 
cesarius, 829. 
glareolus, 829. 
—. britannicus, 
830. 
norvegicus, 829, 830. 
skomerensis, 829., 


827, 


Falcinellus, 651. 

Falco 
pereyrinus, 1049. 

Falga 
hermione, 366, 367. 

Farcimia, 490. 
oculata, 462, 489, 535 

Felis 
caracal, 1098. 
concolor, LOO1. 
ocreata, 1098. 
pardalis, 1091. 
tigrina, LUOL. 

Flabellaris, 474. 
cuspidata, 475. 
flabeliata, 475. 
ligulatum, 475. 
multisertata, 475. 
roborata, 475. 
triseriata, 475. 
(Craspedozoum)  ligz- 

latum, 475. 
(—) rohorata, 475. 
(Menipea)  cuspidata, 
534. 

Flustra, 494, 529. 
abyssicola, 474, 476. 
cecilii, 508. 
mamillaris, 497. 


Francolinus 
infuscatus, 1052. 
Fredericella, 429, 431, 


444, 447. 
sultana, 427, 438, 459, 
441, 448, 457. 
Fregata, 661, 1045. 
Fulica 
leucoptera, 1052. 
Fulmarus, 670. 


Gadus 
eglefinus, 578. 
pollachius, 637. 
Galago 
maholi, 814. 


JNDEX OF SCIENTIFIC NAMES. 


| 
| 


Galbula 
albivostris, 1063. 
rufiventris, 1063. 

Galeopithecus, 631. 

Gallinago 
celestis, LOD4. 

Gallinula 
chloropus, 1052. 
phenicura, 10d2. 

Gallirex, 639. 
jJohustoni, 641, 


Gallus 
gallus, 1051, 1052, 
1098. 


varius, L099. 
Gambusia, 979, 
1000. 
affinis, 982, 984. 
annectens, 992. 
bimaculata, 993. 
bonita, 993. 
caudovittata, 982, 986. 
caymanensis, 982, 990. 
dominicensis, 982, 989, 
990, 1018. 
dovit, ‘982, 986. 
episcopi, 992. 
Jasciata, 995. 


991, 


gracitior, 982, 983, 989, 


1018. 
gracilis, 984, 999. 
holbrookti, 982, 988, 
990. 


humilis, 984. 
infans, 998. 
Jonesti, 993. 


melanopleura, 982, 988. 


melanosticta, 982, 987. 


nicaraguensis, 982, 983, 


985. 


nigropunctata, 982,983, 


987. 
nigroventralis, 992. 
nobilis, 98d. 
oligosticta, 

990, 1018. 
parismina, 992. 
patruelis, 982, 984. 
picturata, 981. 
punctata, 982, 

986. 


982, 


983, 


puneticulata, 982, 987. 


rhabdophora, 997. 
senilis, 982, 983, 98d. 
speciosa, 984. 
terrahensis, 993. 
tridentiger, 992. 
turrubarensis, 992. 
umbratilis, 998. 
wrayt, 
1018. 


988, 


982, 983, 988, 


Gammarus, 426, 431,434, 
437. 

pulex, 427. 
Gasterocoma, 897, 
Gasterosteus 

aculeatus, 428. 
Gavia 

stellata, 394. 
Gazella 

bennetti, 
Gecinus 

vittatus, 1063. 
Gemellaria 

loricata, 465, 466. 
Gemellipora 

glabra striatula, 507. 


898. 


1093. 


protusa, 462, 506, 
53). 

striatula, 507. 
Genetta 

dongolana, 549. 
Gennzeus 

horsfieldi, 542. 
Geocichla 

citrina, 1066. 
Geopelia 


cuneata, 1056. 
tranguilla, 1056. 
Geophaps 
plumitera, 1056. 
Geothelphusa, 924. 
Gephyrophora 
polymorpha, 521. 
Giraffa 
camelopardalis 
quorum, 542. 
Girardinus, 978, 980. 
caucanus, 1013. 
caudomaculatus, 999. 
creolus, 1002. 
denticulatus, 1002. 
Jormosus, 995. 
garmant, 1002. 
guppyt, 1008, 1010. 
theringit, 999. 
januarius, 999. 
lutzi, 996. 
metallicus, 1001, 1002. 
pleurospilus, 997. 
presidionis, 996. 
reticulatus, 1008. 
uninotatus, 1000. 
versicolor, 1018. 
zonatus, 999. 
Girella 
nebulosa, 369. 
Girellops, gen. nov., 369, 
373. 
nehulosus, 369, 373. 
Glareola, 651. 
pratincola, 1055. 


antt- 


INDEX OF SCIENTIFIC NAMES, 


Glaridichthys, 978, 980. 
falcatus, 1000, 1002. 
latidens, 1002. 
torralbasi, 1000, 1002. 


uninotatus, LOOU, LO02. 


Glaridodon 
Januarius, 999. 
latidens, 1002. 
wuninotatus, 1000. 

Glossiphonia, 427. 

Glossostoma 
aterrimum, 1025. 

Glyptolepis 
paucidens, 957. 

Grantessa, 704, 

798. 
compressa, TA2. 
erecta, T52. 
erinaceus, 

az 
flamma, 750, lige 
glabra, 751, 752 
glacialis, 753. 
hastifera, 711, 752. 
hirsuta, 7H0-752. 
hispida, 711, 750-752. 
intusarticulaza, J11, 

Toll, Tass 
kitkenthali, 753. 
lanceolata, 752. 
murmanensis, 7)3. 
nitida, 72. 
elanitoy7 WO 


“all, 


pluriosculifera, 752. 
poculum, T\1, 752. 
polyperistomia, 711, 


(52, 753. 
preiwischt, 753. 
sacca, 711, 

703. 
simplex, 753. 
spissa, 759. 
stauridea, 753, 789. 
sycilloides, 753. 
thonpson, 753. 
zanzibarensis, 703. 
(Ainphoriseus) pocu- 

lum, TAO. 

Grantia, 704, 738, 

749, 750, 751, 

766, 768, 769, 

798, 805. 
aculeata, 761. 
arabica, 773. 
asconoides, 760. 
atlantica, 199. 
botryoides, 721. 
— nen 725, 
brevipilis, TH9. 
canadensis, 759. 
capillosa, TH9. 


lod 


750, 751 


723. 


=t) 


744, 
7TdA, 
781, 


Grantia 
chartacea, 759, 789. 
ciliata, 745. 
clathrus, 725. 
clavigera, T59. 
cliftont, 724. 


comoxensis, 759. 

compressa. T11, 757, 
759, 760, 761, 787, 
789, 795. 


cupula, 761. 

ensata, 763. 
extusarticulata, 761. 
fistulosa, 770. 
Joliacea, 760. 
genuina, 711, 760. 
gracilis, 760, 796. 
hodgsoni, T6V. 


indica, 711, 713, 761. 
intermedia, 759, 760, 
804. 


invenusta, TAL. 
labyrinthica, 762. 
lacunosa, 722. 
levigata, 761, 793, 
796. 
lieberkithniti, 723. 
lobata, 760, 795. 
longipilis, T60. 
mirabilis, 761. 
monstruosa, 760. 
multicavata, 725. 
nivea, 773. 
pennigera, 760. 
phillipsii, 761. 
pulchra, 726, 727. 
scotti, 760. 
singularis, 761, 794. 
solida, "734. 
striatula, 786. 
strohbilus, 761. 
subhispida, 748. 
tenuis, 760. 
tessellata, 749. 
truncata, 765. 
tuberosa, 761. 
urceolus, T61. 
virgultosa, 746. 
vosmaeri. 711, 760. 
Grantilla, 751, 798, 804. 


hastifera, 752, 756. 


quadriradiata, 76, 
797. 
Grantiopsis, 766, 798. 


cylindrica, 762, 763. 
infrequens, 11, 769, 
Tol. 
Grus, 698. 
australasiana, 1053. 
Gryponyx 
africanus, 638. 


XXVil 


Guancha, 790. 
blanca, 724, 790. 
Guire 
piririgua, LOST. 
Gulapinnus, 1000. 
Guttera 
cristata, 1051. 
edowardi, 1051. 
pucherant, 1051. 
Gymnoschizorhis, 641. 
Gymnothorax 
dovti, 373. 
meleagris, 373. 
Gypaétus, 645. 
barbatus, 1049. 


Heematopus 
ostralegus, 1055. 

Halcampa, 968. 
chrysantheilum, 

ale 
clavus, 969. 
fultoni, 970, 972 
medusophila, 970. 
septentrionalis, 969. 

Halcampoides 
abyssorum, 969. 

Haleyon 
pileata, 1060. 
rufa, 1060. 

Haliaétus 
leucegaster, 1049. 

Halysis 
diaphana, 482. 

Haplochilus 
melanopleura, 988, 
melanops, 984. 

Haswellia 
auriculata, 512. 
australiensis, 462, 511, 

OUZA520} 
coronata, 512. 
gracilis, 512. 
grandipora, 512. 

Hebius, gen. nov., 424. 

Heliconius 
galanthus, 343. 
— subrufescens, 

367. 

Helicops 
angulatus, 424, 
leopardinus. 425, 
modestus, 425. 

Heliornis, 678. 
Fulica, 1054. 

Hemeschara 
gigantea, 518, 

Hermathena 
owen, 300, 

Herpetodryas 
carinatus, 1034. 


970, 


343, 


367. 


XXVIII 


Herpetosuchus, 629. 
granti, 626. ~ 
Hesperia 
parvipuncta, 366. 
Hesperornis, 394, 396. 
regalis, 395. 
Hetrandria, 980. 
affinis, 984. 
colombianus, 996. 
cubensis, 1002. 
Jasciata, 995. 


formosa, 977, 994, 995. | 


holbrookii, 983. 
lutzi, 996. 

minor, 1003. 
nobilis, 985. 
occidentalis, 1018. 
petruelis, 984. 


Heteropegma, 728, 737, 
790. 
nodus-gordit, 706, 736- 
738, 790. 
Heteropia, 798. 
compressa, 752. 


erecta, 72. 
glomerosa, 711, 754. 
macera, TOD. 
patulosculifera, 752. 
pluriosculifera, 752. 
polyperistomia, 758. 
ramosa, TO4. 
rodgeri, 754. 
simplex, 754. 
spissa, 753. 
Heterusia 
cingala, 614. 
Hexelasma 
sp., 847. 
arafurae, 847. 
aucklandicum, 
841, 843, 847, 
854. 
corolliforme, 847. 
velutinum, 847. 
Hierococcyx 
varius. 1057. 
Hierofaleco 
candicans, 542. 
Himantodes 
cenchoa, 1035. 
Himantopus 
nigricollis, 1055. 
Hincksiella, 483. 
Hippopodina, 515. 
feegensis, 514. 
Hippoporina, 515. 
Hippothoa 
distans, 462, 501. 
divaricata, 462, 501. 
Hirundo, i069. 
rustica, 1066, 1067. 


840, 
848, 


Holocentrum 

punctatissimum, 3873. 
Holocentrus 

caudovittatus, 379. 
Holoporella, 461, 

509, 529. 

albirostris, 463, 

522, 523, 537. 

aperta, 463, 522. 

bispinata, 522, 523. 

celosia, 521. 

cidaris, 521. 

columnaris, 463, 505, 

621. 

pigmentaria, 509. 
Holoptychius, 957. 
Holoscalpellum, 850. 
Homalocranius 

alticola, 1036. 

coralliventre, 

1038. 

melanocephalum, 1030. 

Homandra, 717, 720, 
788. 

Jalcata, 790. 
Hometta, 791. 
Homoderma, 744, 

802 
sycandra, 


716, 
748. 
Howesia, 628, 632. 
Hydra 
viridis, 427. 
Hydrocherus 
hydrocherus, 1092. 
Hydrophasianus 
chirurgus, 1055. 
Hydrophis 
fasciatus, 414. 
Hydrornia 
alleni, 1052. 
Hydrothelphusa, 
923, 924. 
agilis, 915, 922. 
Hyla 
‘ arborea, 1028, 1024. 
baudintt, 1023. 
maxima, 1022, 
rubra, 1026. 
Hylobates 
lar, 813. 
Hylodes 
conspicillatus, 1025. 
palmatus, 1025. 
Hymenolepis, 564. 
Hyperodapedon, 628. 
Hypocrinus, 894, 897, 
899, 903. 
miilleri, 896, 900, 913. | 
piriformis, 896, 900, 
905, 907, 910-913. 


508, 
516, 


1035, 


791, | 
742, 


918, 


1028. 


INDEX OF SCIENTIFIC NAMES. 


Hypocrinus 
pyriformis, 896, 905. 


schneideri, 895, 896, 
900, 901, 904, 910, 
913, 

Hypodictyon 

longstaffi, 44, TAT, 

TOM 


Hypograntia, 791. 
extusarticulata, 761. 
hirsuta, 752. 
infrequens, 762, 

TM. 
intusarticulata, 753. 
medioarticulata, 753. 


763, 


Hypotnidia 
philippensis, 1052. 
Hypoxanthus 
rivolt, L063. 
Hyracotenia 


hyracis, 552. 


Tanthia 
rufilata, 814. 
Tbis, 690. 
Ichthyoteenia, 562. 
Icterus 
jamaicat, 1065. 
mesomelas, 1104. 
mississipprensis, 1104. 
Tetinia, 1050. 
mississiptensis, 1049. 
Inermicapsifer 
capensis, 568. 
Ischnognathus 
lineatus, 424. 
Ithomia 


bolivart, 341, 367. 


Jacana 

jacana, 1055. 
Julis 

umbrostigma, 371. 


Kebira, 798, 800. 
uteoides, 78d. 
Kellogella 
oligolepis, 373. 
Ketupa, 651. 
Kuarrhaphis, gen. nov., 
767, 780, 798, 803. 
cretacea, 775, 780. 
Kuhlia 
boninensis, 374, 3875, 
378. 
cerulescens, 875-377. 
caudovittata, 375, 378, 
379. 


Kuhlia 


humilis, 369, 374, 375, 


379, 380. 
maculata, 378. 
malo, 378-381. 


marginata, 874, 875, 


317, 378. 


mutabunda, 3869, 373- 


375, 381. 
prozima, 380. 
rupestris, 314-377. 
— hedleyi, 379. 


sandvicensis, 369, 374, 


379, 381. 
sauvagii, 375, 377. 
splendens. 374, 379, 

379, 380. 
sterneckit, 381. 
teniura, 375, 381. 
urvillit, 375, 377. 
venura, 381. 


Labrichthys, 368. 
fuentesi, 371, 375.. 
luculenta, 371. 

Labyrinthodon 
leptognathus, 997. 

Lacerta 
viridis, 546. 

Lachesis 
atrox, 1037. 
brachystoma, 1037. 
lanceolatus, 1037. 
monticellii, 1037. 
mutus, 814, 1037. 
punctatus, 1037. 

Lagenipora, 508. 
borywt, 511, 
lucida, 511. 
nitens, 511. 
rota, 462, 510, 511. 
socialis, O10, 511. 

Lagothrix 
poppigt, 1091. 

Lalocitta 
lidthit, 1104. 


Lamontia, 768, 778, 798. 


zona, 179. 
Lampanyctus 
crocodilus, 1097. 
Lanius 
excubitor, 1066. 
Larus 
argentatus, 1055, 
ridihundus, 105d. 
Leander, 428. 
(Palemon)  squitla, 
427. 
Lebistes, 981, 1010. 
pecilioides, 1008. 


INDEX OF SCIENTIFIC NAMES, b--db:4 


Lebistes 
reticulatus, 978, 1007, 
1008. 
Lechriorchis 
inermis, 933, 936. 
primus, 934. 
validus, 933, 935, 936. 
Leevthiocrinus, 896. 
adamsi, 899, 900, 913. 
Lelapia, 739, 740, 784, 
798, 800, 804. 
australis, 711, 785. 
Lepralia, 461. 
adpressa, 515. 
circumcineta, 515, 
cleidostoma, 517, 518. 
— mermis, 463, 517. 
clivosa, 503. 
crassa, 519. 
cucculata, 528. 
dorsiporosa, 518, 519. 
Jeegensis, 463, 514, 
Silisy, lays}oy, 
Foliacea, 503. 
hippopus, 515, 524, 
inornata, 507. 
japonica, 518. 
longipora, 510. 
margaritifera, 503. 
mont ferrandi, 506. 
occlusa, 489, 519. 
pallasiana, 518, 515. 
pertusa, 502. 
porcellana, 517. 
prelonga, 515. 
prelucida, 5198. 
rectilineata, 515. 
striatula, 463, 507. 
syringopora, 504, 529, 
turgescens, 507. 
turrita, 463, 516, 537. 
venusta, 506. 
wasinensis, 463, 516, 
ole 
(Mucronella). pre- 
lucida, 517. 
Lepreus 
| tricolor, 1091. 
Leptodactylus 
pentadactylus, 1025. 
Leptodira 
albo fusca, 1035. 
Leptognathus 
annulata, 1037. 
spurrelli, 1036, 1038. 
temporalis, 1037. 
Leptophilus 
crumeniferus, 1046. 
| Leptophis 
bilineatus, 1034. 
occidentalis, 1035. 


Leptoptilus, 668, 672, 
678, 677-679, 681, 
685, 685, 686, 699. 

argala, 654. 
crumeniferus, 654. 
Leptorhaphis, gen. noy., 
980, 998. 
infans, 998. 
Leptosia 
xiphia, 616, 
Lerodea 
phyllas, 361. 
rupilius, 360, 367. 
Leucaltis, 712, 736, 738, 
757, 798, 800, 804. 
bathybia, 773. 
— mascarenica, ‘737, 
738. 
clathria, 706, 710, 728, 
736-738, 790. 
crustacea, 770. 
Horidana, 734. 
— australiensis, 734. 
helena, 774. 
impressa, 774. 
nausicae, TTA. 
pumila, T74. 
solida, 734. 
(Heteropegma), 804. 

Leucandra, 709, 731, 733, 
736, 738, 758, 766, 
168, 779, 781, 783, 
798, 804. 

alcicornis, 777. 
— cespitosa, 777. 
amorpha, 772. 
ananas, 769. 
anfracta, 772. 
anguinea, 769, 
anomaa, 769, 
apicalis, 72. 
armata, 769, 
aspera, 730, 769, 774, 
775, 795. 

_ australiensis, 711, 769, 

803. 
balearica, 772. 
bathybia, 773. 
bomba, 732. 
brumalis, 774. 
bublosa, 772. 
callea, 772. 
caminus, 769, 789. 
capillata, 770. 
cartert, 734. 
cataphracta, 777. 
cirrata, 770. 
cirrhosa, 770. 
claviformis, 770. 
coimbre, 772. 
compacta, 770. 


XXK 


Leucandra 
conica, 772. 


crambessa, 770, 776, 
789. 

— callea. 772. 

erosslandi, 772. 


crustacea, 77). 
cucumts, 735, 778. 
cumberlandensis, 770. 
curva, 774. 

cyathus, 736. 
cylindrica, 772. 
donnant, 770. 
echinata, 711, 770. 
egedii, 770, 790. 
elongata, 777. 
falcigera, 770. 
fernandrica, 772. 
fistulosa, 770, 790. 
Frigida, 774. 
gelatinosa, 774. 
gemmipara, 770. 
gladiator, nae 

gossei, 710, 791. 
heathii, 772. 

helena, 774. 

hiherna, 770. 

hirsuta, 770. 

hispida, 711, 771. 
impressa, 774. 

infesta, 769, 771, 803. 
inflate, 774. 
innominata, T74. 
intermedia, (74. 
johnstonii, 773. 
joubini, 772. 
kerquelensis, 771. 
lendenfeldi, T7\. 

levis, TTA. 

lohata, 773. 

loricata, 772. 
lunulata, 771. 
masatierre, 771. 
meandrina. T1\. 771. 
minima, T11, 771. 
minor, 772. 

multifida, 773. 
multiformis. 771. 
nausicae, T7T4. 

nived, 773. 
ochotensis, 
ovata, 773. 
pailida, 771. 
pandora, 774. 
panicea, 77 


775. 


75. 
phillipensis, T11, 771. 
plate, 772. 
prava, Hbitexs 
pulvinar, 771, 792. 
pumila, 774, 790. 
pyriformis, 773. 


Leucandra 
rodrigueztt, 773. 
rudifera, 773. 
saccharata, 734. 
sagittata, 774. 
schauinslandi, 774. 
solida, 787. 
spissa, 173. 
stylifera, 776. 
taylori, 773. 
teluim, 774. 


thulakomorpha, 711, 
771. 

typica, 773. 

vaginata, 771. 

valida, 771. 

verdensis, 774. 

villosa, 771. 

vitrea, 773. 

wasinensis, 772. 

(Leucetta) pandora, 
785. 

(Leucortis) pulvinar, 
78). 


Leucaseus, 716, 717, 732, 
736, 786, 798, 800. 


clavatus, 10, 730, 
Veil 

insignis, 710, 731. 
simplex, 710, 730, 
Tile 


Leucetta, 709, 731, 735, 
738, 757, 758, 768, 
783, 798, 800, 804. 

carteri. TH. 
chagosensis, 710, 753. 
clathrata, 724. 
corticata, 739. 
expansa, 710, 734. 
floridana, 734. 
haeckeliana, 739. 
homoraphis, 734. 
imperfecta, 739. 
infrequens, T34. 
microraphis, 710, 734. 
pandora, 774. 
— anomala, 769. 
— intermedia, 774. 
-— loculifera, 777. 
primigenia, 730, 732, 
734, 789, 790-792, 
796. 
— microraphis, 734. 
prolifera, 710, 734, 
796 
pyriformis, T10, 734. 
sagittata, T74. 
sambucus, 739. 
schauinslandi, 774. 
solida, 734. 
trigona, 734. 


INDEX OF SCIENTIFIC NAMES. 


| 
| 


| 


— 


Leucetta 
vera, 739. 
(Teichonella) prolifera, 

762. 

Leucettaga, 768, 798. 
loculifera, V77. 
loculosa, 749. 

Leucettusa, 732, 728, 

757, 768, 798, 800, 
804. 
corticata, 738, 739. 
dictyogaster, 710, '736, 
738, 739. 
haeckeliana. 733, 739. 
imperfecta, 739. 
sambucus, 739. 
vera, 138, 739. 
(Leucetta) haechkeliana, 
738. 
(—) vera, 738. 

Leucilla, 733, 736, 768, 

778, 728, 798. 
amphora, 730, 783. 
australiensis, 711, 

783. 
capsula, 784. 
connexiva, TAR, 
crosslandt, 774. 
echinus, 784, 792. 
intermedia, 769, 

803. 
leuconides, 784. 
nuttinyt, 784, 793. 
oxeodraginifera, T34. 
princeps, 711, 783, 734. 
prolifera, 734. 
proteus, 784. 
saccharata, 784. 
uter, 784, 793. 
wasinensis, 772. 

Leuckartea, 791. 
natalensis 748, 791. 

Leucogvpsa, 778, 791 

795. 
algozensis, 786. 
gosset, 770, 791. 

Leucomalthe, 731, 768, 

798, 800. 
bomba, 732. 

Leucometra, 791. 
primigenia, 791. 

Leuconia 
anfracta, 772. 
armata, 769. 
balearica, 77 
cirrhosa, 770. 
cotmbre, 772. 
compacta, 770. 
crucifera, 768. 
dura, 734. 
echinata, 770. 


| 
=I 
— 


y) 


ae 


Leuconia 
erinaceus, 752. 
Sernandensis, 772. 
Jistulosa australiensis, 
769. 
Sruticosa, 734. 
glomerosa, 754. 
hispida, 771. 
johnstonii, 773. 
— australiensis, 783. 
levis, 774. 
lendenfeldi, 771. 
lobata, 773. 
loricata, 772. 
masatierreé, 771. 
minor, 772. 
multifida, 773. 
multiformis, 771. 
— amorpha, 172. 
— capillata, 770. 
nivea, T34. 
ovata, 773. 
platei, 772. 
prava, 773. 
pumila, 774. 
pyriformis, 773. 
rodrigueztt, 773. 
rudifera, 773. 
somesi, 723. 
spissa, 775. 
stylifera, 776. 
typica, 773. 
vitrea, 773. 
Leucopsila, gen. noy., 
775, 768, 798, 803. 
stylifera, 776. 
Leucopsis, 792. 
pedunculata, 726, 731, 
792. 
Leucortis, 792. 
anguinea, 769. 
elegans, 771. 
pulvinar, 771, 792. 
Leucosarcia 
picata, 1056. 
Leucosolenia, 705, 708, 
TG, 7, ZN), "eat, 
798, 812. 
agassizti, 724. 
ameboides, 720. 
angulata, 721. 
arachnoides, 721. 
armata, 721, 788. 
atlantica, 72). 
bella, 709, 721, 742. 
ilanca, 724, 790. 
botryoides, 721, 
723. 
botrys, 721. 
canariensis, 720, 723, 
72+, 787, 788, 796. 


INDEX OF SCIENTIFIC NAMES. 


| 


Leucosolenia 

cancellata, 724. 

caroli, 724. 

cavata, 710, 720, 724. 

cerebrum, 724. 

cervicornis, 721. 

challengeri, T24. 

charybdea, 724. 

clarkit, 721. 

clathrata, 724, 728, 
789. 

clathrus, 725, 

complicata, 709, 721, 
799. 

confervicola, 721. 

contorta, 721. 

convallaria, 7295. 

corallorhiza, 721. 

cordata, 721, 787. 

coriacea, 710, 725, 793- 
796, 799. 

darwinii, 725. 

decipiens, 725. 

densa, 721, 796. 

depressa, 710, 725. 

dictyoides, 729. 

discoveryt, 722. 

dubia, 722. 

echinata, 722. 

echinoides, 722. 

eleanor, 722. 

fabricii, 722. 

falcata, 709, 720, 722, 
790. 

Jalklandica, 725. 

flexilis, 725. 

fragilis, 722. 


W217. 
goethet, 725. 
gracilis, 725, 
793. 
grantit, 725. 
hermesi, 722. 
himantia, 726. 
hispidissima, 722. 
horrida, 722, 788. 
incerta, 722. 
intermedia, 724. 
irregularis, 722. 
japonica, 726. 
lacunosa, 722. 
lamarckhit, 726, 788. 
laminoclathrata, 727. 
laxa, 722. 
lieberkithnii, 723. 
loculosa, 726, 788, 794, 
796. 
lucasi, 709, 718, 720, 
723, 742. 


Proc. Zoou. Soc.—1913, No. LXXIV. 


| 


OM! 
Leucosolenia 
macleayt, 726. 
minchint, 723. 
ninoricensis, 726. 
multiformis, 726. 
nansent, 724. 
nitida, 723, 793, 794. 
osculum, 726. 
panis, 723. 
pedunculata, 726, 792. 
pelliculata, 710, 726. 
philiipina, 726. 
pinus, 728. 
poterium, 726. 
primordialis, 726, 727, 
788, 793. 
protogenes, 726. 
proxima, 710, 727. 
psammophila, ‘727. 
pulcherrima, 710, 727. 
pulchra, 727. 
reticulata, 723. 
reticulum, 723, 792. 
robusta, 723. 
rosea, 727. 
sagittaria, 727. 
sceptrum, 727. 
sertularia, 723. 
spinosa, T27, 
spongiosa, 727. 
stipitata, 727. 
stolonifer, 709, 718, 
720, 723: 
sulphurea, 727. 
sycandra, T48. 
tenutpilosa, 723. 
tenuis, 723. 
thamnotdes, 72°2. 
trépodifera, 728. 
uteoides, 729. 
variahilis, 710, 7238. 
ventricosa, 709, 720 
724. 
vesicula, 727, 739. 
vitrea, 710, 727. - 
wilsont, 727, 777. 
(Ascandra) falcata, 
720. 


? 


Leuculmis, 792. 
echinus, 784, 792. 
Leucyssa, 767, 798, 

803. 

eretacea, 780. 

tncrustans, 759. 

-— lichenoides, 779. 

— villosa, 780. 

spongilla, 779. 
Lichenocystis, 897. 
Lichenopora, 461. 
Limenitis 

cealidasa, 614. 

74 


XXX]1 INDEX OF SCIENTIFIC NAMES. 


4 
Limia, 981, 1010. | Mamosa | Menipea 
arnoldi, 1014, 1016, elegans, 1092. | flagellifera, 473, 474, 

1018. Mastigina, 799. 476. 
caudofasciata, 1014, Medon Suegensis, 474. 

1017, 1018. barbata, 786, 792. gracilis, 473. 
couchiana, 1008. wmberbis, 786. occidentalis, 473. 
cubensis, 1014. »Megaleema patagonica, 473, 534. 
dominicensis, 1014, virens, 1063, 1064. | portert, 473. 

LOTS; LOM Megalichthys, 957. smithii, 4738. 
formosa, 1012. Megalotis ternata, 473, 475. 
heterandria, 1014, cerda, 558. Menura 

1017, 1018. Megapogon, 755, 769, superba, 1065, 1067. 
matamorensis, 1011. 798, 803. Mergus 
nigrofasciata, 1014, crispatus, 767, 768. serrator, 1047. 

1015, 1018. cruciferus, 767, 768. | Merlia, 792. 
ornata, 1014, 1016, poilicaris, 767, 768. normant, 792. 

1018. raripilus, 767, 768. | Merops 
pecilioides, 1011. | _villosus, 767, 768. |  sp., 615. 
versicclor, 1009, 1014, | Megistias |  apiaster, 1060. 

1017. cantho, 363, 367, | -philippensis, 1060. 
vittata, 1014. Melanerpes | Merula 

Limnea, 427, 437. flavifrons, 1091. |  tamaulipensis, 1.066. 
peregra, 434, 439. | Melanitis | Mesites, 645, 702. 
Limosa | ismene, 614. Mesocestoides 
lapponica, 1054. | Melanocetus, 1097. ambiguus, 549. 
Liophis | johnsonii, 1096. | Mesosemia 
albiventris, 1035. | Meleagris albipuncta, 350, 367. 
Liothrix gallopavo, 1051. Mesosuchus, 628, 632. 
luteus, 1066. Meliceritites |  brownt, 619, 627, 632, 
Lipostomella, 792. | magnifica, 505. | 633. 
capsuia, T84. |  royana, 505. Metopiana 
elausa, 734, 792. | _vielbanci, 505. peposaca, 1047. 
Lithonina, 739. | Melopsittacus Micropholis, 956. 
Lophius, 960. | wndulatus, 1058. Micropora 
piscatorius, 1097. Membranipora, 529. articulata, 490, 492. 
Lophoceros | armata, 459, 462, 486— | ratoniensis, 490, 
erythrorhynchus, 106). | 488, 534, 535. _ Microporella 
Lophortyx | catenularia, 462,488. | ciliata, 463, 523. 
douglasi, 1052. | eraticula, 475. | Microtus 
Lophura lineata, 475. agrestis, 834, 838, 
nobilis, 10938, | magnilabris, 498. 839. 
Lorius mamillaris, 497. — exsul, 830-834, 
donvicella, 1058. marginella, 488. | 839. 
Lota | melontha, 461. | — levernedii, 834. 
vulgaris, 428. | monostachys, 488. | —macgillivrati, 822. 
TLoxomma, 958-958, 961. | nigrans, 488. 831, 832, 834, 839. 
Lutra panhoplites, 487. | — neglectus, 833, 834. 
leptonyx, 1092. savartii, 462, 486, | arvalis, 838, 839. 
Lycaon | 535. | corneri, 838. 
pictus, 538. unicornis, 475. hirtus, 834. 
Menipea, 472. — bailloni, 834. 
aculeata, 474. orcadensis, 838. 
Macacus benemunita, 475. sandayensis, 838. 
silenus, 1103. | buskii, 473, 534. sarnius, 838. 
Machetes cervicornis, 473, 474.  ~ | Mimo 
pugnax, L054, |  cirrata, 473-475. dumontt, 1065. 
Macropora clausa, 475. Mimus 
cribrilifera, 496. crystallina, 478, orpheus, 1066. 
Macropus, 592. | ATA. | Minchinella, 739, 798, 
Malapterurus cyathus, 474, 554. 800. 


electricus, 1093. flabellum, 475, | damellosa, 710, 740. 


INDEX OF SCIENTIFIC NAMES. 


Mitua 
tuberosa, 1051. 
Mlea, 792. 
dohrnii, 771, 792. 
Mobiusispongia, 792. 
parasitica, 792. 
Mollia 
tubereulata, 507. 
Mollienisia, 978, 981. 
elongata, 1010, 1013. 
formosa, 1010, 1012. | 
gracilis, LO10, 1012. 
Jonesit, 993. 
latipinna, 1010, 1011. 
occidentalis, 1010, 
1013 
petenensis, 1010, 1012. 
sphenops, 1009, 1010, 
1012. | 
spilurus, 1010, 10138. 
Molo 
apella, 865, 367. 
herea, 365. 
humeralis, 365. 
nebrophone, 364, 367. 
Molothrus 
fringillarius, 1094. 
Molpastes 
hemorrhous, 614. 
Momotus 
lessoni, 1060. 
subrufescens, 1060. 
Monacanthus 
cirrifer, 373. 
Monieza, 569. 
Monopylidium, 
872. 
macracanthum, 571. | 
rostellatum, 568, 876, | 
877. 
unicoronata, 877. 
Moronopsis 
argenteus sandvicensis, 


567, 


fuscus, 377. 

sandvicensis, 881, 
Mueronella, 461. 

vultur, 518, 519. 
Murrayona, 789, 740, 

798, 800. 

phanolepis, 710, 741. 
Mus 

musculus, 838. 
Musophaga, 639. 

violacea, 641. 
Mustela 

erminea ricine, 829. 
Mycteria, 699. 

americana, 1046. 
Myiarchus | 

tyrannulus, 10G5. 


Myopsittacus, 1059. 
monachus, LO58. 
Myriozoella, 520. 
Myriozoum 
australiense, 511. 
coarctum, 520. 
marionensis, 520, 521. 
simplex, 520. 
subgracile, 520. 
truncatum, 520. 
Myripristis 
pralinius, 373. 
Myrtillocrinus, 898. 
americanus, 899, 
élongatus, 899. 
Mysis 
chameleon, 427. 


Naia 
bungarus, 1093. 
nigricollis, 1093. 
Nannopterum 
harrisi, 393, 899, 401, | 
A02. 


| Nanocrinus, 897, 898. 


Nanodes 
(Lathamus) discolor, 
1058. 
Napeogenes 
hemisticta, 340, 367. 
Nardoa 
arabica, 726. 
canariensis, 724. 
labyrinthus, 725. 
pelagica, 752. 
reticulum. 728, 792. 
rubra, 724. 
spongia, 725. 
spongiosa, 727. 
sulphurea, 724. 
Nardoma, 793. 
nitida, 793. 
Nardopsis 
gracilis, 721, 798, 
horrida, 722. 
Nardorus, 793. 
primordialis, 793. 
Nascus 
phintias. 358. 
phocus, 358. 
Necturus. 403, 


Nellia, 49v. 


tenella, 489. 


_ Neophron, 661, 


Nephelis, 427. 


| Nicoria 


nasuta, 1030, 
Niltava 

grandis, 814, 
Notamia 


bursaria, 465, | 


XXXi1ll 


Nothura, 1051. 

maculosa, 1050. 
Notodela 

leucura, 814, 
Nototragus 

melanotus, 1092. 
Nycticorax, 649, 656, 679, 

686, 699. 
gardeni, 654, 1045, 


1046. 
violaceus, 1045. 
Nyetidromus, 1065, 
1072. 
albicollis, 1062. 
Nymphicus 


uveensis, 1058. 


Oceanodroma, 1044. 
Octacnemus, 440. 
Octodon 
deqgus, 1095. 
Octorchis, 970. 
Ocydromus 
australis, 10D2. 
Gidemia 
negra, 1O4T. 
Ciidicnemus, 861. 
bistriatus, 861, 863. 
edicnemus, 861. 
scolopax, 1055. 
CEnus 
nausiphanes, 363, 367. 
Oligodon, 424. 
Olindias, 969. 
Olynthella, 793. 
coriucea, 799. 
Olynthium, 798. 
nitidum, 723, 793. 
splendens, 723. 
Olynthus, 719, 793, 
798. 
cathus, 722. 
guancha, 724. 
hispidus, 72). 
pocillum, 721, 
simplex, 726, 793. 
Oncorhynchus, 576. 
gorbuscha, 572, 
608, 610. 
keta, 572, 577, 604, 
605, 610. 
kisutch, 572, 599-602, 
610. 
masu, 572, 
nerka, 572, 586, h88— 
590, 592, 593, 610. 
guinnat, 572. 
tschawytscha, 572, 
B75, 585, 594, 
597, 598, 610. 


74" 


602, 


574, 
o95, 


XXXIV 


Oochoristica, 557, 567, 
568, 571, 876. 
wagenert, 874, 875. 


Opisthocomus 

cristatus, 1052. 
Oreicola 

Serre, 814. 
Oriolus 

galbula, 1066. 
Orizomys 


longivaudata, 1092. 
Ornithorhynchus, 1072. 
Ornithosuchus, 629, 631. 

taylori, 625, 626, 630, 

633. 
woodwardi,, 620, 624- 
626, 6380, 633. 
Ortalis 

araucuan, 1104. 

ruficauda, 1051. 
Ortygospiza 

atricollis, 1093. 

Ortyx 
virginianus, 1052, 
Orycteropus, 886, 888, 
889, 892, 893. 
Osearella, 738. 
Osinotreron 

bicincta, 1056. 
Osteolepis, 957. 
Osthimosia, 510. 

zanzibariensis, 

508, 537. 
Ostinops 

decumanus, 1065. 
Otiditzenia, 564. 

Otis 

tarda, 1054. 
Ovis 

musimon, 1094. 
Oxynaspis, 938, 942, 

948. 
Oxyrhopus 
clelia, 1035. 


462, 


Pachynus 
brachyurus, 1058. 
Palemon, 930. 
dolichodactylus, 
926. 
hildebrandti, 925, 928, 
931, 932. : 
hilgendorfi, 926. 
ide, 927, 928. 
lepidactylus, 925, 926. 
mooret, 929. 


925, 


ritseme, 925, 927, 
928. 


sintangensis, 928. 
(Kupalemon) 7itseme, 
927. 


| 
| 


Palzemon 
(Macrobrachium) Jepi- 
dactylus, 926. 
(Parapalemon) do- 
lichodactylus, 926. 
Palxornis 
fasciata, 1058. 
Palxosyops, 886, 893. 
Palamedea 
cornuta, 1047. 
Palmicellaria, 461. 


Paludicella, 429, 431, 
444, 446, 448, 
449. 


articulata, 427, 489, 
441, 443, 447, 456. 
ehrenbergii, 441. 
Pamphila 
angularis, 366, 
Pamphoria, gen. nov., 
980, 1003. 
scalpridens, 1008. 
Pamphorichthys, gen. 
noy., 980, 1003. 
minor, LOO. 
Pancerina, 571. 


| Pandion, 1069, 1072. 


haliaétus, 1050. 
Papilio 

agamemnon, 616. 

aristolochie, 614, 617, 
618. 

demoleus, 615. 

hector, 614, 615. 

jason, 615, 618. 

nomius, 615. 

parinda, 614. 

polytes, 614-616, 618, 
619 


— cyrus, 615, 616. 

— polytes, 614-618. 

— romulus, 614-616. 
Papio 

anubis, 1090. 
Paragambusia, 982. 

nicaraguensis, 985. 
Paraleucilla, 735, 798. 

cucumis, 778. 
Paramecium, 428. 
Paraphysa 

manicata, 1093. 


| Parathelphusa, 925. 


tridentata, 923. 
Parotia 

lawestt, 1065, 1066. 
Passerculus 

savanna, 1091. 
Pauxis 

galeata, 1051. 
Pavo 

nigripennis, 1098. 


INDEX OF SCIENTIFIC NAMES. 


Peachia 
boeckii, 966, 967. 
carnea, 966, 967. 
fultoni, 970. 
hastata, 963, 964, 966, 
967, 970, 971. 
koreni, 966, 967. 
quinquecapitata, 963, 
966, 967-969, 972. 
tricapitata, 964, 967. 
triphylla, 966, 967, 
972. 
undulata, 966. 
Pectinatella, 431. 
Pelargopsis, 1065. 
Pelecanus, 661, 688, 698, 
703, 1045. 
Penelope, 1052. 
pileata, 1104. 
purpurascens, L051. 
Pennella, 1078, 1089, 
1090. 
Pericharax, 731, 758, 
768, 798, 800. 
cartert, 735, 778. 
— heteroraphis, 735. 
— homoraphis, 734, 
735. 
heteroraphis, 710, 735. 
peziza, T10, 735, 
780. 
polejaevi, 739. 
Pericrocotus 
brevirostris, 814. 
Peripatoides 
orientalis, 612, 613. 
suteri, 611. 
woodwardi, 611, 612. 
Perisama, 348. 
barnest, 347, 867. 
Peristomella, 521. 
Petalognathus 
nebulata, 1085. 
Petalosoma 
amazonum, 981. 
cultratum, 981. 
Petalurichthys, 981. 
Petaurus, 631. 
Petralia, 488. 
castanea, 519. 
chuakensis, 463, 518, 
519, 535. 
dorsiporosa, 518. 
ellerii, 519. 
Japonica, 463, 518, 
519 


thenardii, 519. 

undata, 519, 

vultur, 519. 

— armata, 463, 518, 
519, 535. 


INDEX OF SCIENTIFIC NAMES. 


Petralia 
(Escharella) bistnwata, 
519. . 
(Mucronella) castanea, 
519. 
(—) magnifica, 519. 
(—) porosa, 519. 
Petrostroma, 798. 
schulzei, 740. 
Phaéthon, 661, 1045. 
Phalacrocorax, 395, 
1045. 
albiventris, 401, 402. 
auritus, 401, 402. 
bicristatus, 596. 
carbo, 400. 
dilophus, 402. 
magellanicus, 400, 402. 
pelagicus, 400-402. 
penicillatus, 398-400, 
402. 
punctatus, 402. 
urile, 400, 402. 
vigua, 401, 402. 
Phalloceros, 978-980. 
caudomaculatus, 999, 
1001, 1018. 
Phalloptychus, 978, 980, 
999 


januarius, 998, 999, 
1018. 


Phaps 

chalcoptera, 1056. 

elegans, 1056. 
Philander 

laniger, 1094. 
Philomedusa 

vogtii, 969, 972. 
Phlogeenas 

cruentata, 1056. 

luzonica, 1056. 
Pheenicopterus, 670, 671, 

681, 687, 703. 

antiquorum, 1047. 
Phonicosia, 508. 
Phrynonax 

JSasciatus, 1034. 
Phyciodes 

dora, 345, 367. 

niveonotis, 847, 367. 

phlegias, 346, 367. 
Phyllium 

erurifolium, 546. 
Phyllobates 

chocoensis, 1028. 

pratti, 1026. 
Phyllomedusa 

hypochondrialis, 1025. 
Phyllopezus 

africanus, 1055. 
Physa, 427. 


| Plotus, 668, 


Picolaptes 

affinis, 1065. 
Pieris 

diana, 356. 

limona, 356, 367. 
Pitangus 

sulfuratus, 1065. 
Pitta 

strepitans, 1065. 
Plagiosternum, 956. 
Planorbis, 427, 437. 
Platalea 

leucorodza, 1047. 


Platemys 
platycepha'a, 1104. 
Platycercus 
eximius, 1058, 1059. 
Platypcecilus, 980. 
couchianus, 1003, 
1004. 


maculatus, 1001, 1004, | 


1005. 

perugi@, 1014. 

tropicus, 1013. 
Platystethus 

cultratum, 379. 
Plectroninia, 740, 798. 

deansti, 741. 

halli, TAL. 

hindet, 741. 


| Plegadis 


Jalcinellus, 651, 1047. 
691, 
695, 703. 

anhinga, 396, 397, 654, 
655, 702. 
levaillanti, 655. 
Plumatella, 429, 4381, 
436, 488, 448. 
coralloides, 449, 450. 
emarginata, 452-454. 


— muscosa, 435, 441, | 


452-455, 457. 
— spongiosa, 453, 455. 
fruticosa, 449, 452- 
454. 


fungosa, 427, 449-452, | 
454 


== coralloides, 437, 
441, 449-451, 454, 


457. 


polymorpha, 450, 452, 


454, 
punctata, 452. 
repens, 450, 452-454, 
457. 
spongiosa, 455. 
Podargus, 1065, 1069, 
1072. 


cuviert, 1062. 
nacunda, 650. 


693, | 


XXXV 
Podica 
| senegalensis, 1054. 
| Podiceps 


cristata, 1044, 
| minor, 1044. 
Podoa 
surinamensis, 1054. 
Peecilia, 981, 999, 1002. 
amates, 1013. 
amazonica, 1006. 
| branneri, 978, 1005, 
1007. 
couchiana, 1003. 
cuneata, 1010, 1013. 
decemmaculata. 1000. 
dominicensts, 1015, 
1017. 
| elongata, 1018. 
| _feste, 996, 
gracilis, 1000. 
heteristia, 1007, 1008. 
lineolata, 1011. 
| maculata, 1004. 
melanogaster, 1015. 
|. melanopleura, 988. 
| 
| 


multilineata, 1011. 

occidentalis, 1013. 

pare, 978, 1005, 1006, 
1009. 

petenensis, 1012. 

picta, 1005, 1007-1009, 
1018. 

pittieri, 997. 

presidionis, 996. 

reticulata, 1008. 

retropinna, 997. 

salvatoris, 1013. 

schneideri, 1005. 

sphenops, 1012. 

spilonota, 1013. 

spilurus, 10138. 

surinamensis, 1006. 

tenuis, 1013. 

unimaculata, 1006. 

vittata, 1014. 

vivipara, 978, 
107-1009. 

— pare, 1006. 

| Peciliopsis, gen. 
996. 980, 1010. 

isthmensis, 996-998, 
1018. 

| dutzi, 996. 

| pittiert, 996, 997. 
pleurospilus, 996, 

997 

presidionis, 994, 996. 
retropinna, 996, 997. 

Pceocephalus 
Fuscicapillus, 1094. 
meyert, 1058. 


1008, 


noy., 


XXXV1 


Polejna, 793. 
telum, 774. 
uter, 793. 


Pollicipes, 840, 842, 852 


957, 958, 941, 947. 


aucklandicus, 841, 847, | 


848. 

rigidus, 946. 
Polyborus, 661. 
Polychrus 

gutturosus, 1032. 
Polyodontophis 

bivittatus, 428. 
Pomacentrus 

inornatus, 3870, 873: 

genkinsi, 370, 371. 

ntomatus, 370. 
Porella 

acutirostris, 503. 

plana, 603. 
Poricellaria, 492. 
Porina 

coronata labrosa, 511. 
Porphyrio 

porphyrio, 1052, 1053. 
Porzana 

carolina, 1052. 
Potamon 


bombetokense, 916, 919. 


edule, 924. 
fluviatile, 923-925. 


goudoti, 919-921, $28, 


924. 
grandidieri, 916, 919. 
humbloti, 916, 919. 


madagascariense, I15— | 


917, 921, 923-925. 


methuent, 920, 923 
924, 982. 
(Parathelphusa) 
tongilense, 925. 
(Potamon)  goudoti, 


915, 920. 


(--) madagascariense, 


915, 916. 
(—) methueni, 915, 
920: 
(—) pittarellii, 925. 
(Potamonautes) de- 
pressum,s 925. 
Precis 
iphita, 614. 
Presbytis 
entellus, 1108. 
Priapella, gen. nov., 
992, 979. 
bonita, 993, 994. 
Priapichthys, gen. nov. 
991, 979, 995, 995. 
annectens, Y91, 992 
S94, 


INDEX OF SCIENTIFIC NAMES. 


? 


a 


an= | 


d 


Priapichthys 
episcopt, J92., 
nigroventralis, 992. 
parismind, 991, 992. 
tridentiger, 992. 
turrubarensis, 992. 
Procellaria, 1044. 
Prosycum, 793. 
primordiale, 726, 
simplicissimum, 793. 
Proteus, 403, 
Protopterus 
ethiopicus, 542. 
Pseudocordylus 
microlepidotus, 548, 
1091. 
Pseudoflustra 
solida, 518. 
Pseudolabrus 
inscriptus, 371. 
Pseudomonacan thus 
ayraudi, 372. 
paschalis, 372, 373. 


Pseudopeecilia, gen. nov., 


995, 980. 
Teste, 994, 996. 
Pseudotantalus, 661. 
ibis, 654, 702, 1046. 
Pseudoxiphophorus, 
ioe 
bimaculatus, 993, 994. 
Jonesii, 993. 
pauciradiatus, 993. 
terrabensis, 993. 


Pseudupeneus 
multifasciatus, S73. 
Psophia 


erepitans, 1058. 
leucoptera, 1053. 
obscura, 1053. 
Pterocella, 483. 
Pterocles 
alchata, 1056. 
Pteroglossus 
inseriptus, 1063. 
Pteronymia 
fumida, 341, 367. 
godmani, 341. 
simplex, 341. 
Pteroplax, 955, 956. 
Ptilorhis, 651. 
Ptychobothrium 
belones, 552. 
Ptyonius 
estocephalus, 959. 
Pudu 
pudu, O91. 
Pygosceles 
papua, 1091. 
Pyrrhula 
nepalensis, 1091. 


Querquedula 


castunea, 1047. 


Rallus 


celebensis, 1052. 
longirostris, 1052. 
maculatus, 1052. 


Rana 


capito, 1092. 
holsti, 1028. 
palmipes, 1026. 


Raphiceros 


melanotis, 539. 
sharpet, 539. 


Raya, 960. 


Recurvirostra 


avocetta, 1050. 


Retepora, 523, 529. 


axillaris, 527. 
cellulosa, 489, 503, 
524, 
contortuplicata, 524. 
denticulata, 463, 526, 
586. 
elongata, 524. 
Jormosa, O24. 
gelida, 524, . 
hippocrepis, 524, 
hirsuta, 463, 528, 524, 
527. 
imperatt, I24. 
ancrustata, O24. 
inordinata, 524. 
Jjermanensis, 463, 526. 
lapralicides, 524. 
monilifera, 524, 525. 
— wmbonata, 524, 
536. 
novezealandie, 524. 
philippensis, 527. 
procellana, 525. 
— laxa, 525. 
producta, 463, 525, 
536. 
stnwosa, 524. 
solanderia, 524. 
tessellata, 524. 
tubulata, 468, 526. 
wmnbonata, d24. 


Reteporella 


dendroides, 526. 


| Rhabdodermella, 793. 


nuttingt, 784, 793. 


Rhamphastos 


discolor, 1068, 1064. 


Rhamphoceelus 


brasilius, 1065. 


Rhawphostomella, 520. 
Rhea, 663. 


americana, 1043, 


INDEX OF SCIENTIFIC NAMES. 


Rhinochetus, 645, 651, 

Jubatus, 1054. 1092. 
Rhinthon 

luctatius, 361, 367. 
Rhizodopsis, 957, 958, 

962. 

Rhizodus, 957, 958, 962. 
Rhyacornis 

fuliginosa, 814, 
Rhynchea 

capensis, 1059, 
Rbynchosaurus, 628, 
Rhynchotus, LO5L. 

rufescens, L050. 
Rbynchozoon 

bispinoswin, 523. 

longirostre, 523. 

profundum, 525. 

— laminatum, 463, 

523, 536. 

Rhytidoceros 

undulatus, 1061. 
Rollulus 

roulroul, 1052. 


Safole, 374. 
Salarias 
arenatus, 372. 
Salicornaria, 404. 
Salicornia, 494. 
Salmo 
fario, 594. 
gairdneri, 572, 576, 
577, 605, 606, 610. 
salar, 574, 579, 581, 
582,607.  - ‘ 
Sareorhainphus 
gryphus, 1091, 
Sauropatis 
chloris, 1060. 
sancta, 1060. 
sordida, 1060. 
vagans, 1060. 
Savignyella, 482, 
Sceorhynchus 
ruficeps, 814. 
Scalpellum, 840, 851, 
946, 947. 
sp., 82. 
aucklandicum, 
842. 
cret@, 937, 941, 942. 
magni, 842, 945. 
michelottianum, 853. 
— gassinensis, 853. 
quadratum, 853. 
subplanum, 842, 848, 
854. 
tuberculatum, 937, 938, 


9403, D44. 


841, 


Sealpellum 
| wulgare, 945. 
| zancleanum, 8d0. 


(Arcosealpellum) wngu- | 


| latwm, 850, 853, 854. | 
| 


| Sceloporus 
spinosus, 1092. 

| Schismopora, 509, 510, 

| Schizoblastus, 900, 

Schizoporella, 459. 

| auriculata, 505, 515, 
biaperta, 509, 508. 

| biturrita, 520. 

| cecil, 508. 

| cerustacea, 520. 

galeata, 505, 506. 
harmsworthit, 50d. 
levigata, 506. 
levata, 506. 

| linearis, 504, 505, 515. 

| longirostris, 505. 

| marsupifera, 506. 

| 


montferrandt, 462, 
506. 
nivea, 462, 502-506, 


1 a8, 687. 


— wasinensis, 504, 506. 


pachnoides, 506. 
pertusa, 462, 502. 
polymorpha, 520. 
ridleyi, 505. 
rimosa, 506. 
sanguinea, DUD. 
stmplex, 50d. 
spongites, SO. 
striatula, 507. 
subimmersa, 515, 
triangula, 615. 
tuberosa, 520. 
tumida, 506. 
unicornis, 462, 501, 
504, 505. 
— errata, d0d. 
venusta, DUG. 
viridis, dV. 
vulgaris, 50d. 
Schizotenia 
haymanni, 867. 
Schultzierinus, 898. 
Scillwlepas, 937, 938. 
carinata, 947, 948. 
ornata, 948. 
parone, 948, 
Scincosaurus 
crassus, 959. 
Sciurus 
| saltuensis bonde, 814. 
Sclerodomus, 522, 
| Scleromochlus, 651. 
| taylori, 629, 632. 
Scoliocrinus, 897, 898. 


XXXVI1 


Scolopax 
rusticula, 1Od4. 
Scopus, 645, 646, 648- 
652, 661, 671-674, 
676, 682-709, 703. 


umbretia, 650, 699, 
701, 1045. 
Seruparia 


chelata, 466. 
diaphana, 482. 
Serupocellaria, 459, 481. 
annectans, 477. 
bertholettit, 474. 
cervicornis, 462, 
477, 554. 
ciliata, 477. 
clypeata, 477, 480. 
delilii, 474. 477. 
diadema, 477. 
jerox, 462, 474, 476, 
534. 
incurvata, 476, 477. 
inermis, 473, 475. 
jolloisit, 473, 474, d34. 
macandret, 462, 474, 
476-478, 534. 
mansueta, 474, 476, 
479. 
marsupiata, 475. 
minuta, 77. 
obtecta, 474, 477, 479. 
occidentalis, 474. 
ornithorhynchus, 
477. 
pilosa, 462, 478, 534. 
portert, 474, 
pusilla, 478. 
reptans, 474. 
scabra, 473-475. 
scruped, 477, 478. 
seruposa, 474. 
— dongolensis, 474. 
serrata, 475, 479. 
smitti2, 473. 
varians, 479. 
WasiNensis, 
534. 
Seuticella, 483. 
Seypha, 704. 
coronata, 793. 
ovata, 769. 
Scyphognathus, 629, 631. 
Selenidera 
maculirostris, 1063, 
1093, 1095, 1096. 
Serpentarius, 661, 1048, 
1069. 
Sertularia, 494. 
Seymouria, 955. 
Simia 
satyrus, 1092, 


474, 


474, 


462, 479, 


XXXVIll 


Siniopelta, 510. 

Siphonactinia 
boeckii, 966. 
tricapitata, 966. 

Siren, 403. 

Sistrurus 
miliarius, 10938. 

Sitta 
cinnamomeiventris, 

814. 

Smittina, 461, 515. 
sp., 463, 514. 
longtrostris, 514. 
nitida, 513. 
oculata, 514. 
trispinosa, 508. 

— bimucronata, 513. 
— nitida, 518. 


— protecta, 468, 513. 


— spathulata, 463, 
513. 

tropica, 468, 514. 
Solea, 960. 
Solenidium, 794. 

mitidum, T94. 
Soleniscus, 794. 

loculosus, 794. 
Solenula, 794. 

coriacea, 794. 
Sorex 


araneus, 824, 825, 838, | 


839. 
— araneus, 825. 
— castaneus, 823-826, 


828, 839. 
— tetragonurus, 825, 


828, 838, 839. 


grantti, 822, 824-826, | 


Spongia 


inflata, 748, 769, 774. 


nivea, T7193. 

panicea, 769, 775. 

pulverulenta, 769. 

urceolus, T61. 
Spongilla 

lacustris, 427, 4389, 

973. 

Stagonolepis, 629. 
Starncenas 

cyanocephala, 1056. 
Steatornis 

caripensis, 1062. 


| Steganoporella 


minutus, 826-828, 838. | 
Speotyto 

cunicularia, 1062. 
Spherium 

(Cyelas), 427. 
Spheniscus 


demersus, 1044, 1085. 


Sphenodon, 632. 
Sphenophorina 
singularis, 761, 794. 
Sphenophorus 
singularis, 794. 
Spilotes 
pullatus, 425, 
Spongia, 794. 
ananas, 769. 
botryoides, 721. 
ciliata, 704, '745. 
complicata, 721. 
compressa, 704, 760. 
coriacea, 725. 
coronata, 7495. 
foliacea, 760. 


| Styriodes, gen. nov., 361. 


alveolata, 499. 
buskit, 498, 501. 
connexa, 499. 
lateralis, 499. 


magnilabris, 462, 468, | 


489, 498-500, 536. 
simplex, 499. 
sulcata, 199. 
truncata, 499. 
tubulosa, 499. 

Stenorhina 
degenhardti, 1036. 


| Stentor, 428. 
| Stilesia, 552. 


Stirparia, 458, 465. 


annulata, 468, 469. 
dendrograpta, 462,468— 
471, 533. 


exilis, 462, 468, 471, 


DSoe 
glabra, 468, 471. 
zanzibariensis, 462, 
469, 470, 534. 
Streptoconus, 791, 794. 


australis, 744, 745, 794. 


Streptophorus 
atratus, 1021, 1034. 
Stringops, 1059. 
habroptilus, 1058. 
Strix 
castanops, 1091. 
flammea, 651, 
1062. 
Strongylopora, 483. 
Struthio, 679, 1048. 


lyco, 361, 367. 
Sula 

bassana, 395, 1045. 

capensis, 1085. 
Sycaltis 

conifera, 743. 

glacialis, 753. 

leuconides, 784. 

ovipara, 782. 

perforata, 786. 


1061, 


INDEX OF SCIENTIFIC NAMES. 


| 


Sycaltis 


testipara, 782. 


Sycandra, 777, 798, 802. 


alcyoncellum, 746. 
— virgultosa, 749. 
ampulla, 'TA4. 
— alopecurus, 744. 
— petiolata, 747. 
arborea, 746. 
arctica, T44. 
— maxima, TAT. 
— polaris, 747. 
aspera, 744. 
barbadensis, 745. 
borealis, 745. 
capillosa brevipilis, 
759. 
— lengipilis, 760. 
ciliata, 745. 
— lanceolata, 746. 
— ovata, 747. 
clavigera, 759. 
coacta, 745, 
compressa, 760. 
— clavigera, 759. 
— foliacea, 760. 
— lobata, 760. 
— pennigera, 760. 
coronata, 745. 
— commutata, 745. 
— tubulosa, 749. 
elegans, 745. 
— formosa, 746. 
— tessellata, 749. 
glabra, 764. 
— ensata, 763. 
— rigida, 764. 
helleri, 746. 
humboldtii, 746. 
hystrix, 764, 765. 
inconspicua, 746. 
lobata, 760. 
parvula, 747. 
pennigera, 760. 
quadrangulata, 747. 
— tesseraria, 749. 
quadrata, 748. 
ramosa, 748, 791. 
ramsayt, 748. 
raphanus, 748. 
raphanus proboscidea, 
747. 
— procumbens, 747. 
— tergestina, 748. 
schinidtii, 748. 
setosa, 748. 
staurifera, 748. 
tabulata, 748. 
tuba, 749. 
utriculus, 749. 
villosa, 749. 


INDEX OF SCIENTIFIC NAMES. 


Sycandra 
villosa impletum, 746. 
Syeantha 


tenella, 744, 745, 748, | 


Use 

Sycarium 

ampulla, 744, 795. 

rhopalodes, 760. 

vesica, 748. 

villosum, T49. 
Sycetta, 744, 758, 798, 

801, 802. 

asconoides, 743, 760. 

conifera, 748. 

cupula, 761. 

primitiva, 743, 796. 

sagittifera, 748. 

stauridea, 753, 789. 

strobilus, 761. 
Sycidium 

gelatinosum, 795. 
Sycilla 

chrysalis, 782. 

cyathiscus, 782, 798. 

cylindrus, 782. 

urna, 782. 
Sycinula, 790. 

aspera, 795. 

egedii, 770. 

penicillata, 769. 
Sycocystis 

oviformis, 745, 795. 
Sycodendrum 

ramosum, 745, 795. 
Syecodorus, 798. 

hystrix, 764, 765, 796. 
Sycolepis, 779. 

incrustans, TSO, 79). 

palvinar, 77). 
Sycometra, 795. 

compressum, 795. 
Sycon, 704, 716, 720, 733, 


TAZ, (43, 790, 757, | 


758, 798, 802, 803. 
alopecurus, 744. 
ampulla, 744, 798, 
antarcticum, 744, 796. 
arcticum, 744. 
asperum, 744,746, 769. 
australe, '745, 794. 
barbadense, 745. 
boomerang, 710, 745, 

803. 
boreale, 745. 
caminatum, 745. 
carteri, 710, 745. 
ciliatum, 745, 795, 

796. 
coactum, T45. 
commutatum, 745, 
compactum, 745, 


Sycon 


compressum, 760. 
coronatum, 745, 798. 
dendyt, 763. 
eglintonensis, 745. 
elegans, 745, 748, 789. 
ensiferum, 713, 746, 
ioe 
formosuna, TA6. 
gelatinosum, 710, 746, 
78%, 795. 
giganteum, 710, 746. 
heileri, 746. 
humboldtii, 746, 748. 
impletum, 746. 
inconspicuum, T46. 
tncrustans, 746. 
karajahense, 746. 
kergquelensis, 740. 
lambei, 746. 
lanceolatum, 746. 
lendenfeldi, 710, 747, 
748. 
lingua, 747. 
longstaffi, T47, 791. 
maximum, TAT. | 
minutum, 747. 
mundulum, 747. 
munitum, 747. | 
ornatum, 747. 
ovatum, 747. 
parvulum, 747. | 
pedicellatum, T47. 
petiolatum, 744, 747. 
polare, TAT. | 
proboscideum, 747. 
procumbens, 747. 
protectum, T47. 
guadrangulatum, “47, | 
748, 795. 
ramosum, G48, 791, 
795. | 
ramsayt, 710, 748. 
raphanus, 710, 744, | 
748. 
schmidti, 748. 
schuffneri, 748 | 
setosum, 710, 748. | 
stauriferum, 748. 
subhispidum, 748. 
sycandra, 748, 791. 
tabulatum, 748. 
tenellum, 748, 795. 
tergestinum, 748. 
tessellatum, 749. 
tesserarium, 749. | 
tuba, 749. | 
tubulosum, 749. 
verum, 710, 718, 749. 
villosum, 749. 
virgultosum, 749. 


XXX1X 


Sycon 
(Acyoncelllum) gelati- 
nosum, 787. 
Syconella 
proboscidea, 747. 
quadrangulata, 747, 
795. 
Syeophyllum 
lohatum, 799. 
Sycorrhiza 
corallorhiza, 721. 
coriacea, 799. 
Sycortis 
levigata, 761, 79d. 
lingua, TAT. 
sycilloides, 753. 
Sycortusa 
levigata, 761, 796. 
Sycothamnus 
Sruticosus, 734, 796. 
Syculmis, 798. 
synapta, 783. 
Sycum 
alopecurus, T44. 
areticum, TA4. 
ciliatum, 796. 
clavatum, 749. 
giganteum, 745. 
lanceolatum, 746. 
lingua, 760. 
ovatum, 747. 
petiolatum, 744, 747. 
primitivus, 196. 
procumbens, 747. 
tergestinum, 748. 
Sycurus, 796. 
primitivus, 796. 
Sycute, gen. nov., 763, 
798. 
dendyi, 757, 763. 
Sycyssa, 798. 
huxleyi, 767. 
Sylvia 
nisorta, 1093. 
Sylvicapra 
coronata, 1090. 
Synnotum 
aviculare, 
466. 
contorta, 
532. 
pembaensis, 462, 465, 
532. 
Synute, 798. 
pulchella, 711, 764. 


462, 465, 


462, 466, 


Tenia 
bifaria, 570. 
cucumerina, Ho. 
genette, d49. 


xl 


Tenia 
marchali, 568. 
platydera, 549. 
Tainsia, 1099, 
Talegalla 
lathami, 1051. 
Tanagra 


sayaca, 1065. 


Tantalus, 688-691, 693- | 


698, 7U3. 
loculator, 1046. 
Tarroma, 796. 
canariense, 796. 
Tarropsis, 796. 
coriacea, 796. 
Tarrus 
densus, 721, 796. 
labyrinthus, 725. 
reticulatus, 723. 
Tatusia, 1099. 
Taxocrinus, 911. 
Teichonella 
labyrinthica, 762. 
prolifera, 734, 762, 796. 
Teichonopsis, gen. nov., 
761, 796, 798. 
labyrinthica, 711, 762. 
Telemiades 
mnemon, 857, 367. 
Tenthrenodes, 796. 
antarcticus, 796. 
scotti, 760. 
Tephreops, 368, 369. 
richardsontt, 370. 
Terpsephone 
paradisi, 614. 
Tethrenodes 
artarcticus, 744. 
scottt, 744. 
Tetrabothrium, 562. 
Netrarhynchus 
tetrabothrius, 551. 
Tetrastemma, 428. 
Teuthis 
umbra, 373. 
Thairopora 
manillaris, 462, 497. 
Thalamoporella, 501. 
mamillaris, 497. 
Thalassoma 
purpurewm, 373. 
Thanatophis 
sutus, 1037. 
Thaumantias 
(Phialidium), 970. 
Thecla 
amphrade, 355, 367. 
dolylas, 353. 
guapila, 354, 367. 
hypocrita, 302, 367. 
Boo 367, 


leos, 358, 


INDEX OF SCIENTIFIC NAMES, 


| 


Thecla 
melma, 894, 367. 
politus, 305. 
subflorens, 353, 367. 
Thecotetra, 7388. 
loculosa, 796. 
Thinocorys 
sp., 1055. 
—rumicivorus, 1O92. 
Thylacinus, 892. 
Tima, 970. 
Tinnuneulus 
alaudarius, 1049. 
Titanolepas, subgen. noy., 
943. 
tuberculata, 938, 943, 
944, 946-948. 
Tomeurus, 979. 
gracilis, 98. 
Toxus, 980. 
riddlei, O02. 
Trachinus 
draco, 636. 
Trachyboa 
houlenger?, 1034, 108 
gularis, 1034. 
Trachypoma 
macracanthus, 373. 
Trematosaurus, 956, 
Tribonyx . 
mortiert, L052. 
Trichogaster 
Jasciatus, 542. 
Trichoglossus, 1059. 
nove-hollandie, 1058. 


Trichogypsia, 767, 798, 
805. 

incrustans, 779, 780, 
795. 


villosa, 779, 780, 795. 
Triclaria 

cyanogastra, 1104. 
Tripleurocrinus, 898. 
Trogon 

atricollis, 1063. 

puella, 1068. 
Tropidonotus 

vibakari, 423, 424, 

vittatus, 424. 


Trypostega 

venusta, 462, 506, 507. 
Tubucellaria 

cereoides  chuakensis, 


462, 512. 
Susiformis, 462, 512. 
opuntioides, 494. 
zanzibariensis, 

512. 

Tubulipora, 461. 
Turacus, 639. 
corythaia, 1057, 


462, 


Turacus 
donaldsont, 641. 
Jischeri, 641. 
merian, 641. 
Turdus 
tristris, 1066. 
Turnix 
dussemiert, 10d). 
Tyrannus 
melancholicus, 10698. 


Unio, 428. 

Upupa 
epops, 1061. 

Urceolipora 528. 

Urocordylus 
reticulatus, 959. 
wandesfordii, 959. 

Urogalba 
puradisea, 1063. 

Uromys 

ae she 

Ute, 754, 798 
oe ae. 
capillosa, 759, 764. 
chrysalis, 782. 
ensata, 763. 
glabra, 764. 
papillosum, 786. 
rigida, 764. 
spencert, 764. 
spiculosa, 711, 764. 
syconoides, 711, 764. 
utr ees ee 764. 
viridis, 786, 787. 

Utella, 764. 
hystriz, 796. 

eee gen. noy., 766, 

58, 798. 

gente 766. 


Vanellus 
vanellus, 1050. 
Vespa 
cincta, 618. 
Victorina 
steneles stygiana, 348, 
Bo 
Vittaticella, 
505. 
buskei, 485. 
contet, 484. 
elegans, 462, 484, 485, 
533. 
elegans zanzibariensis, 
462, 484, 485, 533. 
(Catenaria), 484, 
Viverra 
genetta, 549, 


465, 483, 


Vivipara 

(Paludina), 428. 
Vorates 

cotiso, 364, 367. 
Vorticella, 428. 
Vosmaeria 

corticata, 776, 777. 

gracilis, 760, 76. 


Vosmaeropsis, 798, 803. 


connexiva, 756. 
cyathus, 756. 
dendyi, 711, 756. 
depressa, 711, 759. 
macera, 711, 75d. 
primitiva, 711, 756. 
sericatum, 756. 
wilsoni, 711, 735. 
Vulpes 
chama, 392. 


(Alopex) lagopus, 391. 


Wagnerella, 797. 


INDEX OF 


| 


SCIENTIFIC 


Wagnerella 
borealis, 797. 


Xanthichthys 

lineopunctatus, 373. 
Xenopeltis 

wnicolor, 415, 417. 
Xenopus 

miillert, 1093. 
Xenorhynehus, 688, 692- 

694, 696, 703. 

Xiphophorus, 978, 980. 

bimaculatus, 993. 

brevis, 1005. 

gracilis, 998. 

guenthert, 1004, 1005, 

helleri, 1004, 1008. 

Jjalape, 1004. 

montezume, 1004, 
Xiphophorus 

rachovti, 1005. 

strigatus, 1004. 


NAMES. 


Yphtbima 
ceylonica, 614. 


Zamenis 
gemonensis, 933. 
grahani, 1092. 

Zaudidura 
carolinensis, 1056. 

Zeugmatolepas, gen. nov., 

9338. 
eret@, 942. 
macklert, 937, 
941, 942, 948. 

Zonurus 
giganteus, 548. 

Zy gonectes 
atrilatus, 984. 
brachypterus, 985. 
inurus, 985. 


938, 


INDEX 


OF 


ILLUSTRATIONS. 


Acanthistius fuscus, Pl. LV. p. 368. 

Acmepteron poasina, Pl. LIT. p. 839 

Adeonella platalea, Pl. LXXIIT. p. 458 ; 
Fig. 82, p. 530. ; 

Adeonellopsis crosslandi, Pl. LXXIII. 
p. 458. 

Aetea unguina, Pl, LXIV. p. 458. 

truncata, Pl. LXIV. p. 488. 

fix sponsa, Fig. 181, p. 1048. 

Agalychnis calcarifer, Pl. CII. p. 1019. 

spurrelli, Pl. CIII. p. 1019; 
Fig. 177, p. 1025. 

Alfaro amazonum, Fig. 169, p. 993. 

Amblystoma tigrinum, Figs. 75, 76, 
pp. 406, 407. 

Anadia vittata, Pl. OVIT. p. 1019. 

Anampses pulcher, Pls. LVILII., LIX. 

. 68. 

Das breviceps, Pl. CVIT. p. 1019. 

Anoplotherium, Figs, 150-157, pp. 879— 
882, 886, 887, 890. 

Archonias intermedia, Pl. LII. p. 539. 

Ardea, P|. UXXXIII. p. 644. 

Ate canace, Pl. LIV. p. 339. 


Baleniceps rex, Pls. UXXX., LXXXI. 


p. 644; Figs. 119-132, pp. 646- 
695. 

Balenoptera borealis, Pl. CXII. 
p. L073. 


brydei, Pls. C1X.-CXIII. p. 1073. 

Batrachiderpeton lineatum, Pls. XCV1., 
XCVII. p. 949; Figs. 162-166, 
pp- 950-954. 

Belonesox belizanus, Fig. 169, p. 990. 

Bicellaria chuakensis, Pl. LXVIII. 
p- 458. 

Bicidium equoree, Pl. XCVIIL. p. 963. 

—— parasiticum, Pl. XCVIII. p. 963. 

Bifaxaria vagans, Pl. LXV. p. 458. 


Bolla machuca, Pl. LY. p. 339. 

sodalis, Pl, LIV. p. 339. 

Brachyrhaphis rhubdophorus, Fig. 169, 
990. 

Brettia tropica, Pl. LXLV. p. 458. 


Browniella africana, Pl. LXXIX. 
p- 619. 
Bubalis caama_ selbornei, Fig. 135, 
p- 820. 


Bufo hypomelas, Pl. CII. p. 1019. 
Bugula robusta, Pl. LX1X. p. 468. 
Bugulopsis peachii, Pl. LUX1X. p. 458. 
Butleria lethea, Pl. LIV. p. 339. 
lysis, Pl. LIV. p. 339. 
Butorides cyanurus, Fig. 120, p. 650. 


Caberea ellisit, Pl. LXIX. p. 458. 

Cecilia intermedia, Fig. 174, p. 1020. 

palmeri, Fig. 175, p. 1021. 

Calantica (Titanolepas) tuberculata, 
EAROXC VER peor 

Cancroma, Pl. LXXXII. p. 644. 

Canda retiformis, Pl. LXIX. p. 458. 

Canis antarcticus, Figs. 70, 71, 73, 74, 
pp. 889, 387, 389, 390. 

latrans, Figs. 70-72, 74, pp. 885, 
387, 388, 390. 

Carystus subrufescens, Pl. LIV. p. 359. 

Catargynnis dryadina, Pl. L. p. 339. 

Catenaria diaphana, Pl. LXILYV. p. 458. 

Cellaria gracilis, var. _ tessellata, 
Pl. LXVII. p. 458. 

wasinensis, Pl. LX VII. p. 458. 

Ceraterpeton  vreticulatus, Hig. 167, 
p. 958. 

Cervus axis, Fig. 134, p. 816. 

canadensis, Fig. 184, p. 816. 

dama, Fig. 134, p. 816. 

— claphus, Fig. 134, p. 816. 


xliv 


Cervus eustephanus ? 
pp. 548, 544. 

leucurus, Fig. 134, p. 816. 

muntjac, Wig. 134, p. 816. 

unicolor, Fig. 134, p. 816. 

Charis subtessellata, Pl. LILI. p. 339, 

turrialbensis, Pl. LILI. p. 339. 

Chauna chavaria, Big. 179, p. 1041. 

Chlidonia cordieri, Pl. LXY. p. 458. 

Cinosternum spurrelli, Pls. CV., CVI. 
p. LOL9. 

Cnesterodon decemmaculatus, Fig. 172, 
p. 1001. 

Cobalopsis latonia, Pl. LIV. p. 339. 

Cobalus lateranus, Pl. LITT. p. 339. 

laureolus, P\. LIV. p. 339. 

— nigrans, Pl. LIV. p. 339. 

pindar, Pl. LIV. p. 339. 

Corrachia leucoplaga, Pl. LILI. p. 3839. 

Cyclosemia subcerulea, Pl, LIV. 

. 039. 

Cynocephalus anubis, Pls. LXXXVILI.- 

LXXXIX. p. 809. 


Figs. 


Dasypus. See Ewphractus, 

Dendrobates aurotenia, Fig. 178, 
p- 1029, 

— paraensis, Fig. 178, p. 1029. 

—— tinctorius, Fig. 178, p. 1029. 

— yar. chocoensis, Pl, CLY. 
p. LO19. 

war, caer, I OIL 

p- 1019. 


Diagram of Peroneal Muscles in Birds, 
Vig. 190, p. 1071. 

Diplodidymia complicata, TAL, JLRS WIE, 
p- 458; Fig. 79, p, 491. 

Diplopylidium genette, Figs. 
pp. 560, 561, 563, 565, 566. 

Dipylidium caninum, Fig, 88, p. 999. 

dongolense, Figs. 85-87, 
pp. 550, 553, 594, 556. 

Dissura, Pl, LXXXI. p. 644, 


90-94, 


309. 
1OL9, 
IPM 


Echelatus lucina, Pl. LLY. p. 
Hlaps microps, Pl, CVIILI. p. 
Enantia lua costaricensis, 
p. 399, 
Epinosis angularis, Pl. LMI. 
Eresia cela, Pl. L. p. 339, 
sticta, Pl, L. p. 339, 
Hubagis hecuba, Pi. LI. p. 339. 
Eudocimus  longirostris, Fig. 
p. 650. 
Eugonodewm edicnemi, Figs. 141-148, 
pp. 862, 864-866, 868, 870-873. 
Euparkeria capensis, Pls. UXXV., 
LXXVL., LXXVIII., LXXIX. p. 619. 
Eupetoieta poasina, Pl. LI. p. 389. 


LI. 


p. 039, 


1120) 


| 
| 
| 
| 
| 


INDEX OF ILLUSTRATIONS. 


Huphractus — seacinctus, 195, 

pawlO2 

villosus, Figs. 193, 194, pp. 1109, 
1101. 

Euptychia agnata, Pl. L. p. 339, 

drymo, Pl. L. p. 339. 

Eurygona leucon, Pl. LI. p. 339. 

leucophryna, Pl. LIL. v. 339. 

matuta, Pl. LI. p. 339. 

mystica, Pl. LI. p. 339. 

Evotomys alstoni, Fig. 137, p. 829. 


Fig. 


Falco peregrinus, Fig. 182, p. 1049. 
alga hermione, P|. LIV. p. 389. 
Farcimia oculata, Pl. LXVITI. p. 458. 


Hlabellaria (Menipea)  cuspidata, 
Pl. LXIX. p. 458. 

Fredericella  sultana, Pl, UXIII. 
p- 426. 


Gadus eglefinus, Fig. 98, p. 578. 

Gambusia dominicensis, Pl. 
p. 977 ;_Fig. 169, p. 990, 

gracilior, Pl. XOIX. p. 

Fig, 168, p. 983. 

holbrookit, Fig. 169, p, 990. 

nicaraguensis, Fig. 168, p. 983. 

nigropunctata, Fig. 168, p, 983. 

oligosticta, Pl. XCIX. p. 977; 

Fig. 169, p. 990. 

punctata, Hig, 168, p. 983. 

senilis, Big. 168, p. 983. 
wrayi, Pl. XCIX. p. 
Fig. 168, p. 983. 

Gemellipora protusa, Pl. LXX. p. 458, 

Girardinus  metallicus, Fig. 172, 
p. 1001. 

Girellops nebulosus, Pl. LVIT. p, 368. 

Gryponyx africanus, Pl, UXXIX, 
p. 619. 


XCIX. 


O77 ; 


ime 


Heliconius galanthus, ab. subrifescens, 
Pl. LIL: p. 339. 

Hermathena oweni, P\. LILI. p. 339. 

Heternadria formosa, Fig. 170, p. 994. 

Hexelasma aucklandicum, Pl, uXXXV. 
p. 840; Figs. 139, 140, p. 848. 

Holoporelia albirostris, Pl, LXXIIT, 
p. 458. 

Homalocranium coralliventre, Pl. CV ILL, 
p. 1019. 

Hyla maxima, Fig, 176, p. 1028. 
Hypocrinus piriformis, Pl, XC, p. 894; 
Fig. 160, p. 907. 
schnetderi, Pl, XC. p. 
Figs. 158, 159, pp. 901, 904, 


894 ; 


Intromittent Organ in the Peeciiiines, 
Figs. 168-173, p. 983, ete. 
Tthomia bolivari, Pl. L. p. 339. 


INDEX OF ILLUSTRATIONS, 


Kuhlia cerulescens, Big. 68, p. 376. 
—— humilis, Fig. 69, p. 380. 

—— mutabunda, Pl. LVI. p. 368. 
splendens, Fig. 69, p. 380. 


Labrichthys fuentesi, Pl. LVIII. p. 368. 
Lebistes reticulatus, Fig. 1738, p. 1009. | 
Lechriorchis inermis, Pl. XOILT. p. 933. 
Lepralia feegensis, Pl. UXX. p. 458. 
turrita, Pl. LX XIII. p. 458. 
wasinensis, Fig. 81, p. 517. 
Leptognathus spurretli, Pi, 
p- L019. 
Leptoptilus, Fig, 131, p. 686. 
Lerodea rupilius, Pl. L1V. p. 389. 
Limia arnoldi, Pl. CI. p. 977. 
caudofasciata, Pl. OL. p. 977. 
heterandria, Pl. Cl. p. 977. 
—— nigrofasciata, Pl. CL. p. 977. 
ornata, Pl. Cl. p. 977. 
versicolor, Fig. 173, p. 1009. 


CVIII. 


Megalema virens, Fig. 187, p. 1064. | 
Megistias xantho, Pl. LIV. p. 339, | 
Membranipora armata, Pls. LXVIL., | 
LXXI. p. 458. 
savartit, Pl. LXXI. p. 458. 
Menipea buskii, Pl. LXIX. p. 458. 
cyathus, Pl. UXIX. p. 458. 
patagonica, Pl. LXIX. p. 458. 
Mesosenia albipuncta, Pl. LILI. p. 359. | 
Mesosuchus browni, Pls. LUXXVIIL., 
LXXIX. p. 619. 
Microtus agrestis exsul, Fig, 138, p. 832. 
-macgilivrati, Fig. 1838, | 
p. 892. 


Mollienisia sphenops, Fig. 173, p. 1009. 
Molo apelia, Pl. LILI. p. 839. 
nebrophone, Pl. LLY. p. 339. 


Nannopterum harrisi, Pl. UXT. p. 393. 
Napeogenes hemisticta, Pl. L. p. 339. 
Nyeticorax, Fig, 131, p. 686. 

gardeni, Fig. 180, p. 1046. 


Enus nausiphanes, Pl. LIV. p. 339. 


Oncorhynchus gorbuscha, Fig. 116, 
p. 603. 
keta, Fig. 117, p. 605. | 
—— hkisutch, Figs. 112-115, pp. 599- | 
602. 
nerka, Figs. 103-108, pp. 586, 


590, 592, 593. | 
tschawytscha, Figs. 95, 96, 102, | 
109-111, pp. 574, 575, 585, 595, 


597, 598. 
Oochoristica, Fig. 149, p. 875. 


| 


Ornithosuchus taylori, Pl. LXXVII. | 
p- 619. | 
—— woodwardi, Pls. DLXXVIL.- | 


LXXIX. p. 619. 


xlv 


Osteomalacia in a young Anubis 
Baboon, Pls. LXXXVII.-LXXXIX. 
p- 855. 

Osthimosia zanzibariensis, Pl. LX XIII. 
p- 408. 


Palemon hildebrandti, Pl. XCII. p. 914. 

Paludicella articulata, Pl, LXII. p. 426. 

Parathelphusa tridentata, Fig. 161, 
HO23: 

Parotia lawesti, Fig. 189, p. 1066. 

Pavo nigripennis, Fig. 192, p. 1098. 

Peachia quinguecapitata, Pl. XCVIII. 

963. 


Pelecanus, Pl. UXXXIII, p. 644. 

Perisauma barnesi, Pl. LI. p. 389. 

Peroneal muscles in Birds, Figs. 179- 
190, p. 1041 ete. 

Petralia chuakensis, Pl. UXX. p. 458. 


vultur, var. armata, Pl. LXX. 
p. 458. 
Phalacrocorax albiventris, Pl. UX1. 
p. 993 


auritus, Pl. LXT. p. 3938. 
—— dilophus, P\. LXI, p. 393. 
magellanicus, Pl, LXI. p. 395. 
pelagicus, Pl. LXI. p. 393. 
pemcilatus, Pl. LXI. p. 393, 
—— punctatus, Pl. LXT. p. 393. 
urile, Pl. LXTI. p. 3898. 
vigua, Pl. LXT. p. 393. 
Phatloceros caudomaculatus, 
p. 977; Fig. 172, p. 1001. 
Phalloptychus ganuarius, Pl. C. p. 977; 
Fig. 171, p. 998. 
Phyciodes dora, P\. LI. p. 339. 
niveonotis, Pl. LILI. p. 839. 
phlegias, P|. L. p. 339. 
Pieris limona, Pl. LIL. p. 339. 
Platycercus eximius, Fig. 185, p. 1059. 


JPL. GL 


Platypecilus maculatus, Fig. 172, 
p. LOOL. 

Plotus, Pl. LXXXIII. p. 644. 

Plumatella emarginata, var. juscosa, 


Pl. LXITI. p. 426. 

Sungosa, var. coralloides, Pl. UX111. 

p. 426. 

repens, Pl. LXIIT. p. 426. 

Podargus nacunda, Fig. 120, p. 650. 

Pecilia pare, Big. 173, p. 1009. 

-pieta, Pl. ©. p. 977; Fig. 173, 
p- L009. 

—— vivipara, Fig. 173, p. 1009. 

Peciliopsis isthmensis, Pl, C. p. 977; 
Fig. 171, p. 998. 

presidionis, Fig. 170, p. 994 

Pomacentrus inornatus, Pl. LVIII. 
p. 368. 

Porphyrio porphyrio, Fig. 183, p. 1053. 

Potamon fluviatilis, Fig. 161, p. 928. 

madagascariense, Fig. 161, p. 923. 


xlvi 


Potamon methuent, Pl. XCI. p. 914. 

Priapella bonita, Fig. 170, p. 994. 

Priapichthys annectens, Fig, 170, p. 994. 

Pseudomonacanthus paschalis, Pl. LX. 
p: 368. 

Pseudopecilia feste, Fig. 170, p. 994. 

Pseudoxiphophorus bimaculatus, Fig. 
170, p. 994. 

Pteronymia fumida, Pl. L. p. 839. 

Tel, 


Retepora LXXII. 


p. 458. 
monilifera, var. 
LXXIT. p. 458. 
producta, P). LXXII. p. 458. 
Rhanphastos discolor, Fig. 188, p. 1064. 
Rhinthon luctatius, Pl. LIV. p. 889. 
Rhyncozoon profundum, var. laminatum, 
Pl. LXXITI. p. 458. 


denticulata, 


umbonata, Pl. 


Salmo gairdneri, Figs. 97, 118, pp. 577, 
606. 

salar, Figs. 99-101, pp. 579, 581, | 
582. 

Scales of Salmon, ete., Figs. 95-118, 
pp- 574-606. : 

Scalpellum subplanum, Pl. UXXXVI. 
p- 840. 

(Arcoscalpellum) ungulatum, PI. 
LXXXVI. p. 840. 

Schizoporelia nivea, Pls. LXX., LX XIII. 
p. 458; Fig. 80, p. 505. 

Scopus wmbretta, Pl. UXXXT. p. 644; 
Figs. 120, 132, pp. 650, 695. 

Scrupocellaria cervicornis, Pl. LXIX. 
p. 458. 

—=—= ORE: 
p. 458. 

jolloisii, Pl. LXIX. p. 458. 

macaudret, Pl. LXVILI. p. 458. 

pilosa, Pl. LXVIII. p. 458. 

wasinensis, Pls. LXVIII., LXIX. 
p. 458. 

Selenidera 
p. 1096. 


Pls, LXVINI, LXIX. | 


Tales, ht, 


maculirostris, 


INDEX OF ILLUSTRATIONS. 


Skulls of Storks, ete., Pls. LXXXI.- 
LXXXITII. p. 644. 
Sorex araneus castaneus, Pl. LXXXIV. 
p. 821. 
grantit, Pl. LXXXIV. p. 821; 
Fig. 136, p. 825. 
Sponges, Phylogeny of 
Fig. 183, p. 798. 
Steganoporella magnilabris, Pl. LXXII. 
p. 458. 
Stirparia 
p. 408. 
-—— evilis, Pl. LXVI. p. 458. 
zanzibariensis, Pls. 
LXIX. p. 458. 
Stringops habroptilus, Fig. 184, p. 1058. 
Strix flammea, Fig. 186, p. 1061. 
Styriodes lyco, Pl. LIV. p. 339. 
Synnotum contorta, Pl. LXIV., p. 458. 
penbaensis, Pl. LXLV. p. 458. 


Calcareous, 


dendrograpta, Pl. LXVI. 


LXVIIL, 


Tantalus, Pl. UXXXITI. p. 644. 
Telemiades mnemon, Pl. LIV. p. 339. 
Thecla amphrade, P|. LIL. p 359. 
guapila, Pl. LIT. 3389. 
hypocrita, P|. LI. p. 339. 

leos, Pl. LIL. p. 339. 

mela, Pl. LIT. p. 539. 

—-— subflorens, P\. LAI. p. 339. 


Victorina steneles, ab. stygiana, Pl. LI. 
p. 339. 

Vittaticella elegans, Pl. LXV. p. 458. 

-——., var. zanzibariensis, Pl. LXV. 
p. 458. 

Vorates cotiso, Pl. LIM. p. 339. 


Xenopeltis unicolor, Figs. 77, 78, pp. 418, 
417. 

Xenorhynchus, Pl. UXXXT. p. 644. 

NXiphophorus hellert, Fig. 172, p. 100i. 

Zeugmatolepas mockleri, Pl. XCIV. 
p. 937. - 

Zonurus giganteus. Pl. LXXIV. p. 548. 


PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STRUET. 


PROCEEDINGS 


OF THE 


GENERAL MEETINGS FOR SCIENTIFIC BUSINESS 


OF THE 


ZOOLOGICAL SOCIETY 
OF LONDON. 


1913. 


PART III. 


CONTAINING Paces 339 ro 860, witH 40 PuLaTEs 
AND 73 TEXT-FIGURES. 


| 
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SEPTEMBER 1913... =~ -. “© 


PRINTED FOR THE SOCIETY, 
SOLD AT ITS HOUSE IN REGENT’S PARK. 
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Olas [Price Twelve Shillings.] 


LIST OF CONTENTS. 


1913, Part III. (pp. 339-860). 


EXHIBITIONS AND NOTICES. 
Page 
The Sxcrrrary. Report on the Additions to the Society's Menagerie during the month of 
March 1913 2... .... sass Bsa ie aime waavercietceiere Ere de anlastoddntoeiadac sos sees oe O42 


Mr. H. J. Eiwss, F.B.S., F.Z.S. Exhibition of, and remarks upon, the head of an Asiatic 
Wapiti. (Text-figs. 83 & 84.) ......... Boe Bis ike dio Shc lols ae Me hate onctalo rele ts eG erin 542 


Mr. R. H. Burnu, M.A., F.Z.S. Exhibition of malformed antlers of an Axis Deer (Cervus 
LIS) stares onus Rew seaes saicteiniois eioiete dosed ded caaiced so 36-06 elstnlae\olnis love loteenaena 545 


Mr. E. G. Bouumnarr, F.Z.8. Exhibition of living specimens of the Leaf-Insect (Phylliwm 
Crurifolium) ..20+...0.03- aia treorelexe iio setae: nai sie Wiolneleeeen anole Gis iejnefecsie cv eieteae -.. 546 


Mr. E. G. Bovutrzncer, F.Z.8. Exhibition of a melanistic specimen of the Green Lizard _ 
(Lacerta viridis) ..c.secsecsse “oN opibooaconD OGus aya TelswaR anetlalatetenerepctonetererr eje basse OAG 


Dr. R. Broom, C.M.Z.8. Exhibition and Description of a new Species of Golden Mole .. 546 


Dr. R. Broom, C.M.Z8. etiaisad of the female and young of the §. African Lizard, 
Zonurus giganteus. (Pl. LXXIV.).......... wie clobatei quate se tobetareteisiere 18 eiaislcletel ates 548 


The SzcreTary. Report on the Additions to the Society's Menagerie during the month of 
PNT OUD ccna rere eae ra Sod nce ne EEIOAD oth So's seueneiaeks aterete Shoo 5 oe esses. 813 


The Rev. T. R. R. Steszine, M.A., F.RS., F.Z.8., notice of Prof. F. E. Schulze’s 
‘Nomenclator Animalium’ .......... OARS Aero ECC aT CREDO Cocke Beer oS 814 


The Liprartan. Dates of publication of early parts of the Society’s “Transactions” .... 814 


Mr. R. E. Hotvine. Exhibition of a number of antlers, skulls, and phoiographs, illus- _ 
trating variations in the growth of the Antlers of Deer. (Text-fig.134.)............ 815 


Mr. D. Seru-Surru, F.Z.8. Exhibition of the Eggs and Young of tne Mikado Pheasant 
(Calophasis mthado) os-ceecccrevcccrsvereere cs eettennees aa venga gin ates sue ee 818 


PAPERS. 
26. New Species of Rhopalocera from Costa Rica. By Wiuttam Scuavs, F.ZS. 


(Pls. L.-LIY.) eeerteeoreeeneenn ® @eceovce tC eChCeeeeee et ee oveseeeeeeeosevese e@oeeeecsece 339 
97. A Collection of Fishes made by Professor Francisco Fuentes at Haster Island. 

By C. Tarz Ruaan, M.A., F.Z.8S. (Pls. LV.-LX.) ccccee seca ce ee ceneeeenes Abaoos clas 
98. A Revision of the Fishes of the Genus Kuhklia. By C. Tats Ruaan, M.A., F.Z.8. 

(Text-figs. 68 & 69.) ecVeepeeecetreeet®Oce P®soooep ee ee oe Fe eee @Cs.oececrecece tee e@ @¢sece 374 
29, The Affinities of the Antarctic Wolf (Canis antarcticus). als R. I. Pococs, F.R.S., 

E.L.S., F.Z.8., Curator of Mammals. (Text-figs. 70-74.) . ernaharereieneverere -. 882 


Contents continued on page 3 2 Wetine 


Sh tastes teat 


DY Aso is vSi2 ell, Ik, 


Horace Knight del.et lith. West, Newman chr. 


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(6p) 


COSTA RICAN BUTTERFLII 


PAS ASS Jel, Jbl, 


Horace Knight del.et lith. West, Newman chr. 


COSA IRUNCYAIN] © 12) 1O/ DIE IS. Ly aS). 


IWASS), wue)ikSy, IM. IL. 


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West, Newman chr. 


Horace Knight del.et lith. 
GOSIWA WRIGAIN IBUPaa RUE Is S. 


Horace Knight del.et lith. 
COSTA 


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PIA. Sr lhe ele ee 


West, Newman chr. 


IBNAyS) US) hee}, IPM IL IDW/ 


West,Newman chr. 


Horace Knight del.et lith. 


COSTA RICAN BUTE RE iE SS: 


PROCEEDINGS 


OF TIE 


GENERAL MEETINGS FOR SCIENTIFIC BUSINESS 


OF THE 


ZOOLOGICAL SOCIETY OF LONDON. 


PAPERS. 


26. New Species of Rhopalocera from Costa Rica. 
By Wicu1am Scuavs, F.Z.8. 


| Received February 17, 1913: Read March 4, 1913.] 
(Plates L.-LIV.*) 


SYSTEMATIC :— 


Subfam. DanaInz. 


Napeogenes hemisticta, sp. n....... 

Pteronymia fumida, sp. n. 
godmeni, nom. nov. 

Ithomia Bolsa, Sp. We...-.. aseeeeee 


Subfam. SaTyRiInz. 


Huptychia drymo, sp. n. ............ 
5 agnata, Sp. N..........-.. 
Catargynnis dryadina, sp. Ni. ...... 


Subfam. HELIconInz. 


Heliconius galanthus, ab. sub- 
FPO AS OCIS,, B)D> TAG)\ Ye eeappaenareees5o5c 


Subfam. NyMPHALIN”. 


Hupetoieta poasina, sp. n. ......... 
WER ESUCISUZELE,| SPs IS vs eesseasss se eese 
~, wether When 65 it 
Phyciodes dora, sp. n. ...... 
Ay phlegias G.&S., Qin ; 
Fa niveonotis Btl. 
Pym. 


fake 


INDEX, 


Page 


346 


Eubagis hecuba, sp. n. 

Perisama barnesi, sp. n. 

Victorina steneles, ab. stygiana, 
ab. nov. 


Fam. ERYcINID2. 
Eurygona mystica, sp. 0. .....0...++ 


oD UGHEOIOS JVs We sce aoc oopo0s 
op leucophryna, sp. N....... 
op MAtULA, SP. Nl. ........+-s- 


Mesosemia albipuncta, POE Te. cos soc 
Hermathena oweni, sp. 0. 
Corrachia, gen. nov. soe 
leucoplaga, sp. Ibe.» de 

Chari is subtessellata, sp. N.......... 
5 turrialbensis, SPSMeg. swans 


Fam. Lycx#NID2. 


Thecla hypocrita, sp. n. eats 
5 GOTO PIDIHEOD, Ss. oosens ssp see 
Sian gUCOSH SDs buccuemereeere aw Aaas 
6p COATES SNo Do ono cab ess as cdbonc 
cg) GROOT S Ds Wo. oon codossoseo>: 
50 COM OURHOE SH, Do soon secace 


* For explanation of the Plates see p. 367. 


Proc, Zoou, Soc.—1913, No. XXIV, 24 


340 MR. WILLIAM SCAAUS ON 


INDEX (continued). 


Page | Page 
Subfam. Prerin x. | Lerodea? rupilius, sp.n. ......... 360 
Archonias intermedia, sp. 0. ...... 355 | Simio des: eee TON g Goins plas oe 
JEG PIS UOROME, D6 oasseaccncoeccsna BD | Rh nth oe ae athe Ae Ste ie 361 
Acmepteron poasina, sp. n.......... 356 | 6 ae LOY ed OtUS; SD ee aia 362 
Enantia lua costaricensis, subsp.n. 357 | ~?7“"S Se Sy SPs eens ; 
| 5 aureolus, sp. n............. 862 
Kam. ElEspeRriip =. | ap lateranus, SDs Weacoxsonodecy By 
my  FOOREIBIP, Fe Wo covecsscacpad a GhO2) 
Telemiades mnemon, sp.n. .......... 357 | Cobalopsis latonia, sp. n. ......... 363 
Nascus phintias, sp. n................ 3858 | Ginus nausiphanes, sp. un. ......... 363 
Cyclosemia subcerulea, sp... ... 3808 | Megistias xantho, sp. n. ..-... 0.0... 363 
Echelatus lucina, sp.n. ............ 858 | Vorates cotiso, sp.n. .............. 864 
ANA CONDGICO, Fide Ds socscocsoconcenbenccn GOS) || Lalo nebrophone, SO Ua ancnak cad 364 
Bolla sodalis, sp. n. .................. 859 | Molo? apella, sp. n................... 365 
»  machuca, sp. nD. ............... 859 | Carystus subrufescens, sp. n....... 365 
Butleria lethea, sp.n................. 860 | Hpinosis, gen. nov. .................. 3866 
53 UISISS (305 Wood0 006 seo cndananee 360 | Falga? hermione, sp.n. ............ 366 


The following new species are the result of a recent visit to 
Costa Rica, where nearly three years were spent in collecting. 
1081 species of Rhopalocera were taken, 409 more than were 
previously known from that country, and of these I am here 
describing 54 speciesas new. With the Heterocera I was equally 
successful, taking 4000 species, whereas only 560 are recorded in 
the ‘ Biologia Centrali-Americana’ from Costa Rica, to which 
number 87 were added by Pittier and Biolley in Part 11. of the 
‘Invertabrados de Costa Rica.’ 


Subfam. DANAIN#. 


NAPEOGENES HEMISTICTA, sp. n. (PI. L. fig. 1.) 


g. Antenne black on basal half, apical half yellow-brown. 
Head black; whitish points on frons and neck behind; a white 
medial line on vertex. Collar and thorax black; a greyish 
medial line on thorax and small cluster of similar scales behind. 
Abdomen slate-brown above, yellow-white below. Fore wings 
black ; basal third of inner margin and shorter shade above 
submedian rich brown ; spots semihyaline yellowish white ; two 
spots near end of cell obliquely placed ; a spot beyond cell and an 
elongated spot above it on costa ; an outer row of five spots, the 
one between veins 3 and 4 elongated, oblique; seven marginal 
spots, the one at apex preceded by a small elongated spot on 
costa. Hind wings orange-brown, the costa to discocellulars 
grey ; apical portion beyond it black, its inner edge inbent to 
vein 3 or before it, then outbent to near margin, which becomes 
narrower, the black not reaching anal angle; five marginal 
spots ; subterminal spots above and below vein 5, sometimes 
almest obsolescent. Wings below similar; no brown at base of 
fore wing; the costa of hind wing black. 

Expanse 54 mm. 

Hab. Cavillo, Guapiles. 

In British Museum. 


COSTA RICAN BUTTERFLIES, 341 


PTERONYMIA FUMIDA, sp.n. (PI. L. fig. 3.) 


3. Antenne black, apical fourth orange. Head and thorax 
black-brown; a white circle around eyes. Abdomen dark slate- 
brown; a whitish vertical line below terminally. Fore wings 
semihyaline smoky black, the veins, costal margin, inner margin 
to median, and termen narrowly brown- black : faint yellow ich 
spots, one at end of cell close to median; an outangled post- 
medial series of spots, the one between veins 2 and 3 elongated, 
suffusing with the lowest of marginal spots, of which ee 
are fave. Hind wings reddish ‘bow, semihyaline, the margins 
black, more broadly so on outer margin, the inner mar gin very 
finely black; traces of a yellowish snibbenaniie! shade at apex. 
Wings below duller; two white spots at apex of fore wing ; 
mar ginal white spots on hind wings, paired on interspaces. 

Expanse 57 mm. 

Hab, Carillo. 


PTERONYMIA GODMANT, nom. nov. 


This form is the Pteronymia simplea of the Biologia, but not 
P. simplex Salv.; the discocellular line is narrow and curved ; 
apex of hind wings below usually without spot, sometimes ane, 
never two as in P. “simplex Salv. 


Found generally in Costa Rica above 3000 ft. 


ITHOMIA BOLIVARI, sp.n. (PI. L. fig. 2.) 


3. Head black spotted with white. Collar dark reddish 
brown. Thorax black shaded with grey. Abdomen black above, 
ventrally white. Fore wings black; the cell to near end, which 
is filled by the broad oblique fascia from costa, the space between 
veins 2 and 3 to near termen, a large terminal spot between 2 
and 3, and a subapical fascia from costa to vein 3 hyaline, the 
fascia mostly shaded with white. Hind wings hyaline, the veins 
and outer margin narrowly black; the costa grey-black. Fore 
wings below: inner margin fuscous grey; dark spaces brown 
edged with black, more broadly in cell and at apex; base of costa 
black; 4 apical white spots, the one on costa minute. Hind 
wings below: margins brown edged with black; a broad yellow 
streak at base of costa ; veins black; small paired whitish spots 
on three interspaces below apex. 

Expanse 45 mm. 

Generally distributed along the Caribbean coast. 

In British Museum. 


Subfam. SATyRINE. 


EUPTYCHIA DRYMO, sp.n. (PI. L. fig. 6.) 


¢. Body and wings dark brown, the disc of wings lighter 
brown, especially on “fore Wings ; the two lines of randerside 
faintly indicated ; a terminal black line, presen by a faintly 
greyish shade, aol fuscous marginal tener a subterminal fuscous 


24% 


342 MR. WILLIAM SCHAUS ON 


wavy line on hind wings. Fore wings below yellowish buff; the 
two lines bright reddish brown, vertical, the inner line not 
reaching costa; costal margin shaded with grey; a subterminal 
dull brown shade interrupted by an ocellus below costa, black, 
circled with yellow, and containing a bluish-white point; the 
termen greyish crossed by two fine lines, faintly wavy, and a 
terminal fuscous line. Hind wings below lilacine grey tinged 
with pale buff; the two lines parallel, slightly inbent; a large 
ocellus near apex, surmounted by a very small ocellus, and with 
a yellow spot below it; a large ocellus between veins 2 and 3, 
containing two bluish points with a yellow spot above it, out- 
wardly broken; the ocelli edged and connected by a dull brownish 
shade; termen grey, the two marginal lines more deeply wavy, 
shaded with reddish brown at anal angle; a terminal fuscous line. 

Expanse 37 mm. 

Hab. Guapiles. 

Near FZ. confusa Stgr., and #. labe Butl. 

In British Museum. 


EupTycHIa AGNATA, sp.n. (PI. L. fig. 4.) 


3. Wings brown; traces of antemedial and medial darker 
lines. Fore wings: an even marginal darker shade, faintly 
darker edged; the apical spot of underside faintly indicated. 
Hind wings: a large subterminal fuscous spot between veins 2 
and 3; the apical spot of underside indicated; subterminal, 
marginal, and terminal darker lines. Wings below pale grey ; 
the antemedial and medial lines dark reddish brown. Fore 
wings: the two lines vertical; a broad darker brownish-grey 
outer shade; the apical ocellus black, edged with yellow and 
containing a black point; dark subterminal and marginal lines ; 
a fine fuscous brown terminal line. Hind wings: the two lines 
slightly inbent, the outer one angled near inner margin; ocelli 
black, broadly edged with yellow and containing a white point ; 
darker grey shading around ocelli; a small ocellus at apex above 
vein 6, and a large one below it; a small yellow spot below 
vein 5; a larger yellow spot below vein 4 suffusing with the 
large ocellus below vein 3; the subterminal and marginal lines 
dark brownish grey, becoming reddish brown on inner margin ; 
terminal line fuscous brown. 

Expanse 48 mm. 

Hab. Guapiles. 

Near H. antonoé Cramer, found in the same locality and 
E. juan Stgr. 

CATARGYNNIS DRYADINA, Sp. n. (PI. L. fig. 5.) 

3. Body fuscous, clothed with brown hairs; a white lateral 
streak on palpi. Fore wings brown; a vertical outer row of 
white spots on interspaces, broadly edged with darker brown, 


the spots on costa and at submedian fold smaller, followed 
throughout by a broad lighter brown shade, and preceded by a 


COSTA RICAN BUTTERFLIES. 343 


similar triangular shade between veins 5 and 6. Hind wings 
darker, especially terminally ; an outer row of small white spots 
on interspaces ; cilia partly white on interspaces. Fore wings 
below dull olive-brown to beyond cell, its outer edge outcurved 
from costa, uneven; outer third pale buff shaded with dark 
ervey; the white spots as above broadly edged with dull dark 
brown ; a broad subterminal fuscous brown line, lunular between 
veins 5 and 7, obsolescent on costa; some fine dark strize on 
apex ; a terminal interrupted fuscous brown shade. Hind wings 
below dull olive-brown, mottled with white antemedially and 
beyond medial fascia, crossed by darker strie; the medial fascia 
defined by darker edging, broad, outbent in cell to end, downbent 
towards inner margin; the small outer white spots broadly edged 
with fuscous brown, preceded and followed by fuscous brown 
shadings ; the outer margin deeply crenulate from vein 4 to anal 
angle, less so towards apex. 
Expanse 72 mm. 


Hab. Volcano Turrialba, at 8000 feet. 


Subfam. HELICONINA. 


HELICONIUS GALANTHUS, ab. SUBRUFESCENS, ab. nov. (PI. LIII. 
fig. 7.) 


Differs from 17, galanthus Bates on the underside of hind wings; 
the medial space below subcostal black, the fascia being reddish 
brown, limited by a postmedial broad black line, followed by a 
narrow brown shade having its outer edge incurved on interspaces. 
In H. galanthus the medial space is black edged above and below 
with reddish brown. 

A single specimen taken on the slopes of Turrialba. Belongs 
to the group with greyish costa on hind wings. 


Subfam. NyMPHALiIn#&, 
EUPETOIETA POASINA, sp.n. (PI. LI. fig. 2.) 


2. Wings brownish buff. Fore wings: the base olive-brown 
irrorated with black; an antemedial black line in cell, and one 
below it, outset ; a pale spot near end of cell, broadly edged with 
black ; postmedial line black, oblique from costa, inbent, lunular 
below vein 4, angled on veins, followed by a straighter, narrower, 
fuscous line; an outer row of black spots on interspaces, suffusing 
on costa with subterminal fuscous line, which is lunular, inwardly 
toothed on veins; termen fuscous, with a pale interrupted 
lunular marginal line. Hind wings: base darker brown with a 
black line on discocellular, limited by the black medial line, which 
is incurved opposite cell, angled at vein 4, followed by a fuscous 
straighter line; a black shade at base and below cell ; four outer 
black spots between veins 2 and 6; subterminal line lunular 
from veins 6 to 3, otherwise straighter; the pale marginal line 
broader. Fore wings below light brown to postmedial; an 
antemedial black line in cell; the cell spot paler edged with 


344 MR, WLLLIAM SCHAUS ON 


black except on subcostal; space beyond postmedial brownish 
buff, the costa and termen shaded with greyish green; outer 
spots below veins 2, 3, and 4; on costa a dark shade, inbent and 
preceded by some white; subterminal line brown. Hind wings 
below mottled brownish buff and dull green, the veins and termen 
whitish grey; a curved antemedial whitish line above cell; 
medial line fine, fuscous, inwardly edged with whitish grey ; 
subterminal ike fine, brown, outwardly edged w ith whitish ; 
cilia white with brown points at veins. 

Expanse 46 min. 

Hab. Poas. 

In British Museum. 


EReEsIA stricta, sp.n. (PI. L. fig. 7.) 


¢. Antenne black tipped with yellow. Head, collar, and 
thorax black ; a white streak on palpi; two white spots on frons, 
and one on vertex. Abdomen brown above, laterally irrorated 
with white; underneath white with a ventral black line. Fore 
wings black, the spots yellow-white, partly greyish ; a round spot 
at end of cell ; postmedial streaky spots irom costa to vein 4, 
outcurved, and two spots between 2 and 4, inset, vertical; outer 
and marginal spots, both slightly inset between veins 3 and 4, 
the marginal spot obsolescent at apex. Hind wings reddish 
brown ; a black shade along subcostal, expanding from middle of 
wing, forming a broad margin, narrowing to anal angle; marginal 
white spots at apex. Fore wings below: the spots larger, yellow- 
white; some brown shading at base of inner margin. Hind 
wings below similar, but the costa also black with a short yellow 
streak at base; large marginal white spots. 

@. Antenne yellow except at base ; a brown streak below cell 
on basal half. Hind wings with the margin much more narrowly 
black, and no black shading on subcostal, except subterminally ; 
marginal white points from below the apical spots. Hind wings 
below: the costa broadly black; a short yellow streak at base; 
traces of fuscous subterminal spots; the marginal spots tri- 
angular. 

Expanse, 3 53 mm., 2? 60 mm. 

Some males have the costa of hind wings entirely black; the 
species is allied to #. pecilina Bates, but the apex is more bluntly 
produced, and the termen more oblique. 

In British Museum. 


Eresta cara Druce. (PI. L. fig. 8.) 

@. Hnt. Monthl. Mag. xi. (1874) p. 37, fig.; B. C.-A. Lep. 
Rhop. vol. 1. p- 187 (tab. xxi. figs. 2, 3). 

6. Fore wings black; a reddish-brown fascia through entire 
cell, downbent ‘between veins 2 and 3 to near termen, with a 
small black spot on it close to vein 3 postmedially, and shaded 
with yellowish terminally; a small subterminal reddish-brown 
spot just below vein 2; an oblique paler fascia postmedially from 


COSTA RICAN BUTTERFLIES. 345 


osta to vein 5, and large subterminal spots between 4 and 6; a 
costa t 5, and large subt 1 spots betw 4 and 6; 
small round marginal spot, tinged with whitish above vein 4, 
and one below it. Hind wings black; a broad reddish-brown 
fascia from inner margin to near apex; a similar marginal line, 
interrupted by veins, the apical spot paler. Underneath similar 
to female, but all the spots yellow imstead of white; the brown 
fascia on fore wings brighter brown, separated by a black line 
from the large subterminal spot. Hind wings below with the 
fascia as above but tinged with yellow, otherwise agreeing in 
markings with the female. 

Expanse 45 mm. 

Found on the Caribbean coast only. 


PHYCIODES DORA, sp.n. (PI. LI. fig. 9.) 


@. Palpi dark brown, whitish grey below. Head dark brown. 
Body fuscous with fine yellowish irrorations. Fore wings: basal 
half black, its outer edge projecting between veins 4 and 5, and 
less so outcurved between vein 2 and submedian, followed by a 
yellow shade from costa to vein 4, and a yellow spot between 
veins 2 and 3; the base with some yellowish irrorations, leaving 
a geminate antemedial outbent line clear black; two yellowish- 
brown spots across end of cell, suffusing ; a reddish-brown line on 
discocellular ; a medial small yellow spot above submedian ; outer 
half bright brown, irrorated with black subterminally, interrupted 
by round yellow spots edged with black, one below vein 2 and 
another between 4 and 5; three smaller yellow spots from above 
vein 6 to costa; termen fuscous preceded by a sinuous yellow- 
brown line. Hind wings black irrorated with yellow-grey, 
leaving clearer black lines on basal third; a brown line on 
discocellular; a postmedial bright brown fascia from costa to 
vein 2; large outer spots formed by yellow edging, inwardly 
rounded, outwardly straight; these spots fill the entire inter- 
spaces ; a subterminal wavy yellow line; a terminal yellow line 
at anal angle. Fore wings below brighter, the base greyish; a 
yellow-grey shade between the antemedial black lines; the cell 
spots broader forming a band; the yellow shades beyond dark 
space more extended ; the termen brown; a grey shade at apex. 
Hind wings below: base to medial grey-brown, darkest in cell, 
whitish lilacine above cell; a subbasal and an antemedial lunular 
brown line, partly followed by the other less distinct lines; the 
medial line dark brown, inbent on vein 4, outset between vein 2 
and submedian ; postmedial space dull brown followed by a series 
of spots, except on costa, the spot above vein 4 round, fuscous 
brown, the spot below 4 triangular, the others more lunular, 
mostly edged with geminate lilacine lines; a subterminal geminate 
lunular dark brown line; termen lilacine white from apex to 
below vein 6, then dark brown, shaded with still darker brown. 
between spots and lunular line, 

Expanse 39 mm. 

Hab. Poas. 


346 MR. WILLIAM SCHAUS ON 


PuycropEs pHLEGIAS G.&S. (PI. L. fig. 9.) 

-g. B.C.-A. Lep. Rhop. vol. ii. p. 680 (tab. eviii. figs. 21, 22). 

@. Body and wings above fuscous brown, the body and base of 
wings irrorated with ochreous. Fore wings: spots white; an 
oblique line across cell near end, followed by a large white spot 
between veins 2 and 3, and a few white scales above 3 and below 
2; a postmedial oblique broad white line from costa to vein 4; a 
small outer spot on costa; a quadrate spot between veins 4 and 5, 
and a larger spot above submedian, extending just above vein 2 ; 
traces of small whitish spots between veins 5 and 7; traces of a 
fine outcurved subterminal line, broad and well marked from 
vein 4 to near vein 2. Hind wings: a broad postmedial white 
fascia from vein 7 to inner margin, followed below vein 4 by a 
line of ochreous irrorations; subterminal line slightly lunular, 
fine, white on costa, then ochreous. Fore wings below: base and 
costa mottled pale brown and grey; a broad antemedial fascia 
defined by edging of dark outcurved lines; the white line at end 
of cell downbent to inner margin; the space between postmedial 
and outer spots fuscous; apex lilacine grey; a broad terminal 
dark brown shade between veins 4 and 6; termen more narrowly 
brown at tornus; subterminal line fuscous brown. Hind wings 
below lilacine grey, broadly shaded with white postmedially 
between veins 2 and 7; basal half crossed by fine brown lines ; 
the postmedial better defined, incurved on costa, and followed 
there by a brownish shade; a fine lunular outer line, with a series 
of faint greyish spots beyond it; subterminal line geminate, 
lunular, filled in and preceded by a dark grey shade from vein 7 
to vein 2; termen dark brown from above vein 5 to below vein 2. 

Expanse 39 mm. 


Hab. Peralta. 


PHYCIODES NIVEONOTIS Btl. & Druce. (PI. LIII. fig. 10.) 


6. Cist. Ent. i. (1872) p. 100, fig.; B.C.-A. Lep. Rhop. 
vol. i. p. 200 (tab. xxi. fig. 27). 

@. Body above fuscous. Wings fuscous brown, Fore wings: 
a large white spot just beyond middle from submedian to near 
vein 2, edged with yellow-brown, irrorated with black ; the outer 
broad white line from costa to vein 5 similar; small subterminal 
similar spots between wings 4 and 6, and a yellow-brown spot 
below vein 4. Hind wings: a yellow-brown line on discocellular ; 
a line of similar irrorations beyond postmedial fascia which has 
its edge also finely yellow-brown; subterminal line brownish, 
faintly indicated towards inner margin. ore wings below light 
brown; an antemedial fascia defined by dark lines; two whitish 
spots near end of cell; a brown spot on discocellular edged with 
black; a broad black postmedial shade from costa, outecurved to 
near tornus; a subterminal lunular dentate darker brown line ; 
the spots all whitish. Hind wings below: base grey-brown 
crossed by brown lines; the whitish fascia obscured by brown 


COSTA RICAN BUTTERFLIES. 347 


irrorations and crossed by a fine line; outer margin broadly 

brownish, irrorated partly with grey; an outer row of fuscous 

grey spots on interspaces ; a marginal lunular dark brown line, 
Expanse 30 mm. 


Hab, Juan Vinas. 


EUBAGIS HECUBA, sp. n. (PI. LI. figs. 4, 5.) 


¢. Body above fuscous, whitish underneath. Wings bluish 
green. Fore wings: apex and outer margin black, expanding at 
vein 4, its inner edge obliquely outbent to vein 2, below which it 
is very finely black. Hind wings: a marginal and a terminal 
black line, suffusing at apex. Wings below white. Fore wings: 
costa grey-brown with a white streak at base ; subcostal medially 
red-brown ; a fine blue line and red-brown streak below subcostal 
to beyond cell; a red-brown streak on discocellular, edged with 
metallic blue and then black; a broad medial black fascia from 
end of cell, and an antemedial black fascia inbent across cell, 
interrupted by median and narrowest below it; a broad black line 
above vein 3 from cell, to a similar oblique line which extends 
from costa to brown terminal space; another black line from 
vein 3 is obliquely downbent to a black tornal space ; the tornus 
itself whitish; the termen from vein 2 to just above vein 6 
reddish brown, its inner edge fuscous, upbent to costa, leaving a 
white apical space. Hind wings: lines reddish brown partly 
edged with fuseous; an inbent basal line; a medial fascia 
expanding to close to inner margin, and then downbent, enclosing 
a white space from subcostal; a broad subterminal line, its outer 
edging downcurved near apex, forming a marginal fuscous line 
partly mottled with reddish brown; anal angle reddish brown 
where all the lines suffuse ; some metallic blue scaling near anal 
angle. 

@ similar below. Fore wings above green; a white spot in 
cell above vein 2; a black shade below subcostal expanding at end 
of cell, followed by a large postmedial white spot oblique from 
costa to just below vein 4; a large postmedial vertical spot between 
submedian and vein 3; outer margin broadly black ; a subterminal 
small white spot between veins 2 and 3, and smaller spots above 
and below vein 5, also a small inset spot near costa. Hind wings 
green; a postmedial broad white space from costa to below 
vein 5; a broad black subterminal line, and a similar marginal 
line; the termen brown-black with a fine green line near anal 
angle, the lines all suffusing in a brown shade at apex. 

Expanse, ¢ and 2 40 mm. 

fab. Carillo, Guapiles. 

Allied to #. vicaria Bates. 

In British Museum. 


PERISAMA BARNESI, sp. n. (Pl. LI. fig. 10.) 


@. Palpi black broadly streaked with white. Body black 
above, some white scaling around the eyes. Wings black. Fore 


348 MR. WILLIAM SCHAUS ON 


wings: some dark blue irrorations at base, along veins, and in 
cell; a postmedial curved greenish-blue fascia from costa to tornus, 
its edges uneven. Hind wings: some scattered blue irrorations ; 
the outer margin greenish blue; a terminal brown-black line ; 
cilia white. Fore wings below black; the cell to near end and a 
space below it crimson; a postmedial curved series of metallic 
blue spots ; termen white, broadest at apex, crossed by a fine 
marginal black line. Hind wings below grey-brown shading to 
whitish on termen; costa narrowly crimson; outer small round 
black spots on interspaces; a subterminal and a terminal dark 
brown line. 

Expanse 43 mm. 

Hab. Carillo. 

This species is named after J. Barnes, who captured the only 
specimen met with. 

This is the first Perisama recorded from Central America. 


‘VICTORINA STENELES, ab. STYGIANA, ab. noy. (Pl. LI. fig. 6.) 


d. Fore wings fuscous brown, darkest medially ; a large pale 
green spot on inner margin close to middle, almost reaching 
vein 2; a very small outset spot above vein 2; a small white spot 
medially on costa ; subterminal whitish spots, larger and better 
defined near tornus. Hind wings fuscous brown; base darker 
followed by a broad pale green fascia and a medial fuscous fascia, 
all stopping at submedian; the inner margin white; an outer 
row of oval green spots on interspaces preceded by fuscous 
shadings ; veins terminally darker edged. Underneath more like 
the typical form. 

Expanse 82 mm. 

Hab. Avangarez. 


Fam. ERYCINIDA. 


Eurycona Mystica, sp.n. (PI. LI. fig. 8.) 


@. Palpi white. Frons white divided by a brownish line. 
Vertex black. Collar and thorax brown. Abdomen above 
brighter brown, underneath white. Fore wings fuscous brown : 
the cell and basal half of wing below it bright brown; a paler 
rust-brown shade between veins 2 and 4, not reaching outer 
margin. Hind wings rust-brown, the costal margin broadly 
fuscous brown. Wings below pale ih ownish grey, ‘Tne postmedial 
line vermilion finely “edged with dark grey. Fore wings: the 
postmedial line slightly outbent, straight ; a subterminal narrow 
vermilion shade ; a terminal yellow-brown shade. Hind wings: 
the postmedial line vertical to below vein 2, then curved and 
wayily inbent to inner margin; a subterminal lunular dentate 
brownish shade from costa to below vein 3, followed by marginal 
points between veins 4 and 7, between 3 and 4 bya large black 


spot outwardly edged with white; below vein 2, irregular brown 


COSTA RICAN BUTTERFLIES, 349 


lines terminating in black points followed by white shading ; 
termen from above vein 2 to anal angle vermilion, 

Expanse 22 mm. 

Hab. Rio Grande, San Mateo. 


KuRyYGONA LEUCON, sp.n. (PI. LI. figs. 1, 3.) 


3. Palpi, lines on frons and tarsi pale buff. Head fuscous. 
Thorax and base of abdomen dark brown, abdomen otherwise 
bright brownish red above ; underneath greyish buff. Fore wings 
faintly angled at vein 4, fuscous ; a brown-red shade at base from 
cell to inner margin; a similar streak at base below subcostal. 
Hind wings brownish red; the costal and outer margins to 
vein 2 black, the inner margin fuscous brown. Wings below pale 
greyish brown, the medial line brownish red finely edged with 
black. Fore wings: medial line slightly angled at vein 4, 
terminating at submedian fold; an outer vertical dark brown 
line, not so well marked. Hind wings: the medial line incurved 
between veins 4 and 2, deeply downcurved below 2; subterminal 
greyish lunules from costa to vein 4, edged with dark brown; 
between 4 and 3 the lunule is larger, yellowish, followed by a 
black spot, outwardly edged with white ; the inner brown shading 
expands between 3 and 2, and is followed by a dark wavy line ; 
from vein 2 to inner margin are three yellow-brown streaks 
outwardly irrorated with black and terminating in white points 
before a fine marginal fuscous line; cilia reddish brown except at 
apex. 

@. Fore wings fuscous; a large orange space from middle of 
cell and well beyond it, not reaching costa, or below vein 2. Hind 
wings dark brown ; the outer margin outcurved from veins 4 to 2, 
shaded terminally from 4 to anal angle with brownish red. 
Wings below hght yellow-brown, the medial line as in male, but 
brighter red ; the outer line on fore wings also red; the lunules 
on hind wings heavier, geminate, the marginal shadings faintly 
marked, 

Expanse, ¢ 26 mm., 2 29 mm. 

Hab. Turrialba. 


KURYGONA LEUCOPHRYNA, sp.n. (PI. LIII. fig. 8.) 


3d. Frons white. Vertex and body fuscous; some brown 
shading on abdomen dorsally. Wings brown. Fore wings: costal 
margin, front of cell, apex, and outer margin more narrowly 
tinged with fuscous. Hind wings: costal and inner margins 
greyish ; a subcostal fuscous shade. Wings below silvery white. 

Expanse 27 mm. 

Hab. Cachi. 


Near L. cataleuca, but larger aud brighter brown. 


EuRYGONA MATUTA, sp. n. (PI. LI. fig. 7.) 


g. Frons white. Vertex and body above fuscous brown, 
underneath white. Fore wings fuscous, the base in and below cell 


350 MR. WILLIAM SCHAUS ON 


shaded with brown; an orange wedge-shaped spot close to cell, 
between veins 2 and 4. Hind wings brown, the outer margin 
shaded with fuscous ; the costa whitish ; the inner margin grey- 
brown; cilia on inner margin and apex white. Wings below 
white; a very fine pale olive-brown postmedial line, almost 
vertical, bluntly curved below vein 2 on hind wings. Hind wings: 
a small black marginal spot between veins 3 and 4; three black 
points from vein 2 to inner margin, and a few subterminal black 
irrorations. 

Expanse 21 mm. 

Hab. Juan Vinas. 


MESOSEMIA ALBIPUNCTA, sp. n. (PI. LIII. fig. 12.) 


¢. Thorax and base of abdomen metallic blue, abdomen other- 
wise and head fuscous; underneath grey. Wings vivid blue. 
Fore wings: a white point near end of cell ; outer margin broadly 
black, widest on costa, its inner edge outbent and curved near 
tornus. Hind wings: termen black, the veins close to it streaked 
with black; a subterminal black spot at apex, suffusing with 
black costal margin ; in another specimen the spot continues as a 
line to inner margin. Wings below grey-brown, the lines darker 
brown. Fore wings: the two lines before ocellus parallel, outbent 
from subcostal ; ocellus large, a white central point, and two outer 
points, circled with yellow; two lines beyond ocellus, close 
together, diverging slightly towards inner margin ; postmedial 
space grey irrorated with metallic blue from near costa to vein 3, 
limited by a fine greyish yellow line, outbent from costa, at sub- 
median approximated by asimilar subterminal line. Hind wings: 
the antemedial and postmedial lines straight, geminate; the 
medial shade partly linear interrupted by a smaller ocellus, black, 
circled with yellow and containing two white and blue points; a 
broad outer shade; a subterminal line; termen dark shaded. 

Expanse 27-30 mm. 

Hab. Guapiles, Esperanza. 


HERMATHENA OWENI, sp.n. (PI. LIII. fig. 9.) 


g. Head and thorax black clothed with greyish-white hairs. 
Abdomen grey, with transverse black segmental shades. Wings 
white, the base mottled with black with traces of a subbasal 
whitish line; spots black, two beyond cell; a medial spot below 
vein 2; subterminal quadrate spots above and below vein 6, also 
above and below vein 3; apex black; spots at end of veins ; 
terminal interspaces between veins 2 and 3, and 4 and 6 mottled 
with black. Hind wings: terminal spots at veins; subterminal 
quadrate spots above and below vein 7. Underneath similar ; 
the basal mottling showing through from above; the terminal 
markings on interspaces entirely black. 

Expanse 42 mm. 

Hab. Costa Rica. 

Named after Prof. Owen of Madison, Wis., who has kindly 
given me this unique specimen. 


COSTA RICAN BUTTERFLIES. 301 


CoRRACHTA, gen. nov. 


2. Fore wings: outer margin straight at apex, then rounded ; 
vein 3 close to lower angle of cell; 4 from lower angle; disco- 
cellular incurved, slightly oblique; 5 from just above middle ; 
6-10 stalked from upper angle, 10 before 6; 11 from close to end 
of cell anastomosing with costal vein. Hind wings as long as fore 
wings; cell elliptical; vein 3 before end; 4 from end; 5 from 
middle of discocellular ; 6 and 7 stalked. Abdomen stout. 

Type of genus, C. lewcoplaga Schaus. 


CoRRACHIA LEUCOPLAGA, sp.n. (PI. LITT. fig. 2.) 


@. Body and wings brownish slate-colour. Fore wings: a 
broad medial white fascia, slightly outcurved, from subcostal to 
submedian vein, a whitish streak on costa above it between veins 
10 and 11. Hind wings: marginal fuscous lunules, faintly edged 
with greyish. Wings below grey. Fore wings: the white fascia 
followed by a broad outcurved fuscous shade to some whitish sub- 
terminal points. Hind wings: termen irrorated with whitish ; 
small subterminal fuscous spots preceded by whitish shadings. 

Expanse 35 mm. 

Hab, Palmital. 

This peculiar species will probably be found to have a very 
dissimilar male. 


CHARIS SUBTESSELLATA, sp.n. (PI. LITT. fig. 11.) 


Body above fuscous. Fore wings fuscous, the base tinged with 
olive-brown ; a white spot in cell; an outer white spot between 
veins 2 and 3, and a smaller one between 5 and 6; a small post- 
medial spot above vein 6. Hind wings dark olive-brown; cilia 
white between veins 5 and 6; a yellow-grey marginal streak at 
anal angle. Fore wings below reddish brown, markings bluish 
white partly edged with black ; a streak at base of costa; a basal 
streak below subcostal ; an antemedial fascia in cell, and one 
below it outset ; a medial fascia in cell, with a white streak above 
subcostal, and a buff streak above costal vein; an outset medial 
fascia below cell, all of them narrower towards median, the two 
below cell suffusing on submedian; a fine black line on disco- 
cellular followed by small spots from costa to vein 4; two costal 
spots beyond, superposed, and small black spots on interspaces 
below; an interrupted subterminal white line preceded by a white 
point above vein 6,a large one below it, black spots below 5 
and 4, and a still larger white spot between 3 and 2; a marginal 
silvery blue line; a fine terminal black line; cilia fuscous. 
Hind wings below reddish brown except on outer third ; a white 
triangular spot on costa near base, and a quadrate medial spot ; 
an antemedial white spot in cell, and transverse line across end 
of cell; two white spots below cell suffusing with a white streak 
below vein 1; a black line on discocellular, followed by a white 
fascia from veins 7 to 4, and from vein 2 to inner margin; black 
points in some of the reddish-brown spaces, which are also partly 


352 MR. WILLIAM SCHAUS ON 


black-edged ; a postmedial reddish brown shade from costa to 
vein 4, followed by a macular black line, and then by a broad 
grey-brown shade crossed by a fine black line; postmedial space 
below vein 4 greyish brown to vein 1, then reddish brown on 
inner margin, followed by greyish brown shading from vein 4 to 
vein 1, and buff shading on inner margin, these shadings out- 
wardly edged with black and followed by a silvery opalescent 
shade; outer margin dull dark brown outwardly edged with 
opalescent scaling, and containing small fuscous spots below 
vein 4 and vein 3; the tails and a terminal buff line from vein 4 
to anal angle edged on either side with black; from vein 4 to 
apex the terminal line is reddish brown, preceded at apex by a 
short white line. 

Expanse 24 mim. 

Hab. Cavrillo. 

Near C. chrysus Cr. 


CHARIS TURRIALBENSIS, sp. n. (Pl. LIII. fig. 13.) 


3g. Body above fuscous. Wings dark brown, with antemedial 
and postmedial fuscous shadings on interspaces. Fore wings: 
cilia at tornus orange-red. Hind wings: a black line on disco- 
cellular ; a subterminal greyish buff shade from vein 6 to anal 
angle, crossed by a black macular line, preceded below vein 3 by 
a large brown spot, some reddish brown shading on inner margin 
at angle. Fore wings below orange-red ; a pale buff triangular 
spot on costa near middle with its apex on median at vein 3, 
edged with silvery blue; similar metallic scaling on costa near 
base, and two inbent lines across cell; below cell and vein 2, 
medial, postmedial, and subterminal pale greyish shades, the 
intermediate red portions edged and suffused with black ; a black 
line on discocellular, followed by metallic spots; below vein 5a 
macular black line; outer pale greyish spots, followed above 5 
and 6 by silvery streaks, preceded by silvery spots and some black 
edging, below vein 5 the greyish spots are inwardly edged with 
black, outwardly with metallic scaling and black edging; a 
marginal metallic line interrupted by veins. Hind wings below 
orange-red, divided by metallic antemedial, medial, and _post- 
medial irregular lines, with some short black lines and spots in 
reddish portions; outer margin broadly pale greyish irrorated 
with brown; a marginal metallic line finely edged outwardly 
with black, and preceded by some black below veins 4 and 3; a 
terminal black line, also edging tails. 

Expanse 21 mm. 

Hab. Tarrialba, 6000 feet. 


Fam. Lyc £#NIDS&. 


THECLA HypocritA, sp.n. (PI. LIT. fig. 6.) 


$. Body dark brown mottled with bluish grey hairs and 
irrorations. Fore wings fuscous brown, except a pale blue space 


COSTA RICAN BUTTERFLIES, 353 
occupying basal half of cell, space below to inner margin, and 
just above vein 2 to near subterminal space; a pale brown patch 
of coarse scales at end and beyond cell. Hind wings pale blue; 
outer margin fuscous brown; inner margin whitish, becoming 
fuscous towards anal angle; cilia brown tipped with grey. 
Fore wings below fuscous; costal margin broadly whitish, post- 
medially to vein 4; some brown mottling at base of costa; a 
brown medial fascia to median and end of cell enclosing a white 
line on discocellular ; an outer and a subterminal oblique broad 
brown line to vein 5; outer margin narrowly white crossed by a 
fine marginal brown line. Hind wings below yellowish white, 
crossed by broad brown lines; a fuscous basal space, outangled 
on median ; the inner margin broadly irrorated with fuscous and 
pale brown; medial and postmedial lines vertical, angled at 
vein 2 and inbent to inner margin; a subterminal brown shade 
suffusing with postmedial from veins 5 to 2, then continuing to 
anal angle; a broken marginal brown shade; anal angle slightly 
lobed, no tail. 

Expanse 19 mm. 

Hab. Tuis, Juan Vinas. 

A specimen from Cordoba, Mexico, has the lines below partly 
greyish, but undoubtedly belongs to this species. 

Near 7’. dolylas Cr. 

In British Museum. 


‘THECLA SUBFLORENS, sp. n. (PI. LIT. fig. 9.) 


gd. Antenne black ringed with white. Body above fuscous, 
underneath white; some green scales on frons. Wings dark 
purple-slate colour. Fore wings: the costa, apex broadly, 
and outer margin fuscous brown. Hind wings: the margins 
narrowly fuscous brown. Wings below rich green. Fore wings 
paler tinged, except on costa and apex ; the inner margin dark 
grey. Hind wings: a faint paler line on discocellular; a similar 
irregular outer line, black, outwardly edged with white from 
vein 2 to inner margin; a few red scales at anal angle, sur- 
mounted with black and white; a terminal black line at anal 
angle. 

Expanse 20 mm. 

Hab. Poas. 


THECLA LEOS, sp.n. (Pl. LII. fig. 7.) 


3. Palpi black and white. Head, collar, and patagia black ; 
white lines around eyes. Thorax and abdomen above metallic 
blue, underneath whitish. Wings brilliant dark blue. Fore 
wings: apex and outer margin black, narrowest at tornus; a 
large black patch filling the end of cell from before middle 
followed beyond cell by a small slate-coloured spot. Hind wings : 
the termen narrowly black; the inner margin grey. Wings 
below clear bright brown, Fore wings: outer line fuscous brown, 
outwardly edged with white, faintly outbent, slightly Iunular ; 


354. MR. WILLIAM SCHAUS ON 


inner margin greyish; a faint dark marginal line at tornus. 
Hind wings: outer line fine, fuscous, outwardly edged with 
white, more broadly on costa, vertical from veins 8 to 6, appearing 
incurved on account of the broader white edging, slightly outset 
below 6 and below 5, inset below 4, outset below 3, inset and 
horizontal below 2, downbent on submedian, angled and upbent 
to inner margin; a fine subterminal black line from vein 3 to 
inner margin, and darker spots at apex; a marginal red spot 
with black point and terminal white line between 2 and 3; a 
black spot with white irrorations below vein 2; a black spot at 
anal angle surmounted by a reddish shade. 

Expanse 24 mm. 

Hab. Guapiles, La Florida. 

I do not know any species with which I can compare this. 

In British Museum. 


THECLA MELMA, sp.n. (PI. LIT. fig. 4.) 


6. Palpi black, the base of joints white. Body black; some 
white on vertex, and white lines close to eyes. Wings deep blue. 
Fore wings: costa, apex, and outer margin black, the apical 
space reaching to near cell; a black line on discocellular. Hind 
wings: margins very narrowly black, slightly broader at apex. 
Wings below bright greyish bistre. Fore wings: an outer 
vertical fine fuscous line, outwardly edged with white from near 
costa to vein 2; a terminal fine brown line; cilia fuscous. 
Hind wings: a white line on discocellular; outer line black out- 
wardly edged with white, outset on costa, almost vertical from 
veins 7 to 4, then inset and downbent to vein 2, forming the 
usual W to inner margin; subterminal lunules indicated by 
ereyish shading from vein 7 to vein 2, followed between 2 and 3 
by a large red space, outwardly broken by a triangular black 
space ; some reddish scales above vein 3; below vein 2, marginal 
white irrorations ; a terminal white line from vein 5 to sub- 
median ; anal lobe black inwardly edged with white. 

Expanse 27 mm. 

Hab. Guapiles; also Bugaba, Panama. 


THECLA GUAPILA, sp. n. (PI. LII. fig. 1.) 


3d. Thorax and base of abdomen metallic blue, abdomen 
otherwise fuscous above, yellowish white below. Head brown, 
with white lines at eyes. Wingssilky blue. Fore wings: costal 
margin, apex to near glandular spot, and outer margin, narrowly 
at tornus, black; a small black spot at end of cell followed by 
an oval brownish glandular spot. Hind wings: apex narrowly, 
and cilia black. Wings below dark brownish slate-colour. Fore 
wings: inner margin ‘oreyish ; a fine outer fuscous line, almost 
vertical from below costa to vein 2, outwardly faintly pale- -edged ; 
faint traces of a dark subterminal line. Hind wings: outer “line 
interrupted, dark brown, outwardly edged with white, inbent on 


COSTA: RIGAN BUTTERFLIES. 355 


costa, vertical from veins 7 to 2, but inset between 3 and 4, angled 
below vein 2, and inbent to inner margin; subterminal darker 
spots, edged with whitish irrorations, except at apex, inset be- 
tween veins 2 and 3, followed by a red marginal spot, containing 
a small black spot ‘outwardly ; ; below vein 2 only a marginal 
brown spot irrorated with white; below submedian a line in- 
wardly edged with white and parallel with outer line; a terminal 
white line between veins 2 and 3; anal lobe black. 

Expanse 28 mm. 

Hab. Guapiles. 

In British Museum. 


THECLA AMPHRADE, sp.n. (PI. LIT. fig. 10.) 


@. Wings fuscous brown, the hind wings, and inner area of 
fore wings below cell glossed with slate-blue; cilia light brown. 
Hind wings: the terminal black line inwardly edged with white 
near anal angle; a small red-brown spot at anal angle; tails 
black tipped with white. Wings below light brown, the post- 
medial line dark brown outwardly edged with black and then 
white, very indistinctly on fore wing. Fore wings: the line 
vertical from costa to vein 2; Beha indistinct paler subterminal 
shade, crossed by a faint dark line; a terminal fine dark line. 
Hind wings: the line vertical to vein 4, constricted and slightly 
oblique to vein 2, upcurved and outangled on submedian; a sub- 
terminal paler shade as on fore wing, the faint line crossing it 
more distinct followed on inner margin, between veins 2 and 3 and 
just above 3, by reddish shades; a black marginal spot between 
veins 2 and 3 and at anal angle; a black spot irrorated with 
white below vein 2; the terminal dark line inwardly edged with 
white. 

Expanse 23-25 mm. 

Hab. Guapiles. 

Near 7’. politus H. Druce. 

In British Museum. 


Subfam. PrerInz. 


ARCHONIAS INTERMEDIA, sp. n. (PI. LIT. fig. 8.) 


3. Palpi yellow, somewhat fuscous at base. Head and thorax 
black ; three whitish points on vertex. Abdomen brown ; sub- 
lateral yellow line. Fore wings black, markings yellow-white ; 
an oblique spot at end of cell, medially constricted ; two diffuse 
spots beyond cell; elongated postmedial spots above and below 
vein 3, and geminate streaks below 2, all followed by marginal 
spots, ‘small at tornus, elongated towards costa ; a yellow and 
reddish streak along inner margin, not reaching ‘base or tornus. 
Hind wings bright brownish red, ‘the costal and outer margins 
broadly black, the inner margin finely so; the veins black; cell 
edged with black above and below; marginal small white spots. 
Underneath duller, the hind wings browner, with a brown up- 


Proc. Zoo. Soc.—1913, No. XXY. 25 


356 MR. WILLIAM SCHAUS ON 


curved streak on basal half of costa; the subcostal and median 
broadly shaded with black ; the marginal white spots larger. 

2. The markings on fore wings reduced, except the marginal 
spots; the hind wings reddish brawn, the costa with only a black 
shade medially ; the marginal spots lve ger, yellow, irrorated with 
reddish. Underneath similar to the male, 

Expanse, ¢ 60 mm., 2 65 mm. 

Hab. Carillo. 

The fore wings are like A. dismorphites Btl., the hind wings like 
A. nigrescens G. & 8. All these forms will no doubt be found to 
gradually intergrade. The males are difficult to capture, flying 
very high in a circle, and rarely settling. 

In British Museum. 


PIERIS LIMONA, sp. n. (PI. LIT. fig. 5.) 


3g. Antenne black tipped with white. Head and thorax 
fuscous, the latter clothed behind with white hairs. Palpi white, 
the third joint streaked with black. Abdomen fuscous grey 
above, whitish below. Wings white, the veins broadly edged 
with chalky white. Fore wings: a fuscous grey streak on basal 
third of costa ; a small black spot on discocellular ; apex narrowly 
black; marginal fuscous shades, not reaching tornus. Hind 
wings: cilia black. Fore wings below: a large black spot on 
discocellular ; apex greyish. Hind wings below yellowish white, 
the veins dark. 

©. Fore wings above with a large black spot on discocellular, 
and a fuscous streak in cell at subcostal reaching it; the apex and 
termen more broadly black, its inner edge somewhat lunular ; 
underneath the black margin replaced by a pale lilacine orey 
shade. Hind wings below “Vilineine white, the veins brown ; a 
postmedial br oneal shade just beyond cell across veins 3 and 4. 

Expanse, ¢ 75 mm., 9 78 mm. 

Hab, Limon. 

Near P. diana Feld. 

In British Museum. 


ACMEPTERON POASINA, sp.n. (PI. LIT. fig. 3.) 


g. Antenne black. Palpigrey. Head and collar dark brown. 
Thorax and abdomen fuscous irrorated with white. Fore wings 
black ; median, fold, submedian, and vein 2 all basally streaked 
with white ; spots llacine colhthe ¢ a medial streak on costa, spot 
beyond cell, an outer spot above vein 3, and one on costa, also. a 
small subterminal spot above 6; sometimes traces of spots on 
either side of upper discocellular ; cilia greyish brown. Hind 
wings iridescent silky brown tah a large ‘medial dull brown 
spot; from cell behind to inner margin and termen at vein 5 
black, crossed by a grey fascia; cilia yellow. Wings below as in 
A. cinerascens Salv., but the markings all more intense, the anal 
angle and inner margin bright yellow crossed by purple-brown 
strie, 


COSTA RICAN BUTTERFLIES. 357 


2. Wings black, the markings white. Fore wings: an oblique 
fascia from middle of costa consisting of coalescing spots; three 
oblique spots from costa before apex; the outer half of inner 
margin with a white streak. Hind wings: a large white space 
occupying the greater portion of the wing, outangled on vein 5, 
its hind edge straight; the base of cell and a streak below it 
black. Fore wings below: a white space on basal half from 
within cell to inner margin; costa basally lilacine; the medial 
fascia as above but with a yellow streak on costa; apex lilacine 
white, shaded with yellow on costa; termen finely yellow, pre- 
ceded by light brown and greyish white shading. Hind wings 
below white: the base, outer half from inner margin to vein 4, 
and terminally between 4 and 5 lilacine white irrorated with 
brown; the shade in cell extending somewhat, and upbent to 
costa. ; subterminal brown irrorations from costa to vein 6; inner 
margin narrowly bright yellow, termen finely similar pr eceded by 
& narrow inter rupted brown shade. 

Expanse, ¢ 60 mm., 2 67 mm, 

The male can be distinguished from 4. cinerascens Salv. by the 
narrow grey fascia on hind wings above. The female differs in 
the broader black margin, the lilacine shading, and brown irrora- 
tions on hind wings. aN cinerascens female has the hind wings 
below bluish white with faint traces of darker shadings. If the 
two forms ogcurred in different localities I should not separate 
them specifically, but they fly together and no intergrades were 
found. 

In British Museum. 


ENANTIA LUA COSTARICENSIS, subsp. n. (PI. LIT. fig. 2.) 


3. Like #. lua Hew., but the inner edge of black margin on 
hind wings near apex is straight, not dentate ; underneath the 
spots are ‘lar ger and more confluent. The wings are also broader. 

Expanse 62 mm. 


Hab. Cachi. 
Fam. HESPERIIDA. 


TELEMIADES MNEMON, sp. n. (PI. LIV. fig. 1.) 


3. Palpi white below, black above with yellow scales at tip of 
second joint. Head and collar black mottled with yellow. Thorax 
and abdomen brown. Fore wings dark brown, the base to near 
middle thickly irrorated with yellow-brown hairy scales ; similar 
sealing subterminally, expanding at tornus, also a few similar 
scales” beyond cell; medial space below cell tinged with blue- 
black ; spots semihyaline yellowish ; a large spot in end of cell, 
its outer edee incurved ; a narrow vertical postmedial spot be- 
tween veins 2 and 3, and a small spot above 3, slightly outset ; 
three small outer spots below costa, and two subterminal between 
veins 4 and 6. Hind wings with only the costal margin, a post- 
medial macular shade, and terminal spots fuscous brown, other- 
wise overlaid with yellow-brown hairy scales. Fore wings below 

25* 


358 MR. WILLIAM SCHAUS ON 


dull fuscous brown; the inner margin tinged with grey ; some 
yellow scaling at base, below costa, and in cell. Hind wings 
below dull brownish yellow; the outer margin broadly dark 
brown ; some brownish-yellow scaling subterminally from vein 3 
to inner margin. 

Expanse 28 mm. 

Hab. Zent district. 


NASCUS PHINTIAS, sp. n. 


3. Palpi white tipped with black above. Head and collar 
fuscous mottled with olive-brown ; white points laterally on head. 
Thorax and abdomen shaded with orange-brown. Fore wings 
fuscous brown, the base and inner margin shaded with orange- 
brown ; spots semihyaline whitish; a spot at end of cell, out- 
wardly produced in front; a large postmedial spot between 
veins 3 and 4, and a smaller one below 2; an outset large spot 
between veins 3 and 4, and a point between 4 and 5; a curved 
outer series of spots between veins 6 and 10. Hind wings 
orange-brown, the costal margin broadly fuscous, the termen 
narrowly so; a fuscous line on discocellular; a broader post- 
medial line not reaching inner space. Wings below brown- 
black. Fore wings: the base shaded with yellow-white; short 
whitish streaks above cell-spot. Hind wings: the basal half and 
inner margin for two-thirds pale yellow ; a thick black line on 
discocellular. 

Expanse 55 mm. 

Hab. Guapiles. 

The female of this species is figured in the Appendix to the 
‘Biologia’ as a variety of WV. phocus Cr., but the discovery of the 
male proves it to be a distinct species. 


CYCLOS#MIA SUBCHRULEA, sp.n. (Pl. LIV. fig. 14.) 


Body fuscous brown. Wings dark greyish brown, the markings 
fuscous brown ; a postmedial and marginal broad line, somewhat 
macular, also a similar medial line on hind wings. Fore wings: 
the costa and termen broadly shot with green, the veins more 
vividly so towards apex, but only noticeable in a certain light; a 
large black spot in cell at end, faintly edged with paler brown, 
and containing two bluish-white points. Fore wings below dark 
brown shot with green; the inner margin shaded with grey; a 
fuscous shade in end of cell; a postmedial fuscous shade. Hind 
wings below bright blue; the costa, apex narrowly, and cilia 
dark brown. 

Expanse 31 mm. 

Hab. Sixola, Guapiles. 

In British Museum. 


ECHELATUS LUCINA, sp.n. (PI. LIV. fig. 8.) 


Body fuscous brown, the vertex and collar glossed with green. 
Wings fuscous brown. Fore wings glossed with purple ; an ante- 


COSTA RICAN BUTTERFLIES. 359 


medial and a postmedial fuscous shade, the latter slightly out- 
curved below costa, vertical below vein 4. Hind wings with 
fainter purple gloss, except a streak below cell from base to 
termen ; a fuscous shade at end of cell, and a similar postmedial 
shade. Wings below paler brown, the lines as above, dark brown ; 
the antemedial on fore wings replaced by a dark brown shade 
from near base to middle, except on inner margin, which is whitish 
grey at base. 
Expanse 37 mm. 


Hab. Guapiles. 


AYE CANACE, sp. n. (PI. LIV. fig. 17.) 


3. Body and wings fuscous brown. Fore wings: a semi- 
hyaline point medially between vein 11 and costal vein ; a similar 
outer point between veins 8 and 9, and one below 8 slightly inset. 
Hind wings: outer margin broadly blue from vein 6 to inner 
margin ; cilia fuscous brown. Fore wings below duller. Hind 
wings below blue ; inner margin whitish blue ; costal margin and 
apex to vein 6 dull fuscous brown; a similar downbent streak 
from vein 2 to anal angle. 

Expanse 23 mm. 


Hab. Carillo. 


Boa SODALIS, sp.n. (Pl. LIV. fig. 12.) 


2. Palpi brown mottled with buff and white. Body fuscous 
brown. Wings brown slightly irrorated with lighter brown and 
buff. Fore wings: a fuscous shade on discocellular followed by a 
faint paler brown shade; three outer white hyaline spots below 
costa, straight and outbent; three similar postmedial vertical fine 
streaks between submedian fold and vein 4; a faint subterminal 
paler brown shade cut by darker veins. Hind wings: a faint 
subterminal paler brown shade narrower than on fore wings. 
Wings below duller brown ; the subterminal shade on fore wings 
narrow; hind wings irrorated with buff-brown; a small pale 
brown spot in cell, a streak on discocellular, some small post- 
medial spots near inner margin, and the subterminal spots all 
very indistinct. 

Expanse 29 mm. 

Hab. El Alto. 

Nearest B. cylindus G. & $.; smaller, the spots also smaller ; 
the outer spots in a straight line and oblique. 


Boia MAcHUCA, sp. n. (PI. LIV. fig. 4.) 


3. Body above fuscous brown; some greyish white hairs in 
front of antenna; some yellowish white hairs on abdomen. 
Throat white; fore femora grey. Abdomen below greyish - 
brown. Fore wings fuscous brown; outer margin faintly paler, 
forming a subterminal line with darker marginal spots on inter- 
spaces ; three white points outwardly between veins 6 and 9. 


360 MR. WILLIAM SCHAUS ON 


Hind wings dark brown ; large postmedial and marginal fuscous 
brown shades, all indistinct and confused. Fore wings below 
dark grevish brown ; a broad darker postmedial shade; the costa 
tinged with iridescent green; the white points as above. Hind 
wings below dark brown; some white irrorations on basal half, 
and more thickly aiong inner margin; traces of a darker post- 
iwnedial shade, and small marginal spots. 

Expanse 26 mm. 

Hab. San Mateo. 

In British Museum. 


BUuTLERIA LETHHA, sp.n. (PI. LIV. fig. 15.) 


3. Palpi black mottled with yellowish hairs. Body above 
black ; some orange hairs on abdomen; abdomen below yellow. 
Fore wings fuscous brown ; a large orange-yellow spot just beyond 
middle between veins 2 and 4, extending into end of cell, and 
slightly below vein 2; a small oblique similar spot outwardly 
below costa; cilia dark brown. Hind wings black; cilia and a 
large spot occupying middle of wing orange-yellow. Fore wings 
below deep yellow: cell to near end black, extending at base 
below cell; a broad black fascia beyond cell, not reaching costa, 
outbent above vein 4, and downturned to inner margin at tornus. 
Hind wings below yellow; some black scaling at base of inner 
margin. 

Expanse 20 mm. 

Hab. Poas. 

Near B. lalage Godm. 

In British Museum. 


BUTLERIA Lysis, sp.n. (PI. LIV. fig. 2.) 
3. Body above black: abdomen below yellow. Wings dark 


brown, spots yellow. Fore wings: a spot between veins 2 and 3, 
just beyond middle, surmounted by a triangular spot in cell; a 
point between veins 3 and 4 postmedially, and a short oblique 
line outwardly below costa. Hind wings: alarge spot in middle 
of wing from within cell. Fore wings below fuscous; costal 
margin yellow-brown, the extreme edge black; termen finely 
yellow expanding at apex; the yellow discal spot extending 
below vein 2 and expanding along submedian. Hind wings 
below whitish yellow, the veins brown except the discocellular 
and base of vein 5, the brown on veins 2 and 4 suffusing some- 
what towards cell; cilia brown. 

Expanse 23 mm. 

Hab. Vurrialba. 

Near b. faula Godm. 


LERODEA ? RUPILIUS, sp.n. (Pl. LIV. fig. 10.) 


2. Palpi pale brown tipped with white. Body above and 
wings dark brown; body below whitish grey. Wings: spots 


COSTA RICAN BUTTERFLIES. 361 


semihyaline white; outer half of cilia greyish white with small 
dark brown spots at veins. Fore wings: a large spot at end of 
cell, and one postmedially between veins 2 and 3. a smaller out- 
set spot between 3 and 4; three outer small. spots obliquely 
between veins 6 and 9; a more opaque spot above submedian 
beyond middle. Hind wings: a postmedial spot below vein 6, 
crossed by vein 5. Fore wings below similar, the costa irrorated 
with yellowish and white, the apex and termen to vein 2 with 
white; the spot above submedian replaced by a large white 
shade. Hind wings below thickly irrorated with greyish white ; 
small whitish spots antemedially and postmedially below costal 
vein; a siailae spot at end of cell; a small postmedial spot 
between veins 2 and 3, and a linear spot below vein 2; the spot 
below vein 6 as above ; the postmedial spots connected by a fine 
brown shade, and a similar faint shade below end of cell. 

Expanse 36 mim. 

Hab. Guapiles. 

The only species I know approaching this is one in the Godman 
Collection labelled ‘“ phyllas Mab.” Without a male the position 
is uncertain. 


STYRIODES, gen. nov. 


3. Antenne two-thirds as long as costa. Fore wings: costal 
margin slightly sinuous; apex acute; termen rounded below 
apex ; vein 2 from middle of cell; 3 near lower angle; 4 and 5 
from lower angle, the latter upcurved at base ; 6 to 9 evenly 
apart ; 10 and 11 wider spaced ; a downcurved brand below 
median from vein 3 to below vein 2; a small medial brand above 
submedian. Hind wings: anal angle slightly lobed; vein 2 
beyond middle of cell; 3 and 4 from lower angle apart; 6 and 7 
from upper angle. 

Type of genus, Styriodes lyco Schaus. 


SrYRIODES LYCO, sp.n. (PI. LIV. fig. 19.) 


3. Body and wings above fuscous brown. Fore wings: brands 
black; cilia tipped with yellow-ochre, wider at tornus. Hind 
wings: cilia tipped with yellow-ochre, except at anal angle. 
Wings below dark reddish brown, the inner margin of fore wings 
dull dark brown; a fuscous ‘borin shade in ical behind and 
below it. 


Expanse 28 mm. 
Hab. Poas, 


RHINTHON LUCTATIUS, sp.n. (Pl. LIV. fig. 9.) 


3. Body above black, the head shaded with dark metallic 
green ; abdomen below with two yellowish white lines. Wings 
fuscous brown. Fore wings: a very small greyish white post- 
medial spot between veins 2 and 3, and one between 3 and 4 
outset. Wings below not quite so dark. Fore wings: the two 
spots whiter, better defined; a postmedial greyish brown shade 


362 MR. WILLIAM SCHAUS ON 


below vein 2. Hind wings: postmedial buff points between 
veins 2 and 4. 

Expanse 44 mm. 

Hab. Guapiles, also Santa Rosa, Mexico. 

In British Museum. 


CoBALUS NIGRANS, sp.n. (PI. LIV. fig. 5.) 


3. Palpi and thorax fuscous grey mottled with pale yellow ; 
abdomen below with two fine whitish lines. Wings black-brown. 
Wings below duller. Fore wings: a faint greyish white post- 
medial line between veins 2 and 3, and a small outset spot 
above 3; tornus shaded with fuscous grey. Hind wings: an 
orange point in cell at end; similar outer points between veins 3 
and 4, and just below 6. 


Expanse 30 mm. 
Hab. Guapiles. 


CoBALUS LAUREOLUS, sp. n. (PI. LIV. fig. 3.) 


3. Body fuscous clothed with dark brown hairs; the palpi 
mottled with some whitish yellow hairs; abdomen below greyish 
brown. Fore wings: veins medially tinged with fuscous. Hind 
wings: basal half rather darker shaded. Wings below paler 
brown. Fore wings: basal third except costa shaded with 
fuscous; a triangular shade on costa near apex preceded by faint 
greyish points between veins 6 and 9; tornus faintly paler 
shaded. Hind wings: base, a postmedial fascia, and large mar- 
ginal spots from apex to vein 2 darker brown ; a terminal darker 
brown line. 

Expanse 35 mm. 

Hab. Guapiles. 


CoBALUS LATERANUS, sp. n. (PI. LIII. fig. 3.) 


3S. Palpi fuscous inwardly shaded with white. Body and 
wings fuscous brown. Fore wings: spots semihyaline, white; a 
large irregular spot at end of cell; a large postmedial spot be- 
tween veins 2 and 3, and a small outset spot above 3; three small 
outer spots below costa, outbent, the largest between veins 6 and 
7; a small postmedial white spot above submedian. Fore wings 
below fuscous, the costa, apex, and termen shaded with olive- 
brown; the spot above submedian larger. Hind wings below 
dark olive-brown ; a white spot in end of cell; a white point 
subterminally between veins 3 and 4. 

Expanse 32 mm. 

Hab. San Mateo. 

The fore wings above are like C. gabina Godm. 


CoBALUS PINDAR, sp. n. (PI. LIV. fig. 18.) 


3. Palpi olive-yellow irrorated with brown. Body and wings 
above dark brown; abdomen below yellowish bistre. Fore 


COSTA RICAN BUTTERFLIES. 363 


wings: spots semihyaline whitish brown ; a large spot between 
veins 2 and 3, its outer edge incurved ; a small spot between 3 
and 4; three small spots between veins 6 to 9 in a straight out- 
bent line from costa ; cilia tipped with bistre grey. Hind wings: 
a yellow postmedial hyaline point between veins 3 and 4. Fore 
wings below fuscous brown, the costal margin broadly, and cell 
shaded with yellow-brown, the spots as above; a whitish post- 
medial shade above submedian. Hind wings below brown; a 
small faint fuscous shade at discocellular, and similar spots post- 
medially from veins 2 to 7, the one between 3 and 4 showing the 
semihyaline point. 

Expanse 29 mm. 

Hab. Port Limon, Juan Vinas. 

In British Museum. 


CoBALOPSIS LATONIA, sp.n. (PI. LIV. fig. 16.) 


3. Body fuscous brown; palpi above whitish grey; abdomen 
below pale buff. Wings dark brown. Fore wings: a fine 
short whitish inbent streak below discocellular ; a small white 
postmedial spot between veins 3 and 4, also above and below 
vein 7. Hind wings: the veins fuscous. Fore wings below 
fuscous in cell and ‘postmedially below vein 3, the inner margin 
irrorated with greyish ; costa, apical space, and termen yellowish 
brown. Hind wings below yellowish brown; a fuscous shade near 
inner margin expanding at anal angle; five bluish white post- 
medial spots from veins 2 to 7. 

Expanse 33 mm. 

Hab. Juan Vinas, Cachi. 

In British Museum. 


CENUS NAUSIPHANES, sp.n. (PI. LIV. fig. 6.) 


3. Palpi mottled whitish grey and dark brown. Body fuscous 
brown ; abdomen below whitish grey; legs yellow-brown. Wings 
dark brown. Fore wings: the brand fine, black, inbent from 
vein 3, and downcurved to submedian ; some yellowish scaling on 
costa ; cilia tipped with grey. Hind wings: cilia yellow-grey. 
Fore wings below fuscous at base, in, and below cell; costa yellowish 
brown ; terminal half dull brown. Hind wings dull brown; a 
broad medial whitish shade, widest on costa and not reaching 
inner margin which is simply irrorated with whitish grey. 

Expanse 25 mm. 

Hab. Poas. 


In British Museum. 


MEGISTIAS XANTHO, sp. n. (PI. LIV. fig. 13.) 


3. Body fuscous mottled with brown haus. Wings fuscous’ 
brown. Fore wings: a yellow-brown streak on costa from base 
to just beyond middle ; a similar streak above submedian medially 
extending towards base; a few similar scales outwardly beyond 


364 MR. WILLIAM SCHAUS ON 


cell, outbent along vein 6, vertical to below 5, inbent, expanding, 
forming a spot between 4 and 3, and a larger spot between 3 
and 2; cilia light brown spotted with black. Hind wings: some 
yellow-brown hairs at base and on inner margin ; a broad similar 
postmedial shade not reaching margins ; the spots on cilia narrow, 
barely traceable. Fore wings below fuscous ; veins on costa, and 
terminally, also a streak in cell, yellowish ; a postmedial yellowish 
spot between veins 3 and 4, a vertical outer streak below costa, 
and streak subterminally from veins 6 to 4; a terminal yellowish 
line; base of cilia and spots black. Hind wings below greyish 
tinged with lilacine, the veins yellowish; an outcurved dark 
brown medial fascia, and large terminal spots, not reaching inner 
margin. 

Expanse 20 mm. 

Hab. Juan Vinas. 

In British Museum. 


Voraves cotiso, sp.n. (Pl. LIII. fig. 6.) 


3. Body above and wings dark brown ; throat whitish ; thorax 
below greyish; abdomen below yellowish white. Fore wings: 
costal margin to beyond middle bright olive-brown ; a similar 
streak above and below submedian to middle, followed above 
submedian by a small yellow spot; a yellow streak below median 
from veins 2 to 8; a small spot between 3 and 4; an outer yellow 
point between veins 6 and 7. Hind wings shaded with bright 
brown, and olive-brown hairs. Fore wings below fuscous brown ; 
yellow marking between veins 2 and 4 as above ; costa and veins 
terminally yellow, the streaks longest at apex. Hind wings 
below dull dark brown, the veins all yellow; a streak through 
cell, and a broad yellow streak below cell and vein 2. 

Expanse 24 mm. 

Tab. San Mateo. 


MoLo NEBROPHONE, sp. n. (PI. LIV. fig. 7.) 


S$. Palpi orange-brown tipped with black. Body fuscous 
partly clothed with golden-brown hairs; abdomen below yellowish 
with a black ventral line. Fore wings black, except base of 
costal margin to near middle, cell, inner margin to beyond 
middle, postmedial spots from submedian to vein 4, a small outer 
spot above 4, and spots above and below vein 7, which are orange- 
brown; a slight fuscous streak in cell. Hind wings orange- 
brown; costal margin broadly black with a medial inbent pro- 
jection; inner margin broadly black ; a broad black streak below 
cell to middle of wing; a fuscous shade on termen towards anal 
angle. Fore wings below deep yellow; some black on base of 
cell; a broad black shade below cell, upbent on discocellular ; 
termen black from vein 2 to inner margin; apex shaded with 
reddish, the outer spots at vein 7 larger. Hind wings 
below yellow on basal half; the inner margin broadly black 


COSTA RICAN BUTLERELIES. 365 


irrorated with yellow-brown; outer half reddish with oblong 
outer yellow spots from vein 6 to below vein 2; subterminal 
fuscous shading between veins 2 and 4; a darker streak just 
beyond cell between 4 and 6. 

2 fuscous brown ; a small yellow spot at upper angle of cell ; 
traces of some postmedial yellowish spots. Fore wings below 
fuscous; costal margin yellowish ; apex tinged with dull reddish ; 
« yellow spot at end of cell; a large postmedial yellow spot below 
vein 2, and smaller spots above 2 and 3; small yellow outer spots 
above and below vein 7. Hind wings below as in male, the ter- 
minal space duller red, limited by a yellow streak before the 
fuscous inner space. 

Expanse, d 27 mm., 9 33 mm. 

Hab. Cayrillo. 

The female from the Bejuco river, Panama. 

Near MW. herewa Hew. 


Moto? APELLA, sp. n. . (Pl. LITT. figs. 4, 5.) 


3. Palpi, head, collar, and patagia orange-brown. Thorax 
and abdomen fuscous, the latter with orange-brown hairs 
laterally; underneath yellowish. Wings black, the cilia and 
markings orange-brown. Fore wings: the costal margin broadly 
orange-brown, entering cell, narrowing towards black apical 
space; a fascia from vein 8 narrow and outangled between 
veins 5 and 6, then broader and expanding to middle of inner 
margin, cut by black veins. Hind wings: base and inner margin 
clothed with orange-brown hairs; a broad postmedial fascia, not 
reaching apex or inner margin. Wings below brownish yellow ; 
some terminal black points. Fore wings: the outer margin 
broadly darker with subterminal black shadings from vein 3 to 
inner margin; a black space on basal third below cell; a fine 
black line on discocellular.. Hind wings: antemedial, postmedial, 
and terminal darker shadings on interspaces. 

2. Wings above tinged with brown, the fascia much reduced, 
more macular, yellowish; the costa of fore wings with some 
yellowish irrorations. Wings below dark olive-brown, the veins: 
paler ; the postmedial fascia on fore wings broad, as in male, but 
yellower. Hind wings with postmedial dull yellowish spots om 
interspaces. 

Expanse, ¢ 37 mm., @ 45 mm. 

Hab. Cavrillo. 

The male above is very much lke J/.? humeralis Mab., but 
differs considerably on the underside of hind wings. 

In British Museum. 


CARYSTUS SUBRUFESCENS, sp.n. (PI. LIV. fig. 11.) 
2. Palpi whitish buff. Head light brown. Palpi shaded with 


dull reddish brown. Abdomen fuscous brown, the hairs at base 
dull greyish green. Wings fuscous brown. Fore wings: costal 


366 MR. WILLIAM SCHAUS ON 


margin at base shaded with reddish; spots semihyaline whitish 
yellow ; a streak in end of cell above median, and a spot below it 
to vein 2, its outer edge incurved; an elongated postmedial spot 
above vein 4, and a smaller spot below it; small outer spots 
between veins 6 and 8; a small yellowish spot above submedian 
just beyond middle; cilia yellowish at tornus. Hind wings: 
cilia tipped with pale yellow. Fore wings below black; the 
costal margin, apical space beyond cell, and termen to vein 2 
brownish red ; a yellowish white shade medially above submedian. 
Hind wings below brownish red; veins finely pale yellow; a 
fuscous shade from base before fold, expanding at anal angle. 

Expanse 36 mm. 

Hab. Guapiles. 

Near C. commodus Plotz, distinguished by the pale veins on 
hind wings below. 


EPInosis, gen. nov. 


S$. Palpi upturned, hairy. Antenne more than half the length 
of fore wing, thickened towards end which is curved and pointed. 
Wings broad. Fore wings: outer margin evenly curved; vein 2 
from middle of cell; 3 close to lower angle; 4 from lower angle; 
5 from below middle of discocellular, which is slightly oblique ; 
6, 7, 8 close together; a narrow brand below median from 
vein 3 to 2. Hind wings: veins 3 and 4 from lower angle 
slightly apart ; 6 and 7 from upper angle. 

Type of genus, Pamphila angularis Moschl.= Hesperia parvi- 
puncta Méschl. fide Godman, (Pl. LITT. fig. 1.) 

A common species in Costa Rica not mentioned in the ‘ Biologia’, 
and agreeing perfectly with other specimens from the Guianas. 


FAuGa ? HERMIONE, sp.n. (Pl. LIV. fig. 20.) 


¢. Palpi black, the second joint with a lateral white shade. 
Body above fuscous with orange-brown hairs; abdomen below 
yellowish white with a fine interrupted medial and outer black 
line. Fore wings black; base of costa shaded with brown ; sub- 
costal broadly orange-brown, not reaching apex, beyond cell with 
short upbent lines; a broad postmedial orange-brown shade 
inbent from vein 4 to submedian, its edge uneven, and joined by 
a similar streak below cell; cilia orange-brown partly spotted 
with black, entirely black at apex. Hind wings and cilia orange- 
brown, the margins black; a black streak from base near inner 
margin. Wings below orange-brown. Fore wings: a black 
streak through cell, and on interspaces above and below vein 5; 
tornus black ; subterminal black spots on other interspaces, and 
streaks on costa towards apex. Hind wings: a terminal black 
line; a streak below cell and vein 2, and one at apex ; subterminal 
black spots above and below vein 3. 


COSTA RICAN BUTTERFLIES. 


The female has the orange - brown on fore wings 
restricted. 

Expanse 26 mm. 

Hab. Carillo. 

In British Museum. 


367 


more 


EXPLANATION OF THE PLATES. 
Pirate L. 
Fig.1. Napeogenes hemisticta. Fig. 6. Huptychia drymo. 
2. Ithomia bolivari. 7. Hresia sticta. 
3. Pteronymia fumida. 8. 6 CHa Bx 
4, Huptychia agnata. 9. Phyciodes phlegias, ?. 
5. Catargynnis dryadina. 
Pxuate LI. 
Fig. 1. Hurygona leucon, 3. Fig. 6. Victorina steneles, ab. stygiana. 
2. Hupetoieta poasina. 7. Hurygona matuta. 
3. Hurygona leucon, 2. 8. BS mystica. 
4. Eubagis hecuba, 6. 9. Phyciodes dora. 
5. 35 sp oF 10. Perisama barnesi. 
Prate LII. 
Fig. 1. Thecla guapila. Fig. 6. Thecla hypocrita. 
2. Enantia lua costaricensis. Yo by 9 UGOBs 
3. Acmepteron poasina, 2. 8. Archonias intermedia, 3. 
4. Thecla melma. 9. Thecla subflorens. 
6. Pieris limona, 2. 10. 4, amphrade. 
Prats LIII. 
Fig. 1. Epinosis angularis. Fig. 8. Eurygona leucophryna. 
2. Corrachia leucoplaga. 9. Hermathena oweni. 
3. Cobalus lateranus. 10. Phyciodes niveonotis, 2. 
4, Molo? apella, 3. 11. Charis subtessellata. 
Suns 33 2 12. Mesosemia albipuncta. 
6. Vorates cotiso. 13. Charis turrialbensis. 
7. Heliconius galanthus, ab. sub- 
rufescens. 
PuatEe LIV. 
Fig. 1. Telemiades mnemon. Fig. 11. Carystus subrufescens. 
2. Butleria lysis. 12. Bolla sodalis. 
3. Cobalus laureolus. 13. Megistias xantho. 
4. Bolla machuca. 14. Cyclosemia subcerulea. 
5. Cobalus nigrans. 15. Butleria lethea. 
6. Enus nausiphanes. 16. Cobalopsis latonia. 
7. Molo nebrophone, : 17. Ate canace. 
8. Echelatus lucina. 18. Cobalus pindar. 
9. Rhinthon luctatius. 19. Sturiodes lyco. 
10, Lerodea ? rupilius. 20. Falga ? hermione. 


368 MR. C. TATE REGAN ON 


27. A Collection of Fishes made by Professor Francisco 
Fuentes at Haster Island. By C. Tare Reaay, M.A., 
F.Z.8. 

[Received March 3, 1913: Read April 8, 1913.] 


(Plates LV.—LX.*) 


INDEX. Page 
Acanthistius fuscus, sp. n. seechipiaeetaeieeaee aca ae MeO OS, 
Ganellopsicemims te teenrcre teen ccichen sieoteeriinos riences tee 369 
Wabrichehysmfuentest spe sre acs-ccuneesseasdekeeee ee eee OL 
AlGMOIDSES FOUNALOWP, So Wo goo cc0800 28000 a00000 900160700000 860 006 371 
Pseudomonacanthus paschalis, sp. ne ...... 0. ccc ceecee eevee =—872 
IBAChySteliusronientale spo meee ereeeeeae cee OTS 


Easter Island lies in 27° 6'S. and 109° 17' W.; it is very isolated, 
distant about 2500 miles from the coast of Chile and nearly 
1000 miles from Ducie, the nearest island of the Paumotu 
Archipelago. A small collection of fishes made at Easter Island 
in April, 1911, by Professor Fuentes, is of considerable interest. 
Tt includes examples of eleven species : seven belong to genera 
or species of the tropical Pacific, three are new and are related 
to species known from Norfolk Isiand or from New South Wales, 
and one belongs to a new genus whose affinities are with Zephreops — 
of New South Wales and the southern coasts of Australia. No 
Chilean species are represented, and it is of interest to note that 
in the case of Chilean genera, Acanthistius and Labrichthys, 
relationship is with the Norfolk Island species and not with the 
Chilean ones. 

SERRANID &. 


1, AcANTHISIIUS FuscUs. (PI. LV.) 


Acanthistius cinctus (non Giinth.) Kendall & Radcliffe, Mem. 

Mus. Comp. Zool. xxxv. 1912, p. 107. 
‘¢ Capuco.” 

Depth of body 2% in the length, length of head 22. Snout 12 
as long as diameter of eye, which is 54+ in the length of head ; 
interorbital width 7. Lower jaw a little projecting : maxillary 
extending to below middle of eye, its width distally nearly ? 
diameter of eye; occiput, interorbital region, cheeks, and opercles 
covered with ciliated scales; rest of head naked, or with minute 
imbedded scales; 9 or 10 gill-rakers (and some rudiments) on 
lower part of anteriorarch. Scales strongly ciliated, 105 in a longi- 
tudinal series above the lateral line, 55 in the lateral line, 17 
from base of spinous dorsal to lateral line. Bioveal XIII 15; 
fourth to sixth spines longest, # length of head, ee soft 
rays. Anal III 8; second : spine longest and strongest, - + length 


* For explanation of the Plates see p. 373. 


EPO VOM pe ly exe 


A.A, Searle, del et lith. Huth,imp. 
PSEUDOMONACANTHUS PASCHALIS. 


Pent 


ae 
eee 


ers 


“meant 
AERA t 


ie Hy ye 3 
iui Is 


ty 


P7196. Pl Lix. 


AH. Searle, del et Lith. Huthimp. 
ANAMPSES PULCHER. 


& 
“Ss Gil waned! ONT 


aittmwWitlad SpAHoOAINVNV 


kg 
td 
Oo 
KS 
Ma 
Q 
4 
Z 
4 
By 
G 
o 


oe 


ISH LNANY SAH LAO aVY 1 


iE 
ty 
isp) 
a 
b 
Au 
-: 
a 
lem 
E 


‘des “qa 


‘THAT ‘Tad @16ET SZ a 


aes 
en aN 
3 i ‘ ene i Rares Peete ume er 
BUC Se A teed hou te pe age ; eral lessee at 
Pee ents: et ‘3 ng Pease 7 i z Ss aeeh lar ented fy 
Saye ka <4 ' :? 
Rei 


Nae 


ata, 


eeu 


12 Zl Qls), Pw IVAN 


AH. Searle del et lith, ee Huth imp, 


GIRELLOPS NEBULOSUS. 


1 Zor), WSN) IE, IENWAL; 


A.H.Searle,del et lith. Huth,imp. 
KUHLIA MUTABUNDA. 


J 4 hSy, LOS), JELLY, 


A.H.Searle del. etlith. 


Huthimp. 
AGAN Pols LoS PUSGuS: 


FISHES FROM EASTER ISLAND, 369 


of head. Pectorals and pelvics subequal, a little more than 
length of head. Caudal rounded. Dark brownish. 

A single specimen, 230 mm. in total length. 

This species is very closely related to A. cinctus Giinth. from 
Norfolk Island, described from a specimen of nearly. the same 
size. Comparison shows that 4. cinctus differs in the larger head 
(22. in the length), higher dorsal fin (longest spines } the length 
of head), longer second anal spine (3 the length of head), longer 
and stronger maxillary (extending to below posterior } of eye, its 
width 7 the diameter of eye), and the banded coloration. 


Die 


KUHLIIDA. 


2. KuHLIA MUTABUNDA Kendall & Radcliffe. (Pl. LVI.) 


Mem Mus. Comp. Zool. xxxy. 1912, p. 105, pl. iii. f. 1. 

‘“¢ Mahori.” 
Depth of body 23 in the length, length of head 31. Snout 2 
diameter of eye, which is 22 in length of head; interorbital width 
33. Lower jaw a little prominent; maxillary extending to below 
anterior + of eye; cheek with 4 series of ctenoid scales; angle and 
lower edge of preoperculum finely denticulated; 25 gill-rakers on 
lower part of anterior arch. 50 scales in the lateral line, 6 between 
sheath at base of anterior dorsal spines and lateral line, 13 from 
lateral line to pelvic fin. Dorsal X11; fourth and fifth spines 
longest, 4 length of head; last 3 length of head, a little shorter than 
seventh, considerably shorter than longest soft rays. AnalIII 11; 
third spine a little longer than second, as long as tenth dorsal ; 
base of fin a little more than $ length of head or 11 its distance 
from caudal. Pectoral 2 length of head. Caudal deeply forked. 
Caudal peduncle 12 aslongas deep. Silvery, back darker ; caudal 
blackish posteriorly. — 

A single specimen, 190 mm. in total length. 

Related to A. humilis De Vis, from Queensland and the Fiji 
Islands, and to K. sandvicensis Steind., known from the Sandwich 
Islands and Society Islands. 


GIRELLIDS. 
GIRELLOPS, gen. nov. 
Related to Tephrwops, but with uniserial teeth. 


3. GIRELLOPS NEBULOSUS Kendall & Radcliffe. (Pl. LVIL.) 
Girella nebulosa Kendall & Radcliffe, Mem. Mus. Comp. Zool. 
RAV, OO ple lites 2 pl. tive tt. Vo 2. 
‘“* Maahaki.” 
Depth of body 22 in the length, length of head 4. Diameter 
of eye 4 in length of head, interorbital width 2. Scales on cheeks, 


temporal region, and upper) edge of operculum; rest of head 
naked, Teeth in jaws fixed, uniserial, incisor-like, the anterior 


370 MR. C. TATE REGAN ON 


ones truncated, the lateral ones obtusely tricuspid; palate toothless ; 
maxillary extending to below nostrils. Depth of preorbital 2 
diameter of eye. Gill-membranes broadly united, forming a fold 
across isthmus. Dorsal XVI 12, scaly at the base ; spines stout, 
subequal from the fifth to the last, which is 3 the length of head 
and a little shorter than the anterior soft rays. Anal III 10; 
second and third spines subequal, 4 length of head; second or 
third soft rays + as long as head. Caudal slightly emarginate. 
Pectoral as long as head. Seales ciliated, 78 in a longitudinal 
series, 12 or 13 between sheath at base of anterior dorsal spines 
and lateral line. Dark greyish, with traces of darker spots and 
markings. 

The above description from a specimen of 270mm. A young 
example, 43 mm. in total length, has the same number of scales 
and fin-rays, but all the teeth are tricuspid, and the markings 
obscurely indicated in the adult take the form of 7 dark cross- 
bars and scattered spots on the body, and longitudinal bands on 
the dorsal fin. The nearest ally of this species appears to be 
Tephreops richardsonit, which ranges from New South Wales to 
Western Australia. 


CARANGIDS2. 


4, CARANX CHEILIO Snyder. 


Caranx guara (non Bonn.) Kendall & Radcliffe, Mem. Mus. 
Comp. Zool. xxxv. 1912, p. 99. 
“« Poopo.” 
Originally described from a specimen 30 inches long obtained 
in the market at Honolulu; the example now recorded from 
Easter Island is 12 inches long. 


PoMACENTRID A. 


5. Pomacentrus inornatus. (PI. LVI. fig. 1.) 


““ Cototi.” 
2 Pomacentrus niomatus De Vis, Proc. Linn. Soc. N. 8. Wales, 
vill. 1883, p. 451. 
2 Pomacentrus inornatus Seale, Bull. U.S. Fish. Bureau, xxv. 


UQOS; jos BBs 

Pomacentrus jenkinst Kendall & Radcliffe, Mem. Mus. Comp. 
Zool. xxxv. 1912, p. 132. 

Depth of body 14 in the length, length of head 32. Snout 
a little shorter than eye, the diameter of which is 33 in the 
length of head; interorbital width 3. Mouth subterminal. 
Head scaly, except lips and extremity of snout; 2 series of scales 
on cheek and 1 on lower limb of preoperculum. Preorbital 
entire; suborbitals and preoperculum denticulated. 28 scales 
in a longitudinal series, 20 in the lateral line, 3 between middle of 
spinous dorsal and lateral line. Dorsal XIII 16; spines subequal 


FISHES FROM EASTER ISLAND. By! 


from fourth to last, which is 4 length of head; middle soft rays 
nearly as long as head. Anal II 13. Pector: i] as long as bead, 
Caudal deeply emarginate. Caudal peduncle much deeper than 
long. Brownish; scales black-edged; vertical fins blackish ; 
a small blackish spot above axil of pectoral. 

A single specimen, 100 mm. in total length. 

De Vis’s description is so incomplete that it is impossible to 
identify his species with certainty. He described it as probably 
from the South Seas, and Seale has recorded this, or a related 
species, from Samoa. P. jenkinsi; from the Sandwich Islands, as 
described and figured by Jordan and Evermann, seems to have 
a broader and more convex interorbital region, deeper preeorbital, 
and less deeply emarginate caudal fin. 


LABRIDZ. 


6. CHEILIO INERMIS Forsk, 
“ Ure-ure.” 
A species of wide distribution, ranging from Africa to the 
Western Pacific. 


7. JULIS UMBROSTIGMA Riipp. 


Sa acojues 
This also occurs heowlivont the tropical Indo-Pacific. 


8. LABRICHTHYS FUENTESI, sp. n. (Pl. LVIIT. fig. 2.) 
Pseudolabrus inscriptus (non Richards.) Kendall & Radcliffe, 
Mem. Mus. Comp. Zool. xxxv. 1912, p. 137, pl. v. f. 2, pl. vi. f. 1. 


** Cootea,” 

Depth of body 23 in the length, length of head 33. Snout 132 
as long as diameter of eye, one is 5d in the length of head 
and a little less than the interorbital width. Upper jaw with a 
posterior canine on each side and a pair of strong anterior 
canines; lower with 2 pairs of anterior canines; cheek with 
5 series of scales. 25 scales in the lateral line. Dorsal IX 11; 
seales extending on to basal part of fin. Anal IIT 10. Pectoral 
2 length of head. Caudal subtruncate. Body with about 
8 irregular cross-bars ending in blotches on basal part of dorsal 
fin ; head and upper part of body with numerous small pale 
spots or vermiculations; soft dorsal and anal with broad 
marginal dark bands including numerous pale spots with dark 
centres ; base of pectoral fin blackish. 

A single specimen, 165 mm. in total length. 

This species is nearest to L. luculenta NRichards., from Norfolk 
Island and Australia. 

9. ANAMPSES PULCHER, sp.n. (Pl. LVIIJI. fig. 3 & Pl, LIX.) 

“ Mouri.” 

Depth of body 2? in the length, length of head 34. Diameter 

Proc. Zoot. Soc.—1913, No. XX VI, 26 


BD MR. C. TATE REGAN ON 


of eye 7 in length of head, 25 in fenguln of snout. Interorbital 
region strongly convex, its width + the length of head. <A 
naked strip running backwards from ‘occiput to origin of dorsal 
fin. 28 scales in the lateral line. Dorsal 1X 12. Anal IIT 12 
Pectoral # the length of head. Caudal subtruncate. Each scale 
with a blue spot surrounded by a dark ring; head with blue 
horizontal stripes or series of spots; one through upper part of 
eye; another from end of snout through lower edge of eye, a 
third across the cheek ; vertical fins with spots like those on the 
body and with a narrow pale blue or white margin; base of 
pectoral blackish. 

A specimen of 205 mm., from Haster Island, differs from one 
of the same size of A. cwruleopunctatus Riipp. of the tropical 
Indo-Pacific, especially in the broader and more convex inter- 
orbital region, the incompletely scaled nape, and the different 
coloration of the head, A. cewruleopunctatus having vertical 
stripes running from eye to suboperculum. 

A second example of A. pulcher, 165 mm. in total length, 
from Tahiti (Wragge), is extremely similar to the type, but has 
the interorbital region less convex, and the naked median strip 
on the nape less elongate, not quite reaching the dorsal fin. 


BLENNIIDS. 


10. SALARIAS ARENATUS Bleek. 


Alticus striatus (? Cuv. & Val.) Kendall & Radcliffe, Mem. Mus. 
Comp. Zool. xxxv. 1912, p. 134. 


‘¢ Patuki,” 


A species of the Kast Indian Archipelago and Islands of the 
South Pacific. 


BALISTID. 


11. PSEUDOMONACANTHUS PASCHALIS, sp. n. (PI. LX.) 
“ Coreba.” 

Seales small, irregular, covered with minute spines. Depth 
of body 27 in the length. Snout produced, with upper profile 
a little convex, more than 3 times as long as eye and a little less 
than 4 the length of the fish. Lower angle of gill-opening below 
anterior margin, base of pectoral below middle of eye. Dorsal 
spine nearly as long as snout, above posterior 4 of eye, antero- 
posteriorly compressed, with a series of 24 small barbs on each 
side; soft dorsal of 35 rays, elevated anteriorly; anal rays 34. 
Caudal rounded. Greyish, with numerous scattered small dark 
spots. 

A single specimen, 300 mm. in total length. 

This species is related to the Australian P. ayraudi Quoy & 
Gaim. 


FISHES FROM EASTER ISLAND. 373 


EXPLANATION OF THE PLATES. 
Prater LV. 


Acanthistius fuscus. 
PLATE LVI. 
Kuhlia mutabunda. 


PuatE LVII. 
Girellops nebulosus. 
(a) Teeth. (6) Young. 
Prater LVIII. 


Fig. 1. Pomacentrus inornatus. 
2. Labrichthys fuentesi. 
3. Anampses pulcher; head of specimen from Tahiti. 


Puate LIX. 
Anampses pulcher. 


Prats LX. 


Pseudomonacanthus paschalis. 


APPENDIX. 


[It was not until the above was in the press and the plates 
had been drawn, that I saw Kendall and Radcliffe’s recent memoir 
on fishes from the eastern tropical Pacific. I have altered my 
paper only by taking their names for the new species of Kuhlia 
and Girellops and by inserting references to their work when 
our determinations did not coincide. They obtained nine of the 
eleven species, all but Anampses pulcher and Pseudomonacanthus 
paschalis. They also got a number of other species from Easter 
Island mostly known previously from the tropical Pacific, viz. 
Myripristis pralinius Cuv. & Val., Holocentrum punctatissimum 
Cuv. & Val., Decapterus sancte-helene Cuv. & Val., Cyphosus 
cinerascens Forsk., Pseudupeneus multifasciatus Quoy & Gaim., 
Thalassoma purpurewm Forsk., Teuthis umbra Jenkins, Kellogella 
oligolepis Jenkins, Alticus variolosus Cuv.& Val., and Xanthichthys 
lineopunctatus Hollard. ‘The identification of a species of 
Monacanthus with the Japanese M. cirrifer Schleg. seems open 
to question, and still more so the determination of an Eel as 
Gymnothorax dovi Giinth., as the type from Panama appears to 
differ in many respects from the specimens from Easter Island, 
which, from the data given, seem more like the Indo-Pacific 
G. meleagris Shaw. 

Finally, the resemblance to Norfolk Island is increased by the 
addition of two species—the one, 7’rachypomamacracanthus Giinth., 
apparently not distinct from the Norfolk Island form, the other 
a Bathystethus, requiring a new specific name. 


BAtHYSTETHUS ORIENTALE, Sp. 0. 


Platystethus cultratum (non Schneid.) Kendall & Radeliffe, 
Mem. Mus. Comp. Zool. xxxv. 1912, p. 97, pl. ii. £. 3. 
Closely related to B. cultratum, but head longer, 4 in length 
26% 


374 MR. C. TATE REGAN ON 


of fish (42 in B. cultratum), eye larger, 35 in length of head or 
nearly equal to its distance from edge of lower limb of pre- 
operculum (43 in head and 14 in its distance from lower edge 
of preoperculum in ZB. cultratwm), and fin-rays more numerous. 
Dorsal VIII, I 27; anal III 32 (VIII, I 23-25 and IIT 28-30 
in B. cultratum). 

The type from Easter Island measures about 200 mm. in total 
length (to end of middle caudal rays), the examples of B. culiratum 
from Norfolk Island that I have examined 170 to 180 mm.| 


28. A Revision of the Fishes of the Genus Kuhlia. 
By C. Tate Regan, M.A., F.Z.8. 


[Received March 4, 1913: Read April 8, 1913.] 
(Text-figures 68 & 69.) 


INDEX. Page 
Kuhlia cerulescens; sp. 1. ......0.0-deccevee eee eeesse ee 316 
RO TOAUGOG On CID 1G ceosudeededbaeecbsocdecoue onc nooosanmncas, | SU 
TRG, SDUCTIIGIS, SDs We 306600.39500500000 000 cens0n000000acc00a | SI) 


I have been led to undertake a revision of the species of the 
Indo-Pacific genus Auhlia Gill, owing to the difficulty I 
experienced in determining the correct name for a fish of this 
genus from Haster Island (Kuhlia mutabunda). 

To the synonymy of Auhlia as given by Boulenger (Cat. Fish, 
i. p. 36) may be added Boulengerina (non Dollo) Fowler (Proce. 
Acad, Philad. 1906, p. 512), and Safole Jordan (Proc. U.S. Nat. 
Mus. xlii. 1912, p. 655). 

These names are intended to distinguish the more silvery 
and more strictly marine species with more numerous gill-rakers 
(Safole) from the spotted forms that enter rivers and have fewer 
gill-rakers (Kuhlia). There is but little justification for such 
a generic division. . boninensis is intermediate between the 
two groups in the number of gill-rakers; A. marginata may 
sometimes be unspotted, and in the form and coloration of the 
caudal fin is nearer to K. humilis than to A. rupestris, and 
K. splendens, a silvery, unspotted species, with numerous gill- 
rakers, is described below from the fresh-waters of Rodriguez. 


Synopsis of the Species. 


I. Lateral line of 40 to 45 scales (+ 4 or 5 on base of caudal 
fin); 16 to 19 gill-rakers on lower part of anterior 
arch ; body usually spotted. 

A. Caudal fin slightly emarginate ; middle rays } or ¢ as 
long as longest; lobes rounded. Dorsal X 11. 
Anal III 10. : 

1. Maxillary extending to below middle of eye (young) 
Oe loenv Oral (Exh) Goo coconaons xono00900800020 940009 0005000 

2. Maxillary not or barely reaching middle of eye in 
adult. 


1. rupestris. 


FISHES OF THE GENUS KUHLIA. 375 


Upper opercular spine nearly as strong as lower ; dorsal origin 
in advance of pelvies.., 53 Seavstea asec harseceenpeierae : 
Lower opercular spine ins the nornsteg agra origin above 
base of pelvics .......... Job soapp>aanmounghooehseMCLA OCA IOce 


BG. Caudal fin more Heonly 6 emar roinattes suave rays $ 
to 2 as long as longest ; lobes pointed. Dorsal X 
10-12, Anal III 11-13. 


Maxillary extending beyond middle of eye ............0000:0020 4. wnvilliz. 
Maxillary not or barely reaching middle of eye .................. 5. marginata. 
II. Lateral line of 46 to 56 scales (+ 4 or 5 on base of caudal 
fin) ; no spots on body. 
A. 21 or 22 gill-rakers on lower part of anterior arch ... 6. boninensis. 
B. 24 to 29 gill-rakers on lower part of anterior arch. 
1°) Dorsal xe 13=145 > Amal WW V2=W3) eoe..s 22k seis 7. caudovittata. 
2. Dorsal X11. Anal III 11; caudal fin plain, usually 
with dark posterior margin. 
a. Middle caudal rays more than + as lone as 


longest ; 4 scales from sheath at base of spinous 
dorsal to nearest part of lateral line. 


Kye a little shorter than postorbital part of head .............. 8. splendens. 
Hye as long as postorbital part of head ................60.0.se0cc0 ee 9. humilis. 
6. Middle caudal rays less than 3 as long as longest ; 
5 or 6 scales from sheath at base of spinous 
dorsal to nearest part of lateral line. 
Anal nearly as long as ne much longer than caudal 
peduncle eer. s-s-- ener ... 10. sandvicensis. 


Anal } length of head, a en ener: thant aida Peannicls .. IL. mutabunda. 


8. Dorsal X 9-11. Anal III 10-11. Caudal fin with 
five blackish bands, one median and two on each 
WO bey ee Se: i Settee do astiiec adem. satis satcnasth aceoncs eee bas MUNG. 


bo 


. cerulescens. 


1. KUHLIA RUPESTRIS. 


Oentropomus rupestris Lacep. Hist. Nat. Poiss. iv. pp. 252, 
273 (1802). 

Kuhlia rupestris (part.) Bouleng. Cat. Fish. i. p. 36 (1895).* 

Kuhlia rupestris hedleyi Ogilby, Proc. Linn. Soc. N. 8. Wales, 
Sul, WD 06 WO 

Depth of body 23 to 3 in the length, length of head 23 to 33. 
Snout ?# to 13 diameter of eye, which is 37 to 54 in length 
of head; interorbital width 3 to 38. Maxillary extending to 
below middle of eye (young) or beyond. 16 to. 19 gill-rakers on 
lower part of anterior arch. 40 to 44 scales in lateral line, 4 or 
43 from scaly sheath at base of spinous dorsal to highest part of 
lateral line, 9 to 11 thence to ape of pelvic fin. Dorsal X 11 ; 
fourth or fifth spine longest, 1 to more than 2 length of head ; 
last as long as sixth, seventh, or eighth, 7 to more than 2 2 length 
of head. Anal III 10; third spine usually shorter than tenth 
dorsal; base a to 4 length of head, longer than caudal peduncle. 
Pectoral 4 to = length of head. Caudal peduncle as long as, or 
longer than, deep. Bluish silvery ; each scale usually with a 
dark spot ; each lobe of caudal and soft dorsal with a blackish 


* The synonymy is as given by Boulenger, except for Dules fuscus of Sanvage, 
which is a distinct species. 


376 MR. C. TATE REGAN ON 


or brownish spot in young, these spreading to form broad marginal 
bands in adult ; soft dorsal and upper and lower margins of 
caudal often narrowly edged with white ; soft vertical fins usually 
with one or more series of spots. 

Indo-Pacific, from Africa to the Paumotu Archipelago. Twenty- 
six specimens, 85 to 400 mm. in total length. : 


2. KUHLIA CHERULESCENS, sp.n. (Text-fig. 68.) 

Kuhlia rupestris (part.) Bouleng. Cat. Fish. i. p, 36 (1895). 

Depth of body 3 in the length, length of head 3}. Snout ? dia- 
meter of eye, which is 32 in length of head; interorbital width 33. 
Maxillary extending to below middle of eye; upper opercular 
spine nearly as strong as lower; 7 gill-rakers on lower part of 
anteriorarch. 41 scales in lateral line, 44 from sheath at base of 


Text-fig. 68. 


Head of Kuhlia cerulescens. 


spinous dorsal to highest part of lateral line, 95 thence to base of 
pelvic fin. Dorsal X 11; origin in advance of base of pelvies ; 
fifth spine longest, 2 length of head; last as long as seventh, 
3 length of head. Anal III 10; third spine 3 length of head ; 
base ?# length of head or 12 as long as caudal peduncle. Pectoral 
less than #2 length of head. Caudal slightly emarginate, with 
rounded lobes. Caudal peduncle 1+ as long as deep. Bluish 


FISHES OF THE GENUS KUHLIA. 377 


silvery; a dark spot at base of each scale ; cheeks and opercles 
spotted ; soft dorsal and anal with longitudinal bands or series of 
spots; caudal with broad dark band posteriorly. 

Solomon Islands. 

A single specimen, 290 mm. in total length, from Stirling 
Island. 

This species is near K. sawvagii, but has the coloration of 
K. rupestris. Sauvage’s figure of the former agrees with the 
example in the British Museum collection, and differs from the 
type of K. cerulescens in the much smaller upper opercular spine 
and the more posterior origin of the dorsal fin. 


3. KUHLIA SAUVAGII, sp. n. 

Dules fuscus (non Cuv. & Val.) Sauvage, Hist. Madag., Poiss. 
p. 149, pl. xv. f. 4 (1891). 

2 Moronopsis fuscus Steind. Sitzungsb. Akad. Wien, Ixxxu. 
1881, p. 240. 

Depth of body 22 in the length, length of head 33. Snout 
2 as long as diameter of eye, which is 3 in the length of head and 
equal to the interorbital width. Maxillary extending to below 
anterior + of eye; lower opercular spine much stronger than 
upper; 16 or 17 gill-rakers on lower part of anterior arch. 44 
scales in lateral line, 4 between sheath at base of spinous dorsal 
and arch of lateral line, 10 thence to base of pelvic fin. Dorsal 
X 11; origin above base of pelvics; fifth spine longest, 2 length of 
head ; last nearly as long as seventh, 4 length of head. Anal IT 
10; base nearly as long as head or nearly twice as long as caudal 
peduncle ; third spine less than 3 length of head. Pectoral ? 
length of head. Caudal slightly emarginate. Caudal peduncle 
a little longer than deep. Silvery, back darker; sides with 
scattered rounded or erescentic reddish-brown spots; soft dorsal 
and anal spotted ; caudal with dark membrane and pale rays. 

Madagascar. 

The above description is based on a single specimen, 150 mm. 
in total length, from Imerina, Madagascar, which evidently 
belongs to the same species as the example of 300 mm., also from 
Madagascar, figured by Sauvage. This larger fish has, of course, 
shorter spines, a smaller eye, ete., and in it the maxillary almost 
reaches the vertical from the middle of the eye. 

Dr. Pellegrin has kindly examined the types of Dules fuscus 
Cuv. & Val., two examples only 4 inches long, and writes that the 
maxillary extends slightly beyond the vertical from the middle of 
the eye. There can be little doubt but that these are specimens 
of K. rupestris. 


4, KUHLIA URVILLII. 


Kuhlia urvillii Bouleng. Cat. Fish. i. p. 38 (1895). 

This species is based on a coloured figure in Dumont D’Urville’s 
‘Voyage au Pole Sud,’ which represents a fish of 160 mm. very 
similar to K. marginata, but with the last dorsal spine longest of 


378: MR. C. TATE REGAN ON 


all, 2 the length of head, and with the maxillary extending to 
below the posterior part of the eye. 


5. KUHLIA MARGINATA. 


Dules marginatus Cuv. & Val. Hist. Nat. Poiss. ii. p. 116, 
pl. 1. (1829). 

Dules maculatus Cuv. & Val. op. c. vol. vii. p. 475 (1831). 

Dules malo Cuv. & Val. t. ¢. p. 479. 

Dules mato Lesson, Voy. Coquille, Zool. ii. p. 223 (1831). 

Dules leuciscus Jenyns, Zool. Beagle, Fish. p. 17 (1842). 

Kuhlia marginata Boulenger, Cat. Fish. i. p. 38 (1895) 
(with synonymy). 

Kuhlia malo Kendall & Goldsborough, Mem. Mus. Comp. 
Zool. xxvi. 1911, p. 280, pl. 1. f. 2. 

Kuhlia maculata Kendall & Goldsborough, ¢. c. p. 281, pl. 11. 
ity The 

Depth of body 22 to 31 in the length, length of head 8 to 3. 
Snout 4 to ? diameter an eye, which is 22 to 34 in length of 
head ; interorbital width 3 to 32. ee extending to below 
anterior 1 of eye. 16 to 18 gill-rakers on lower part of anterior 
arch. 40 to 45 scales in lateral line, 4 or 44 from sheath at base. 
of spinous dorsal to highest part of lateral line, 8 or 9 thence to 
base of pelvic fin. Dorsal X 10-12; fifth spine usually longest, 
4 to 2 length oe head ; last as long as sixth, seventh, or eighth, 
Pi to more than 2 2 length of head. Anal III 11-13; third spine 
shorter than tenth dor sal; base nearly as long as ‘head, nearly 
twice as long as caudal ’ peduncle. Pectoral 2 to # 2 length of 
head. Caudal moderately emarginate, the middle rays 4 to 2 
as long as longest. Caudal peduncle longer than deep. Silvery, 
usually dark spotted above lateral line and sometimes below it ; 
soft dorsal and anal dark edged, at least anteriorly, often dark 
spotted basally; caudal with dark posterior margin and with 
dark spots or a band parallel to posterior edge ; soft dorsal, anal, 
and upper and lower margins of caudal often narrowly edged 
with white ; pelvics sometimes spotted. . 

Malay Archipelago and South Pacific. 

Numerous examples, 75 to 160 mm. in total length, including 
the types of D. leuciscus, kindly lent to me by Mr. L. Doncaster. 

Some examples: are almost covered with spots, others are 
silvery with scarcely a trace of spots, the variation in this respect 
recalling our British Trout; probably the silvery examples are 
marine, whilst the strongly spotted ones may have been for a 
long time in fresh water. 


6. KUHLIA BONINENSIS. 


Kuhlia caudovittata (part.) Bouleng. Cat. Fish. i. p. 41 (1895). 
Dules marginatus boninensis Fowler, Proc. Acad. Philad. vin. 
1906, p. 510, fig. 


Depth of the body nearly 3in length, length of head 33. Snout 


FISHES OF THE GENUS KUHLIA. 379 


2 diameter of eye, which is 3 in length of head; interorbital 
width 33. Maxillary extending to below anterior 7 of eye. 
22 gill-rakers on lower part of anterior arch. 52 scales in lateral 
line, 5 from sheath at base of spinous dorsal to arch of lateral 
line, 12 thence to base of pelvic fin. Dorsal X 11; fifth spime 
longest, a little more than } length of head; last as long as 
eighth, 4 length of head. Anal II] 12; base as long as head 
or 14 as long as caudal peduncle. Pectoral 3 length of head. 
Caudal forked, the middle rays 3 as long as the longest. Caudal 
peduncle 13 as long as deep. Silvery ; back darker ; caudal with 
dark posterior edge and pale intramarginal band. 

Kastern Pacific. 

Here described from a specimen of 190 mm. from Tahiti: 
this is not so deep in the body as Fowler’s type, a specimen of 
about 115 mm. from the Bonin Islands. Otherwise, making 
allowance for differences due to size, the two examples agree 
perfectly. 


7. KUHLIA CAUDOVITTATA. 


Holocentrus caudovitiatus Lacep. Hist. Nat. Poiss. iv. pp. 332, 
367 (1802). 

Dules caudovittatus Cuv. & Val. Hist. Nat. Poiss. iii. p. 117 
(1829) and vii. p. 475 (1835); Sauv. Hist. Madag., Poiss. p. 151, 
pl. xvii. f. 3 (1891). 

- Kuhlia caudovittata (part.) Bouleng. Cat. Fish. i. p. 41 (1895). 

Evidently related to K. humilis, sandvicensis, etc., but dis- 
tinguished by a small head, the subequal ninth and tenth dorsal 
spines, and the increaséd number of soft rays, 13 or 14 in the 
dorsal, 12 or 13 in the anal. 

Madagascar and Mauritius. 

Total length 225 mm. 


8. KUHLIA SPLENDENS, sp. n. (Text-fig. 69 B.) 
Kuhlia mato (part.) Bouleng. Cat. Fish. i. p. 40 (1895). 


Depth of body 24 to 22 in the length, length of head 3 to 33. 
Snout 2 to 2 diameter of eye, which is 3 in length of head; 
interorbital width about 34. Maxillary extending to below 
anterior 1 of eye. 26 or 27 gill-rakers on lower part of anterior 
arch. 48 to 50 scales in lateral line, 4 from sheath at base of 
spinous dorsal to arch of lateral line, 11 thence to base of pelvic 
fin. Dorsal X 11; fourth or fifth spine longest, nearly = length 
of head: last as long as or longer than eighth, nearly 3 length of 
head. Anal III 11; third spine a little shorter than tenth 
dorsal; base a little shorter than head or 12 as long as caudal 
peduncle. Pectoral 2 length of head. Caudal deeply emarginate, 
but middle rays more than 2 as long as the longest. Caudal 
peduncle longer than deep. Silvery; back darker ; caudal with 
blackish posterior edge. 

Four specimens, 120 to 180 mm. in total length, from 
Rodriguez (fresh water) and Mauritius. 


380 ' MR. C. TATE REGAN ON 


A small stuffed specimen from South Africa also belongs to 
this species. 


Text-fig. 69. 


mo\\ 


Heads of (A) Kuhlia humilis and (B) Kuhlia splendens, in each case 
from a specimen 120 mm. in total length. 


9. Kuuuia HuMILIS. (Text-fig. 69 A.) 


Kuhlia humilis De Vis, Proc. Linn. Soc. N. 8. Wales, ix. 
1884, p. 396; Ogilby, Ann. Queensland Mus. x. 1911, p. 46, pl. vi. 
ie dl. 

Kuhlia malo (part.) Bouleng. Cat. Fish. i. p. 40 (1895). 

Kuhlia proxima Kendall & Goldsborough, Mem. Mus. Comp. 
Zoolexxvin | QUES 282 replete ta: 2. 


Depth of body 22 to 3 in the length, length of head 3 to 3}. 
Snout 3 to 2 diameter of eye, which is 2} to 2 in length of head ; 
interorbital width 31 to 3}. Maxillary extending to below 
anterior } of eye.. 25 gill-rakers on lower part of anterior arch. 
46 to 50 scales in lateral line, 4 from sheath at base of spinous 
dorsal to arch of lateral line, 10 or 11 thence to base of pelvic fin. 
Dorsal X 11; fourth or fifth spine longest, 2 length of head ; 
tenth a little shorter than seventh, more than ? length of 
head. Anal JII 11; third spine as long as or a little longer 
than second, as long as tenth dorsal; base of fin nearly 3 as long 
as head or 1} as long as caudal peduncle. Pectoral 3 length of 
head. Caudal deeply emarginate, but middle rays more than 4 as 
long as longest. Caudal peduncle 14 as long as deep. Silvery ; 
back darker; caudal with a rather broad dark posterior margin. 

Queensland ; Fiji Islands. 

Three specimens, 90 to 120 mm. in total length, from Fiji, are 
undoubtedly A. prozima, but this seems to be a synonym of the 
Queensland A. humilis. The species is scarcely distinct from the 
preceding, but has a larger eye, the anal fin a little shorter, and 
the blackish margin of the caudal fin broader. 


FISHES OF THE GENUS KUHLIA, 381 


10. KUHLIA SANDVICENSIS. 


Moronopsis argenteus, var. sundvicensis Steind. Sitzungsb, Akad. 
Wien, xxiv. 1876, p. 205. 

Moronopsis sandvicensis Steind. ib. xevi. 1887, p. 56, pl. i. f. 1. 

Kuhlia malo (part.) Bouleng. Cat. Fish. i. p. 40 (1895). 

?Kuhlia wenura Jord. & Everm. Bull. U.S. Nat. Mus. xivi. 
1836, p. 1015. 

Depth of body 22 to 24 in the length, length of head 31 to 32. 
Snout 2 to 4 diameter of eye, which is 22 to 3 in length of head ; 
interorbital width 31 to 4, Maxillary extending to below anterior 
edge or anterior 1 of eye. 25 to 28 gill-rakers on lower part of 
anterior arch. 48 to 52 scales in lateral line, 5 or 6 from sheath at 
base of spinous dorsal to arch of lateral line, 11 to 13 thence to base 
of pelvic fin. Dorsal X 11; fifth spine longest, 3 to length of 
head; last a little shorter than seventh, + to 4 length of head. 
Anal III 11; third spine as long as tenth dorsal; base of fin 
nearly as long as head or 12 to 14 as long as caudal peduncle. 
Pectoral 2 length of head. Caudal deeply forked, middle rays 
less than 4 as long as longest. Caudal peduncle 14 to 1} as long 
as deep. Silvery; back darker; caudal with narrow blackish 
posterior edge; dorsal and anal sometimes similarly margined. 

Eastern Pacific. 

Six specimens, 90 to 240 mm. in total length, from the Sand- 
wich Islands and Tahiti. 


11. KUHLIA MUTABUNDA. 


Kuhlia mutabunda Kendall & Radcliffe, Mem. Mus. Comp. 
Zool xexmye Ol, p05) poles teal 

This species from Easter Island, described above (p. 369, 
Pl. LVI), differs from A. sandvicensis especially in the shorter 
anal fin. 


12. KUHLIA TENIURA. 


Dules teniurus Cuv. & Val. Hist. Nat. Poiss. ii. p. 114 
(1829). 

Kuhlia teniura Bouleng. Cat. Fish. i. p. 39 (1895) (with 
synonymy). 

Kuhlia sterneckii Steind. Sitzungsb. Akad. Wien, cvii. 1898, 
p. 461, pl. 


This species is at once distinguished by the coloration of the 
caudal fin, with five black bands, a median longitudinal and two 
on each lobe. I am unable to regard Steindachner’s A. sternechit 
as distinct, for the relative length of the ninth and tenth dorsal 
spines varies considerably, and they are equal in 4 or 5 of the 
20 specimens I have examined. 

The species ranges throughout the Indo-Pacific, from Africa to 
the Revillagigedo Islands. 


382 MR. R. I. POCOCK ON 


29. The Affinities of the Antarectie Wolf (Canis antarcticus). 
By Re Pocock, ERS. eS ie ZS.) Curatenol 


Mammals. 
{Received and Read April 8, 1918.] 
(Text-figures 70-74.) 
INDEX. Pave 
1. Published views on the affinities of C. antarcticus............... 383 
2. Skull characters of C. antarcticus and C. latrans ............... 3884 
3. Hxternal characters of C. antarcticus and OC. latrans ......... 391 


The story of Canis antarcticus has been told by Darwin *, by 
Hamilton Smith 7, and more recently by Mr. Rupert Vallentin = 
from first-hand information, and by several authors indebted 
either to Darwin’s or Hamilton Smith’s account, or to the 
accounts of travellers who visited the Falklands before Darwin’s 
time. References to the literature down to 1890 may be found 
in Mivart’s ‘Monograph of the Canide,’? published in that 
year. According to Mr. Vallentin, Canis antarcticus became 
extinct in 1876, without leaving a trace of its former existence 
in the Falkland Islands; and since all the known material of the 
species appears to be preserved in London and Paris, I have 
attempted to supply the want expressed by Allen §$ by figuring a 
skull of one of the specimens in the British Museum. I have 
not, however, given detailed measurements of the skull, because 
these may be found in Mivart’s monograph and in the paper by 
Huxley mentioned below. 


Some six or seven years ago, when trying to identify some 
South American dogs exhibited in the Zoological Gardens, I took 
the incidental opportunity of looking at the skulls of a few of 
the species of Neotropical Canidze contained in the» British 
Museum, to learn, if possible, something of their affinities to 
one another and to the better known species inhabiting North 
America and the countries of the Old World. Amongst the 
species examined were Canis antarcticus, the so-called Wolf of 
the Falkland Islands, and Canis latrans, the Coyote or Prairie 
Wolf, which ranges roughly from Canada to Mexico. The 
examination was made without any intention on my part of 
adding to the literature of the subject, with which I was only 
acquainted in a very general way; and after satisfying myself 
that C. antarcticus was related to certain Neotropical forms, 
of which C. thous (= cancrivorus) may be taken as an example, 
and that the affinities of C. latrans lie with some of the so-called 
jackals and wolves of the Old World, I was contented to let the 
matter rest. 


* In Waterhouse’s Zool. of H.M.S. ‘ Beagle,’ Mammalia, p. 7, 1839. 

+ In Jardine’s Nat. Libr., Mammalia, ix. p. 252. 

{ Manchester Memoirs, xlviii. p. 45, 1904. This paper is quoted hy 
Mr. Lydekker, and some of the interesting and puzzling points connected with 
C. antarcticus are discussed in ‘The Field,’ Oct. 1, 1904. 

§ Rep. Princeton Univ. Exped. Patagonia, iii. pt. 1, p. 153, 1905. 


me THE ANTARCTIC WOLF, 383 


But in the summer of 1912, Lreceived for review from the Editor 
of ‘ Nature’ a copy of Dr. R. F. Scharft’s volume, ‘ Distribution 
and Origin of Life in America,’ 1911; and when I found it 
definitely stated therein that C. antarcticus is closely related to 
C. latrans, and when I saw the obvious difficulties in which 
Dr. Scharff was involved in his attempt to explain, on geo- 
graphical grounds, this singular aftilation, I ventured to reassure 
him by remarking, in effect, that his belief was devoid of morpho- 
logical foundation. 

Now, an author who compiles a volume on zoology of the size 
and scope of the ‘ Distribution and Origin of Life in America’ 
cannot be expected to verify all the statements of earlier and con- 
temporary writers. Nor in the present instance could Dr. Scharff 
be justly criticised for not travelling to London to examine for 
himself the preserved material of C. antarcticus, of which, I take 
it, there is no specimen in Dublin. Very naturally, therefore, 
he trusted to the verdict of others, and promptly replied to my 
remark with a request for my reasons for making it. But since 
I could not ask the Editor of ‘ Nature’ to give me the necessary 
space for justifying the statement I had made, I pledged myself 
to do this elsewhere, and the matter that follows is an attempt 
to redeem that promise. 

The acknowledged source of Dr. Scharfi’s opinion about the 
mutual affinities of C. antarcticus and C. latrans was the following 
passage in Mr. Lydekker’s ‘ Geographical History of Mammals,’ 
1896 :—“ Of the two indigenous mammals, the most remarkable 
is the Falkland Island Wolf (Canis antarcticus), which differs 
markedly from all the Canide of the mainland and is apparently 
closely allied to the North American Coyote (C. latrans)” (p. 140). 
I therefore wrote and asked Mr. Lydekker if he would kindly 
tell me his reasons for this conclusion, and he informed me that 
he took it from Prof. Huxley’s classic paper upon the cranial and 
dental characters of the Canidz, published in the ‘ Proceedings’ of 
this Society, 1880, pp. 238-288. Upon looking up this paper 
I find the following passages referring to the two species under 
discussion and bearing upon the question at issue :— 


(1) ..... But sometimes there is a well-defined though com- 
paratively narrow sagittal area, from the centre of which 
a low sagittal crest rises. This is well seen in some 
Jackals, and especially in C. antarcticus (p. 250). 

(2) In the large size of the upper molars..... C’. antarcticus 
presents the closest approximation to some specimens of 
C. latrans (p. 266). 

(3) From the range of variation of C. cancrivorus it can hardly 
be doubted that the examination of more extensive 
materials will prove the existence of an uninterrupted 
series of gradations from C. vetulus to C. antarcticus and 
C. jubatus (p. 266). 

(4) Seven crania of C. latrans, when measured, exhibit a con- 
siderable range of variation, though probably less than 
a larger series would show. But, as they are, I must 
confess myself unable to find an important break in the 


384 MR. R. I. POCOCK ON 


series of gradations of cranial and dental structure 
between Canis latrans and C. antarcticus on the one 
hand, and C. latrans and C. occidentalis on the other. 
..... IT may further remark that I can discern no 
difference of the slightest importance between skulls of 
C. latrans and those of some of our domestic dogs 
(pp. 272-273). 

(5) In the genus Canis we have .....as a lowest section the 
species of the C. cancrivorus and C. vetulus type (an- 
swering pretty much to the Aguarra dogs of Hamilton 
Smith), the Sacaline section (C. aureus, C. anthus, 
C. mesomelas, C. antarcticus, C'. latrans), and the Lupine 
section (C. dwpus and all its varieties) (p. 286). 

Whether these paragraphs justify Mr. Lydekker’s statement * 
that C. antarcticus differs markedly from all the Canidz of the 
mainland of South America and is apparently closely allied to 
C. latrans, and Dr. Scharff’s extension of this to the effect that 
C’. antarcticus is certainly closely related to C. latrans, must be 
left to individual judgment. 

Paragraph 1 merely points out one resemblance between 
C. antarcticus and some jackals. Paragraph 2 similarly points 
out one resemblance between the two species, but contains no 
suggestion of affinity between them. Paragraph 3 may be 
interpreted as suggesting affiliation between the extreme forms 
of South American Canide represented by C. vetulus and 
C. jubatus, with C. antarcticus lying midway between them. 
Paragraph 4 is more precise and states that there is no im- 

ortant structural break between C. antarcticus and C. latrans, 

and that the latter similarly intergrades with C. occidentalis 
and C. familiaris. Paragraph 5, on the contrary, definitely 
associates C. antarcticus and C. latrans, and at the same time 
severs the former from the group typified by C. vetulus and the 
latter from the group typified by C. lupus or occidentalis, an 
arrangement not easy to reconcile with the views expressed by 
paragraphs 3 and 4, 

After reading Prof. Huxley’s paper rather carefully for 
enlightenment on this subject, I must confess that I cannot 
form any clear idea as to his views of the athnities of the species 
he discussed, except in a broad sense. 

If the substance of paragraphs 4 and 5 afford some justifi- 
cation for Mr. Lydekker’s declaration respecting the relationship 
between CO. antarcticus and C. latrans, it must be admitted that 
paragraph 3 does not support the contention that C. antarcticus 
is quite unrelated to the species of Canide inhabiting the South 
American mainland. However that may be, the conclusions 
forced upon me by the examination of five crania of C. ant- 
arcticus and twelve of C. latranst in the British Museum and 

* Tn the article in ‘The Field’ (Oct. 1, 1904), above referred to, Mr. Lydekker 
evinces less assurance on these points; but he evidently could not bring himself to 


reject the authority of Huxley’s opinion. 
+ I use this term in its old-fashioned and broad sense, disregarding the species or 
subspecies recently dismembered from C, latrans by American systematists. 


THE ANTARCTIC WOLF. 


Text-fig. 70, 


L By 
Z ; 


SUNY Aer 


AOS 


NN 


NY) 


Bia 


NY 


tr 


) 


TMT 


A. Dorsal view of posterior part of skull of Canis latrans. 
B. Dorsal view of skull of C. antarcticus. ; 
0.¢., occipital crest; s.c., sagittal crest; s.a., sagittal area. 
The figures of the skull of C. latrans are from a specimen (¢) in the British 
Museum from Assiniboia (2.8.22), and those of C. antarcticus trom a specimen in 


the British Museum (69,2%24.3). ] 


386 MR. R. I. POCOCK ON 


the Museum of the College of Surgeons are:—(1) that C. ant- 
arcticus and C. latrans are not closely allied; (2) that C. antarcticus 
is more nearly related to the C. thous (= cancrivorus) group of 
South American Canidee than to C. latrans; (3) that C. latrans 
must be affiliated with such Old World species as C. pallipes, 
C. lupaster and C. anthus, and not with C. antarcticus. The 
first and third of these conclusions are borne out by the 
external characters of the two species concerned. My reasons 
for these conclusions are as follows :— 

The sagittal area and sagittal crest—As Huxley and Mivart 
have shown, the skull of C. antarcticus has a well-marked lyriform 
sagittal area which, according to the evidence of available crania, 
persisted throughout life, although in one of the three specimens 
in the British Museum it is decidedly narrower than in the two 
others. In the skulls of C. latrans that I have seen there is no 
distinct lyriform sagittal area, but in adult skulls there is a 
median cariniform sagittal crest varying in height with age. 
Even in two young skulls, in both of which the sphenoidal and 
occipital sutures are open, while one still retains a milk canine 
behind the permanent canine, there is no lyriform sagittal area. 
The significance of this depends upon the fact that the young of 
many species of Canidee of corresponding age or older show a 
stronger or weaker lyriform area corresponding with the sinuosity 
of the upspreading temporal muscle on each side, although in the 
young of no species of dog in which the adult possesses a carini- 
form sagittal crest does the lyriform sagittal area show, I believe, 
the development and definition it exhibits in the adult of 
C. antarcticus. However that may be, if C. antarcticus and 
C. latrans were closely related, we should at least expect to see 
a well-defined lyriform sagittal area in the skulls of subadult 
individuals of C. latrans killed before the temporal muscles had 
reached the summit of the cranium. But, as has been said, this 
area is remarkable for its indistinctness in immature skulls of 
that species. 

The occipital crest.—In C. antarcticus the occipital crest, when 
viewed from above, is transversely truncated and not angular; 
when viewed from the side it only overhangs the vertical portion 
of the supraoccipital to a small extent; and when viewed from 
behind it forms a truncated angle. In C. latrans this crest is 
angularly produced backwards in the middle line, overhangs the 
occipital area to a much greater extent, and is more acutely 
angled from behind. It varies in shape and development in this 
species, but never, so far as I have seen, resembles that of 
C. antarcticus (text-figs, 70 & 71). 

The malar bone. Th Canis antarcticus the anterior portion of 
the malar bone is marked by a strong masseteric ridge traversing 
approximately the middle of its ane surface; the sinless ior edge 
of the bone close to the maxilla is expanded convexly to afford 
additional support to the masseter muscle ; its upper edge close to 
the maxilla is somewhat out-turned, forming a very appreciable 
hollow on the subjacent portion of the eee above the first 


6 


a 


THE ANTARCTIC WOLF. 387 


molar tooth. In C. latrans the masseteric crest of the malar is 
low down on its external surface, the inferior edge of the bone is 
scarcely at all expanded, so that the area for the attachment of 


Text-fig. 71. 
BoLa: 


A. Occipital region of skull of Canis antarcticus. 
B. Occipital region of skull of C. latrans. 


S.a., sagittal area; s.c., sagittal crest. 


the masseter is much narrower than in (. antarcticus, and the 
upper edge of the malar is not noticeably out-turned, so that the 
hollow on the maxilla beneath it is less pronounced (text-figs. 72 
& 73, pp. 388-9). 

Upper carnassial tooth.—In C. antarcticus the antero-external 
Proc. Zoot. Soc.—1913, No. XX VII. 27 


388 MR. R. I. POCOCK ON 


cusp has the front border more rounded and the very fine crest 
that runs down it is defined on the inner side by a very indistinct 
groove. The antero-internal cusp is wider and rises further hack 
and has no distinct little crest running inwards towards the 


Wits 


I, 


Yd 


wd 


ae 


1] 


tyre 
A, 
f 
I 
/ 


Yi 


in ee 
il 
J My 
f) 
LE 


Z4 
ey 


if 


m., masseteric ridge or malar bone. 


Text-fig. 72. 


Side view of skull of Canis latrans. 


antero-external cusp. In C. latrans the crest traversing the 
anterior edge of the antero-external cusp is more pronounced 
and is defined by a distinct groove, the two combining to make 
the edge of this cusp more cutting than in C. antarcticus. The 


THE ANTARCTIC WOLF. 389 


antero-internal cusp is narrower and set distinctly more forwards 
than in C. antarcticus, and there is a delicate crest running along 
its surface towards the base of the antero-external cusp (text- 
fig. 74, A, B, p. 390). 

Lower carnassial tooth—The main cusp is higher and more 
pointed in C. antarcticus than in C’. latrans, and the little cusp at 
its base on the inner side is much lower, so that it stands on a 
little higher level than the internal cusp of the talon. In C.latrans 


m., masseteric ridge or malar bone. 


Mexct=tio~ Wa. 


Side view of skull of Canis antarcticus. 


this cusp is comparatively high up the main cusp of the tooth and 
is considerably above the inner cusp of the talon (text-fig. 74, 
CDM p9390): 

There are other minor differences both in the skull and teeth. 
The palatine bones, for instance, extend farther forwards with 
relation to the upper carnassials, and the margin of the posterior 
nares is also farther forwards with relation to the posterior 


molars in C. latrans than in C. antarcticus: the incisor teeth are 
21 * 


390 MR. R. I. POCOCK ON 


smaller and the crowns of the cheek-teeth are higher with relation 
to their breadth in C. antarcticus than in C. latrans. But apart 
from these, the principal differences mentioned above are quite 
sufficient to disprove the claim that the two species are closely 
related. According to modern standards of classification they are 
subgenerically, if not generically, distinct. 

But the characters above described tell us more than that. 
Taking C. latrans first, it is obvious that in the cariniform sagittal 
crest, the angularly produced occipital crest, the position of the 
masseteric ridge on the malar bone, and in the points alluded to 
in connection with the upper and lower carnassials, the species 
falls into line with the large wolves like C. occidentalis and lupus, 


Text-fig. 74. 


A. Vertical view of upper carnassial of Canis latrans. 
B. Vertical view of upper carnassial of C. antarcticus. 
C. Internal view of Jower carnassial of C. latrans. 

TI). Internal view of lower carnassial of C. antarcticus. 


and with (. pallipes and C. lupaster, which, according to fancy, 
may be called large jackals or small wolves. These resemblances 
explain Mivart’s dismissal of the cranial and dental characters of 
C. latrans with the remark, ‘‘The skull possesses no distinctive 
characters, nor have we been able to detect any in the shape of 
the teeth.” 

On the other hand, the skull of C. antarcticus, with its lyriform 
sagittal area and truncated occipital crest, agrees in the main with 
the skulls of certain species or subspecies of South-American dogs 
in the British Museum labelled C. thous (=cancrivorus), rudis, 
sclatert (=microtis), parvidens, urostictus, gracilis, and fulvipes. 
And in the skull of a dog, perhaps referable to C. gracilis, which 


THE ANTARCTIC WOLF. 391 


eame from Mar del Plata and died in the Gardens, the above- 
described crests on the upper carnassial are not better developed 
than in C. antarcticus, and the masseteric ridge on the malar bone 
shows a decided approximation to the condition seen in that 
species. This latter character is still better marked in the skull 
of another South-American dog, the exact locality of which is 
unknown, but which was a different species * from the Mar del 
Plata example, and the crests on the carnassial exhibit the same 
feebleness of development. But it may be noted that in both 
these skulls the positions of the cusps on the upper and lower 
carnassials are more latrans-like than antarcticus-like, so that in 
this respect at least they serve to bridge over the difference 
between those two species; a fact in keeping with the idea that 
C. antarcticus is a specialised form of the group of South- 
American dogs above alluded to, but specialised in a direction 
away from that taken by C. latrans and its allies. 

The external characters of C. antarcticus and CO. latrans also 
afford no justification for the claim of close relationship between 
them. In the first place the ears of C. antarcticus are very small, 
smaller indeed comparatively, I believe, than in any wild species 
. the dog family, with the exception perhaps of C. sclatera, 

C. (Nyctereutes) procyonoides, and Vulpes (Alopex) lagopus. In 
C. latrans, on the contrary, they are as large as in most, at all 
events, of the species of Canis. An idea of their length in the 
two species may be gathered from the measurement of a specimen 
of each of approximately the same size given by Mivart, the ear 
of OC. antarcticus being 6°5 cm. (= about 24 inches) and that of 
C. latrans 14 em. (=about 54 inches), or more than twice as long. 
It may be added that the measurement of 2 inches 9 lines assigned 
by Waterhouse to the ear of (. antarcticus confirms Mivart’s 
statement. 

As regards colour ©. latrans varies from grey to greyish fawn, 
mixed with black above, and shows the characteristic clouded or 
patchy coloration caused by the running together of the bands of 
the individual long coarse hairs of the ‘back and sides seen in so 
many of the so- called wolves and jackals. One of these long 
coarse hairs, pulled at random from a skin, measured about 
3 inches long, the black tip being 3 inch (12 mm.) and the 
whitish area below it 1 inch (25 mm.). The whole of the ventral 
surface from the chin to the root of the tail is usually white or 
whitish, and always apparently markedly paler than the back and 
sides, though sometimes the continuity of the light tint is inter- 
rupted on the throat by an infusion of fawn. There is no dark 
patch above the hock on the hind leg, and the tail matches the 
back approximately in colour throughout, the tip and the gland- 
spot being blacker than the rest. 

In C. antarcticus the coat is thick and soft, and comparatively 
short, with none of the long coarse hair seen in C. latrans. One 


* The South-American dogs of this groun are in such 2 systematic muddle that 
it is very difficult to identify specimens without a complete revision of the whole 
series, 


392 ON THE ANTARCTIC WOLF. 

of the longish hairs pulled at random from the back measured 
only 14 inches (37 mm.) in length, the dale apical tip being 
1 inch (6 mm.) and the pale ‘eral below it + inch (3 mm.) in 
length. The prevailing colour of the body is brown, relieved by 
the. fine speckling due to the narrow pale band on the individual 
hairs just described. The lower side is white only on the 
posterior portion of the belly and on the upper end of the throat, 
the-chin and lower jaw being white stained with a fuscous tint. 
Apart from the areas described the ventral surface is brownish. 
There is, moreover, as Mivart said, a fuscous patch above the 
hock of the hind leg and the tail is particoloured, its basal 
portion being like the back, its tip white, and the intermediate 
area blackish, the colour of this area gradually blending proximally 
with the brownish basal portion, but- being quite black distally 
and sharply defined from the white ter minal area, 

Perhaps it may rightly be claimed that these differences in 
the length of the ears and in colour do not count for much in 
themselves. That may be so. Nevertheless, if C. antarcticus and 
C. latrans were only known from their skins, it is quite certain 
that the latter would be placed in the same category with such 
species as C. pallipes and C. lupaster, and that C. antarecticus 
would be excluded therefrom. The latter would be difficult to 
classify; but there is one significant colour-feature connected 
with the species. ‘This is the presence of the dark patch above 
the hocks; and the interest of this lies in the circumstance that 
it is a very common feature in various species of the smaller 
South-American dogs and occurs in some of the species of Vulpes, 
hike V. chama*. 

There is one other little point that may be referred to. 
Darwin says he was informed that the cries of C. antarcticus 
resembled those of the South-American species C. azare. I have 
never heard C. azare bark or howl, but the keeper in the Gardens 
informs me that examples of wild dogs from Mar del Plata and 
Cordova, which are closely allied to and perhaps only racially 
distinct from C. azare, bark after the manner of foxes. On the 
whole, however, they are silent dogs in captiv ity, and, like the 
foxes, never succumb to the temptation of joiming in the howling 
concerts in which the dingos, jackals, prairie wolves, and laree 
wolves in the Gardens indulge. and lhe they seem unable to 
resist contributing to. Personally I believe that voice in mammals 
is often a good guide to affinity; and, in the present case, the 
voices of C. antarcticus and C. latrans bear out wy opinion of 
the relationship of these species to others, shown by structural 
characters. 

Finally, if the conclusions above put forward are correct, 
Huxley’s classification, expressed in par. 5 (p. 384), must be 
emended by transferring C. antarcticus to the lowest section of 


* The presence of this patch in some of the primitive Canide is well worth more 
attention than it has received. I do not know what it may mean, any more than I 
know what the pale area behind the shoulder, observable in many Canidx, both wild 
and domesticated, may mean. 


“WCIOVEODOUOVIVHA AO WTTIHLVd 


‘onoyud “Sg mM a 


UXT “iel Cll “S “A al 


ON AVIAN OSTEOLOGY. 393 


dogs, containing C. thous (=canerivorus) and vetulus, and by 
uniting his Sacaline and Lupine sections. Possibly these sections 
contain groups worthy of subgeneric, if not of generic, recog- 
nition: C. antarcticus and C. thous, for example, may be sub- 
generically or generically separated by the structure of the 
mandible; but I do not see how C. latrans is to be distinguished 
other than specifically from such forms as C. pallipes, anthus, or 
even lupus. 


30. On the Patella in the Phalacrocoracide. 
By Dr. R. W. Saurezpr, C.M.Z.S. 


[Received April 14, 1913: Read May 20, 1913.] 
(Plate LXL*) 


While recently employed in preparing a detailed account of the 
skeleton of Harris’s flightless Cormorant (Vannopterum harrisi), 
in which the osteology of that species is compared with that of 
a number of others of the family, I became interested in the 
morphology of the patellz of those birds. 

Many years ago J published a number of papers on the skeleton 
in the Cormorant, in some of which the patella of the Phala- 
crocoracide was referred to and figures given of it. But my 
material, at the time to which I refer, was very limited— in fact 
J think there were but the skeletons of one or two species of 
those birds at hand, and this included the skeletons of two or 
three young ones. 

In the higher groups of birds the patella, when present, is 
usually small, and offers but little of value to the avian taxonomist. 
This, however, is not the case when we come to examine into the 
osteology of some of the groups occupying lower positions in the 
system, and especially is this true of many of the Pygopodine 
forms and their allies near and remote. 

Owen paid but scant attention to this bone of the skeleton in 
Aves, devoting less than half a paragraph to it, thus :—‘‘ The 
chief of the sesamoid bones in the hind limb is the patella: it is 
of unusual size in the Penguin, is ossified from two centres, and 
articulates with the procnemial process of the tibia: it coexists 
with the long rotular process in the Loon, fig. 34,7; it is large 
and of an angular form in the Musk-duck (Biziwra): in the 
Merganser the patella is largest and deeply notched; in the Coot 
it iselongate. In most aer al birds a patella is wanting je hhere 
is no reference made here either to a Grebe or a Cormorant! and 


* For explanation of the Plate see p. 402. 

+ Shufeldt, R. W. “Osteology of the Cormorant,” Science, Dec. 7, 1883, p. 739 ; 
Feb. 8, 1884. vol. iii. No. 53, p. 143; ibid. Apr. 18, 1884, No. 63, pp. 474, 475. 

oe Gonter ning some of the forms assumed hy the Patella in Birds, » Proc. U.S. Nat. 
Mus. 1884, vii. pp. 324-331. Numerous text-figures. 

 Osteology of the Steganopodes,” Mem. Camegie Museum, Pittsburgh, Pa., Apr. 
1903, vol. 1. No. 3, Art. 3, pp. 15-70. Plates and many text-figures. 

t Owen, Richard. Comp. Anat. and Phys. of Vertebrates, vol. ii. p. 88, London 
1866. 


394 DR. R. W. SHUFELDT ON 


Professor Owen evidently considered the smal, scale-like bone 
found in the tendon of the extensor femoris, at its insertion, 
to be the patella in that bird. When I first examined this 
question, this was likewise my opinion, and in a paper on the 
patellee in birds, published in 1884 in the ‘ Proceedings of the 
United States National Museum,’ I so figured it (fig. A, p. 328) 
for Colymbus septentrionalis (Gavia stellata). This opinion was to 
some extent qualified later on, when I stated, with respect to the 
Loons (“ Urinatoride”) that they possess ‘‘ only a very small, 
flake-like sesamoid, which occurs in the tendon of the extensor 
femoris muscle at its insertion, and probably the true patella has 
coossified in the adult with the elongated cnemial process of the 
tibio-tarsus”*. Possibly some avian osteologist has published on 
this subject, but if so, | have not seen the work; and never 
having been so fortunate myself as to have come into possession 
of the skeleton of any Loon, secured at the right time to demon- 
strate the exact composition of the tibio-tarsus in that bird, I am 
still in doubt on the question. However, it is quite possible— 
indeed quite likely—that the true patella in Loons (Gavia) is, in 
the adult, completely codssified with the great elongate cnemial 
process of their tibio-tarsi. The moulding of the patella on the 
back of this process in Grebes, especially in very old birds, is 
wonderfully close,—so close in some ligamentous preparations as 
to deceive the eye upon casual examination. 

To settle this interesting poimt—if it has not as yet been 
settled—will require the examinations of the skeletons of Loons 
including those of individuals of the genus of all ages. 

At present [ am inclined to think that the patella in Gavia, 
in the adult, has been indistinguishably fused with the cnemial 
process of alg tibio-tarsus, for the reason that it is on the road to 
such a fate in the Grebes ( tolymbidee), and that in all such birds 
as Hesperornis—an ancient ancestor of the Loons—the Penguins, 
the Cormorants, and some others, the patella is very large. 

Then, finding it large in a Gr abe, one would naturally look for 
the same in eh forms as Loons, especially when one considers 
the relationship of these two families. 

The statements now being made are, in a way, prefatory, leading 
up to what I have to say on the patella of the Cormorants. It 
must be borne in mind in this connection that the patella in 
Penguins is very large, and grooved obliquely across the anterior 
face for the tendon of the ambiens muscle 7. 


* “Concerning the Taxonomy of the North American Pygopodes based upon their 
Osteology,” Jour. Anat. & Phys. London, Jan. 1892, pp. 199-208. The lines quoted 
are trom page 202. In this paper I also give two figures of the skeleton of the thigh 
and leg of a Grebe, in which the patella is included. One of these figures was repro- 
duced by Coues in his fifth edition of the ‘ Key,’ without acknowledgment (vol. 11, 
p. 1052, fig. 712). 

+ Coues, E. “Material for a Monograph of the Spheniscide,” Proc. Acad. Nat, 
Sci. Phila. xxiv. 1872. 

Watson, Morrison. “Anatomy of the Spheniscidie” (Rep. Scient. Results of 
Vovage of H.M.S. ‘ Challenger,’ Zoology, vol. vii. pl. vii. figs. 9 & 10, 1883). 

Shuteldt, R. W. Proc. U.S. Nat. Mus. 1884, p. 326, fig. 1. Reproduces from Watson 
figures of patellze of Aptenodytes pennantii and Ludyptes chrysocome. Comments 


AVIAN OSTEOLOGY. 395 


Personally I have not examined the patella in Hesperornis with 
the view of substantiating Marsh’s statement that it is ‘‘ perforated 
by a large foramen for the tendon of the ambiens muscle, agreeing 
in this respect with the patella of the Gannet (Sula bassana 
Briss.).” His several figures of the patella of Hesperornis regalis 
do not show this “large foramen” (‘Odontornithes,’ p. 23, 
plates xv. and xx. figs. 1-3), while I have yet to see the patella 
of Sula bassana presenting any such perforation for the ambiens 
muscle as Marsh describes. 

However this may be, the fact remains that very radical dif- 
ferences exist with respect to the fate and disposition of the 
patella and cnemial process of the tibio-tarsus in Grebes, Loons, 
Hesperornis, Penguins, Gannets, Cormorants, and many other 
different species of water birds. Great differences also exist in 
the morphology of this sesamoid and the aforesaid apophysis. 

This brings us to a point where the patella of the Phalacro- 
coracidee may be discussed. For this purpose I have before me 
the skeletons of several species of Cormorants, all belonging to 
the Collection of the U.S. National Museum, to which institution 
I am indebted for the loan of them. It will not be necessary to 
name these species here or to give their museum numbers, as this 
information is fully set forth in the explanation of Plate LXI., 
which appears at the end of this article. As to the 19 figures on 
the plate, they are reproduced, natural size, from my own grouping 
and photograph,—the latter having been made direct from the 
specimens. 

Among avian osteologists the general opinion has prevailed that 
the patella in Cormorants is a big trihedral one, in some way 
“perforated” for the ambiens muscle. 

Garrod found the ambiens muscle present in the family Phala- 
crocoracide ; and, in noting that Meckel had not done so, he states 
that “it is peculiar in that [in the Cormorant] it runs through 
the substance of the large triangular patella, in a bony canal” *. 

In another paper this gifted ornithotomist is still more positive, 
and in speaking of the “‘ Steganopods,” he says: ‘“ The ambiens is 
of fair size; it deeply grooves the large ossified patella ; and some 
of the fibrous ligament overlapping this groove shows traces of 
ossification ; so that in aged birds the groove may be converted 
into a foramen, as is always the case in Phalacrocorax, where the 


made thereon. Thereisacurious slip in this paper with respect to the description of 
the patella of the Gannet (Sula bassana). On page 362 I state that “ Professor Marsh 
tells us that the patella of Sula is pertorated by a large foramen for the passage of 
the tendon of the ambiens muscle.” On pages 329 and 330 I quote the entire para- 
graph from Marsh’s ‘ Odontornithes’ (p. 93) where he gives this description of the 
patella of Sula and states on page 327 that “I fail to find any such foramen in the 
patella of the specimen of Suda before me, although it has a shallow, oblique groove 
across its anterior face that seems to correspond with the one described when speaking 
of the patella of the Penguin.” This statement is supported by a correct figure of 
the patella in Sula bassana (No. 16643, Coll. U.S. Nat. Mus.), and I must believe 
that the description left us by Prof. Marsh is another of those curious slips of his m 
the osteology of modern birds. 

* Garrod, A. H. “On Certain Muscles of the Thigh of Birds, and on their value 
in Classification,” Pt. I, P. Z, 8. 1873, pp. 636, 637 (footnote). 


396 DR. R. W. SHUFELDT ON 


thus formed foramen is far from superficial” *. This is a very 
interesting statement, and will be commented upon further on in 
this paper. 

Forbes, in describing the anatomy of Biziwra, makes the state- 
ment that “The ambiens muscle is large, and peculiar in that its 
tendon perforates the large-sized triangular patella, just as it does 
in Phalacrocorax and the extinct Hesperornis” y. 

When referring to this sesamoid in the Cormorants, Coues says, 
in the fifth edition of his ¢ Key’ (p. 961), that “ There is a bulky, 
free patella, coexistent with a short cnemial apophysis or rotular 
process of tibia, but perfectly distinct therefrom, as in Grebes.” 
This somewhat ambiguous description is made clearer by an illus- 
tration of the patella in ‘“‘ Phalacrocorax bicristatus,’ which he 
borrowed from a previous paper of mine. Nothing is said with 
regard to the ambiens or the “ perforation ” in the patella, so this 
note is quite useless for present purposes. 

From my own various accounts of the patella in Cormorants 
I select the following as best suited to throw light upon the 
facts I propose to here set forth. 

In my above-cited article on the patella in birds (Proe. U.S. 
Nat. Mus. 1884, p. 330, fig. 7) IT remark: “In No. 41 of ‘Science,’ 
I presented a lateral view of the leg-bones of a Cormorant (Phala- 
crocorax), Showing the form of the patella in these birds. The 
Same specimen is given here, only an anterior view is chosen 
instead of the lateral one. 

‘“This form is a particularly interesting one, and I am not 
aware of any bird at present that can show a similar condition of 
the parts in question. 

‘On the face of it, it almost looks as if a patella had developed 
of a size equal to the rotular process, and subsequently the two 
became thoroughly united, and formed one large patella, articu- 
lating as shown in the drawings I have made of it. Ido not say 
that this is actually the case, nor were the young of this spe- 
cimen, which also belong to the collections at the Smithsonian 
Institution, of a proper age to determine exactly the manner in 
which this great bulky patella was developed.” 

Apart from the suspicion expressed in this last paragraph, 
which will be put into more exact terms further on, there are but 
two opinions extant, at present,in regard to the development of 
the patella in the Phalacrocoracide. They are the opinions of 
Garrod and of Forbes, and have been set forth in their own words 
above. 

There is but one construction that can be placed upon Forbes’s 
view of a Cormorant’s patella, and that view cannot be success- 
fully defended. It will be noted above that, in his calling 


* Garrod, A. H. “Notes on the Anatomy of Plotus anhinaa,” P. Z.S. 1876, 
p. 340, pl. xxviii. figs. 5, 6. The figures give the patella, indicating the position of 
the foramen. 

+ Forbes, Wm. Alex. “ Note on some points in the Anatomy of an Australian 
Duck (Biziura lobata),” P. Z. S. 1882, p. 457. 

I regret to say that I have never had the opportunity of examining the patella in 
this duck, and I doubt if we have a skeleton of it in the United States. 


AVIAN OSTEOLUGY. 397 


attention to the fact that the tendon of the ambiens muscle 
passes through the patella in Cormorants, he uses the word 
“* nerforates ” "(emphasizing it by the use of italics). Surely 
could not have meant that the tendon really did “ perforate ” 
bore its way through the patella as the Cormorant grew sail 
developed? And, after finding its way through the “bone (or 
cartilage) that it duly made attachment at its point of insertion ¢ 
No such thing has ever occurred in anatomical development, and 
Forbes was altogether too good an anatomist to have set up any 
such ridiculous explanation. This being true, and disregarding 
entirely any such theory or such another possible suggestion as 
that, in subadult life, the patella of a Cormorant, forming first 
in elementary cartilage, does, when in that condition, surround 
the tendon of the ambiens muscle, and afterwards, when the 
patella has completely ossified in old birds, the aforesaid tendon 
runs through a foramen, which passes through the middle of 
it. Barring this, too, as I say, as being a totally untenable sug- 
gestion, we are left with but two other possible ways by means of 
which the ambiens comes to pass through the foraminal canal in 
the patella of Cormorants. 

Tn discussing these, we are at once confronted with Garrod’s 
opinion in regard to it—an opinion which had never, apparently, 
occurred to Forbes. Garrod took the view that, inasmuch as 
in anhinga (Plotus) the anterior face of the patella is somewhat 
deeply grooved, and as ligament stretches across that groove to 
confine ‘the tendon of the ambiens muscle which passes through 
it, and as this same ligament sometimes ‘“‘ shows traces of ossi- 
fication,’—hence in Cormorants this also takes place, and the 
fibrous ligament in them becomes, in old individuals, thoroughly 
ossified, and we thus find the tendon of the ambiens passing 
through the bone. 

Such an opinion will not hold for an instant in the face of the 
necessary material to examine into it. 

Now in some Cormorants the patella is very large and thick, 
and the aforesaid foraminal passage, passing through it from side 
to side, is found but slightly in advance of the middle and above 
the centre of the bone; so that, had it been a “groove” in the 
young bird spanned by fibrous ligament, and this ligament sub- 
sequently ossified in the adult, that groove must have been a very 
deep one, and the patella in old birds would, through its form 
alone, after ossification was complete, exhibit the manner as to 
how it had come about. Moreover, in connection with this it is 
very clear that, were the foramen in the patella in Cormorants 
formed by a groove in front of it being covered over by fibrous 
ligament in the subadult bird, which ligament later in life ossified, 
that ossification would be smooth on its anterior face, which is by 
no means the case, as one may see by a study of the figures in the 
plate accompanying this paper. For instance, a fibrous ligament, 
stretching across a deep, narrow groove as a retaining band for 
the tendon of a muscle, would not, in ossifying, take on any other 
shape beyond that possessed by the flat ligamentous band; for 


398 DR. R. W. SHUFELDT ON 


instance, as I say, such as has occurred in the patella shown in: 
Pl. LXI. fig. 11 and others. It must stand to reason, then, that 
Garrod entertained an untenable opinion in this matter. 

Coming to the second way in which ‘this foramen could have 
been formed, it is clear that, in young and subadult individuals, 
the tendon of the ambiens muscle could be confined by an 
additional piece of bone or cartilage being placed in front of it, 
and this added part subsequently ossify and not only form, in 
the adult, a foraminal passage for the aforesaid muscle or its 
tendon, but largely add to the bulk of the patella. It might, too, 
—from the thoroughness of the coéssification—cause the atrophy 
of the muscle and its tendon at that part of the course of the 
latter across the front of the true patella; and such a result would 
be brought about by the entire sealing up of the foraminal 
passage, which not only actually takes place in the patelle of 
some Cormorants, but again proves Garrod to have been wrong 
when he stated that such a foramen or foraminal passage was 
always present in the patella of Cormorants. See Pl. LXI. 
fig. 3—-Phalacrocorax penicillatus—where it has been sealed up 
entirely, leaving not so much as a trace or a suspicion of its ever 
having been present there. 

In my opinion, the foraminal passage for the ambiens muscle, 
passing transversely through the patella in the Phalacrocoracide, 
when present, 1s formed as set forth below, which formation can 
be demonstrated by the material figured in the plate accompanying 
this paper. When the foraminal passage is entirely absent it 
has been absorbed, atrophy of the ambiens probably having 
ensued. 

It would appear that in young Cormorants the ambiens passes 
in a groove, of the required depth only, obliquely across the 
anterior face of the patella, which latter is very closely adpressed 
against the posterior surface of the long enemial process of the 
tibio-tarsus,—a condition that persists throughout life in Grebes 
(Colymbide). Codssification between the patella and the upper 
two-thirds of the cnemial process of the tibio-tarsus, in the case 
of the growing Cormorant, next sets in, which, owing to the 
morphology of the parts involved, would, in time, depreciate 
the action of the knee-joint. This is clear when we come to 
consider the origins and insertions of the various muscles about 
the knee-joint anteriorly, posteriorly, and laterally *. These, 
during their continued action in locomotion—especially during 
the act of swimming—tend to overcome the aforesaid danger, 
militating against the complete freedom of action of the knee- 
joint. Codssification steadily proceeds ; the individual grows ; a 
constant tugging is exerted during locomotion at the tibio-tarsal 
enemial apophysis. This eventually results in its becoming 
dissociated from the bone to which it belongs along a transverse 
line, at a short distance above the level summit of the tibio- 
tarsal shaft, thus leaving the lower third of the cnemial 


* Shufeldt, R. W. ‘The Myology of the Raven,’ p. 187, figs. 51-53, London 
1890. 


AVIAN OSTEOLOGY. 399 


‘process where it occurs in all birds when such an apophysis is 
present *. 

In the Phalacrocoracide, then, the patella is a compound bone, 
consisting of the true patella posteriorly, and the upper two- 
thirds, more or less, of the cnemial process of the tibio-tarsus 
anteriorly, which latter has become dissociated and thoroughly 
codssified with the former. By this union, a foraminal passage 
is formed for the ambiens muscle, and this, in some species, may 
gradually, but entirely, disappear, whereupon, in time, the muscle 
meets its usual fate. 

For very obvious reasons, the patella is very large in Harris’s 
flightless Cormorant (Vannopterum harrisi); indeed, in so far as 
I have been enabled to discover, itis larger in this species than in 
any other existing form of the entire family (Pl. LXI. figs. 1 & 2). 
It has an average height of 25 millimetres, the longest diameter 
of its base being 19 or 20 millimetres, and is nearly a square in 
outline. Posteriorly, it is flat and triangular, the acute angle 
being above. Above its middle, and nearer its posterior surface 
than its anterior margin, it is pierced, transversely, by the foramen 
for the ambiens, which is of considerable size. Externally, its 
exit is almost flush with the surface of the bone, while on the 
inner aspect it is situated at the base of an extensive concavity. 
Anteriorly, this compound bone exhibits a uniform longitudinal 
excavation which, when the patella is articulated as in life, is the 
continuation with the similar excavation between the pro- and 
ectocnemial processes of the tibio-tarsus. On the anterior face 
of this patella, at the lower-internal angle, there is a transverse, 
triangular facet which is intended for articulation with the 
superior border of the cnemial crest of the tibio-tarsus. In fig. 2 
we have the mesial or inner aspect of the left patella of 
NV. harrisi, from the same skeleton (No. 19719), where the 
perforating foramen for the ambiens is seen at the base of the 
aforesaid extensive concavity. A study of these two patelle 
reveals the nature of the composition of the bone as a whole, as 
described above. 

Phalacrocorax penicillatus possesses only a medium - sized 
patella, which is elongate and wedge-shaped. It has the general 
form of the patella in WVannopterum: but the transverse fora- 
minal passage for the ambiens has been entirely absorbed,—not 
a vestige of it remaining in an individual of the age to which 
this patella belonged. The union of the two parts composing the 
bone is very complete, and all traces of their origin have been 
entirely obliterated,—that is, beyond the anterior rotular channel, 
and the here extensive facet below it for articulation with the 
cnemial process of the tibio-tarsus. 

This is a most excellent example of the patella in a Cormorant 

* This is the part referred to by Coues in his ‘Key’ (5th ed. vol. ii. p. 961), 
where he describes the patella in the Phalacrocoracide in the following words: 
“There is a bulky free patella, coexistent with a short cnemial apophysis or rotular 
process of tibia, but perfectly distinct therefrom, as in Grebes.” It is very evident 


from this definition that Coues had never given the patella of a Cormorant any very 
close study, 


AOO DR. R. W. SHUFELDT ON 


showing the conditions described as they occur in a very old 
bird (figs. 3 & 4), and they are equally well shown in the patella 
of P. punctatus (figs. 5 & 6), where the foraminal passage has 
likewise entirely disappeared. The rotular channel in front is 
very narrow and pointed above, as in a Grebe or Loon, and there 
is a rounded notch on the superior border of the cnemial process 
of the tibio-tarsus, which marks the limitation, externally, of the 
facet for articulation with the patella on that border (fig. 6, 
where this “notch” is plainly seen). This patella gives barely 
any hint as to the two parts of which it is composed. 

This is to a less degree true of the patella of Phalacrocorax 
magellanicus (figs. 7 & 8); for here again we find the foraminal 
passage in its very last stages of ultimate absorption, while the 
continuation of the rotularl channel on the anterior face of the 
patella, and the evidence of this part of the patella once having 
belonged to the tibio-tarsus, is very complete (fig. 7, viiol 
should be compared with fig. 6, the former in no way recalling 
the form of these bones in the Grebe). 

Phalacrocorax urile (Nos. 19655 & 18982) has, in old individuals, 
a rather bulky patella (figs. 9, 10, & 11), in which, in the 
specimens selected, the foraminal passage is reduced to capillary 
proportions, and the sutural traces of the elements composing 
it have become nearly obliterated. Fig. 10, which is from the 
right limb of P. wrile, No. 18982 of ale Wollleetion of the U.S. 
National Museum, shows the minute entrance to the foraminal 

assage; while in fig. 11, it being the left limb from the same 

skeleton, the opposite opening is in view,— posterior to two 
other foramina which occur here. Fig. 11 is given on the plate, 
to the exclusion of the patella of Phalacrocor ax carbo, for the 
reason that in no Cormorant, other than P. wrile, do we find, 
on this anterior view, a better example of the interenemial 
channel on both the big sesamoid and the tibio-tarsus, and the 
line between them where they eventually parted company. 

Phalacrocorax carbo (No. 18850, Coll. U.S. Nat. Mus., not 
figured) has a patella that is an interesting one in several 
particulars. On its imner aspect the surface is flat and smooth, 
the minute opening of the almost entirely absorbed foraminal 
canal being situated far posteriorly upon it. In size and general 
form the patella of this Cormorant is much like the one shown 
in figs. 18 & 19 of P. pelagicus; while it likewise possesses 
characters peculiarly its own. It has the usual trihedral form 
and makes rather more than the average articulation with 
the tibio-tarsus in extent, especially on the proximal margin of 
the procnemial process. Its posterior face is triangular and flat, 
while the external one is pierced near its middle by the outer 
opening of the foraminal passage. Anteriorly, the usual inter- 
enemial channel is present, it being continuous with the same on 
the tibio-tarsus. On the patella, it is unusually narrow, and 
faces very much to the outer side. 

Phalacrocorax carbo (No. 18851) has the patella almost 
exactly like that in P. uwrile (figs. 10 & 11.) The former, 


AVIAN OSTEOLOGY. 40] 


however, in this particular individual, has completely lost the 
foraminal passage, while otherwise the morphology is quite the 
same. 

One of the most interesting forms of the Patella among 
Cormorants is found in Phalacrocoraa vigua (figs. 12 & 13); for 
it is not only very small and chunky, but the facet on its 
infero-anterior aspect for the tibio-tarsus is relatively, as well as 
actually, large. On the inner aspect, the big foraminal opening 
is situated at the base of a fossa, as in No ‘annopterum harris, 
while on the external side it is sical again, and has, leading 
down from it to the lower border of the bone, a deep groove, 
which not only indicates the suture between the two elements 
composing this bone, but apparently the continuation of the 
groove for the ambiens as well. 

Passing to Phalacrocorax auritus (No. 19262), the patella of 
which species is shown upon two views in figs. 14 & 15, we find 
that that sesamoid is rather small for a Cormorant of its size. 
In it, the foraminal passage has gone entirely, while the bone, 
upon the other hand, shows better than any other Cormorant’s 
patella [ have examined, the two parts of which it is composed. 
These are very plainly to be seen in the figures,—indeed, equally 
as well as in the specimens themselves. 

There is a large patella in Phalacrocorax albiventris (No. 18487, 
figs. 16 & 17), where again we find, in the adult bird, ‘ihe 
foraminal passage almost obliterated, having been reduced to a 
capillary calibre throughout. Sutural traces between the bone 
are faint, while anteriorly the rotular excavation is broad and 
deep, being but slightly wider at the top than it is at the distal 
border. 

A most instructive patella is found in Phalacrocorax pelagicus 
(No. 19032, figs. 18 & 19). Once more we find the foraminal 
passage reduced to a very small calibre, though the entrances on 
the inner and outer surfaces of the bone are still in evidence, 
though very small. What is to be specially noted among other 
things is, however, the position of the foraminal aperture on 
the inner aspect (fig. 18); this is but 4 millimetres from the 
posterior margin of the patella, while it is 9 millimetres from 
the anterior margin, which is sufficient to render the theory of 
the “anterior” groove being filled in by an ossified fibrous liga- 
ment—oridieulous! 

In its amalgamation and coossification of its parts, this patella 
became to a large degree twisted,—-a twisting that cannot well 
be appreciated in the ficures. Still, the bone fits most poeetcly 
on the superior border of the cnemial process of the tibio-tarsus 
its lower border, antero-posteriorly, being no thicker than is that 
of the leg- pone, while everything in “its morphology plainly 
indicates its origin. On the front of the patella, the rotular 
channel is soune 0 shallower than usual, and is as broad above 
as it is below, where it, in all particulars, directly continues the 
corresponding channel separating the pro- and _ ectocnemial 
apophyses of the tibio-tarsus. 


402 ON AVIAN OSTEOLOGY. 


The obliquity of the foraminal passage for the ambiens is 
easily made out, its higher aperture being on the inner aspect of 
the bone, and situated, as I have pointed out above, far back 
from the antero-internal border of the bone. To best appreciate 
the fact that this sesamoid in P. pelagicus is formed as I have 
demonstrated it to be in Cormorants of all species, it should be 
viewed from above rather than from either side or in front. 


EXPLANATION OF PLATE LXI. 


[All the figures are reproductions of photographs made by the author direct from 
the specimens. Each bone is of natural size, and from an adult individual. They 
are from skeletons in the Collections of the U.S. National Museum, the Museum 
numbers of which are given under the figures. | 


Fig. 1. Right patella of Nannopterum harrisi; antero-lateral aspect. (No. 
19719.) 

. Left patella of NV. harrisi; mesial or inner aspect. (No. 19719.) 

. Right patella of Phalacrocorax penicillatus, antero-lateral aspect, or the 
same view in which fig. 1 is shown. (No. 18535.) Note that the 
foraminal passage for the ambiens has been entirely absorbed. 

. Left patella of P. penicillatus ; inner aspect. (No. 18535.) 

. Left patella of Phalacrocorax punctatus; inner aspect. (No. 18282.) 

. Left tibio-tarsus of P. punctatus ; anterior view, with the patella articulated 
in situ. (No. 18282.) 

. Right tibio-tarsus, anterior view, of Phalacrocorax magellanicus, with 
patella articulated in situ. (No. 18438.) 

. Left patella of P. magellanicus; inner surface. (No. 18438.) Note that 
the exit of the foraminal passage points to the fact that it is im the 
very last stages of its ultimate disappearance. This is an interesting 
link in the chain of evidence on the real composition of this bone in the 
Phalacrocoracide. 

9. Proximal extremity of tibio-tarsus and fibula of right pelvic limb of 
Phalacrocorax urile; adult. (No. 19655.) Outer aspect, with patella 
and femur articulated in sitz. 

10. Inner aspect of the bones composing the knee-joint of P. wrile; partly 
ligamentous. (No. 18982.) Right pelvic limb. 

11. Left patella, tibio-tarsus and fibula (superior moieties), anterior aspect ; 
left pelvic limb of P. wrile. (No. 18982.) Note the complete union 
and the continuation of the rotular channel of the tibio-tarsus on the 
anterior face of the patella. 

12. Right patella of Phalacrocorax vigua; outer surface. (No. 18479.) 
Note groove leading down from the foraminal opening. 

13. Left tibio-tarsus of P. vigua (No. 18479), with patella articulating 
im situ. 

14. Bones of the knee-joint of Phalacrocorax auritus (“P. dilophus,” 
No. 19262.) Right pelvic limb, inner aspect. Femur somewhat 
displaced, but patella properly articulated. 

15. Bones of the knee-joint of P. awritus (No. 19262) ; left pelvic limb, outer 
aspect and articulated in situ. Note the line of demarcation between 
the true patella and the anterior part of the bone, which originally 
belonged to the tibio-tarsus. 

16. Anterior aspect of the proximal extremities of the left tibio-tarsus and 
fibula of Phalacrocorax albiventris, with the patella duly articulated 
above them. (No. 18437.) The rotular channel is very broad in this 
species and rounded above. 

17. Right patella, inner surface of P. albiventris. (No. 18437.) The foramen 
is almost closed up. 

18. Bones entering the knee-joint of Phalacrocorax pelagicus, adult, nat. size. 
Right pelvic limb, outer aspect. Ligamentous preparation with femur 
slightly luxated. Patella in situ. (No. 19032.) 

19. Skeleton of the knee-joint of P. pelagicus. (No. 19082.) Left limb, 
outer aspect; patella i situ. 


do bo 


co sn DO 


ON THE METAMORPHOSIS OF THE AXOLOTL. 403 


31. Experiments on the Metamorphosis of the Mexican 
Axolotl (Amblystoma tigrinum), conducted in the 
Society’s Gardens. By E. G. BouLEncsEr, F.Z.S., 
Curator of Reptiles. 

| Received and Read May 20,1913. ] 


(Text-figures 75 & 76.) 


During the past year I have been experimenting on points relat- 
ing to the metamorphosis of the Mexican Axolotl (Amblystoma 
tigrinum), and have succeeded in obtaining the transformation of 
a number of specimens. ‘This paper deals with the methods 
employed, and gives a detailed account of the external changes 
undergone by the animal during the process, a subject which, 
with the exception of Duméril’s (1) brief account, has not, so far 
as I am aware, been previously treated. 

Before describing my experiments, it may not be out of place 
to give a short account of the history of the animal. Although 
suspected by Cuvier to be but the larva of some unknown air- 
breathing salamander or newt, the Axolotl of Mexico was 
considered for many years to be one of the Perennibranchiata, 
such as Proteus, Vecturus, and Siren. In the year 1863 a number 
of specimens were imported from Mexico to France, where some, 
kept in the Jardin des Plantes in Paris, bred, and the young were 
successfully reared. The conclusion was not unnaturally drawn, 
that the Axolotl, having bred in the branchiate condition, could 
not possibly be anything but a perfect aquatic animal. It was, 
not, however, until over two years later that the subject assumed 
a different aspect, for some individuals of the second generation 
lost their gillsand the dermal folds of the back and tail, developed 
eyelids, and yellow spots on the skin, and taking to land, changed 
into a land salamander, already well known from North America 
as Amblystoma tigrinum. 

A few years later Weismann (2) tried to solve the question 
as to whether it were possible to force the larve, if brought into 
conditions which rendered the use of the gills difficult and that 
of the lungs easy, to change into Amblystomes, and he therefore 
experimented with several broods which were placed in shallow 
water, and thus compelled to breathe air more frequently. 
Although he met with no success Weismann was not discouraged, 
and came tothe conclusion that the failure of his experiments was 
due to his having been unable to bestow the necessary care and 
attention on the animals: he therefore asked Mlle. de Chauvin, 
a lady who had already done much careful experimental work, 
to take a number of larve, just out of the egg, rear them, and 
make an attempt to bring them to the perfect condition. When 
the Axolotls were about 6 months old, Mlle. de Chauvin placed 


Proc. Zoo. Soc.—1913, No. XX VIII. 28 


404 MR. E. G. BOULENGER ON THE 


them in large glass vessels which were so disposed and the 
water so restricted that at one spot only could they dive quite 
under, while everywhere else they came into contact with the 
air. The water was then gradually reduced. Within a few 
days a change took place, the creatures leaving the water in from 
four to fourteen days, the complete metamorphosis following 
about ten days later. Mlle. de Chauvin (3) summarized her results 
as follows :—‘ From what I have said, the correctness of the view 
suggested by Weismann must be established, namely that most 
Axolotl larvee, if not all, complete their metamorphosis, if in the 
first place they come out of the ege healthy and are properly 
fed, and in the second place meet with arrangements which 
force them to change from breathing under water to breathing 
above water.” 

Dr. J. H. Powers (4) at Doane College, Nebraska, has more 
recently conducted numerous experiments on the metamorphosis 
of Nerth American examples of the Axolotl, and he hes come 
to the conclusion that the metamorphosis is not due, as was 
thought by Mlle. de Chauvin, to a direct response to changes in 
conditions of environment, compelling them to resort to aerial 
respiration, but to checked nutrition, and that a careful study of 
Mlle. de Chauvin’s methods and results seems to cast a doubt 
upon the conclusion that enforced air-breathing caused the 
metamorphosis. The following is a passage from Dr. Powers’s 
paper on the subject of this lady’s experiments :—‘‘ Fearing that 
her charges would die, as indeed they sometimes did, she 
always prepared them for the trying ordeal of metamorphosis by 
raising the temperature of the water in which they were kept 
and feeding to the maximum for several days, to which she 
ascribes no other importance than giving the animals increased 
strength. The Axolotls were then brought immediately into 
water sufficiently shallow as to force them, at least part of the 
time, to breathe air. In this latter condition the experimenter 
complains again and again that it was next to impossible to 
induce the Axolotls to take any food whatever. Thus in these 
experiments we have high feeding followed by practical starva- 
tion, and it seems that no control experiments were instituted to 
determine what the effects of over and under nutrition might 
have been with Axolot!s still in abundance of water. Yet most 
interesting is it to note that even the varying factors of nutrition 
seem to have been wholly neglected in the final interpretation of 
the results.” 

Dr. Powers’s field-notes show that metamorphosis occurs 
rarely, if ever, as the result of enforced air-breathing through 
the drying up of ponds, and that in spite of repeated search at 
appropriate times and places, no Axolotls have been found trans- 
forming on the mud of drying ponds. 

Dr. Gadow (5), who not long ago visited the lakes near Mexico 
City in which this creature lives, and where it is said to retain its 
branchiate condition, has been able to refute the theories framed 


METAMORPHOSIS OF THE AXOLOTL. 405 


by various zoologists, as to why the Axolotl does not transform 
in those localities. The reason he gives is that the unfailing 
abundance of food and water, and the immerse hiding- places 
amongst the reeds under the banks, constitute fon these 
Batrachians a real paradise where they remain, in spite of the fact 
that there is nothing to prevent them from leaving the water. 
Dr. Gadow’s explanation appears to be in keeping with the results 
of both Mlle. de Chauvin’s and Dr. Powers’s experiments. 

The conflicting conclusions arrived at made it highly desirable 
that further experiments should be undertaken. The lack of 
success obtained by the majority of those who have attempted to 
force the Axolotl to transform, has led many zoologists to believe 
that the change is due to some congenital disposition, possessed 
only by certain individuals, and that Mlle. de Chauvin and the 
few others who have succeeded in obtaining Amblystomes from 
the larval form were specially fortunate in the choice of their 
subjects. Therefore, when I commenced my experiments, I was 
by no means sanguine of attaining any definite results. 

In August of last year I obtained five Axolotls with very fully 
developed gills and fins, ranging in length from 105 mm. to 
138 mm., and therefore probably from six to nine months old. 

The enforced breathing of air, by the gradual absorption of 
the water, and by the gradual elimination of the necessary amount 
of oxygen from the water, starvation and irregular feeding, and 
increased temperature having all been given or suggested as 
means of obtaining the metamorphosis, sr placed the animals 
under the following conditions :— 


Nes. 1 & 2.—In shallow water so that after the first week 
only the top of the animal’s head and fins remained un- 
covered, thus forcing the creature to make free use of its 
lungs. These I kept at a uniform temperature of between 
75° and 80°. 

Nos. 3 & 4.—Under similar conditions, but at a uniform 
temperature of between 55° and 60°. 

No. 5.—In deep water from which the necessary amount of 
oxygen was eliminated by the gradual substitution of 
boiled water and by the introduction of decomposing 
vegetable matter. 


Asat the time I could not obtain any more Axolotls of the size 
required, I decided to abstain from experimenting with the 
feeding, until I had either obtained negative results with those I 
was about to experiment on or had secured more suitable spe- 
cimens. Nearly a month later I was fortunate enough to obtain 
six further suitable specimens. At the same time some of the 
first brood showed signs of metamorphosing in spite of the fact 
that they had fed with the greatest regularity. I therefore decided 
not to abstain from offering them food as success seemed likely 
without resorting to starvation, and placed the individuals of 
the second brood, which shall be referred to as numbers 6, 7, 8, 9, 

28* 


406 MR. E. G. BOULENGER ON THE 


Text-fig. 75. 


STAGE f/f 


STAGE 2 


elt ot 


STAGE 3 


STAGE 4 


First stages in the metamorphosis of Amblystoma tigrinum. 


METAMORPHOSIS OF THE AXOLOTL. 407 


Text-fig. 76. 


Z 
Ss 
ee ce STAGE 5 
/| nt 
Ih ‘ 
Ss Ze 
STAGE 6 


pee, SINGS 7 


STAGE 8 


Final stages in the metamorphosis of Ainbiystoma tigrinum. 


408 MR. FE. G. BOULENGER ON THE 


10, and 11, under similar conditions to those of brood 1. Nos. 6 
and 7 were forced to breathe air, being placed in shallow water, 
at a high temperature; Nos. 8, 9, and 10 were placed under the 
same conditions at a normal temperature, and No. 11 was placed 
along with No. 5 in foul water, from which the oxygen was 
removed by the addition of boiled water. In all, therefore, I 
had eleven specimens under artificial conditions with the object 
of forcing the metamorphosis, and as a result succeeded in 
bringing six individuals to transform into the Amblystome 
stage, all six specimens being those induced to make free use of 
their lungs by being placed in shallow water; three of these were 
kept at a temperature of between 75° and 80°, three at between 
55° and 60°. The time required for the entire metamorphosis 
was from 12 to 16 weeks, periods which all to a remarkable degree 
exceeded those taken by Mlle. de Chauvin’s specimens. ‘Three in- 
dividuals kept under identical conditions died in from 33 months 
to over 5 months, but not until after having undergone a change, 
which, although considerable, did not necessarily suggest, as will 
be referred to later, that, but for their death, the entire metamor- 
phosis would have taken place. The two specimens kept in deep 
water from which the oxygen was eliminated died in the one 
case after 19 weeks, in the other after 22 weeks. These speci- 
mens, but for a slight reduction in the size of their gills, did not 
undergo any change, the fins being at the time of their death as 
well developed as at the commencement of the experiment. It 
is worthy of note that these specimens, in spite of the fact that 
they must have felt exceedingly uncomfortable, did not rise to 
the surface for air, even just prior to their death, more frequently 
than individuals kept in oxygenated water. Another point of 
interest lies in the fact that both specimens grew rapidly during 
this period, No. 5 from 136 mm. to 155 mm., No. 11 from 
112 mm. to 120mm. The specimens kept in shallow water, in 
the case of those that died, remained stationary, while those that 
transrormed actually decreased im size. 

Text-figures 75 and 76 (pp. 406 and 407) represent eight stages 
through which my Axolotls passed in the course of their 
metamorphosis. 

Stage 1 represents the perfect larval form, the condition which 
all eleven were in before being placed in shallow water. A few 
days after the animals had been placed in shallow water, the gills 
with their fringes began to shrink, and the dorso-caudal fin 
which lopped over to one side, to the left side in all my specimens, 
became reduced in size, the change being plainly visible in from 
a fortnight to three weeks (Stage 2). 

In from 10 to 20 days later the gills were only half their 
normal size, while the fin of the back was represented by a mere 
ridge ; that of the tail, although having undergone further reduc- 
tion, was still well developed on the upper surface and flopped 
over to one side; on the lower surface, however, it showed con- 
siderable reduction (Stage 3). It was not, however, until from 
five to seven weeks later (Stage 4) that the metamorphosis proper 


METAMORPHOSIS OF THE AXOLOTL. 4.09 


took place, when the part of the dorsal fin nearest to the head 
began to be entirely absorbed ; at the same time the head became 
shorter, assuming a more Salamander-like shape, while a swelling 
above the eye foreshadowed the advent of the upper eyelid; the 
gills at this stage measured but from 3 to 5 mm.; the fin of the 
upper edge of the tail, although now much reduced, still lopped 
over to one side; the lower fin, but for a rudiment at the extreme 
end, had entirely disappeared. 

Stage 5 represents the conditions seven to eight days later, 
when the dorsal fin had retreated to the middle of the back; 
the caudal fin on the lower surface had entirely disappeared ; the 
head had quite altered in aspect, having become much shorter, 
giving the eyes the impression of having been shifted forwards. 
At this stage the upper eyelid had become completely formed, 
the lower eyelid being incomplete. 

In from seven to nine days later (Stage 6), the fin of the back 
had receded to a level with the hind limbs, while on the upper 
surface of the tail, where it still lopped over, it measured only 
1 mm. Both eyelids were now formed. At this stage the 
animals raised their heads completely out of the water, and 
the opportunities they were given of getting on land were 
taken advantage of. Stage 7 represents the creatures about a 
week later, when nothing remained of the fin but a mere rudi- 
ment bordering the extreme end of the upper surface of the 
tail. The gills were represented by three knobs. Stage 8, the 
perfect form, the gill slits having closed up and the tail having 
become more or less roundish, was attained five to eight days 
later. In the case of both Duméril’s and Mlle. de Chauvin’s 
specimens the yellow spots appeared several days prior to the 
complete metamorphosis; in all my specimens, however, it was 
not until after they had attained the perfect Amblystome con- 
dition that the yellow spots put in an appearance, these being 
situated on the sides of the body, tail, limbs, and throat; none 
appeared on the back. 

As I was desirous of ascertaining whether the metamorphosis 
could be checked and the shrinking gills and fins made to undergo 
fresh development, and if so, at what stage it could no longer be 
checked, two specimens were taken out of the shallow water and 
replaced in deep water as they reached successive stages in their 
development. No.6 was kept ata temperature of between 75° and 
80°, and No. 9 at a temperature of between 55° and 60°. Both 
these Axolotls were placed in shallow water and then induced to 
breathe air frequently by means of their lungs on October 7th. 
Stage 2 was reached on October 20th; they were then replaced 
in deep water. The gills grew immediately and the fins developed, 
Stage 1 being reached again on October 28th and November 5th. 
They were then once more put into shallow vessels, with the 
result that the gills and fins again began to shrink, Stage 2 being 
attained on November 15th and 20th, Stage 3 on December 5th 
and 26th. On the latter dates they were again transferred to 
deep water, Stage 2 being attained on December 20th and 
January 10th. The metamorphosis was continued on placing the 


* 


* 


410 MR. HE. G. BOULENGER ON THE 


Axolotls in shallow water, Stage 3 being reached on January 
10th and 25th, and Stage 4, the stage when: the front part of the 
fin becomes totally absorbed, on February 12th and 28th. The 
Axolotls were then returned to deep water: the development at 
this stage, however, could no longer be checked and both creatures 
completed their metamorphosis in the water. 

From the latter experiment, showing that when replaced in 
deep water the previous stages were resumed in about half the 
time required to reach them, the conclusion may I think be 
drawn that the natural tendency undoubtedly is for the animal 
to remain an Axolotl, and that compulsion is needed to bring 
about metamorphosis. 

It will be noted that at Stage 4, when the front part of the 
fin becomes absorbed, the metamorphosis can no longer be checked, 
and that this is therefore to be regarded as the critical stage, at 
which the further development must proceed whatever the 
conditions under which the animal is placed. 

During the course of all my experiments, with the exception 
of the last stages of the metamorphosis, the Axolotls fed with 
regularity twice a week, and in this respect [ experienced none 
of Mile. de Chauvin’s difficulties. 

Below will be found detailed accounts of the conditions under 
which my eleven specimens were kept, and of the changes which 
took place under the circumstances. 


Specimen No. 1. 
Placed in shallow water at a temperature of 75°-80°. 


Stage 1. August 17th.—Length 138 mm. 
September Ist. 

September 15th. 

October 17th. 

October 24th. 

November 2nd. 

. November 8th. 

November 14th.—Length 131 mm. 


Total length of period—12 weeks. 


DID OF wb 


Specumen No. 2. 


Placed in shallow water at a temperature of 75°-80°. 


Stage 1. August 17th.—Length 119 mm. 
. September Ist. 

September 15th. 

. November 9th. 

November 16th. 

November 21st. 

November 29th. 

December 4th.—Length 113 mm. 


Total length of period—15 weeks 


BONIS Ot pe bo 


METAMORPHOSIS OF THE AXOLOTL. 41 


Specimen No. 3. 
Placed in shallow water at a temperature of 55°-60°. 


Stage * August 17th.—Length 124 mm. 
ees September 25th. 

. October 10th. 

November 12th. 

. November 20th. 

. November 27th. 

. December 5th. 

. December 11th.—Length 118 mm. 


Total length of period—16 weeks. 


OID ONE wb 


Specimen No. + 
Placed in shallow water at a temperature of 55o=608: 


Stage 1. August 17th.—Length 105 mm. 
Be SD: September 25th. 
» 3 October 10th. 


Died January 22nd.—Length 105 mm. 


Specumen No. 3. 


Placed in foul water from which the oxygen was removed by 
the gradual addition of boiled water. 


* Stage 1. August 17th.—Length 136 mm. 
Died January 4th.—Length 155 mm. 


Specumen No. 6. 


Placed in shallow water at a temperature of 75°-80°, and 
replaced in deep water as each successive stage was reached. 


Stage 1. October 7th.—Length 134 mm. 
» 2. October 20th. 
Replaced in deep water. 
» Ll. October 28th. 


Replaced in shallow water. 

», 2. November 15th. 

», 3 December 5th. 
Replaced in deep water. 

» 2. December 20th. 
Replaced in shallow water. 


,, o January 10th. 
,» 4. February 12th. 


412 MR. E. G. BOULENGER ON THE 


veplaced in deep water. 


Stage 5. February 18th. 

6. February 25th. 

7. March 3rd. 

8. March 10th.—Length 129 mm. 


29 
99 


br) 


Py 


Specimen No. 7. 
Placed in shallow water at a temperature of 75°—-80°. 


Stage 1. October 7th.—Length 98 mm. 
2. October 31st. 
» o& November 25th. 


Died January 29th.—Length 98 mm. 


29 


Specimen No. 8. 
Placed in shallow water at a temperature of 55°--60°. 


Stage 1. October 7th.— Length 96 mm. 

. October 20th. 

. November 10th. 

January 3rd. 

. danuary 10th. 

. January 19th. 

. January 27th. 

. February 4th.—Length 91 mm. 


CON rE eo bt 


Total length of period—164 weeks. 


Specimen No. 9. 
Placed in shallow water at a temperature of 55°-60°, and 
replaced in deep water as each successive stage was reached. 


Stage 1. October 7th.—Length 117 mm. 
» 2. October 25th. 
Replaced in deep water. 

1. November oth. 
Replaced in shallow water. 

2. November 20th. 

3. December 26th. 
Replaced in deep water. 

2. January 10th. 


Replaced in shallow water. 


3. January 25th. 
4, February 28th. 


2) 


METAMORPHOSIS OF THE AXOLOTL. 413 


Replaced in deep water. 


Stage 5. March 6th. 

6. March 13th. 

: 7. March 19th. 
8. March 27th.—Length 113 mm. 


Specimen No. 10. 
Placed in shallow water at a temperature of 55°—-60°. 


Stage 1. October 7th.—Length 105 mm. 
2. October 31st. 
, 3 November 30th. 


Died March 24th.—Length 105 mm. 


99 


Specimen No. 11. 


Placed in foul water from which the oxygen was removed 
by the gradual addition of boiled water. 


Stage 1. October 7th.—Length 112 mm. 
Died March 22nd. ete b20) sonar 


The results of these experiments show, I think, in the first 
place that, in accordance with Mlle. de Chauvin’s observations, 
and contrary to those of Dr. Powers, the Mexican Axolotl will, 
with perhaps a few exceptions, transform into the Amblystome 
stage 1f placed, when about six months old, under conditions 
which force it to make frequent use of its lungs; secondly, that 
starvation, irregular feeding, and temperature have no influence 
on the metamorphosis; thirdly that, as no change occurs when 
the Axolotl is placed in poorly oxygenated water, owing to the 
fact that it will not under the circumstances rise to the surface 
and make use of its lungs, the quantity of oxygen in the lakes 
of Mexico can have little bearing on the explanation of the 
phenomenon of neoteny; and lastly, that there is a critical stage 
in the metamorphosis. 


References. 


ra 


. A. Dumtrit.—Comptes Rendus, lx. (1865), and Nouv. Arch. 
Mus. 11. (1866). 

. A. Weismann.—Zeitschr. wiss. Zool. xxv. (1875). 

. Mute. pe CuAuvin.—Zeitschr. wiss. Zool. xxvii. (1876). 

. J. H. Powrrs.—Amer. Nat. xxxvii. (1903). 

. H. Gapow.—Nature, Ixvii. (1903). 


Ce & 0 


A414. SURGEON J. C. THOMPSON ON THE 


32. Contributions to the Anatomy of the Ophidia. 
By Josep C. THompson, Surgeon, United States Navy *. 
[Received April 25, 1913; Read June 3, 1913.] 
(Text-figures 77 & 78.) 


INDEX. Page 

Suggestion as to the Possible Origin of the so-called Tracheal 
Notes upon the Anatomy of :— 

AGAUSOOHNTEIS. WOIGOUOP ccacasosgcnn 569600 500509886 dovoadsd00b5 vovaen | Sli) 

Cycloconusnliineatusmerce een ee ee ee ee Le 

CRIS VGUCE, OPMGED cos odacsvccssodabooescaysoansssdocososcdexacqese | CHAO) 

(CHROGTUIS: COPTAGDIEEB oso 000000 bodvoebadeco.ceese0 Sag acncoDsbocasendeae , SP 
The Intromittent Organ in :— 

PolyodontophisiOiwittatus cence seeeessctese eee eeee eee Ee 

LUO HUCKONOGIS GUNCUUNFO Bonorecrnensses000250esoo00GscK0aA0aeo8008 43) 

JEAUSCOIDS. CHUGRUGIGES ocaads coogucssoes8 00650005 6300a0 90000 000ccqana | GH) 

Cycloconusilineatuserccn ce ee eet ee eee ee eee EO 

SVad@reS (DOWUCECIS soc conccucoboodesccde dav occ sue sédooeooeovscccdan0c0 4A 
Systematic : 

SET D 14 S MEENA OVE es.c 2c sence teesaeeee eae eR a ee 


Suggestion as to the Possible Origin of the so-called 
Tracheal Lung. 


It is suggested that the tracheal membrane has been developed 
as a device originally contributing to the required flexibility 
of the tube. The cartilaginous rings being interrupted behind, 
or on one side, permit that the structure be subject to a much 
greater pressure, especially when bulky morsels are being 
swallowed, than would be the case were the rings complete. 
The tracheal membrane, being continuous with the lung, appears 
to have afforded a convenient foundation for the extension of 
pulmonary tissue when the need for an increased breathing 
surface has arisen, or when the lung has been encroached upon 
by other organs. 

In the Hydrophide an increased pulmonary area is required 
both to give buoyancy and to enable the sea-snakes to remain 
some time beneath the surface. The most specialized species in 
the genus Hydrophis Daudin, those in the fasciatus group, possess 
a lung that extends nearly to the vent, and a tracheal lung of 
due proportion. 

In the Acrochordine, the members of which are of aquatic 
habits, there is also a highly developed tracheal lung. Owing to 
the abnormal size of the heart the respiratory tissue on the 
trachea is widely separated from the lung itself. 

In the Viperide the elaboration of venom requires a large 
liver. These serpents as a rule have short and thick bodies, and 
the liver invades the region just caudad of the heart. As a 


* Communicated by Dr. F. E. Bepparp, M.A., F.R.S., F.ZS. 


ANATOMY OF THE OPHIDIA. AlL5 


result of this the crowded pulmonary tissue seeks the direction 
of least resistance, which is along the tracheal membrane. 

In recording the position of the viscera it has been found 
advantageous to adopt the following routine with the view of 
correlating the data obtained. The serial number of the gastro- 
stege that underlies the anterior tip or the posterior extremity 
of an organ is taken as the external landmark. This number is 
reduced “to a percentage, the total number of the ventral shields 
in the specimen being used as the base. From this procedure 
there results a set of figures which will materially aid in com- 
paring one species with another or in learning the extent of the 
variation which the individuals of a given form exhibit. 

It appears that whenever a serpent varies widely in an 
important character from one of the natural groups, a further 
investigation regularly results in the finding of several additional 
structural features that are well worth making a matter of 
record. The single species constituting the family of Xenopeltidee 
may be taken as an illustration of this general statement. 


Some Notes wpon Anatomy. 


XENOPELTIS UNICOLOR Reinhardt. (Text-figs. 77 & 78.) 


Specumen. No. 16750, California Acad. Sci. Singapore. 
Female ; total length 480, tail 58 mm. 

Squamation.—Scales in 15 rows anteriorly, and the same 
posteriorly, an oblique series commencing at one gastrostege 


Text-fig. 77. 


ae, 
A B Cc 


Teeth of Xenopeltis unicolor. 


A. Maxillary bone viewed at a right angle to the outer edge, and showing the 
oblique fashion in which each tooth is set in the jaw. 
._B. Profile view of the same teeth, showing the lateral cutting-edges of the cusp. 
C. Palatine tooth viewed from below, and showing a stout and blunt tip which 
exhibits a tendency to assume the arrow-head shape of the cusps. 


terminates at the ninth ventral shield to the rear. Gastrosteges 
177. Anal divided. Urosteges 32 pairs, the second entire. Loreal 
and preocular absent. Postoculars 2. Anterior temporals 2, 
posterior 3. Supralabials 8, the fourth and fifth entering the 


eye. Infralabials 9. 


416 SURGEON J. C. THOMPSON ON THE 


Anatomy.—There are two dental characters in this species that 
are unparallelled among the Ophidia. The most striking is the 
shape of the individual tooth, and the fact that the teeth borne 
on the palatine bone are decidedly the largest. Each tooth is 
set in the alveolar ridge with the anterior surface directed 
forward and outward at an angle of 45° with the long axis of the 
bone. When a tooth is viewed from the anterior surface, it 
appears a trifle constricted at the middle; towards the extremity 
it flares out laterally, and terminates like a blunt spear-head, 
with two cutting-edges that meet at a right angle. When 
viewed from the side, this anterior cutting-edge is seen as a 
narrow, beveled cusp, and from its base the tooth is continued 
backwards as a horizontal process, the length of which is equal 
to the broad diameter of the tooth. The edges of the cusp and 
the tip of the backward—directed point are covered with brownish 
enamel. 

The hypapophyses are present on the anterior vertebre and 
absent on the posterior. They have a moderately long base. 

In the writings of Cope * this form is credited with possessing 
a coronoid bone. By Mr. Boulenger, however, it is placed in 
the category of those in which this structure is absent. Two 
specimens have been examined and no vestige of the bone can 
be found. 

The external landmarks of the principal viscera in terms of 
gastrosteges are as follows :— 


Total number of gastrosteges...... 177 100 p. cent. 
ANDERS O18 INGA GE oasanogageodbooseoqdac 52 29-4 
Wiversambertor tip) s.-5:- ere eeere 64 39°] 

PE DOSLeLIOn end ann. abt 6 116 65°3 
Gall-bladder, middle ............... 129 73 
Kidney, right, anterior tip ...... 146 82°5 

As »  posteriorend ... 155 

BS left, anterior tip ...... 150 85 

ss » posteriorend ... 161 
iDleo-Geacell Waly) Bocdsesonsocbnocsocne 157 88°8 


The heart is large, and its position is indicated on the thoracic 
walls by a distinct bulging. Over the base of the organ there 
are two areas in which the scales are enlarged; each area is 
triangular, with the base along the edge of the ventrals and the 
apex at the upper border of the fourth row. In the first row are 
five scales, each of which is one-third larger than those before or 
behind the heart. The scales in the seven dorsal rows are not 
altered. The increase in the diameter of the body over the heart 
is compensated for by an increase in the size of the scales in the 
outer rows, and not by the interpolation of an additional series. 

The tracheal rings are complete from the glottis to the ninth 
gastrostege. From this point the rings are interrupted, and the 


* Croc., Liz. & Sn. N. Amer 900, p. 731. 


ANATOMY OF THE OPHIDIA. 417 


dorsal wall of the trachea is formed by the tracheal membrane 
which joins the ends of the incomplete rings. From the 9th 
to the 21st gastrostege the ends of the rings are in con- 
tact and the membrane lies collapsed above them. From this 
point to the base of the heart, the membrane widens and the 
ends of the rings are permanently separated. Dorsal to the 
heart the tracheal membrane winds around to the right side of 


Text-fig. 78. 


SS 


Fi SCO 


ise 


Base of the lungs of Xenopeltis unicolor. 


Viewed from below; the object being te display the bridge uniting the two lungs, 
the termination of the trachea, the long free apex of the left lung, the adherent 
apex of the right lung, with its tiny free tip, and anterior to it the portion of 
the tracheal membrane which is lined internally with pulmonary tissue and is 
bordered by a branch of the pulmonary artery. 


the tube, and 2:5 mm. before the trachea enters the lung it 
becomes lined with pulmonary tissue. The two lungs are 
adherent for a distance of 10 mm., the connecting bridge 
extending from the 53rd to the 57th gastrostege. Viewed from 
within, this bridge appears as a septum lined with alveoli and 


418 SURGEON J. C. THOMPSON ON THE 


dividing one lung from the other. There are two perforations 
affording communication between the lungs; the larger, measuring 
2mm. in diameter, is at the termination of the trachea, and the 
smaller, measuring 1 mm. in diameter, is near the posterior 
border of the septum. The trachea ends abruptly 3 mm. behind 
the apex of the heart, and on the lower surface of the bridge. 

The right lung extends from the 49th to the 119th gastrostege. 
Anteriorly, it reaches 8 mm. beyond the border of the bridge; 
it is adherent to the right side of the trachea for the distance of 
6 mm., and has a free apex 2 mm. long. The trachea communi- 
cates with the lung 2°5 mm. behind the point where they join. 

The left lung extends from the 50th to the 77th gastrostege. 
Anteriorly it terminates in an acutely pointed free apex 8 mm. 
long, at the level of the auriculo-ventricular septum. Poste- 
riorly the lung terminates in a blunt cone, and towards the 
end the walls are a trifle thinner and the alveoli larger. There 
are two pulmonary veins. The right courses along the angular 
ventral border of the right lung and enters the anterior inferior 
corner of the auricle. The left commences at the posterior 
border of the bridge and enters the posterior inferior corner of 
the auricle. This vein is the thicker of the two. It is formed 
of two branches that arise along the mesial side of each lung; 
the branch from the right lung is the larger, and it has frequent 
anastomoses with the right pulmonary vein. 

The liver extends from the 64th to the 116th gastrostege. 
There is no trace of segmentation. 

The centre of the gall-bladder is at the 129th gastrostege. 

The right kidney extends from the 146th to the 155th, and 
the left from the 150th to the 16lst gastrostege. Each is 
semidivided into nine irregular lobes. The right kidney is 
supped by two renal arteries, the anterior entering at the 
second and the posterior at the sixth lobe. There is a single 
renal vein leaving the anterior tip of the organ, and this promptly 
unites with its fellow to form a common trunk. 

The lining of the rectum is smooth. The ileo-cecal valve is 
at the 157th gastrostege. ‘The rectal cecum is 9 mm. long, and 
is directed forwards and lies on the right side of the ileum. Its 
walls are thin, and the opening into the cecum is just anterior 
and to the right of the valve. Its lumen was filled with fecal 
matter, though the cecum and rectum were empty. 

Along the dorsal wall of the cloaca there is a longitudinal, 
rounded ridge, which terminates in a papilla 1 mm. in length, 
and at the level of the posterior border of the last gastrostege. 
This ridge is formed by the two parallel tubes of the completely 
divided vagina ; these terminate in two separate orifices at the 
tip of the papilla. The lining of the vagina is finely and 
longitudinally plicate. According to Cope* the vagina in the 
Peropoda is undivided and the walls are nearly or quite smooth, 


* Op. cit. p. 7Q0. 


ANATOMY OF THE OPHIDIA. 419 


and in the Colubroidea the organ is bilobate and the walls have 
deep longitudinal grooves. This form more closely resembles the 
latter. 


CycLocoruUS LINEATUS Reinhardt. 


Specimen. No. 15240. California Acad. Sci. Olongapo, 
Luzon, P.I. Female; total length 503, tail 81 mm. 

Squamation.—Scale-rows 17 anteriorly ; the V row suppressed 
at the 127th gastrostege on the right and the 125th on the left, 
leaving 15 rows which are continuous to the vent. The last 
scale in the row that is dropped is before an enlarged scale 
in the row below. Anteriorly an oblique series of scales 
starting at one gastrostege terminates at the tenth to the rear. 
Gastrosteges 163. Anal entire. Urosteges 41, single, Pra- 
oculars 2, postoculars 2. Anterior temporals 2, posterior 2. 
Supralabials 8; the third to the fifth entering the eye. Infra- 
labials 9; the first to the fifth in contact with the anterior 
geneials. Anterior larger than the posterior geneials. 

Anatomy.—The hypapophyses on the dorsal vertebre are 
continuous throughout the column. On the vertebra above the 
122nd_ gastrostege the base of the hypapophysis occupies 
the posterior one-third of the centrum; the apex is horizontally 
truneate, and extends a trifle posterior to the vertical of its 
centrum. The caudal vertebra at the level of the 12th urostege 
has two triangular hemapophyses the bases of which occupy the 
posterior half of the centrum. The inferior zygapophyses are 
developed as broad wing-like plates, 1-4 mm. long. 

The maxillary bone at the junction of the anterior and middle 
third is bent inwards at an angle of 45°, and the teeth are 
arranged in two groups. The first set consists of 7 teeth that 
are mounted on the anterior one-third of the bone; the first five 
are very small and increase in size posteriorly ; the sixth is 
nearly three times the size of the fifth, and the seventh is larger 
still, being 1:5 mm. long. These teeth are inclined backwards. 
The two groups are separated by an interval of 155 mm. The 
second set contains 13 teeth ; the first nine are small and nearly 
equai; the tenth to the fourteenth increase rapidly until the last 
is twice the size of the ninth. These teeth are inclined towards 
the median line. The anterior tip of the palatine bone reaches 
the interval between the fifth and sixth maxillary teeth; there 
are 16 teeth. The pterygoid bone bears 19 teeth; these are on 
a line that is convex externally ; the middle teeth being three 
times as far from their fellows on the opposite bone as are the 
teeth at the extremes. The dentary bone is also bent inwards at 
an angle of 45°, and has the teeth in two groups. The first set 
is mounted on the inflexed portion of the bone, and consists of 
6 teeth; the first to fourth gradually increasing in size; the 
fifth and sixth strongly enlarged. The two groups are separated 
by an interval which is shorter than the sixth tooth. The second 
set consists of 17 very small teeth. 

Proc. Zoot. Soc.—1913, No. XXIX, — 29 


420 SURGEON J. C. THOMPSON ON THE 


The external landmarks of the principal viscera in terms of 
gastrosteges are as follows :— 


Total number of gastrosteges ... 174 100 p. cent. 
INGE OF INCAVAY 5 coasodvocnscscs0e060r 37 21:3 
JbmyEIe, QVNESOTE UND) oogconsconsdecoass 44 25°3 
7 posterior end ~.......-...... 96 55-1 
Grille Glee dogepdocesbsoosesodaceds 114 65°3 
Kidney, right, anterior tip ...... 142 81:8 
i; }  posterion endy i.) 29 85°8 
As left, anterior tip......... 144 82°8 
a iP posberlom endian tol 86°8 
iileo=ccecalliivalivielec en snc-ce steer 154 88°95 


The tracheal membrane begins at the glottis, and for a short 
distance lies along the right side of the tube. Almost immediately 
it broadens, and at gastrostege 14 it is lined with respiratory 
tissue. At gastrostege 22 the membrane alters its position so 
that it forms the dorsal instead of the right wall of the trachea. 
Well above the apex of the heart the alveoli on the membrane 
have assumed the same size and shape as those in the lung; there 
is no gross difference to indicate the ending of one structure and 
the beginning of the other. The trachea terminates abruptly 
7 mm. posteriorly to the apex of the heart. There is uo rudi- 
mentary lung. Arteriorly the walls of the lung are thick and 
lined for the entire circumference with pulmonary tissue ; 
posteriorly they become thin and terminate in an air-sac. The 
lateral lobes of the liver are distinct. The left lobe is the longer ; 
anteriorly it extends beyond the right lobe 6 mm. and posteriorly 
jl mm. There are four transverse fissures on the left lobe near 
the anterior tip, the rest of the organ is smooth. The anterior 
one-third of the rectum is thickly and regularly plicate; the 
posterior two-thirds are irregularly folded. There is no body to 
the vagina. The organ is bilaterally divided into two separate 
tubes that open into the dorsal wall of the cloaca by two separate 
orifices, which are 2 mm. apart. 

Notes.—This species is described as having scales with apical 
pits, and an eye with a round pupil. Over a dozen specimens 
have been studied and the pits cannot be made out. The pupil 
is usually round in the young and frequently vertically oval in 
the adult. 


CHRYSOPELEA ORNATA Shaw. 


Specimen. No. 16707. California Acad. Sei. Cochin China. 
Female; total length 910, tail 237 mm. 

Squamation.—The number of scale-rows on the body, the 
sequence In which they become suppressed, and the gastrostege 
level at which they terminate on each side may be thus 
presented — 

17 rows, IV row ends, right 137th, left 136th gastrostege, leaving : 
yee VALLE ees, 4 Wadia, op Jashiln 


13. ,, which are cortinuous to the vent. 


ee] 29 


ANATOMY OF TIE OPHIDIA. 421 


Gastrosteges 227, the last shield divided. Anal entire. 
Uvosteges 120 pairs. Supralabials 10; the fifth to the seventh 
entering the eye; on the right the fourth is reduced and fused 
with the posterior inferior angle of the third. Infralabials 10 on 
the left, 11 on the right, the additional shield is between the 
corresponding third and fourth, Anterior geneials larger than 
the posterior, the right in contact with four and the left with 
five infralabials. Gular shields in six pairs, 

Anatomy.—Maxillary bone with 20 teeth, the posterior three 
with a broad shallow groove; the five preceding these are 
also grooved but less distinctly. Palatine bone with 8 teeth, 
increasing in size posteriorly ; the anterior tip of the bone 
reaches to the interval between the second and third maxillary 
teeth. Pterygoid bone with 28 teeth. Dentary bone with 
18 teeth, the anterior enlarged and grooved on the external 
quadrant. 

In terms of gastrosteges the external landmarks of the viscera 
are as follows :— 


Total number of gastrosteges ...... 227 100  p. cent. 
Aexcolshearty Gara ssct-ceiscecat|sce hes 58 25°6 
Miversanterlonibip (reece osece os: 73 32°2 
5) [DOSS Cha VOSS a Soebgacoeeuae. 112 A9-4 
Galll=bladden centre tase. .0.0ssee eee 232 58°2 
Kidney, right, anterior tip ...... .. 183 80°8 
ps oy) posterior endi= sce 201 89 
ee elit era mbenroritipis ween. 196 86°5 
cs ye posterior ends. 214 94-2 
lilewzceecallt valves sae eeees asec aeee 209 92-2 
Waginalcormuay basemen .c eee 216 95 


The cartilaginous rings of the trachea are complete from the 
elottis to the angle of the jaw; from this to their termination 
on the ventral surface of the lung, at the apex of the heart, they 
ave interrupted. ‘The tracheal membrane begins with the semi- 
rings; it is on the right side of the tube, and is continued eaudad 
to the base of the heart. For the greater part of its length it 
comprises more than half the circumference of the tube. At 
its termination it becomes continuous with the pulmonary pleura. 
The elasticity of the rings maintains their free ends in close 
apposition, and the membrane lies in a lax state by the side of 
the trachea. When the windpipe is dilated, the ends of the 
rings are separated and the tracheal membrane becomes a 
functioning part of the organ. Under ordinary conditions the 
air passes only through that part of the tube that is formed by 
the semi-rings. Anteriorly the pulmonary tissue begins at the 
level of the auriculo-ventricular septum. In the lung it is 
confined strictly to the dorsal half of the circumference. 
Posteriorly it is drawn out as a fine streak from which are sent 
out transverse septa ; it terminates on the left side of the air-sac, 
at the level of the 79th ventral. Posteriorly the lung is continued 


as a membranous air-sac; exactly where it ends cannot be 
G ¥ 
a] 


bo 4% 


423 SURGEON J. C. THOMPSON ON THE 


ascertained. There is a small rudimentary lung, a mere vesicle 
without pulmonary tissue. ‘The anterior tip of the liver is not 
divided into lateral lobes ; posteriorly the left lobe is about three 
times as thick as the right, and is 5 mm. longer, The surface is 
smooth, there being no transverse fissures. The ileo-czcal valve 
is at the 209th gastrostege ; there is no cut-off tube or rectal- 
cecum. The lumen of the rectum is longitudinally plicate, with 
faint closely-set transverse ridges. The vagina bifurcates at the 
level of the 216th gastrostege, the walls are smooth. 

Habits.—The observations made by Mr. Shelford * on the aereal 
locomotion that this serpent indulges in afforded one of the most 
interesting field-notes on a reptile that has been published for 
some time. Attention was drawn to the hinge-lines along the 
gastrosteges, and to the habit of retracting the middle section of 
these shields “ so that the snake became deeply concave along the 
ventral surface ”. This serpent has been captured several times 
in Luzon, and the same performance witnessed. If the lung in 
this species had thick walls for its entire circumference, this 
radical change in the shape of the body could not be so readily 
brought about. On the contrary, 1t is so modified that when 
subject to ventral pressure it merely has the lower membranous 
wall stove in and no serious interference with respiration takes 
place. 


CROTALUS CONFLUENTUS Say. 


Specimen. Field No. 8208. U.S. Nat. Mus. Mt. Tamilpais, 
alt. 650 M., California. Female; total length 460, tail 45 mm. 


Squamation.—The number of scale-rows on the body, the 
sequence in which they become suppressed, and the gastrostege 
level at which they terminate on each side may be thus 
presented :— 


25 rows, VI row ends, right 108th, left 160th gastrostege, leaving : 
Me egy Vigbes. on sis Fl okd ees Gla 5 i 

PHI Pio 8 One EIN, 45 G2! bs ia 
20. X ,, intermittent between 124th and 165th __,, 

19 ,, which are continuous to the vent. 


Ventrals 174. Anal entire. Urosteges 21 entire, the first 
and last three paired. Preoculars 2, suboculars 2, postoculars 4. 
Supralabials 14 on the right, 13 on the left side; the reduced 
count due to the fusion of the fourth and fifth shields. 

Anatomy.—The external landmarks of the principal viscera in 
terms of gastrosteges are as follows :— 


Total number of gastrosteges ... 174 100 p. cent. 
Igoe OH INCHED ooooodoonqvsnscdesacoase 70 40:2 
ToihyGIey PyOEVAOVe (HN)8), -benccoanabaoonsos 70 40-2 

py ee postenioncemdiinn rare. ses. 109 62°5 


* “ A Note on ‘ Flying’ Snakes,” P. Z.S. 1906, p. 227. 


ANATOMY OF THE OPHIDIA. 423 


Kidney, right, anterior tip ...... 144 82°8 p. cent. 
> 3, posteriorend ..: 163 
3 left, anterior tip ......... 146 84 
. » . posteriorend ... 166 

Wei@Quaen, GYD eobeogsstanbecsonoddads 164 99-2 


The cartilaginous rings of the trachea are complete from the 
glottis to the 6th gastrostege. At this point the tracheal 
membrane begins on the dor sal quadrant of the tube. Almost 
immediately, ‘while still very narrow, it acquires pulmonary 
tissue, this rapidly increases and becomes continuous with the 
lung, The semi-rings extend along the ventral surface of the 
trachea ; viewed from. within they appear as a narrow gutter; they 
terminate 15 mm. caudad from the apex of the heart. The walls 
of the lung posterior to the heart have a very thin lining of 
pulmonary “tissue ; the alveoli are large and shallow in contrast 
with those in the trachea, which are small, thick, and superimposed. 
In other words, the area of the respiratory tissue in the lung is 
much less than it is in the trachea. The liver is deeply divided 
into a right and left lobe, and these are partially divided by 
shallow and transverse fissures. Anteriorly the left lobe is thick 
and rounded, extending three gastrosteges beyond the right, and 
overlapping the apex of the heart by 1 mm. The peritoneum 
covering the liver and the abdominal walls is pigmented, the 
remainder is plain. The vagina is short, it extends over 10 
gastrosteges, and is bilobate. The cornua are adherent to each 
other at the base for a distance of 15 mm. Externally each 
horn gives the appearance of being composed of two adherent 
tubes, with the oviduct entering the outer. The lining is 
smooth, and is thrown into several longitudinal folds; two of 
these are larger than the rest, and their free edges nearly 
touching tend to divide the lumen of each horn into an inner 
and an outer compartment. 


The Intromittent Organ. 


Polyodontophis bivittatus Boulenger.—The hemipenis is un- 
divided and the sulcus simple. The base is plicate and is followed 
by a spinous zone 4 mm. in length. The spines are about ten in 
number ; there are two placed opposite to the sulcus that are 
much enlarged, being 2°5 mm. in length. The distal 6°5 mm. of 
the organ 1s calyculate. The calyces are very small and their 
borders bear numerous minute spines. The calyces at the border 
of the spinous zone and at the tip are enlarged, the latter have 
simple borders. 

In Cope’s classification the enlarged basal hooks place this form 
in the Natricine. The occurrence of these hooks with a calyculate 
apex represents a divergence in the direction of the Colubrine. 


Tropidonotus vibakari Boie.-—The hemipenis and the sulcus 
spermaticus are undivided. The organ is densely spinous; those 
on the middle are a trifle the larger, diminishing in size towards 


494. SURGEON J. C. THOMPSON ON THE 


the apex and the base. There are a few enlarged basal hooks 
situated close to the suleus, one being on one side and four on 
the other; of these four the one nearest the base is double the 
size of any of the others. The tip of the organ is rounded and 
smooth, and the line of demareation between the bare and the 
spinous areas is sharp. On either side of the smooth area at the 
tip, there are mounted two apical papille, which are one-fourth 
the length of the body of the organ. Each papilla is spinous for 
three-fourths of its circumference, the inner quadrant being 
smooth and continuous with the spineless area at the tip. 

Terminal papille similar to those in this species are of com- 
paratively rare occurrence. Cope has reported them in the genus 
Oligodon Boie, and in the three following species: Jropidonotus 
vittatus Linneus, [schnognathus lineatus Hallowell, and Coluber 
helene Daudin. Hach of these has been set apart by Cope as 
the type of a new genus, established on the ground that the 
intromittent organ differed widely from the type prevailing 
among the nearest allies. 7. vibakari in this respect is clearly 
separated from the Far Eastern representatives of the genus. 
These have been placed in the genus Bothrodytes by Cope, and 
are characterized by having the organ furcate. The organ in 
vibakari most closely resembles that found in Ischnognathus 
lineatus Hallowell, and the figure given by Cope* of this 
species shows that it differs from vibalari in the one detail of 
being less completely covered with spines. Furthermore, this 
serpent is of precisely the same mild-actioned and fearless nature 
that is characteristic of the species of /schnognathus. In this it 
offers a marked contrast with the Far Hastern members of the 
genus Tropidonotus. Over a dozen species of these have been 
captured and every one showed a nervous, irritable, and snappy 
disposition. bain 

To allow this: species to remain in a complex genus like 
Tropidonotus, after it has been found to differ so radically, 1s not 
good taxonomy. On the other hand, to chronicle promptly each 
new anatomical discovery under a new generic caption does not 
materially aid our studies. It is believed that the dilemma can 
be overcome for the time being by strictly adhering to the 
nomenclature contained in the ‘Catalogue of Snakes in the 
British Museum,’ and at the same time by registering a pro- 
visional new generic term. This new term is not to become 
current until the entire group to which it belongs has been 
re-examined from the standpoint of comparative anatomy. 

With this understanding it is suggested that the new genus 
Hebius be established for Tropidonotus vibakari Boie, the salient 
character distinguishing it from all species of Zropidenotus 
being the densely spinous and undivided hemipenis, bearing two 
long apical papillee. 

Helicops angulatus Linneus.—The total length of the hemi- 
penis is 42 mm.; it is divided into two quadripenes, the length 


? 


* Op. cit. pl. xx. fig. 12. 


ANATOMY OF THE OPLIDIA. 425 


of each being 10 mm. The quadripenis is calyculate, and the 
cells are arranged in seven rows that trend obliquely forward 
from the suleus. he distal half of the body of the hemipenis is 
spinous; opposite to the suleus there is a septal fold. The 
proximal half is plicate. At the border of the spinous area and 
close to the sulcus there are two round, flat-topped, and gristle- 
like papille ; these are closely set and one is distal to the other. 
The sulcus is divided at a point 12 mm. from the base, this is 
well within the plicate area. 

Mr. Rosén* writes that the hypapophyses are absent from the 
posterior vertebre in H. modestus Giinther and H. leopardinus 
Schlegel. In this species they are well developed throughout, 
one occurring even on the last dorsal vertebra. 


Cyclocorus lineatus Reinhardt.—The hemipenis is of extreme 
length and slenderness, being 38 mm. long and 2 mm. in diameter. 
It is not divided and the sulcus is simple. The base for the 
distance of 4 mm. is plicate; the remainder of the organ, even 
to the tip, is spinous. Towards the tip there are three slightly 
elevated longitudinal ridges, one 8 mm., the others 5 mm. in 
length. The spines are nearly uniform in size, being about 
°3 mm. to 4 mm.long. Each is mounted on a broad fleshy base, 
and is slightly recurved. When the walls are stretched the 
spines are seen to be arranged in longitudinal series. 


Spilotes pullatus Linneus.—The structure in this species is 
one of the most complicated recorded. The total length of the 
hemipenis from the base to the insertion of the retractor muscle 
is 77 mm. The organ and sulcus spermaticus are undivided. 
From the base to the 20 mm. point are longitudinal plications ; 
the folds are thick, at first straight, later becoming wavy. From 
the 20 mm. to the 38 mm. point it is densely beset with stout, 
blunt spines, each of which is about 2-5 mm. long. From the 
38 mm. to the 54 mm. point on each side of the sulcus are rows 
of flounces, these trend obliquely forward and acquire partitions 
which convert them into rows of calyculi; the cells of these 
calyculi increase rapidly in size, the largest being situated 
opposite to the sulcus. On each side of the sulcus at the 60 mm. 
point are two thick flaps 10 mm. long; these are in the longi- 
tudinal plane, hang free in the lumen of the organ, and are 
covered with small regular ruches. Between one of these flaps 
and the sulcus there is a thin membraneous septum 20 mm. long, 
parallel to the sulcus, and hanging free in the lumen. Towards 
the extremity of the organ the calyculi change to irregular 
flounces; these in turn are converted at the extreme tip into 
minute frills. 


United States Fisheries Steamer ‘ Albatross ’, 
Sausalito, California, April Ist, 1913. 


* Ann, Nat. Hist. (7) xv. 1905, pp. 170-171, fig. 1. 


426 DR. S. F. HARMER ON THE 


33. The Polyzoa* of Waterworks. 
By Sipney F. Harmer, Sc.D., F.R.S., F.Z.8.7 


[Received March 11, 1913: Read April 22, 1913.] 
(Plates LXIL. & LXILL2) 


Inpex. 

Bionomies : Page 
Relation of Polyzoa tothe other Constituents of the Fauna and Flora... 428 
Economic importance of Polyzoa .......... Pu Reet ncc ss ea REO, 

Records of occurrence of Polyzoa in Branch VWiatonworeel Fae oe cs eo OD 

Systematic : 

JEGHORICANG GRRECCOWAEE 450.503 090 009200 048 oEneO6 od 600 SB LOD ecanagedosodasedanannocacca . 4lal 
Fredericella sultana .......... Pe Recn ee ronoselcescucanmonecenteccdcadaacual | (aiats) 
Plumatella fungosa, var. Perales BRE atar toc oni tonicooseconarcoepmog neh aoe 449 
Plimatella emarginata, VAL. MUSCOSH 0.60 ce0cceccveeeceecseceeeeeisecessss en, 452) © 
Development of hibernacula of Puludicella .......0.c.0c1eccccecsee eee eeereceseresere 444 


Morphology of hibernacula and statoblasts 


I. Historical Account. 


It is perhaps not generally recognized in this country that, 
under certain circumstances, Polyzoa may be of great economic 
importance. But it has long been known to those interested in 
the subject, particularly in Germany, Holland, Belgium, and the 
United States, that a group of organisms, among which certain 
Polyzoa play a leading part, may flourish in the pipes of water- 
works which are not provided with an efficient filtering apparatus, 
to such an extent as to give rise to the most serious inconvenience, 
and may indeed finally throw the whole system out of gear. It 
is the object of this paper to show, by means of examples which 
have come under my notice during the last two or three years, 
that this country is by no means exempt from the risks which 
have been experienced in various places abroad. 

The first important contribution to the subject was an ex- 
tremely interesting paper by Kraepelin (85), published under the 
title of “ Die Fauna der Hamburger Wasserleitung” ; although, 
AS Kraepelin himself points out, a list of 18 species of amines 
from the pipes of the same water- supply had been published 
by Petersen some nine years earlier. The occurrence of such 
organisms as Asellus and Gammarus in the water supplied to 
houses in Hamburg was no rare event in that city; while the 
pipes were frequently choked by Eels, ‘ Leitungsmoos” and 
other organisms. Kraepelin undertook a biological investigation 
of the subject, mfluenced partly by the hope of finding blind 
Crustacea among the inhabitants of the water-pipes, to which 


* [In view of the difference of opinion as to whether this Phylum should be called 
Polyzoa or Bryozoa (see Proc. Linn. Soc. 1911, p. 63), I have accepted the preference 
of the author.—Ep1ror. | 

+ Published by permission of the Trustees of the British Museum. 

+ For explanation of the Plates see p. 456. 


BP, 4A. S.IGIS, IPL, IAN. 


0 20 40 60 80 100 iS) 


8. tO). 


5.F. Harmer del. j Bale & Danielsson ,L*4 imp. 


PALUDICELLA ARTICULATA. 


12, Ge S._ IIS), Jet, ILD, 


| | ae. 25. 


S. F. Harmer, del. ; Bele & Danielsson, L*? imp. 


STATOBLASTS OF PHYLACTOLAEMATA. 


POLYZOA OF WATERWORKS. 497 


light had, of course, no access; and by the further consideration 
that he might be able to detect some modifications of structure, 
as compared with the ordinary inhabitants of the Elbe, in the 
animals which were living under these unusual conditions. 
Although the results of the investigation were not of the nature 
which he had anticipated as possible, they were nevertheless very 
surprising; since he was able to demonstrate that some fifty 
genera of animals occurred, often in enormous numbers, in the 
system. The examination was made with the assistance of a 
specially devised piece of apparatus which was screwed on to 
various parts of the water-mains. A column of water was 
allowed to escape through this arrangement, and the animals 
which were contained in it were filtered off. The results obtained 
were very uniform, in whatever part of the system the apparatus 
was used, so that it became clear that the organisms found were 
not merely a fortuitous collection of individuals which had acei- 
dentally been introduced into the supply, but were, on the 
contrary, perfectly normal constituents of a definite fauna in- 
habiting the water-pipes under a pressure of 23-53 atmospheres. 
Among these organisms a conspicuous, and usually the greater, 
part consisted of large masses of what had become known to the 
workmen as ‘“ Leitungsmoos ”—a mass of entangled tubes which 
belonged principally to Polyzoa of the genera Mredericella, Pluma- 
tella and Paludicella, but partly to the well-known Hydroid 
Cordylophora. Sponges (Hphydatia fluviatilis and Spongilla 
lacustris) were frequently found. Colourless forms of Hydra 
were discovered; but, as might have been expected, /. viridis 
appeared to be completely absent. Several species of Freshwater 
Oligocheetes were recorded, as well as “almost incredibly large 
numbers” of Leeches (Glossiphonia and Nephelis). Parasitic 
worms were represented by Lchinorhynchus, found in large 
numbers, in its larval condition, in Asellus; and, in its adult 
condition, apparently derived from Eels. Among the Polyzoa 
the “ mossy” appearance of the masses of tubes above alluded to 
was generally due to Fredericella sultana. Traces of Paludicella 
articulata were generally found ; while Plumatella was represented 
principally by a form described by Kraepelin as P. princeps, var. 
muscosa, and by the Alcyonelloid P. fungosa. Almost every 
sample contained hundreds, or indeed thousands, of <Asellus 
aquaticus, which were found creeping over the Polyzoa “in ugly 
crowds.” The Freshwater Shrimp, Gammarus pulex, was some- 
what less numerous, while various forms of Cladocera, Copepoda, 
and Ostracoda were nearly always present. Leander | Palemon| 
squilla and Mysis chameleon were represented by single finds. A 
small water-mite and a few Dipterous larvee and pup were rarely 
met with. ' 
Among Molluscs the commonest species was Dreissensia poly- 
morpha; but Bithynia tentaculata was frequent, and other fresh- 
water genera—Physa, Limnea, Planorbis, Ancylus, and Spheriwum 
(Cyclas)—were also discovered; while Petersen had previously 


428 DR. S, F. HARMER ON THE 


recorded Vivipara (Paludina), Anodonta and Unio. Planarians 
and a Nemertine (Zetrastenma) were also found, although the 
method employed for collecting the samples was not well adapted 
for the discovery of the smaller organisms. Rotifers and Infusoria 
(Vorticella, Stentor, Paramecium, and Acineta) were, however, 
discovered. 

Even the Vertebrates were not unrepresented in this remark- 
able fauna. The Eel, up to a foot in length, was common. As 
many as six individuals were sometimes present in one sample, 
and Kraepelin estimated that there must have been many 
thousands in the system. Other fishes were less numerous ; 
but the Stickleback (Gasterostews aculeatus) was occasionally 
found ; and on one occasion a Burbot (Lota vulgaris), a foot in 
length, was discovered. A young Flounder, several centimetres 
long, was found in company with the Prawn (Leander) above 
alluded to. 

Summarizing his observations, Kraepelin points out that the 
constituents of the fauna of the water-pipes consist almost exclu- 
sively of forms which feed either on detritus or on one another ; 
that in the absence of ordinary green plants, forms which feed on 
these are absent ; while animals like insects, which require access 
to air, are also absent. For the species which had succeeded in 
adapting themselves to their novel environment the conditions 
were, on the contrary, exceptionally favourable, and indeed more 
favourable than those found normally in Nature. Some of the 
factors specially alluded to, in this connexion, were the absence 
of carnivorous insects, and the existence of large surfaces suitable 
for the attachment of sessile animals. 

Kraepelin proceeds to consider the ultimate source of the food 
of the pipe-fauna; and he points out that this must consist of 
the microscopic organisms and other detritus introduced into the 
system by the constant flow of water from the Elbe. The absence 
of any filtering arrangements, at that time, allowed these particles 
to enter without hindrance. In working up these materials into 
new organic substances the Polyzoa played a part of pre-eminent 
importance; particularly since these animals formed so large a 
proportion of the entire fauna, The carnivorous animals natur- 
ally depended for their supply of food on the organisms which 
were able to make use of the microscopic detritus. Kraepelin 
accordingly arrived at the perfectly legitimate conclusion that if 
the food-supply were to be cut off by the introduction of an 
efficient filtering arrangement, the pipe-fauna would before long 
die out. The soundness of this conclusion was amply demon- 
strated by the cessation of the trouble at Hamburg on the 
introduction of the remedy suggested ; while the experience of 
Water ‘Engineers in many other places has been of a similar 
nature. 

The presence of Polyzoa in water-pipes is, however, not an 
unmixed evil; since, as has been pointed out by several observers, 
if floating organisms exist in the water, the Polyzoa serve the 


POLYZOA OF WATERWORKS. 429 


purpose of removing many of them. But any good they may do 
in this way is more than counterbalanced by other inconveniences. 
One of the most obvious of these is the diminution of the calibre 
ot the water-pipes, which may take place to such an extent as 
seriously to interfere with the circulation. Thus Kemna (99, 
p- 46) mentions a case which had come under his own observation 
of a pipe 60 cm. (about 24 inches) in diameter, which was en- 
circled, on its inner side, by a layer of Polyzoa 15 cm. (6 inches) 
in thickness. It has further been found by Water Engineers 
who have been confronted with this difficulty that, as the Polyzoa 
disintegrate, masses of the tubes are torn off from the walls of 
the pipes and pass into the circulation. They thus pass to the 
smaller pipes of the domestic supply, where they give endless 
trouble by choking the ball-taps and strainers of the meters to 
such an extent as to stop the supply of water. 

These are, however, by no means the only troubles which 
result from the presence of Polyzoa and other organisms in the 
water-supply. It is the habit of Freshwater Polyzoa, in these 
latitudes at least and in the state of nature, to die towards the 
approach of winter. There is evidence that even in the sheltered 
conditions under which they live in water-pipes protected from 
the weather, they are not entirely insensible to seasons; although 
it may be noted that Kraepelin (85, p. 12) points out that many 
of the constituents of the pipe-fauna were found, in the depth of 
winter, at a time when the water containing or ganisms living in 
exposed places was covered with ice and snow, in a condition 
differing but slightly from that in which he had found them in 
the summer. But he adds that the polypides of Pluwmatella were 
all dead at this time (December), although in March the hiber- 
nacula of Paludicella were splitting (as a preliminary to germina- 
tion), and the statoblast-“‘ embryos” of Wredericella were already 
escaping from their shells. Owing to the death of the polypides 
of the Polyzoa, at the approach of winter, a certain amount of 
decomposition of organic material naturally occurs. ‘This sets up 
further troubles, partly by tainting the water and partly by pro- 
viding material which encourages the growth of Bacteria. The 
tainting of the water, which may be due to a combination of 
these two factors, is sometimes very serious. Thus De Vries (90, 
p- 38) records a case of this kind in which even horses would 
not drink the water. 

It is not my purpose to discuss the flora of the filtering- beds 
and water-pipes—a subject of great biological inter est. with 
which I do not feel myself competent to deal—but it is neces- 
sary to refer to an illuminating study of this question which was 
made, in 1890, by Hugo De Vries, in the memoir just referred 
to. ‘The town of Rotterdam is supplied by water from the Maas. 
The waterworks were established in 1874, and at first gave no 
trouble. In the spring of 1877 serious difficulties began to be 
experienced as the result of a sudden invasion of “ Tron- 
Bacteria,” and in particular of a form described by De Vries 


430 DR. S. F. HARMER ON THE 


as Crenothrix Kihniana (=C. polyspora Cohn). A scientific 
Commission was appointed to investigate the matter; and the 
memoir cited was the result. The Bacteria in question, which 
grow associated in long filaments, flourish in the presence of iron 
in solution in the water. Under these conditions, and when pro- 
vided with the requisite organic matter, they deposit the iron, in 
an insoluble form, in the sheath which forms the outer part of 
the filament. Where they are present in sufficient numbers 
they then flake off the inside of the pipes in masses which pass 
into the circulation and are delivered with the water into the 
domestic supply, where they cause great annoyance. This result 
may become a serious calamity by rendering the water extremely 
unpleasant for domestic use, and completely unusable for many 
industrial purposes (De Vries, $0, p. 9). The remains of animals 
and plants which have died in the pipes are said to provide 
the organic matter which is required by the Iron-Bacteria. The 
growth of these organisms is commonly associated with the 
formation of a hard deposit on the inner sides of the pipes; 
and this may give rise to serious trouble by reducing the flow 
of water. 

Tam indebted to my colleague Mr. A. Gepp, of the Botanical 
Department of the British Museum, for calling my attention to 
some of the special points of interest in connexion with Tron- 
Bacteria. A good general account of the subject may be found 
in De Vries’ memoir and in the works of Lafar (98, 04), while 
an extensive bibliography is given on pp. 90-92 of one of 
Kemna’s memoirs (05). One of the special points of interest in 
this connexion is the theory, first brought forward by Winogradsky, 
that the Iron-Bacteria take up iron in solution in the ferrous state 
and obtain their energy by oxidizing it to the ferric state, which 
appears in the form of iron-rust deposited in the sheaths of the 
filaments. The theory has been criticised by some of the more 
recent workers, as by Molisch, who claims to have kept Tvon- 
Bacteria for several generations in iron-free solutions. It has 
been suggested that the deposit of iron-rust in the sheath of 

/renothri« and other forms is of the same general nature as the 
appearance of silica in the cell-wall of a Diatom: and that it has 
not the physiological significance which was attributed to it by 
Winogradsky. But whatever the nature of the process may be, 
the peculiar property possessed by these Bacteria of depositing 
iron in the ferric state is a matter of the utmost practical 
importance to Water Engineers. 

The removal of the pipe-fauna, or the establishment of condi- 
tions under which a fauna of this kind cannot effect an entrance, 
appears to be of great importance in preventing the growth of 
Tron-Bacteria. As De Vries pointed out, these organisms require 
organic matter, which is supplied to them in abundance, in un- 
filtered water, by the disintegration of the constituents of the 
pipe-fauna. An instructive case, which appears to be of this 
nature, is discussed by Kemna in his account of the troubles 


POLYZOA OF WATERWORKS. 431 


which have been experienced at Liverpool. In order to increase 
the supply of this city a dam was constructed at Vyrny, in 
Wales. Filters were installed at Oswestry, thirty kilometres 
from the dam ; and the water was then carried for about twice 
that distance to Liverpool. In this latter part of the system no 
difficulties have been experienced ; but in the part which carries 
the water from Lake Vyrny to the filters, constant trouble has 
been caused by the formation of a mucilaginous layer which 
seriously reduces the calibre of the pipes. In the deposit thus 
formed Kemna states (05, p. 78) that he has found Crenothrix, 
one of the Iron-Bacteria. 

Although De Vries was principally concerned with the study 
of Crenothrix, he did not restrict himself to the consideration 
of the flora of the system. He points out (pp. 34, 35), as the 
result of his observations, that the fauna in the Rotterdam 
water-pipes is closely similar to that which had been found by 
Kraepelin at Hamburg. Among the organisms thus recorded, 
Cordylophora, Paludicella, and two forms of “ Pliumatella”’ [one 
of which may have been F’redericella| were important con- 
stituents. Here, as at Hamburg, deficient filtration was the 
real cause of the trouble. In parts of the system where the 
conditions were favourable, owing to faulty construction of the 
filters, some of the animals occurred in astonishing profusion. 
I may refer specially to certain Crustacea (Asellus and Gam- 
marus), which were found in almost incredible numbers. This 
is brought out sufficiently clearly by the observation recorded on 
p- 65 of the memoir cited, that the bottom of one of the filters 
wes found to be coated by a continuous layer, of almost the 
thickness of a finger, of the feces of Aselluws, while the animals 
themselves occurred in ‘‘ countless millions ” ! 

-De Vries’ memoir gives references to other cases of water- 
works which have at various times suffered seriously from the 
occurrence of Iron-Bacteria or an unduly rich fauna, or both 
combined. Among these [ may mention the “ Berlin water- 
calamity,” in 1878 onwards, which was completely cured by the 
introduction of sand-filtration in 1883 (pp. 9, 14, 15, 37); 
Philadelphia (p. 35); and Lille (p. 37). Further information 
with regard to cases which have occurred in America is given by 
Whipple (10), who mentions Boston, where Polyzoa, especially 
Fredericella and Plumatella, were the most important constituents 
of the fauna. Other cases recorded are Brooklyn (p. 169) and 
Henderson, N.C. (p. 123). In addition to the genera of Polyzoa 
which have already been mentioned as occurring in drinking- 
water, Whipple gives Pectinatella and Cristatella. 

There is some reason to believe that the Corporation of Man- 
chester has suffered from troubles of a similar nature to those 
experienced at Hamburg and Rotterdam. Hickson (04, p. 675) 
has stated that, two years before the date of his Address, the 
mains had become partially blocked by the growth of an organism 
which was described as a “ moss,” but which Hickson had the 


43% DR. S. F. HARMER ON THE 


opportunity of inspecting and thus satisfied himself that it was 
really composed of Polyzoa. 700 tons of this “moss” are said 
to have been removed from the mains by an expensive and 
laborious process ; but the subject was not investigated biologi- 
cally, and I have been unable to ascertain anything more with 
regard to the subsequent history of this case. 

At Paris (cf. Kemna, 05, p. 26) the Molluscan fauna of the 
water-pipes was investigated by Locard, who recorded 44 species, 
belonging to 13 genera. 

At Ypres, in Belgium (Kemna, 05, p. 63), a quantity of shells, 
with masses of Polyzoa and innumerable Infusoria, could easily be 
obtained by opening a hydrant in the streets. 

Mr. 8S. C. Chapman, M.Inst.C.E., the Water Engineer to the 
Borough of Torquay, to whose kindness I am indebted for much 
information on the subject of waterworks, is preparing a paper in 
which he proposes to give an account of the troubles which have 
been experienced in the district supplied by-the system under his 
own charge, and of the result of the enquiries which he has made 
of the Engineers of other waterworks, at home and abroad. ‘This 
paper is expected to appear in the ‘Transactions of the Institu- 
tion of Water Engineers.’ I believe Lam right in saying that it 
will contain valuable information tending to prove the correct- 
ness of the view, which is held by nearly everyone who has studied 
the subject, that the troubles which are caused by the occurrence 
of a rich pipe-fauna can be got rid of by the introduction of an 
efficient system of filtration. 

Enough has been said to show that the occurrence of conditions 
favourable to the growth of Polyzoa and other organisms in the 
pipes of a water-supply is not a matter to be treated lightly. A 
study of the literature shows, moreover, that the Biology of 
waterworks is a subject of great complexity and of very special 
scientific interest. An excellent general account of the problems 
involved has been given by Dr. A. Kemna (99, 05), who speaks 
with special authority since, in addition to his scientifie qualifica- 
tions, he has had the advantage of being the Manager of an 
important Water-Company which supplies the town of Antwerp. 
As the subject is one which has not attracted much attention in 
Zoological literature, | may perhaps be excused for adding a few 
remarks, based principally on Kemna’s account (05), on the 
subject of sand-filtration, which 1s so closely connected with the 
practical side of the question. 

The introduction of sand-filters was due to James Simpson, 
Engineer to the Chelsea Company, who gave evidence on the 
subject before a Royal Commission on the Water-Supply of 
London in 1827-1828, and before a Committee of the House 
of Commons in 1828. Kemna comments (p. 109) on the fact 
that although Simpson’s views on the theory of sand-filtration 
were not altogether sound, he nevertheless succeeded at his 
first attempt in introducing a system which has not been 
materially improved on since. 


POLYZOA OF WATERWORKS 433 


In this system the water is introduced into shallow filtering- 
beds, which are of large superficial area and have a bottom of 
sand through which the water passes before it is admitted to the 
pipes. The efficiency of the filter does not depend on a mechanical 
straining action of the sand, but is a biological process, princi- 
pally confined to the surface-layer of the sand. In this layer 
grows an abundant flora, consisting of Alge and Bacteria; and 
the beneficial action is the result mainly of the presence of 
Diatoms. By means of the slimy or gelatinous investment 
formed by these organisms, the intervals between the sand- 
grains are filled up, thus increasing the effectiveness of the 
filter as a strainer; while the surface-tilm of living organisms has 
a very marked effect in reducing the number of Bacteria, and 
probably affects the character of the water in other ways. As 
the filter continues in action, the number of organisms in its 
floor increases and the surface-film grows in thickness in a corre- 
sponding proportion. The filtering surface also tends to become 
choked by the accumulation of detritus which has been strained 
off, until from these causes it ceases to transmit the water so fast 
as is necessary. The filter then has to be put out of action and 
the surface-film must be removed. The film has to be allowed to 
re-form for two or three days before the filter is in a condition 
to be used once more. . 

The organisms which are of importance in the filtering process 
undergo seasonal variations. Thus at one time of year Kemna 
finds that one species of Diatom is dominant, while at another 
time the effectiveness of the layer depends on some other species 
or group of species. The filter thus needs constant observation, 
since it may readily get out of working order. An interesting 
cease is recorded by Kemna (99, p. 48), who found that Dipterous 
larve (Chironomus) may have an important practical effect. In 
the observation alluded to, the surface-film was inhabited by 
numerous larve, which were living in tubes constructed out 
of sand-grains. So long as the larve inhabited their tubes the 
filter worked well. But when they left the surface-film in order 
to undergo their metamorphosis, their empty tubes formed a 
number of perforations which were naturally not closed by 
Diatoms, ete., and the filter then ceased to work properly. 
Information that the insects were undergoing their meta- 
morphosis was given by the appearance of Swallows circling 
rouud the filters in order to feed on the emerging Diptera. On 
noticing this indication the filter was put out of use and given 
three days’ rest, during which the holes left by the emergence 
of the Chironomus became closed by a new growth of the 
surface-film. 

The practical importance of sand-filtration is well brought out 
by the facts which are cited by Kemna in the two memoirs 
already referred to. One of the most instructive cases is the 
history of the Hamburg Waterworks during the years which 
preceded Kraepelin’s investigation of the pipe-fauna. In 1842 


434 DR. S. F. HARMER ON THE 


a disastrous fire destroyed a large part of the city of Hamburg, 
including the buildings which contained the machinery of several 
distributions of water. The Municipality thereupon took ovei — 
the supply of water; and it is satisfactory to learn that the 
‘English Engineer, Lindley, who is described as the author of the 
pr oject, insisted on having an installation of sand-filters. Partly 
owing to financial Aieniniog! and partly in consequence of the 
objections raised by the manufacturers and vendors of domestic 
filters, Lindley’s recommendation was not followed for many 
years. At last, however, the difficulties were overcome, and the 
construction of filters, which were to be completed in 1894, had 
commenced in 1890. Shortly after this date the water taken 
from the Elbe became contaminated by a temporary encampment 
of Russian emigrants, and cholera broke out in August 1892 
before the completion of the new works. This epidemic, which 
claimed 10,000 victims, was clearly traced to the defect in the 
water-supply, since the neighbouring town of Altona, which took 
its water from the same source but filtered it before using it, 
remained unaffected. The cases which actually occurred in 
Altona were traced to the use of Hamburg water ; while blocks 
of houses in Hamburg which were supplied from Altona escaped 
infection. Mr. 8. C. Chapman informs me that he has heard 
from Professor Kraepelin that the introduction of filtration, 
“through extensive gravel-beds,” was almost immediately suc- 
cessful at Hamburg; the pipe-fauna dying from starvation in 
the course of a few months after the installation of the filters. 

It might naturally be anticipated that the conditions in the 
reservoirs and filter-beds (in the layer of water above the sand) 
would be favourable for the growth of numerous freshwater 
organisms. ‘These are the sources from which the pipes may 
become colonised, if not protected by filters; and it is thus of 
interest to notice that the facts are in accordance with expectation. 
A case of this kind is recorded by Hickson (04, p. 675), where 
great trouble was caused in the supply of the town of Burnley 
by the occurrence of enormous numbers of Limnea peregra 
in the Hecknest reservoir. An even more striking instance is 
described by Kemna (99, pp. 47, 48) from his own experience 
in the Antwerp reservoirs, in 1896. In this case the trouble 
was caused by the extraordinary multiplication of Cladocera, 
which were so numerous that it was necessary to strain all the 
water passing to the filters through screens of wire-gauze. So 
great was the number of these small Crustacea that it was found 
necessary to keep six men constantly employed, night and day, 
in changing the strainers. The quantity of Crustacea thus 
removed was estimated by Kemna at ten tons at least. 

Another striking case, due, however, to defective construction 
of the filters, is tho one recorded by De Vries (90, pp. 56 e¢ seq.), 
where enormous numbers of Gammarus and Asellus occurred in 
the filter-beds and other parts of the system. 

It need not be a matter for surprise if a particular system should 
be found to have been free from trouble due to Polyzoa and other 


Dr 


POLYZOA OF WATERWORKS, 435 


organisms for a long period, and then to have become suddenly 
infected. It is at least theoretically possible that Polyzoa are 
absent in certain waterworks merely because no statoblast or 
embryo happens to have found access to the system; even though 
there may be nothing to prevent these organisms from flourishing 
if they succeed in effecting an entrance. The means of distribu- 
tion of freshwater organisms have frequently been discussed. 
Darwin gives important information on the subject in ‘The 
Origin of Species’, Ch. xiii :—with particular reference to dis- 
persal by water-birds and insects. Other facts of the same 
nature are recorded by H. Wallis Kew, in ‘The Dispersal of 
Shells’ (Internat. Sci. Series, 1893). A short general discussion 
of the subject was contributed by O. Zacharias to the ‘ Biologisches 
Centralblatt,’ ix. Nos. 3 and 4, April 1889. Reference may also 
be made to B. M. Puschkarew’s recent paper, ‘‘ Ueber die Ver- 
breitung der Siisswasserprotozoen durch die Luft” (Arch. f. Pro- 
tistenkunde, xxvii. Heft 3, 1913, p. 323). 

It can hardly be doubted that the method by which Polyzoa 
most commonly colonize a new place is by means of their stato- 
blasts, which from their habit of floating at the surface are 
extremely likely to adhere to some part of the lees, beak, or 
feathers of a wading or swimming bird, or to some part of a 
water-insect. De Guerne (88), struck by the occurrence in the 
Azores of a lacustrine fauna almost entirely composed of European 
types, was led to examine the material adhering to the feet of 
Ducks; and on one occasion (p. 297) he discovered a half stato- 
blast of a Plumatella on the web of the foot of a Wild Duck. 
Similar positive information with regard to the dispersal of fresh - 
water Polyzoa has been published by Garbini, in a paper which I 
have only been able to consult by means of the notices given in 
Vol. ii. (p. 195) of the ‘ Zoologisches Centralblatt’, and (by Matz- 
dorff) in the “Jahresbericht wber die Bryozoen fiir 1897 und 
1898” in the ‘Archiv fiir Naturgeschichte, lxi. Jahrg., ii. Bd. 
Heft 3, p. 181. It appears from the latter notice that stato- 
blasts of a Plumatella were found attached to the beak of a 
Heron (“ Reiher”’). 


IJ. Occurrence of Polyzoa in British Waterworks. 


I turn now to the consideration of the cases which have come 
under my own observation in this country. I am able to record 
five cases, in very different parts of Great Britain, in all of which 
serious trouble has been experienced. I commence by giving 
some account of the occurrence of the organisms, reserving the 
question of the determination of the specimens to Part IIT. of 
this paper. The cases are as follows :— 


(1) Torquay AnD Newron Apzot. 


Towards the end of September 1910, I received samples of a 
Plumatella (P. emarginata, var. muscosa) from Mr. Samuel 


Proc. Zoou. Soc.—1913, No. XXX, 30 


436 DR. S. F. HARMER ON THE 


C. Chapman, M.Inst.C.E., Water Engineer to the Borough of 
Torquay. I am greatly indebted to Mr. Chapman for the 
trouble he has taken in giving me the fullest information with 
regard to this particular occurrence, and in calling my attention 
to similar instances in other places. 

The Torquay case is one of the usual kind, in which no efficient 
system of filtration bad been in operation. In spite of this fact 
no trouble had been given by the growth of Polyzoa or other 
organisms, in sufficient quantity to give rise to anxiety, during 
fifty years. The apparently sudden invasion may perhaps have 
been due to the fact that the Polyzoa in question had not 
previously effected an entrance into the system. The Pluwmatella 
was first observed in pipes which had been in use a little more 
than two years. The pipes were at the bottom of a steep hill, on 
a trunk-main which supplied a large part of Newton Abbot, and 
where the flow of water was constant and of considerable velocity. 
The Polyzoon was soon traced in every direction all over the 
system, both at. Torquay and at Newton Abbot. It quickly 
began to give rise to serious inconvenience, by becoming detached 
from the walls of the pipes and by being carried to the ball-taps 
and the strainers of meters, which became blocked and were thus 
thrown out of work. The trouble became specially acute imme- 
diately after a frost in the early part of 1912. This is entirely 
in accordance with expectation ; the Plwmatella which had been 
growing on the walls of the pipes having no doubt broken up 
during the cold weather, its branches having then passed into the 
flow of water and so having given rise to the choked taps and 
meters which were reported in large numbers. Jn one case men- 
tioned to me by Mr. Chapman, eleven houses in one block of 
buildings were without water owing to this cause. It need 
hardly be remarked that the well-known habit of dying at the 
end of the year gives these freshwater Polyzoa special advantages 
in distributing themselves over a wide area of a system. It is 
only necessary for statoblasts or fragments of branches containing 
them to remain attached to some irregularity on the inside of a 
pipe to make it practically certain that some of them will have 
the opportunity of germinating in the next period of warm 
weather. 

No complete examination of the pipe-fauna has been made in 
the Torquay Waterworks, but a few other animals have been 
observed in them. One of the most conspicuous of these is a 
Freshwater Sponge, which has been found growing on the insides 
of some of the pipes, and has been determined by Mr. R. Kirk- 
patrick as Hphydatia fluviatilis. In some of the samples there 
were large quantities of a Dipterous larva, of a bright red colour. 
These have been determined by Mr. F. W. Edwards as a species 
of Chironomus. The majority of these larve, and perhaps nearly 
all of them, could hardly have succeeded in completing their 
metamorphosis; and it seems probable that, as in the case of the 
Dipterous larve recorded by Kraepelin (85, p. 11), the trans- 


POLYZOA OF WATERWORKS. 435 


formation into the imago, or at least the emergence of the adult 
form into the air, could not take place except by some chance 
combination of circumstances which would hardly occur very 
often. It may be noted that Kraepelin, in commenting (p. 11) 
on the absence of Chironomus-larve from the underground pipes 
of the Hamburg system, points out that, according to his obser- 
vations, these larvee are the most dangerous enemies of freshwater 
Polyzoa, whose delicate branches they tear to pieces. This might 
have an important bearing on the distiibution of the Polyzoa, 
since, even in the most active period of their growth, branches 
containing statoblasts would be liberated by the Chironomus- 
larve, and some of them would come to rest in places where a 
new growth of the Polyzoa would be started. 

In at least one of the samples, considerable numbers of a small 
freshwater Oligochete were found; but [ am not able to deter- 
mine it from the preserved specimens in my possession. 

From the descriptions which have been given to me by 
Mr. Chapman, there can be no doubt of the occurrence in the 
Torquay system of small Gasteropods (perhaps Dithynia or 
Limnea, and almost certainly Planerbis), and of Asellus, Gam- 
 marus, Cladocera, and Copepoda. 1 have not had the opportunity 
of examining specimens of any of these animals. 

I have no certain evidence of the occurrence of [ron-Bacteria, 
but the appearances suggest that these occurred. In some places 
the pipes were found to be coated with a thick black layer of 
“‘ neaty substance,” cemented by iron-compounds. My colleague 
Mr. A. Gepp, who has kindly examined one of the samples, tells 
me that its condition is not such as to make the demonstration 
of Jron-Bacteria certain, but from his experience of other similar 
cases, he has little doubt but that these had really been present. 

It may be remarked that the Torquay authorities have now 
instituted a filtration-system ; and, judging from the experience 
of other water companies, it may be presumed that the trouble 
from which they have suffered will soon be a thing of the past. 


(2) Lorp Powis’s StycHEe Esrare (at Lydbury, North Shrop- 
shire, near Market Drayton). 


In May 1912 I received information, through Dr. Augustus 
Voelcker, of the Royal Agricultural Society, of the occurrence of 
a Polyzoon which had been giving serious trouble near Market 
Drayton. I later received specimens, in the fresh condition, 
through the kindness of Mr. hk. H. Newill, Lord Powis’s agent. 
The specimens 1n question were well-grown examples of Pluma- 
tella fungosa, var. coralloides. In this case a feed-tank received 
its water from a small open pool in which the Polyzoon was 
proved to be growing. ‘The water passed to the tank through a 

strainer, vahrcr did no more than filter off the coarse detritus. 
From the tank the water passed down an incline, in an under- 
ground pipe, to a ram-chamber situated about thirty-three yards 


30* 


438 DR. S. F. HARMER ON THE 


away. The ram isa pump which raises spring-water from another 
source for use in the house and other buildings. From infor- 
mation supplied by Mr. Newill, the Plumatella is found in the 
open pool (from which the sample examined was obtained), in 
the copper strainer at the beginning of the pipe from the pool to 
the tank, in the same pipe, in the feed-tank, in the pipe leading 
from the tank to the ram, and in the ram-house. ‘The middle 
parts of the pipes were said to be less affected than the parts 
near the tank and the ram. Considerable trouble had been 
experienced in 1911; and it is expressly pomted out that the 
Polyzoon ceases to grow about August. In May 1912, when I 
received the sample, the Polyzoon was in vigorous growth. ‘The 
nature of the trouble was of the same kind as at Torquay, and 
consisted in the choking of the pipes and in the blocking of the 
ram to such an extent that it ceased to work. Unsuccessful 
attempts had been made to check the growth by the use of copper 
sulphate. With the Polyzoon were found multitudes of bright 
red “‘ worms,” of a greenish hue when young and dark red when 
mature. They are described as being marked by dark bands, as 
growing to a length of one-half to three-quarters of an inch, and 
of having the habit of changing their skins, “ which float on the 
water.” From these indications there is a high probability that 
the animals in question were Dipterous larve, probably of some 
species of Chironomus, as in the case of the Diptera found at 
Torquay. 

Mr. Newill informs me that sand-filters will be installed in the 
spring of the present year. 


(3) Bartey, 


The first intimation of trouble at Batley was received from the 
Director of the Royal Botanic Gardens, Kew ; but I later obtained 
samples from Mr. J. C. Barrowclough, the Waterworks Engineer 
at Batley, who kindly gave me information on the subject. The 
Polyzoa sent belonged to Fredericella sultana, and they had been 
responsible for trouble of the usual kind—-namely, blocking of 
the meters, strainers, etc. In the sample examined the meter 
had been fixed on Aug. 19, 1912, and was removed on Oct. 16 
completely choked by branches of the Mredericella. The supply 
of these waterworks is obtained from a moorland gathering 
ground on the Pennine Chain, “having large impounding 
reservoirs, but no form of filters.” The trouble had been known 
to exist for a long time, but it had been ascribed to peat and 
other deposits. An examination of the trunk-main had recently 
been made on account of its reduced capacity through incrus- 
tation. The large quantity of iron-compounds found in a dry 
sample of this inerustation points to the presence of Ivon- 
Bacteria, though no certain evidence of their presence was dis- 
covered. It is stated by Mr. Barrowclough that the autumn is 
the time when special trouble is experienced. As in the other 


POLYZOA OF WATERWORKS. 439 


cases, this is no doubt due to the fact that that time of year 
is the period at which the tubes of Polyzoa naturally disin- 
tegrate, with the result that branches are set free and choke 
the meters. 


(4) 

At the end of October 1912 I received, through Mr. 8. C. 
Chapman, a sample from an English waterworks installation, the 
locality of which I am not permitted to mention. It consisted, 
in the main, of a mass of tubes of Paludicella articulata. The 
polypides had already disappeared owing to the lateness of the 
season; but the sample was a particularly beautiful specimen, 
covered with a profusion of hibernacula or winter-buds, which 
were developed on every branch. Among the branches of the 
Paludicella there were disintegrated fragments of Fredericella 
sultana. Mr. Chapman informs me that serious trouble had been 
experienced for some time at the locality in question, although 
the nature of the organisms causing it had not been recognized. 


(5) ABERDEEN. 


At the end of March, in the present year, I received a sample, 
very similar to the last one, from the Aberdeen Waterworks, 
again through the kindness of Mr. Chapman. This consisted 
principally of a tangled mass of Paludicella articulata, composed 
of empty zoccia bearing numerous hibernacula. The sample 
also included a few pieces of Spongilla lacustris (determined by 
Mr. R. Kirkpatrick), crowded with gemmules, a few young speci- 
mens of Limnea peregra (determined by Mr. G. C. Robson), and 
some Dipterous larve. ‘These were referred by Mr. F. W. 
Edwards to a species of Chironomus, which was, however, not 
the same as the species found at Torquay. There were also a 
number of tubes, formed of cemented mud, which appear to 
have belonged to the Chironomus-larvee. 


The following case may also be mentioned, because, although 
not illustrating the occurrence of Polyzoa in waterworks, it has 
an obvious bearing on the importance of filtration :— 


(6) Hampron-on-T'HAMES. 
On May 22, 1912, ‘The Daily Mail’ published a short account 


of the occurrence of enormous numbers of ‘ freshwater mussels ” 
in a 36-inch water main at Hampton-on-Thames. It was stated 
that 90 tons of these animals had been removed from a length of 
a quarter of a mile of this main, the diameter of which had been 
reduced by them from 36 inches to 9 inches. The “ mussels ” 
were examined by Mr. G. C. Robson, who found them to belong 
to Dreissensia polymorpha, a Lamellibranch which has been 
recorded in the pipes of the Hamburg water-supply and elsewhere. 
Mr. H. B. W. Stent, A.M.Inst.C.E., the Resident Engineer of 


440 DR. S. F. HARMER ON THE 


the Metropolitan Water Board, has kindly given me some addi- 
tional information with regard to this case. Without certifying 
the exactness of the above recorded computation of the weight 
of the Molluses removed, he admits the existence of large 
numbers of these animals in the main. He emphasizes the fact, 
however, that no trouble of this kind has been experienced in 
the water which has been admitted to the pipes after filtration. 
The Molluses are found only in the mains which are used for 
conveying water from the Thames to the storage reservoirs, or 
from the reservoirs to the filter-beds. The importance of filteri ing 
the water before admitting it to the pipes is admirably br ought 
out by this case, since the Molluscs in question have been unable 
to get past the barrier of the filter-beds. 


The consideration of the above cases is sufficient to show that 
the risks which might be caused by the entry of animal life into 
the pipes of a water-system are not confined to any particular 
part of the country. The cases which I have examined come 
from Aberdeen in the north to Torquay in the south-west. It 
is obvious that if the water is unfiltered and thus contains micro- 
scopic organisms in sufficient quantity, the chance introduction 
of a few Polyzoon statoblasts or embryos might be followed by a 
growth in the pipes sufticient to interfere very seriously with the 
circulation, and ultimately to choke the taps. It is important 
to notice, as has been so well shown by Kraepelin, that the 
Polyzoa play a leading part in promoting the growth of animal 
life in the pipes. They are specially adapted by the disposition of 
their ciliated tentacles to catch the diatomsand other microscopic 
organisms on which they habitually feed, or, as Kemna* has 
expressed it, they are ‘‘ micr: ophagous.” They serve in their turn 
as the food of larger animals ; and, unless they prepared the food 
in this way, it is hardly likely that animals such as Eels would be 
capable of flourishing in the pipes of a water-system. The great 
quantity of Polyzoa found at Hamburg, Torquay, and elsewhere 
justifies the conclusion that in any discussion of this question 
the occurrence of Polyzoa is a fact of cardinal importance. 
Sponges are perhaps the organisms which rank next in im- 
portance to the Polyzoa in this respect. 

The practical and theoretical interest of this subject has 
received very little attention in this country. Professor S. J. 
Hickson, in his Presidential Addiess to Section D of the South- 
port Meeting of the British Association (04, p. 675), has, how- 
ever, used words to which I can completely subscribe. He states 
“that our ignorance of the general balance of animal and vege- 
table life in the large reservoirs is profound, and that a systematic 
inquiry conducted by competent persons would most certainly 
lead to knowledge which would be of great scientific importance, 
and in the long run remunerative to the community.” 


* Kemna, A., “ Octaenemus, Une Ascidie mégophage,” Ann. Soc. roy. Zool. et 
Malacol. de Belgique, xli. 1906, p. 57 


POLYZOA OF WATERWORKS. 44] 


It is a curious coincidence that, at the same Meeting of the 
British Association at which Professor Hickson’s address was 
delivered, the Presidential Address of Mr. Charles Hawksley to 
Section G (Engineering) dealt specially with water-supply and 
the construction of waterworks. In this address it is stated 
(Report, p. 767) that “most waters obtained from gathering 
grounds are much improved by filtration”; and the mode of 
construction of sand-filters is indicated. But there is no refer- 
ence to the importance of the filter as a means of depriving the 
water of the microscopic organisms which would otherwise supply 
nourishment to Polyzoa and other ‘“ microphagous” animals 
capable of flourishing in the pipes if they are supplied with 
appropriate food im sufficient quantity. 

I am indebted to Mr. 8S. C. Chapman for having put at my 
disposal the result of an extensive enquiry which he has made as 
to the occurrence of Polyzoa and other animals in the waterworks 
of this country. As he proposes to publish the result of his 
enquiries J will not attempt to anticipate him; but he returns 
show that organisms of the kind indicated occur more frequently 
than has generally been supposed, and that the occurrence of 
trouble due to their presence can usually be traced to the want 
of an efficient system of filtration. 


III. Species of Polyzoa found in British Waterworks. 


The species of Polyzoa which I have had the opportunity of 
examining may be referred to the following four species :— 


(1) Paludicella articulata Khrb. 

(2) Fredericella sultana Blamenbach. 

(3) Plumatella fungosa Pall., var. coralloides Allman. 

(4) Plumatella emarginata Allman, var. muscesa, Kraepelin. 


I subjoin a few notes on the synonymy of the species and on 
the specimens which form the subject of this paper. 


(1) PatupiceLia arTicuLATA Khrb. (PI. LXIT. figs. 1-10.) 


Alcyonella articulata Ehrenberg, ‘‘Symbole Physice,” “ Ani- 
malia Evertebrata,” Dec. 1, ‘‘ Phytozoa Polypi,” 1831, fol. a. 

Paludicella Ehrenbergii Dumortier & Van Beneden, ‘“ Hist. 
Nat. des Pol. Comp. d’eau douce,” 11. Partie, Nouv. Mém. Acad. 
Bruxelles, xvi. 1843, p. 38. 

Although Allman, in his classical monograph (56) on the 
Freshwater Polyzoa, accepted the specific name ehrenbergi—and 
has been followed in this respect by many other writers—the 
reasons given by Dumortier and Van Beneden for rejecting 
Ehrenberg’s name are not in accordance with modern principles 
of zoological nomenclature. They state as their motive for intro- 
ducing a new name that “ articulata” refers to a generic character, 
and therefore cannot be used as a specific name. The invalidity 


449, DR. S. F. HARMER ON THE 


of this argument has not escaped the notice of some other 
writers. 

The first specimens examined were received from a waterworks 
system (No. 4, above), which does not permit itself to be men- 
tioned by name, at the end of October 1912. They were remark- 
able for the extraordinary freedom with which hibernacula were 
developed. In correlation with the lateness of the season, the 
polypides had degenerated, and the zocecia were represented 
merely by their empty ectocyst. Nearly every zocecium, through- 
out the whole of the material, bears one or two hibernacula im 
the place of the ordimary lateral daughter-zoecia; and, if it 
happens to be at the end of a branch, a terminal hibernaculum 
as well. These hibernacula (Pl. LXII. figs. 1-10) are, almost 
without exception, of an elongated fusiform shape, although 
varying much in length and, to a jess extent, in breadth. They 
thus differ strikingly from most of the figures of these structures 
which have previously been published. 

The hibernacula were originally described, under the name of 
‘“hybernacles,” by Dumortier and Van Beneden, in the memoir 
cited above (p. 51, pl.i. fig. 1”; pl. ii. figs. 24— 35). Although 
shown as replacing levherell bale, and ahhenefione in the same 
position as in the specimens which have come under my notice, 
they are described and figured as short bodies, not more than 
about one-fifth of the length of the zoecia. They are pointed at 
their free end, and were said to be always strongly compressed. 
In colour they were greyish black. 

Kraepelin, who states (87, p. 76) that the hibernacula do not 
appear to have been found by any observer since Van Beneden, 
describes and figures them (pl. iv. fig. 117) as swollen and ellip= 
soid or irregular, and states that iley are yellowish in colour, 
and that they have a considerable amount of calcareous matter 
in their cuticle. Several of the specimens figured by Kraepelin 
resemble the hibernacula described by Van Beneden in their 
form, but one or two of them are fusiform, although not so elon- 
gated as most of the hibernacula which have come under my own 
observation, Some of them remain attached to the substratum 
when the zocecia, which have meanwhile lost their contents, dis- 
integrate and break up. In the second part of his monograph 
(92, p. 61) Kraepelin refers to the irregular form generally 
possessed by the hibernacula. He co the occurrence in 
them of an elongated polypide-bud (pl. v. fig. 163), which is 
formed early in their development, in re ee for germination 
in the ensuing spring; and he mentions the yolk- like material 
by which the polypide-bud is surrounded. He was unable to 
discover the mode of formation of the hibernacula. 

Levinsen (94, p. 85, pl. viii. figs 24-26) gives figures of the 
hibernacula which are much like those of Kraepelin. 

Wesenberg-Lund ($6, pp. 321, 363, xxiv; pl. iv, figs. 44, 45) 
found the hibernacula in large numbers i in October. They were 
all of the same form and colour as in Kvraepelin’s description, 


POLYZOA OF WATERWORKS. 445 


and were attached to the substratum, none being discovered on 
the erect shoots. Their mode of development was not ascertained. 

Chirica (04, pp. 7, 8, sep.) does not describe the development 
of the hibernacula, but records one or two interesting observa- 
tions. He obtained them, not only at the beginning of the 
winter, but also attached to branches of dead zocecia, floating at 
the surface of the water in April, and he regards this as the 
means by which the species is distributed in the spring. He 
adds that colonies which have resulted from the germination of 
hibernacula develop ovaries and testes at once (cf. also Kraepelin, 
87, p. 86). 

The hibernacula of the material examined by myself are, as a 
rule, very different in form from those which have been described 
by Dumortier and Van Beneden, Kraepelin, Levinsen, and 
Wesenberg-Lund. Although varying much in their proportions, 
the great majority have an elongated fusiform shape, as shown 
in the figures on Pl. LXII. It does not, however, seem to me 
necessary to conclude that there is any specific difference between 
the form examined by me and those of which hibernacula have 
previously been described. Although Paludicella is a light- 
shunning organism (Allman, 56, pp. ae rhis 5), these specimens, 
obtained from the interior of the pipes of a waterworks system, 
must have been growing in absolute darkness, and under condi- 
tions which were otherwise different from the normal habitat of 
the species. It would thus not be surprising if the hibernacula 
were found to show some differences from those growing in a 
normal environment. The conclusion that the specimens under 
consideration should be referred to P. articulata is confirmed by 
the fact that a single branch observed bore two hibernacula (one 
of them shown in fig. 1) which have the form described by 
Dumortier and Van Beneden. 

It remains to be seen, however, whether the hibernacula 
usually developed by Paludicella in this country are typically 
spindle-shaped or not. Mr. C. F. Rousselet has kindly allowed 
me to examine specimens of hibernacula, collected in Norfolk 
by Mr. H. E. Hurrell, in his own collection. These hibernacula 
are all of the general forin described below, although they are on 
the average rather less elongated than the specimens which have 
come under my own observation. Some of them have germinated, 
in the manner described by Dumortier and Van Beneden and 
by Kraepelin. The distal end of the hibernaculum splits into 
two valves, in order to allow of the escape of the tissues of the 
young zocecium which grows out of the hibernaculum. It is of 
some interest to notice, in connexion with what is said below, 
that the stall of the hibernaculum remains unsplit during the 
process of germination. 

Mr. Rousselet informs me that he has never seen hibernacula 
of any other form than the spindle-shape here described. 

The hibernacula observed by me are white in colour, although 
some of them show signs of becoming darker. The material was 


444 DR. S. F. HARMER ON THE 


obtained towards the end of October, and therefore soon after the 
hibernacula had been formed. It is not improbable that they 
would all have become darker as the season progressed. All were 
growing on erect branches of the colony, and most of them were 
quite free, although some show the habit which has been described 
by other observers of attaching themselves to foreign objects ; 
in this case zocecia, or even other hibernacula, of the Paludicella 
itself, or zocecia of the Fredericella with which the species was 
associated. Allusion has already been made to the fact that one, 
two, or even three hibernacula are borne on nearly every zocecium. 
They vary much in length and to a less extent in breadth. 
Leaving out of account the two hibernacula of which one is 
shown in fig. 1, they are nearly all spindle-shaped, although 
occasionally showing some irregularity of form, and vary in 
length from 608 pw (fig. 4) to about 1968. They are usually 
pointed at their free end, the broadest part being as a rule beyond 
the middle of their length. ‘The proximal stalk-like portion is 
generally slender, but varies in breadth from about 72 p to 96 p. 
Measured at their widest part, the width varies from 144 p to 
160. They are usually developed in the position occupied by 
either the lateral or the terminal daughter-zocecia ; but, as shown 
in fig. 7, the hibernaculum may originate more proximally than 
the position normally occupied by a lateral bud; and the bases 
of the lateral hibernacula are not always opposite one another 
(fig. 8). Most of the hibernacula contain an elongated polypide- 
bud, in agreement with Kraepelin’s statement (92), of the form 
which also occurs in the development of the polypide-buds of 
ordinary zoecia. J have not observed any calcareous matter in 
their wall. 

It has often been suggested that the statoblasts of Phylolacto- 
lemata may have been evolved from structures like the hiber- 
nacula of Paludicella (cf. Kraepelin, 87, pp. 163, 167; 92, p. 62). 
But while the mode of origin of the statoblasts, from the funi- 
culus, is well understood in its general outlines, I have not been 
able to discover any account of the development of the hiber- 
naeula, The material at present under discussion enables me to 
make a contribution to this subject, although the facts could have 
been interpreted with more certainty if the specimens had been 
collected slightly earlier in the year. There is, however, enough 
evidence to show that the hibernaculum is not a modified terminal 
yoecium, as is assumed by Braem (90, p. 112) in criticizing 
Kraepelin’s opinion, but is something which is left when the 
distal part of a zocecium has been thrown off. The evidence of 
figs. 2, 3, 9, 10 seems to be unmistakable on this point. In these 
cases the hibernaculum is well formed, with its thick cuticle and 
inclusions of yolk-like material. But although its stalk appears 
to be constituted, in some cases at least, by the proximal part of 
the zocecium, the distal end of the hibernaculum lies freely in the 
cavity of the zocecium, which is prolonged in its normal form 
beyond it. The interpretation which naturally suggests itself is 


POLYZOA OF WATERWORKS. 445 


that the formation of the hibernaculum is inaugurated by a con- 
traction of the endocyst or body-wall from the original ectocyst 
of the zocecium, in the distal part at least, and by the deposition 
of a new cuticle over that part of the hibernaculum which lies 
freely inside the old ectocyst. In some cases, as in fig. 10, 
the hibernaculum has a wall of its own which is distinct 
throughout from the original ectocyst. In the specimen shown 
in fig. 2 a single zoccium has developed two hibernacula, the 
first of which is attached to the parent-zocecium in the position 
of a terminal bud, while the second, or distal hibernaculum, lies 
freely in the cavity and has no base of attachment. It can 
hardly be doubted that the hibernacula represented in figs. 3, 9, 
and 10 would have assumed the appearance shown by most of the 
hibernacula in this material by the loss of the empty caps formed 
by the zocecia within which they have severally been developed. 
IT vegard it, therefore, as probable that the hibernaculum owes 
its terminal position to the loss of the distal part of its own 
zocecium, which, as shown in fig. 10, is not necessarily the 
terminal individual of a branch. 

There are, however, certain appearances, shown in figs. 6-9, 
that are in need of explanation. In these cases most of the 
ectocyst of the zocecium which precedes the definitive hiber- 
naculum has split into two valves, in the manner characteristic 
of a germinating hibernaculum. Like the other zocecia of this 
material, these zowcia are destitute of cellular contents ; the only 
living tissues left in the colonies being those which constitute the 
hibernacula. It is important to consider whether the bivalve 
arrangement indicates that a process of germination has actually 
taken place. In view of the close resemblance between these 
zoecia and a hibernaculum germinating after the end of the 
winter and arrived at the stage of having completed the formation 
of the first regenerated zocecium, it seems legitimate to assume 
that germination of a hiber naculum has Gcourred in spite of the 
fact that the material was collected in the autumn. It would 
appear that a new zocecium has been completely formed, since 
the remains of its tubular orifice are present. But the develop- 
ment of a new colony has stopped short with the formation of a 
single zocecium, and a second process of hibernaculum-develop- 
ment has taken place, resulting in the definitive winter-buds. 
The living material of the regenerated zocecium has passed into 
the hibernaculum, or hibernacula, which have been formed from 
it, though there is no evidence to show whether the transference 
of the material took place en masse or by the more gradual process 
by which a bud is ordinarily supplied with its living contents. 
It is perhaps not impossible that no polypide was formed by the 
regenerated zocecium, but that on the splitting of the hiber- 
naculum which preceded it the contents of that structure passed 
bodily into the newly-formed hibernaculum. Some such trans- 
ference of cellular material may be indicated by fig. 10, where 
each of the hibernacula is seen to traverse the septum which 


44.6 DR. S. F. HARMER ON THE 


occurs between two zowcia. In fig. 8 the definitive hibernaculum 
may be proved by measurement to be of exactly the same length 
as the hibernaculum which is indicated by the split valves in the 
wall of the preceding zoccium. The material of the first hiber- 
naculum would thas! just fill the cavity of the second one. In 
fig. 9, however, the second hibernaculum is distinctly smaller 
than the first one; while in fig. 7, what is assumed to have been 
the first hibernaculum has had to provide the material for two 
new hibernacula. Further observations on material collected at 
a more favourable season will be requir ed to settle these points. 

The great majority of the zowcia in a Paludicella-colony give 
rise to a pair of lateral buds, situated opposite one another and in 
a position which has a definite relation to the orifice of the parent- 
zoecium. It may be significant that in all the cases shown in 
figs. 6-9 there is some abnormality in this respect. In figs. 6 
and 9 only one of the lateral zoccia is indicated; while in fies, 7 
and 8, where two are present, they are not opposite one another, 
and in fig. 7 one of them is much nearer the proximal end of the 
parent- Zo0ecitim than is normally the case. I have observed 
similar irregularities in other zocecia of the same kind which have 
not been figured; and these cases perhaps support the view that 
ger mination of the hibernacula first formed has taken place, since 
it is well known that in young Polyzoon colonies the zocecia first 
developed frequently fail to give rise to the full number of buds 
which are formed by the zocecia in an actively growing branch. 

If the explanation above suggested is correct, the germination 
of the first set of hibernacula may perhaps hese heen. induced by 
the occurrence of warm weather succeeding a cold period, which 
induced the formation of the first set. But the explanation i in 
question is not the only one which can be imagined; and it is 
not inconceivable that as the time of hibernaculum- formation 
approaches, some of the zocecia make an abortive attempt to 
develop hibernacula, and that the bivalve arrangement seen in 
figs. 6-9 implies an unsuccessful effort of this nature. 

In the majority of the zowcia of this material the definitive 
hibernacula have been produced at the first attempt, and the 
zoecia Which respectively precede them show no appearance of a 
bivalved arrangement. But in all cases the zocecia are empty of 
cellular contents, and there has obviously been a centrifugal 
passage of the living material of the colonies into the hibernacula, 
which are finally left as the outermost members of the series. 

It remains to be considered whether these observations have 
any bearing on the question of the homology, if any corres- 
pondence really exists, between the hibernacula of Paludicella 
and the statoblasts of Phylactolemata. It has frequently been 
maintained that statoblasts have been evolved from structures 
more or less resembling the hibernacula of Paludicella, which on 
this view is regarded as related to the ancestral form from which 
the Phylactolemata have been derived. This argument has been 


specially developed by Kraepelin (87, pp. 163, 167; 92, p. 62), 


POLYZOA OF WATERWORKS, 447 


who based his conclusions on the general similarity of the hiber- 
nacula to statoblasts, particularly those of Hredericella in which 
no annulus is present. He did not, however, bring forward any 
evidence to show how the hibernacula are developed. Braem 
(90, p. 112) has disputed Kraepelin’s conclusions, basing his 
objection largely on the assumption that the hibernacula are 
complete, specially modified, terminal individuals of the colony. 
On this assumption there would be a considerable difference 
between them and the statoblasts, which are essentially buds 
produced internally within the body-cavity of a zoccium. 
Braem’s objection loses some of its force if his assumption as to 
their mode of origin can be shown to be incorrect. I think that 
the evidence here brought forward is sufficient to render the 
conclusion that the hibernaculum is typically an end-bud of the 
colony very doubtful. Although the proximal part of its ecto- 
cyst is commonly a modification of the corresponding part of the 
ectocyst of the parent-zocecinm, cases like those shown in figs. 2 
and 10 appear to prove that the hibernaculum may be formed as 
an internal bud-like structure, with a complete cuticle of its 
own; and there is indeed no very essential difference between 
the proximal hibernaculum shown in fig. 10 and the statoblast 
ot a Fredericella. I do not, of course, overlook the fact that the 
statoblasts of Phylactolemata are developed in a peculiar way 
from the funiculus of a polypide. But Paludicella is, in any case, 
only distantly related to the Phylactolemata; and the difference 
between the winter-buds of the two forms is not, apparently, so 
great as has hitherto been assumed by most observers who have 
considered the question. 

When one considers the facility with which freshwater 
organisms, from Protozoa upwards, develop some special cyst- 
like coat which enables them to survive the winter or periods of 
drought, one must no doubt be cautious in assuming that the 
homology suggested is a probable one. But my observations 
seem to show that the hibernaculum of Paludicella, like the 
statoblasts of Phylactolemata, is a special form of winter-bud 
which is developed inside the parent zocecium out of a mass of 
living tissue which becomes surrounded by a chitinous coat ; and 
that thus the two kinds of winter-buds are morphologically com- 
parable. In view of this consideration it does not appear to me 
impossible that the hibernaculum and the statoblast represent 
two different conditions which have been evolved from a common 
starting-point. 

The specimens of this species received from Aberdeen (No. 5, 
above) agree closely with those just described. Numerous hiber- 
nacula of an elongated type are present, although they are 
perhaps more variable in length than in the other sample. 
Some of them are even longer than the longest described above. 

Paludicella articulata las previously been recorded as an 
inhabitant of the pipes of waterworks, both at Hamburg 
(Kraepelin, 85, p. 6) and at Rotterdam (De Vries, 90, p. 25). 


448 DR. S. F. HARMER ON THE 


(2) FReDERICELLA suLTANA Blumenbach. (PI. LXIII. figs. 11— 
14.) 

It is unnecessary for me to discuss the synonymy of this 
Species, since there is a general agreement with regard to it 
among practically all recent writers. Excellent accounts of the 
species are given by Allman (56), Kraepelin (87), and many 
others. It was recorded by Kraepelin (85, p. 6) as one of the 
commonest forms found in the Hamburg Waterworks system ; 
and I suspect that one of the species of “ Pluwmatella” recorded 
by De Vries (90) from the Rotterdam Waterworks may have been 
really Mredericella. 

I have myself found /. sultana in two English waterworks ; 
namely the system (No. 4) in which the Paludicella above described 
occurred and at Batley. In both cases the material was collected 
in October, and the polypides had already degenerated, leaving 
empty tubes containing statoblasts. ‘The tubes show the septa 
which have been described by Allman (56, p. 112), Kvaepelin (87, 
p- 100), and others, at the point where a tube diverges from the 
main stem. The statoblasts are developed in both localities with 
greater profusion than is sometimes the case in this species. 
Kraepelin has commented (pp. 103, 104) on the variability of the 
statoblasts of /. sultana. He states that it is hardly possible to 
find two alike in the same colony; and that they may be reni- 
form, almost quadrangular, or even nearly circular. The average 
length is given by him as 430 p, and the average breadth as 300 p. 
The length varies between 380 and 570; and the breadth 
between 210 p and 370 p. 

Figs. 11-14, taken trom the Batley material, show that my 
own results are in close agreement with Kraepelin’s. The varia- 
tion in form is sufficiently obvious from these figures. The 
average length of nine statoblasts from Batley was 444 py, a 
measurement almost identical with Kraepelin’s. The extreme 
measurements were 416 and 480. The average breadth of 
the same nine specimens was 242 »—somewhat less than Kraepe- 
lin’s figure,—while the extremes were 200 and 272. Some 
were more than twice as long as broad (fig. 14); while the 
statoblast shown in fig. 11 is almost circular. 

Kraepelin states that the funiculus of Yredericella generally 
gives rise to one or two statoblasts, rarely to three. In my own 
material, while many of the zocecia contain a single statoblast, 
and others contain either two or three, one of the tubes in the 
Batley material contains four statoblasts, which certainly belonged 
to one polypide. In several instances it can be seen that the 
statoblasts are closely adherent to one wall of the tube. In cases 
where two tubes are connate, the statoblasts are adherent to the 
common wall which separates the two tubes. In the Batley 
material the statoblasts are almost black in colour, and a large 
proportion of them, although still contained in the zocecia, have 
split into their two valves, between which can be seen a mass of 
living tissue. This isa clear sign that the statoblasts are ready 


POLYZOA OF WATERWORKS, 449 


to germinate. Itis well known that the statoblasts in this species 
ordinarily germinate without leaving the tubes in which they 
have been produced. It need hardly be pointed out that the 
state of the winter-buds, here as in the Paludicella described 
above, is such as to give every opportunity for the dispersal of 
the species throughout the system into which it has effected an 
entrance. In the Batley case at least (and probably in the other) 
the branches examined were not taken from the places where 
they had been growing in the pipes, but had become free by the 
disintegration of the colonies at the approach of winter, and had 


accumulated in the meters and taps to such an extent as to 
block them. 


(3) PLUMATELLA FUNGOSA Pallas, var. coraLLormpes Ailman. 
(Pl. LXIIT. figs. 22—25.) 


I follow Braem (90, p. 5) in regarding Plumatella coralloides of 
Allman as a form of P. fungosa. This seems to be indicated by 
Allman’s fig. 4 (pl. vii.) of the floating statoblasts. It should, 
however, be noted that Annandale, in his recent account of the 
Indian species (11, pp. 217, 218), gives P. coralloides as a synonym 
of P. fruticosa. 

In dealing with representatives of the genus Plumatella one 
is confronted with numerous difficulties; since there is no 
general agreement with regard to the species which ought to be 
recognised. It is worth remembering that Pluwmatella has three 
different kinds of reproductive bodies, namely, the embryos 
produced by the development of the eggs, the floating statoblasts, 
provided with an “annulus” of gas-containing cavities, and the 
sessile statoblasts. We are in almost complete ignorance with 
regard to the question whether the form of the colony depends 
to any extent on the particular kind of reproductive body from 
which it has been developed. As Braem has pointed out (90, 
p. 8; 11, p. 9), experimental evidence on this subject is urgently 
needed ; and it is probable that a well-conceived series of breeding 
experiments carried out with statoblasts and embryos of Plwmatella 
might resolve some of the difficulties which at present exist. 
Braem (90, p. 6) has recorded the important observation that in 
cases Where the remains of the statoblast from which a colony 
has germinated can be detected at the proximal end of the 
colony, the form of that statoblast can sometimes be proved to 
be identical with those which are being produced in the tubes of 
the same colony. ‘There are reasons for believing, as has been 
remarked above, in dealing with Paludicella, that ovaries and 
testes are usually produced early in the summer from colonies 
which have resulted from the germination of statoblasts or 
hibernacula (cf. Kraepelin, 87, p. 85). But it has not been 
sufficientiy ascertained how far, if at all, these sexually mature 
colonies differ from those produced by statoblasts; nor is it 
known whether there are any differences between colonies 


450 DR. &. Fe. HARMER ON THE 


produced from floating statoblasts and those which have developed 
from sessile statoblasts. 

It is believed by most recent authorities that Alcyonella is not 
generically separable from Plumatella. Alcyonella is character- 
ised by its compact form, resulting from the arrangement of its 
crowded, more or less parallel, tubes, more or less at right angles 
to the substratum. It has been suggested that this habit of 
growth is largely, if not entirely, the result of the simultaneous 
germination of a number of statoblasts situated close together. 
But I am not aware of the existence of any experimental 
evidence to show whether a large colony of Alcyonella fungosa 
can be produced by the germination of a single statoblast. Some 
evidence that the completely parallel tubes of the typical /ungosa- 
form owe their arrangement to want of room, is given by the fact 
that the peripheral parts of a fwungosa-colony may have the form 
of the var. coralloides, which is characterised by a looser habit of 
growth. 

The conclusion that “ Aleyonella” is merely a form of Pluma- 
tclla seems to me to be well founded. The only generic distinction 
which can be maintained is the habit of growth; and in this 
character P. fungosa var. coralloides is intermediate between 
“ Aleyonella” and Plumateila. WKraepelin (87) has gone so far 
as to regard P. fumgosa as merely a form of P. repens, which is 
termed by him, on insufficient nomenclatorial grounds, P. poly- 
morpha. In this conclusion he was at one time criticised by 
Braem ($0), who pointed out differences which led him to believe 
that P. fungosa is specifically distinct from P. repens. Braem 
has more recently (11, p. 9) retracted this view, and he now 
regards P. repens and P. fungosa as varieties of one species. 

The specimens which may be referred to the P. fungosa type 
were sent on May 31, and were received alive, with their 
polypides in full vigour. As has been pointed out above, they 
were taken from the pool which supplies the system, and I have 
not seen specimens from the pipes themselves, where great trouble 
has been caused. Mr. R. H. Newill, to whose kindness I owe 
the specimens and as much information as he was able to give, 
has, however, assured me that the specimens which blocked the 
pipes and ram resembled those which had been taken from the 
pool supplying the system. There is thus a strong presumption 
that the specimens examined belonged to the same species as 
those which occurred in the pipes ; a conclusion which 1s rendered 
all the more probable by the fact that this form has a very 
characteristic habit of growth, which Mr. Newill cannot have 
failed to notice. 

IT have bad some difficulty in deciding whether the specimens 
in question should be regarded as typical examples of P. fungosa 
or should be referred to its variety coralloides (= Plumatella 
coralloides Allman, cf. Allman, 56, p. 105). In the more central 
parts of the masses the tubes have the fungosa-arrangement. In 
the more peripheral parts of the material they may be less 


POLYZOA OF WATERWORKS. 45] 


intimately connected with one another, and are therefore like 
the form which has been described as var. coralloides. 

Braem (90, p. 4) has stated that the floating statoblasts of 
P. fungosa (typical form) have an average length of 410 pu, and 
an average breadth of 320 4; the corresponding measurements 
for var. coralloides being respectively 380 « and 300. In my 
own material the average length of eleven floating statoblasts, im 
one series of measurements, was 371 4; and the average breadth 
was 240 ». The extreme measurements were :—Length, 328 y and 
416 w; breadth, 224 and 2644. In another series of measure- 
ments, in which the statoblasts of the compactly arranged tubes 
were distinguished from those of the peripheral, more coralloides- 
like tubes, the average length was 388 w and the average breadth 
rather more than 280 » * for the former; while for the latter the 
length was 358 y and the breadth was 266 ». There is thus some 
difference between the sizes of the statoblasts in different parts of 
the same mass of tubes; the loosely arranged zoccia at the 
periphery having somewhat smaller statoblasts than the more 
centrally placed ones. But even the largest statoblasts agree 
more closely in size with those of var. coralloides (from Braem’s 
measurements) than with those of the typical fwngosa-form (as 
given by the same author). I refer the specimens, therefore, to 
var. coralloides ; and this conclusion is confirmed by the measure- 
ments which I have made of a very typical fungosa-form from 
a pond at Upper Norwood (Brit. Mus, 85.7.28.5-8), in which 
the average length of the floating statoblasts was 431 yu, and the 
average breadth 307 «: in close agreement with Braem’s figures. 

The floating statoblasts of the material from Styche have the 
form shown in fig. 22. They do not vary much in form, although 
in some of them one of the sides is more convex than the other. 
The annulus leaves a considerable part of the central capsule 
exposed on both surfaces of the statoblast. The average ratio of 
length to breadth (11 cases) is 1°543: 1, 

The sessile statoblasts are considerably larger than the floating 
statoblasts. On the attached surface (fig. 23) the central capsule 
gives off an irregular peripheral ring of cementing substance, 
which forms a vertical ridge by which the statoblast is fixed to 
the wall of the tube in which it has been produced. On the free 
surface (fig. 25), a saucer-like shape is produced by the develop- 
ment of a thin marginal collar. This represents the annulus of 
the floating statoblasts, as is shown by its structure. 

In some statoblasts the gas-cells are represented by only a small 
proportion of the number found ina floating statoblast ; and these 
occur in isolated groups, of irregular and obviously inconstant 
form (fig. 25). In other eases, as in fig. 24, the annulus forms a 
complete band, which is almost as well developed as in the floating 
statoblasts. 


* The uncertainty about the breadth is due to the fact that the statoblasts 
measured were not resting accurately on one of their flatter surfaces. 


Proc. Zoon. Soc.—1913, No. XX XI. 31 


452 DR. S. F. HARMER ON THE 


The sessile statoblasts are very variable in shape and size 
(figs. 23-25), but they are all larger than the floating statoblasts 
of the same specimens. The largest one found measured 640 pu 
by 432 p, giving a ratio of 1:-481:1. <A shorter and broader 
one was 560» by 480, with a ratio of 1:166:1. The average 
of seven measurements was 554 p by 434, with a ratio of 
PATS) Bu 


(4) PLumMareLLA EmARGINATA Allman, var. MuscosA Kraepelin. 
(Pl. LXITI. figs. 15-20.) 


In his well-known work on the German Freshwater Polyzoa, 
Kraepelin (87) refers the forms of Plumatelle which have come 
under his observation to three species. These are described 
respectively as P. princeps, P. polymorpha, and P. punctata. 
For the last, with which it is not necessary for me to deal, he 
has adopted Hancock’s name. ‘The first two were renamed by 
him, on the ground (pp. 118, 119) that the older names, from 
which a choice might have been made, have been used in so 
many different senses by previous authors that it would only 
have introduced additional confusion to have made use of any of 
them for the speciesas characterised by him. I do not think it 
necessary to follow Kraepelin in this conclusion. 

P. princeps and P. polymorpha were distinguished prmcipally 
by the form of the floating statoblasts. In P. princeps these are 
relatively long and narrow, and the ratio of length to breadth is 
said to vary from 1:53: 1 to 2°79: 1, the typical ratio bemg given 
as about 18:1. The species thus characterised includes P. emar- 
ginata Allman and P. fruticosa Allman. 

In P. polymorpha the floating statoblasts are broader in pro- 
portion to their length, and the ratio of length to breadth is 
described as varying from 1:1: 1 to 1-42:1; the average being 
about 1°25: 1. Under this name Kraepelin includes P. repens 
and P. fungosa auctt. 

The forms of Plumatella with relatively broad statoblasts are 
represented in the material which I have received from English 
waterworks only by a variety of P. fungosa, which, following 
Braem’s earlier opinion, I have treated above as a species distinct 
from P. repens. Those with elongated statoblasts are represented 
by the material from Torquay, and it thus becomes necessary 
to discuss the question of the name which should be used in 
describing them. 

Braem (90, pp. 9, 10) has brought forward arguments to show 
that P. fruticosa and P. emarginata are distinct species ; and he 
points out the following differences between them :— 

(1) P. fruticosa has relatively slender tubes, which grow into an 
erect form; while P. emarginata has broader tubes, which are 
more adherent to the substratum. (2) The floating statoblasts of 
P. fruticosa are more than twice as long as broad (average ratio : 


POLYZOA OF WATERWORKS. 453 


length to breadth, 2°497: 1 *), while the annulus does not cover 
so much of the central capsule as in the other species. In 
P. emarginata, on the other hand, the floating statoblasts are 
less than twice as long as broad (average ratio: length to breadth, 
1868: 1), while the annulus covers nearly the whole of the 
“upper” side, leaving only a small portion of the central capsule 
uncovered. (3) In P. fruticosa the sessile statoblasts have a broad 
vestigial annulus, and are more than twice as long as broad (ratio, 
2-461: 1), while those of P. emarginata resemble the sessile stato- 
blasts of P. repens, and have no annulus, while they are less than 
twice as long as broad (ratio, 1°312: 1). 

Through the courtesy of the authorities of the Hamburg 
Museum, I have had the opportunity of examining specimens of 
both var. muscosa Kraepelin and var. spongiosa Kraepelin, from 
the original material described by that author. I have figured 
a floating statoblast (fig. 18) and a sessile statoblast (fig. 20) of 
var. muscosa, from the Hamburg material, for comparison with 
the Torquay specimens. The measurements which I have made 
of the statoblasts from Hamburg agree closely with those given 
by Kraepelin, and are as follows :-— 


FLOATING STATOBLASTS. SESSILE STATOBLASTS. 


Length. | Breadth. ANTOIEY | Length. | Breadth. eweraze 


P i ratio, L: B. i it ratio, L: B. 


Var. muscosa ...| 482-496 | 240-280 1:817:1 | 384-592 | 272-352 1°456 : 1 


Var. spongiosa...) 448-480| 256 1833:1 | 416-496 | 320-384] 1:278:1 


| 


Very few floating statoblasts are present in the spongiosa- 
material, in which there are a number of sexually produced 
embryos. 

If Braem’s criticism of Kraepelin’s results is correct, it 
follows from these measurements that both varieties, muscosa 
and spongiosa, ave to be regarded as forms of P. emarginata; 
and in this I am following what I understand to be Braem’s 
opinion. 

The specimens sent from the Torquay Waterworks by Mr. S. 
C. Chapman appear to me to be referable to P. emarginata, 
as understood by Braem, although their floating statoblasts are 
less elongated than in the Hamburg specimens. I find that these 
statoblasts vary in length from 328, to 416 and in breadth 
from 224 to 264, the average measurements (11 cases) 
being :—Length, 371 1; breadth, 240; ratio, L:B, 1-543:1. 


* The ratio is calculated from the measurements given by Braem. 
21s 
ol 


A454 DR. S. F. HARMER ON THE 


The sessile statoblasts have the following measurements :— 
Length, 432 p-448 yp, average 4424; breadth, 320 p-368 u, 
average 336; average ratio, L: B, 1-317: 1. 

If Braem is right in separating P. emarginata and P. fruticosa, 
the proportions of the sessile statoblasts would alone justify the 
reference of the Torquay specimens to P. emarginata rather than 
to P. fruticosa. Iam confirmed in the belief that they belong to 
the former species by the fact that the annulus covers nearly the 
whole of the central capsule on one surface of the statoblast 
(figs. 15-17), as shown by Allman (56, pl. vii. fig. 7) in one of his 
figures, and by Braem (90, pl. i. fig. 12), who has cailed special 
attention to the importance of the character in question. 

In Kraepelin’s series of measurements of the forms regarded 
by him as belonging to P. princeps (87, pp. 112, 113) the average 
ratio of length to breadth is givenas 1°8:1. But he admits a 
considerable variation in this respect, the limits given being from 
2°79: 1 to 1:53:1. It will be seen that my own measurement of 
1:543:1 falls within these extremes, although it is much nearer 
Kraepelin’s lower figure. The highest ratio of length to breadth 
measured by me is 1°666: 1, and the lowest is 1:°333: 1. The latter 
ratio would bring the statoblasts well within the limits given by 
Kraepelin for his P. polymorpha, which includes P. repens auctt. 
It thus becomes necessary to consider whether the Torquay speci- 
mens should be referred to P. repens instead of to P. emarginata. 

I have accordingly compared the Torquay statoblasts with the 
variety of P. fungosa described above and with what I regard as 
a typical form of P. repens, obtained at Cringleford, near Norwich. 
It will be seen from the figures that the floating statoblasts of 
the Torquay specimens (figs. 15-17) are distinctly longer. in pro- 
portion to their breadth, than those of either P. repens (fig. 21) 
or P. fungosa var. coralloides (fig. 22), and that the annulus 
covers much more of the central capsule. It will be remarked, 
moreover, that in the Torquay specimens the annulus extends 
over the central capsule further on one side than on the other 
side of the statoblast, in agreement with P. emarginata var. 
muscosa (fig. 18, from a Hamburg specimen); while in the other 
two forms the annulus is about equally developed on both sides. 
The floating statoblasts of P. fungosa (fig. 22) are distinctly larger 
than those of P. repens (fig. 21). I have not obtained sessile 
statoblasts of P. repens for comparison. Kraepelin denies the 
existence of this form of statoblast in the typical P. repens, but 
Braem (90, p. 6) claims to have found them. In P. fungosa the 
sessile statoblasts (figs. 23-25) are very large, and possess a rather 
well-developed annulus; while they are smaller, and with a more 
vestigial annulus, in the specimens of P. emarginata var. mus- 
cosa, both from Hamburg (fig. 20) and from Torquay (fig. 19). 

I fully admit that the floating statoblasts of the Torquay 
specimens are short as compared with those of more typical 
specimens of P, emarginata; but it appears to me that the result 


POLYZOA OF WATERWORKS. 455 


of the above comparison is to support the conclusion that the 
Torquay material should be referred to that species, and I lay 
special stress on the unequal development of the annulus on the 
two sides of the statoblast. Kraepelin’s var. muscosa was 
characterised by him asa luxuriantly branched form of Pluma- 
tella in which the branches were not cemented together (in this 
respect differing from his var. spongiosa *). The ‘Torquay speci- 
mens seem to me to agree closely with those from Hamburg in 
their mode of growth, and I have accordingly thought it best to 
refer them to the same variety. 


Literature cited. 


1856. Auuman, G. J.—‘‘ A Monograph of the Freshwater Polyzoa.” 
London, Ray Society. 

1911. AnnanpALe, N.—“ Freshwater Sponges, Hydroids, and 
Polyzoa,” in ‘The Fauna of British India.’ London 
(Taylor & Francis). 

1890. Brarm, F.—‘‘ Unt. wb. d. Bryozoen des siissen Wassers,” 
Bibl. Zool., Bd. i. Heft vi. 

1911. Brarm, F.—‘‘ Beitrige zur Kenntnis der Fauna Turkestans:” 
vii. “ Bryozoen,” Trav. Soc. Imp. Nat. St.-Pétersbourg, 
xlii. Lief. 2, Teil 1. 

1904. Currica, C.—‘‘ Notes sur les Bryozoaires de Roumanie,” 
Ann. Sci. de ’Université de Jassy, 111. pp. 4-14 (sep., 
pp- 1-11). 

1888. p—E GuerRNe, J.—‘“‘ Sur la dissémination des organismes 
eau douce par les Palmipédes,” C. r. hebd. des Séances 
et Mémoires de la Soc. de Biol. (Paris), 8 ser., v.-Ann. 
1888, p. 294. 

1890. De Vries, Huco.—‘“ Die Pflanzen und Thiere in den 
dunklen Riumen der Rotterdamer Wasserleitung,” 
Bericht tb. d. Biolog. Untersuchungen d. Crenothrix- 
Commission zu Rotterdam vom Jahre 1887. 

1909. Harrmeyver, R.—“ Bryozoen, Moostierchen,” in A. Brauer, 
‘ Die Stisswasserfauna Deutschlands,’ Heft 19, p. 49. 

1904, Hawxkstey, C.—Presidential Address to Section G (Kngi- 
neering), Southport Meeting, 1903, ‘ Report Brit. Assoc.’ 
pp. 764—, ‘“ Water-Supply.” 

1904. Hickson, S. J.—Presidential Address to Section D 
(Zoology), Southport Meeting, 1903, ‘Report Brit. Assoc.’ 
p. 675, 

1899. Kemna, A.—“ Ta Biologie du Filtrage au Sable,” Bull. 
Soc. Belge Géol. xiii. Ann. 1899, Mémoires, p. 34. 

1905. Kemna, A.—‘“ La Biologie des eaux potables,” Ann. Soc. 
roy. Zool, et Malacol. de Belgique, xxxix. (1904) p. 9. 


* Hartmeyer (09, p.53) describes Kraepelin’s var. spongiosa as a distinct species, 
under the name of Plumatella spongiosa. 


4 


Dt 
lor) 


DR. S. F. HARMER ON THE 


1885. Krarpenin, K.—‘“ Die Fauna der Hamburger Wasser- 
leitung,” Abh. Naturwiss. Ver. Hamburg, ix. 

1887. Krarpenin, K.—‘“ Die Deutschen Siisswasser-Bryozoen,” 
i. “‘ Anatom.-Syst. Teil,” Abh. Naturwiss. Ver. Hamburg, 
x. Festschrift. 

1892. Krarpetin, K.—“ Die Deutschen Siisswasser-Bryozoen,” 
il. “ Kntwickelungsgeschichtlicher Teil,’ Abh. Naturwiss. 
Ver. Hamburg, xii. 

1898. Larar, F.—‘“ Technical Mycology,” vol. i. London, 
Charles Griffin & Co., Ch. xxxiv. pp. 355-362. 

1904. Larar, F.—“ Handbuch der Technischen Mycologie,” 
A Lief. pp. 193—. 

1894, Levinsen, G. M. R.— Zoologica Danica,” 9 Hefte, 
‘* Mosdyr.” 

1896. Wesenperc-Lunp, C.—‘“ Biologiske Studier over Fersk- 
vandsbryozoer,” Vid. Meddel. naturh. Forening Kjoben- 
havn, 1896, p. 249. 

1910. WuiprLe, G. C.—“ The Microscopy of Drinking-water,” 
2nd ed., New York (John Wiley & Sons) and London 
(Chapman & Hall). 


EXPLANATION OF THE PLATES. 
Prate LXII. 


Showing hibernacula of Paludicella articulata from the pipes of an English 
waterworks-system. 

The figures were drawn with a Zeiss A objective, and were then reduced two and a 
half diameters. The scale represents hundredths of a millimetre. 


Fig. 1. Zocecium bearing a hibernaculum of the type described by Dumortier and 
Van Beneden. (Only two hibernacula of this type were found in the 
whole of the material examined.) 

Fig. 2. A zovecium, accidentally folded, which has developed two hibernacula, 
while it bears two other hibernacula (bases shown) in the place of the 
ordinary lateral buds. The preceding zovwcium also bears two lateral 
hibernacula. 

Fig. 3. Zocecium bearing three hibernacula. ‘The distal one is small, and is still 
enveloped distally by the remains of the zocecium in which it has been 
formed. 

Fig. 4. The smallest hibernaculum found. It shows distally the remains of the 
zocecium in which it was formed. The elongated polypide-bud is seen in 
its interior. 

Fig. 5. Zocecium with three hibernacula, each of which shows a polypide-bud. 

big. 6. Basal view of a zocecium which shows two split hibernaculum-valves in its 
ectocyst. 

Fig. 7. A similar zocecium, in basal view, with a terminal and a lateral hiber- 
naculum. The latter is developed nearer the proximal end of the 
zocecium than is usually the case. 

Fig. 8. Side view of a similar zocecium, showing the septa from which lateral 
zocecia have broken off. A fully-formed hibernaculum is present distally. 

Fig. 9. A similar zoecium, bearing a terminal hibernaculum, which is still enclosed, 
distally, in the remains of the zocecium inside which it has been formed. 

Fig. 10. A series of three zocecia, with the proximal end of a fourth zocecium. 
The proximal zoccium has developed a hibernaculum which has passed 
partly through the terminal septum. The third zocecium bears a terminal 
hibernaculum, the proximal end of which partly traverses the corre- 
sponding septum. 


POLYZOA OF WATERWORKS. 457 


Prate LXIILI. 


Statoblasts of various Phylactoleemata (detailed structure of the annulus not repre- 
sented). 


The figures were drawn with a Zeiss C objective, and were then reduced two and a 
half diameters. The scale represents hundredths of a millimetre. 


Figs. 11-14. Fredericella sultana, showing the variable form and size of the stato- 
blasts. Batley Waterworks. 

Figs. 15-17. Plumatella emarginata var. muscosa; floating statoblasts. Torquay 
Waterworks. 

Fig. 15. “ Ventral” view ofa young statoblast. The central capsule is represented 
by the deeper shading. The line concentric with the outline of the 
central capsule, and next inside that line, indicates the extension of 
the annulus on the “ventral” side. The inner circle represents the 
edge of the annulus on the “dorsal” side, as seen through the central 
capsule. 

Fig. 16. “ Ventral” view of an old statoblast. The outline of the central 
capsule is obscured by the annulus, and is not seen. 

Fig. 17. “ Dorsal” view of an old statoblast. The annulus covers nearly the 
whole of this surface of the central capsule. 

Fig. 18. Plumatella emarginata var. muscosa. Hamburg Waterworks (from 
Professor Kraepelin’s material). The annulus covers most of the central 
capsule on this surface of the statoblast. 

Fig. 19. Plumatella emarginata var. muscosa. Torquay Waterworks. Sessile 
statoblast, showing the vestigial annulus; from the unattached surtace. 

Vig. 20. Plumatella emarginata vay. muscosa. Hamburg Waterworks (from 
Professor Kvaepelin’s material). Sessile statoblast ; from the attached 
surface. 

Fig. 21. Plumatella repens. Floating statoblast; from a pond, communicating 
with the River Yare, at Cringleford, near Norwich. 

Figs. 22-25. Plumatella fungosa var. coralloides. Styche Waterworks (from the 
peol supplying the system). 

Fig. 22. Floating statoblast. 

Tig. 23. Sessile statoblast, from the attached surface; showing the cement- 
substance by which it is attached to the inner wall of the zocecium. 

Fig. 24. Sessile statoblast, with well-developed aanulus, the details ef which 
are represented in a small area on the right side of the figure; from 
the unattached surface. 

Fig. 25. Sessile statoblast, with vestigial annulus, which shows remains of the 
gas-containing cavities (only represented at the upper and left parts 
of the figure); from the unattached surface. 


458 MR. A. W. WA'TERS ON 


34, The Marine Fauna of British Hast Africa and Zanzibar, 
from Collections made by Cyril Crossland, M.A., B.Sc., 
F.Z.S., in the Years 1901-1902. Bryozoa *—Cheilo- 
stomata. By ArtHur Wm. Watemrs, F.L.S8., F.G.S8.f 


[Received March 18, 1918: Read April 22, 1913.] 
(Plates LXIV.-LXXIII.¢ and Text-figures 79-82.) 


INDEX. 

Systematic : Page 
JEYPAGHID WROOLGy Bs Wo cacect sao soosse cuapsaedscnanapccassoawacanasocso UBL) 
(SOMMAORLTOFOOUBOOEISUSy S61  covanvoogadsova cbecooceoauncdaenn concen 2 KokS) 
Sig QUIDGOVAR KIOEENG, coagne sooedooad ec degoandeee aoe cebbeacoeanadodepe conceal “Ans 
LOAM OPOR, QOMIEEDSUS,; So Mo roaaco oppasocgenvocsoocooaaccocuseconee FAOY/ 
SISUPIOGHPUR CXOROSONCHPOADEOS, S06 Ws cuodeooodbeaossoodeon coogoonsoouncod | ZAOS) 
Sh CAUPROGT AH DEG. SiDo Ts scoossananneancandos oebeaddocsbadtapaccasocunese  “ffO) 
Scrupocellaria wasinensis, SP. MN.  ...0....0ceceeceeneeereceersereee ADO 
Vittaticella elegans, var. zanzibariensis, NOV. ......... 22.06 485 
Cellaria gracilis, var. tessellata, NOV. .........1..cseseeeeeereese ADS 
Chapasinensisespamcres tte esas secace ics cock e aes eee eae So LOD 
Osthimosia zanzibariensis, SP. MN.  ...... 2-1 sce cee cne eee reeeeesee» 008 
SULT LIU GIS D ca Meee eee ho fe se sbi sae Ghana ie aes SEER eee OEE EE are OL 
TOGO RRUCEL UREISTIGIOSUS S05 Wo epacas sedaddocabcovssdegcoeagoaeaacosceasa | OIL) 
IZQG RHE, QPUGECHOSIS,, SDs Wo sasodatorapesoscoocdacopasouasenncovececes HUIS} 


IE, GOLEM WE CAGE Gy TXOe). “ sa3Boc bon ao0c00cn0 605000 son cob HEnocHoe  OpIlfs) 
Rhynchozoon profundum, var. laminatwin, NOV. ......-..000+. 523 
AdeonellopisKenosslam de SD vnlls-c-barcee-pecer cece meee eee eeeeeee 531 


Senupocelllarid soiree naecee scares danse ec eee ea eee aes 
Catenicelllidias ita asecoe-cesciecs Sos bias. aeewctlene ee uae eee eee CEE ee LOD 
(SMICTPOO. connie ones 66 Baa aaa EE RERBe aca adacpisa ncocdocancburinucroenes ior 
[SADC SOPQNIG “sat aot aonoce ssa nanese reeeateceneMacdCcoccaddakcoacdsanodeams . taNS) 
Calle pon genre acorns) soe aicisasvisins nels stee COs uet epee Soke eaee ee eeae OOO 
LEGG C1100 HOR as EL ears dak iols Gold SE He af BERT RS REE OR LO. 
TEC PRU Gani aes erie es ces oulspacd i deldee GRESHAM ORE EE RE SE OLD 
IW ABKOVAONKOLEEh” “Guocuamoodssuesneooaeemadedeaacaeebocnoae cddonaasscuaccaneo. GPO) 
ARCH XOIACD. Groton na scec Oca one aaee a aeRRER Paar ace eomE anne sespeaneannodaconT ar 


The collection made by Mr. Cyril Crossland in the neighbour- 
hood of Zanzibar contains 76 species or varieties of Cheilostomata, 
and all are from shallow water, in fact with the exception of two 
are from 10 fathoms or under; so that, for a purely shallow-water 
collection, it is a very large one. 


Points of Special Interest. 


(1) In Stirparia the first zooecium of a tuft has the character 
of a primary zocecium (p. 470). 


* [In view of the difference of opinion as to whether this Phylum should be 
called Bryozoa or Polyzoa (see Proc. Linn. Soc. 1911, p. 61) I have not interfered 
with the preference of the author.—KprTor. | 

+ Communicated by Cyriu Crosstanp, M.A., B.Sc., F.Z.S. 

t+ For explanation of the Plates, see p. 532. 


eee iiey ei. LOIN, 


‘ 


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BRYOZOA FROM ZANZIBAR. 


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mers 


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————————— 


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BRYOZOA FROM ZANZIBAR. 


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BRYOZOA FROM ZANZIBAR. 


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BRYOZOA FROM ZANZIBAR, 


BRYOZOA FROM ZANZIBAR. 459 


(2) The nature of the articulation can be used in grouping the 
Serupocellaride, the other characters being the ovicells, vibracula 
(especially the number at a bifurcation), and the band, perhaps a 
spermotheca at the distal end. 

(3) The difference in form and position of the ovaria in Bugula 
and Serupocellaria gives a generic character, and the same is the 
case in other genera (p. 476). The form of the ovaria is likely 
to give us considerable assistance in classification, but as the 
ovaria pass through various stages of development, time and 
caution are requisite to make it fully available. 

(4) In Membranipora armata Hasw. there are two mature 
polypides in most zocecia, and on one side of the zocecium there is 
a long chambered aviculari ium, whereas on the other side there is 
a similar long chamber containing a secreting gland (p. 488). 

(5) In some species of Schizoporella a bar curving towards the 
sinus is seen across the operculum. This is the commencement 
of the lower wall of another zocecial layer, and in many cases the 
distal walls are seen to extend over the operculum (p. 504). 

(6) In Diplodidymia complicata Rss. a small ovum starts im 
a small sac hanging down from the opercular region. Both grow 
large, and the larva ultimately fills up a large portion of the 
zocecium (p. 491). 


Some of the species in this collection were also found in 
Crossland’s Red Sea collections, and my report thereon is quoted 
with a shortened reference *. 

This communication may be considered as part of a series 
dealing with tropical forms, of which three papers on the Red 
Sea etc. have been published ; and I have a small collection from 
Cape Verde, also collected by Crossland. 

From the locality Wasin about 60 species were procured, and 
from Ras Osowamembe about 30 in all. 

Since the examination of this Zanzibar collection was com- 
menced, Levinsen has published his ‘ Morphological and Systematic 
Studies on the Cheilostomous Bryozoa,’ and we have for some time 
been anxious to see his ideas on classification developed. He has 
given the particulars of much valuable detail examination of the 
calcareous parts of the Bryozoa, and doubtless many of the cha- 
racters he deals with will receive increased attention, and our 
knowledge is much advanced by the wealth of observation. 

Unquestionably much of his altered classification will be 
adopted ; but if any of us thought that all our troubles would 
be ended as soon as Levinsen published his results, we find that 
this is far from being the case. We have to examine each new 
grouping, and see whether, when other characters are examined, 
they uphold suggested alterations; also, do other species of the 
groups fall into place? I already see where some modifications 
will be required, and no doubt other workers who have carefully 


* Journ. Linn. Soc., Zool. vol. xxxi, pp. 123-181, 231-256. 


-460 MR. A. W. WATERS ON 


examined Levinsen’s great work are coming to similar con- 
clusions, and in this way our starting from Levinsen’s standpoint, 
and using it as a stepping-stone, should ultimately advance the 
correctness of the classification very much. At present the 
position remains very difficult, since for a large part we are not 
sure what will be accepted. 

I do not want to be misunderstood as giving an adverse 
criticism, and so would add that the task was too great to be 
final, as Levinsen must have fully felt; and we must now be 
alive to the fact that, in the future, classifications will to a 
large extent be based upon the structure of the soft parts, which 
furnish a very large number of characters, some of which are of 
great value, while others will not be found very useful. For 
every external character there are many internal ones of 
importance. 

It must be emphasized that, as I have often said*, characters 
of great value in one group or family are almost useless in the 
next; and all attempts at fixing certain characters as being 
of Al importance, others of secondary importance, and so on 
down the scale lead to no result, but we must get together our 
groups of species based upon as many characters as possible, and 
gradually build from them larger divisions; and this process must 
be slow, but it will be natural, whereas the attempt to work from 
the larger divisions has led to false results. 

As a case in point, Levinsen makes great use of the rosette- 
plates, which I have found in certain cases to give most useful 
results, in other families none at all; and in tabulating Levinsen’s 
results we find that the family character is often uni- or multi- 
porous, with or without pore-chambers, that is to say, the 
character in such cases is of no value for the higher group. 
In those families in which we should have been most glad of 
help, namely, in Membraniporide, Cribrilinide, Microporide, 
Escharide, Smittinide, it is pore-chambers or uni- or multi- 
porous rosette-plates, and in eight other families uni- or multi- 
porous rosette-plates ; also in genera we find the same range, so 
that, while Levinsen’s work in this direction is very valuable, 
care is required lest we attach undue importance to the rosette- 
plates or any one character. 

Where the opercular aperture is not on a level with the frontal 
surface of the zocecium there is often a shelf upon which the 
distal end of the operculum rests, and Levinsen has shown that 
this is a character of some value; but here, again, we must not 
expect too much. The value of most of the characters used by 
Levinsen, even for the main divisions, are still on trial. 

Levinsen has followed 7 Norman in using names given by the 

* Page 71, Levinsen expresses these facts as follows:—‘ The same structural 
feature in different systematic divisions can have a very different systematic 
importance, so that characters which are constant in one genus or family, in 
other corresponding divisions are not always constant even within the species.” 

+ Since I wrote the above, Canu has sharply criticised Norman’s suggested 
alterations of generic names (Rey. Crit. de Paléozool. vol. xvii. p. 49, 1918). 


BRYOZOA FROM ZANZIBAR. 461 


earlier authors in a sense different from that applied to them 
for generations ; and not only that, but generic names now well 
established are shifted to be used instead of other well-established 
names. Cellaria is to be replaced by Cellularia, a name that has 
been used in numerous senses during the lifetime of most of us. 
There is no rule obliging us to revert to old names which were 
made for genera without any adequate and recognisable deserip- 
tion; but even if a rule were being broken, we should continue 
to do so in order to stay the appalling confusion caused by this 
desire to keep alive a name in its doubtfully supposed original 
sense. Cellaria still means for me what it has meant all my 
scientific life. I do not accept Lepralia as meaning JMembrani- 
porella, and this Levinsen also refuses to do and gives an amended 
description of Membraniporella; but I am not prepared to accept 
Aspidelectra, and should place melontha under Membraniporella. 
Nor is the variously used Hscharoides a satisfactory name, seeing 
that it was only given for a subgenus of Cellepora based upon 
characters seen to be useless. Discopora is another genus that 
should have been dropped, as it was quite insufficiently described 
at first ; im fact, the name was given by Lamarck to a Cheilostome 
and by Fleming to a Cyclostome, so that it has been employed for 
all kinds of unrelated things, as Cellepora, Haloporella, Membrani- 
pora, Mucronella, Smittina, Palmicellaria, Diastopora, Lichenopora, 
Tubulipora, etc., and has heen var iously used by paleontologists. 
It has not been proposed to retain Discopora on account of the 
definition, which now tells us nothing, but from what is supposed 
to be the first-mentioned species of an incongruous group. 

Such changes back to discarded genera add much to the diffi- 
culties of those who are closely following all that is written; and 
are not these premature and puzzling changes of names keeping 
back new workers from entering the thinning ranks? Until we 
are sure of the characters, and have enough material to test the 
relationships, we must often gather information round species 
and genera the names of which we know will ultimately be 
changed or disappear. 


I have again to thank Mr. Kirkpatrick for allowing me to 
make frequent comparison with the British Museum col- 
lections. The Plates were mostly drawn before the appearance 
of Levinsen’s work, otherwise they would have been slightly 
differently arranged. Miss Thornely’s paper, ‘“‘ The Marine 
Polyzoa of the Indian ee from H.M.S. Sealark,” Trans. 
Linn. Soc. vol. xv. pp. 137-157, has also appeared during the 
preparation of my aes It materially increases our knowledge 
of the distribution of tropical Bryozoa from moderate depths. 

Other groups collected by Mr. Crossland from Zanzibar have 
already been described by specialists in the Proceedings of this 
Society. 


462 MR. A. W. WATERS ON 


Table of Distribution from West to Hast. 


zg 
on 
Aetearangwina cal per seeteseasee ners: 208 
» truncata Landsb. .... | 465 
Brettia tropica, sp. n. ..................| 405 
Synnotum aviculare Pieper ............) 465 
3 pembaensis, Sp. N............. 465 
is contorta, sp. N................| 466 
| Hucratea chelata Li. ..........0.06..++-..| 466 
Beania spinigera MacG. ...............) 467 
» mirabilis Johnst. ...............,467 
Bicellaria chuakensis, sp... ...+..-.. 467 
Stirparia exilis MacG. ..................| 468 
2” zanzibariensis, Sp. Nn. ...... 469 
3 dendrograpta, sp. 0. 470 
Bugula neritina var. minima Waters 471 
ie robusta MacG. Mates ata AL 
Scrupocellaria ferox Raskeeie oe 476 
5 cervicornis Busk 477 
3 macandrei Busk ...... 477 
5 pilosa Aud. ............ 478 
wasinensis, Sp. 1. 479 
Canda ‘retiformis, Pourtales ......... 479 
Catenaria lafontii Aud. ............... 481 
diaphana Busk ............... 482 
Vittaticella elegans Busk ..| 484 
‘ 5 var. zanzibari- 
ensis, NOV....... 485 
Membranipora savartii Aud. .........) 486 
3 armata Hasw. . 486 
a catenularia Jameson 488 
Farcimia oculata Busk  ............... 489 
Diplodidymia complicata Rss.......... 490 
Chlidonia cordiert Aud. .. 4.92 
Cellaria gracilis var. tessellata nov. | 495 
if WASINENSIS, SP. ND. oe... ees ee 495 
Thairopora mamillaris Lamx..........)497 
Steganoporella magnilabris Busk ...|498 
Cribrilina radiata Moll. ...............)501 
Hippothoa distans MacG................ 501 
7 divaricata Lamx. 501 
Schizeporella unicornis Johust. 501 
ia pertusa TOBE sooosooae 502 
* nivea Busk ... 502 
montferrandt Aud. ...|506 
Gemellipora protusa Yhornely ...... 506 
Trypostega venusta Norm. ....| 506 
Arthropoma cecilii Aud. ...............|508 
Osthimosia zanzibariensis, sp. 0. 508 
Lagenipora rota MacG. sscencee|| G10 
Haswellia australiensis Hasw. ...... 511 
Tubucellaria cereoides var. chua- 
kensis Waters ...... | 512 
Susiformis VOrb. ......)512 
3 zanzibariensis Waters 512 


| Atlantic. 


++ 
++ 


+ 


+++: +4: 
++ 444: 


+4: 


' 


+ 


| British. 


Mediterranean. 


+ ++ 


tte: 


_(elstanle 


<P ap 8 


te 


| Indian Ocean, 


South. 


n.=North. 


8. 


| 
« . 
Sl 6 
ei <a 
Blel/S\eia 
SlE18) 8/2 
3/6/25) e/¢ 
D jlo es |e 
[75] |+| N. Pacific 
sPauat \-+ 
as oF || 
-F|...1...| California. 
Slices + : 
A 4| ( Pacific, 
| yr* Barmah. 
SEalleaa or 
| 
Ge + 
ae +| 
SSP 14 
i albeoe| 
ae leo) ae 
leeaveeeilictal le Loyalty Isl. 
+. | Loyalty Isl. 
4) 4+)4+) 4+ 
J+ }...[ 2]... | 4] 
|] + 
Hel +|...|+ 
+ |...) 4F] oe 
alive ...|.../+] New Guinea. 
alte te 
enlleae \ar| 
+ +/+)+)4+) Brazil. 
+)...) 4]... + 
Nita tal 
+|+ 
ocolfaso|| ar |e 4 
|...|-.-|F|...|4] Bacific. 
aria | 
+i) + 
In seal Ec +| {ane i 
eran “ U S. 
ies les auritius 
50/4 
sar 
| 
je. [+ IF | 
ial 


* This was not mentioned in my Red Sea paper, but I have a piece from a sounding-line 


BRYOZOA FROM ZANZIBAR. 463 


Table of Distribution from West to East (continued). 


| 
: al Lal el 
= \\ASunish cul a ald 
/ | espa ISH ad 
| Vidar S53) |e! | 
beige eels ey 
5 /-a 5) o |= at es 
|g |s2| 2 | @ la ll ilslsis/sin 
faolSie2] S |S Sa ales) 2) ele 
| @ |els Sy 2 |S |@) oO) S| ce! 
(RIA) A |S Oia a & 
Smittina trispinosa var. protecta Th. 513 + + | +N. j+ 
on 3 var. spathulata | | | | 
¥ | | 
WIEKAC%  sanccoeee NSIS ois) soe | saa AB AP iio laellecs ae | 
55 iRayadteteh NITES” 56 406 e6ccee case oll (Weasel fact | 
3 GIs cececndessabcadnbsoooseoccass5oae| (AI! ihe | | | 
| Lepralia feegensis Busk ............... Joe eealiecel| | age che |) ar asfo [ap ifaco, f |fecclacs|| ene WE 
A turrita Smitt ...............,516|-++|...|  ... san. Warley Sh | 
55 wasinensis, Sp. 0. ............/516| | | esi 
5 cleidostoma var.inermis Ort. BILZ/ jleeell ben! woldee 44: oe ey le eetaes +|...| N. Pacific. 
Petralia japonica Busk ...... ......../518]...]... mY de |) aPasto ar iiar ooo lari 
. chuakensis, sp. N. ......-...../518) | | ae (a 
a vultur var. armata, nov. ...|518 | Vegi al 
? Escharoides occlusa Busk ............|519}...)... 568 ot +t Fi) | 
Holoporella columnaris Busk ......... BPE Nosollone| . onc sco. |) Arlo |S llooo P|) 3° 

3 aperta Hincks ............ Bye aileell) Mona + | +N. |+]......]-..]...] Cuba. 

5 (MOPORGPOS 3006.55 oosn0aca6 }522/+)...) ... er || SPS llocclfaoo"a|ls05) Ar 
Microporella ciliata Pallas .............523;)+ +) + del] aeixio. i|arllecs ap| sear 
Rhynchozoon Re var. lami- alec 

natum, nov. spanaco 50 6a| (Oe) | | | 
Retepora hirsuta Busk ..........0...-.. N523 |. + + | 
3 producta Busk ...............| 525 |...| SPs) |fasalfasch Fue) far 
5 denticulata Busk ............ 526 |...| bee Baa leaalenetiaa: +)...| Amboina,Lifu, 
55 Jjermanensis Waters .........,526)}. + ia | Sanduri Isl. 
a tubulata Busk Phun dd Ws le 626 |... Re cee we | EN. IE]. + | 
12 7 52 oa] | : 
Bifaxaria vagans soe Seeaaese 527 |...|...| .-- soe || ap | Gintins SEN 
Adeonella platalea Busk . 31/3) scolleooll 00 Beetllh sapoNe a ldeal lace. seu sea| ieee 3 
Vesela || | ana South Pacific. 
Adeonellopsis crosslandi, sp. n. 531 | awl 
| | | 


Hinceks records Lepralia striatula from Zanzibar. 


AETEA ANGUINA Linneus. (Pl. LXIV. figs. 1 & 2.) 
For synonyms see Miss Jelly’s Catalogue, and add :— 


Aetea anguina Calvet, “ Bry. Mar. de Cette,” Trav. de l’Inst. 
de Zool. de Univ. de Montpellier, ser. 2, mem. 11, p. 8 (1902); 
Jullien & Calvet, Bry. prov. des Camp. de l’Hirondelle, p. 122 
(1903); Thornely, ‘Ceylon Pearl-Oyster Fisheries,” vol. iv. Suppl. 
Rep. xxvi. Polyzoa, p. 108 (1905) ; Robertson, ‘‘ Non-Incr. Bry.,” 
Univ. Calif. Pub.. Zool. vol. ii. No. 5, p. 244, pl. iv. figs. 1-4 
(1905); Norman, “ Polyzoa of Madeira,” Journ. Linn. Soc., Zool. 
vol. xxx. p. 283 (1909); Levinsen, Morph. & Syst. Studies on Cheil. 
Bry. p. 93 (1909); Canu, “ Bry. Helv. de /Egypte,” Mém. de l'Inst. 
EKeyptien, vol. vi. p. 190 (1912); Osburn, R. C., ‘“‘ The Bry. of the 
Wood's Hole Region,” Bull. Fisheries Bureau, vol. xxx. p. 220, 
pl. xxi. figs. 14, 14 @ (1912). 

In the specimens from Wasin I have seen ovicells in the same 
position as those of A. recta, namely, at the top of the tubular 
projection at the back. This dorsal ovicell has now been seen 


4.64. MR. A. W. WATERS ON 


in A. recta from Rapallo* and Naples in the Mediterranean, and 
in the ‘ Belgica’ Antarctic material. Miss Robertson describes 
ovicells on the front, some distance down, but was in doubt as to 
whether the species was a true A. anguina. Whether she really 
had another species before her or not must be left uncertain, but 
certainly the front position as drawn by her is a very strange and 
unexpected one. 

Miss Robertson has confirmed what I wrote about the ovaria 
occurring in the creeping part, and this seems to be the usual 
place ; however, in the present specimens I do not find that the 
polypide extends far into this part, as it sometimes does in A. recta 
and as Miss Robertson describes and figures in A. angwina. 

Smitt, Waters, Jullien, and Robertson have all shown that the 
polypide etc. does not entirely live in the tubular prolongation ; 
but Jullien, although appreciating the fact, called this part the 
peristome or peristomia. Surely the peristome is something 
beyond the operculum and is the part where the polypide is only 
to be seen when extended; so that this term applied to Aetea is 
most unfortunate and misleading. This tubular prolongation 
has been called the neck, and the terminal portion the spoon, but 
no satisfactory name has been given to the creeping portion, which 
is only a part of the zoecium. There are 12 tentacles. 

The diaphragm does not make an infold when retracted, like 
most of the Cheilostomata; and the appearance of sete, which 
has frequently been alluded to, must be caused by a partial 
extrusion of the diaphragm. In some respects Aeteaw approaches 
the Ctenostome Cylindreciwm, but no Ctenostome has an external 
ovicell. 

I do not altogether understand what Levinsen f says regarding 
the ovicells of this species ; for though the wall of the ovicell is so 
thin that the embryo can readily be seen, yet decalcified pre- 
parations and sections have been studied. If Levinsen means to 
suggest that the sacs containing the ova and embryos are only 
accidentally at the termination and might adhere in any position, 
then this is not the case, as ] have now seen a large number, perhaps 
hundreds, always in exactly the same position, and see no reason 
why we should not speak of them as ovicells. One section shows 
the zocecial wall bulging out and the ovum partly in this portion, 
which is the commencement of the ovicell £. 

Loc. Arctic; Atlantic; Mediterranean; Gulf of Manaar; 
Zanzibar (Hincks); 8. Africa; Australia; New Zealand, Tas- 
mania; Pacific ¢ (Robertson); Tristan da Cunha (‘ Challenger’). 
Wasin, Brit. E. Africa, 10 fath. (500 $), collected by Crossland. 

Fossil. Upper Tert. Italy (Veviani), Helvetian of Egypt (Canv). 


* “ Bryozoa from Rapallo,” Journ. Linn. Soc., Zool. vol. xxvi. p. 5, pl. i. figs. 1-5 
1896). 

+ Morph. & Syst. Studies on the Cheil. Bry. p. 93 (1909). 

{ Since the above was written, Prof. R. C. Osburn has confirmed the existence of 
ovicells in the position described, having found numerous such ovicells in specimens 
from Fish Hawk Station: see “ Bryozoa of the Wood’s Hole Region,” Bull. of the 
Bureau of Fisheries, vol. xxx. Document No. 760, p. 220 (1912). 

§ These and similar numbers are Crossland’s registration numbers. 


BRYOZOA FROM ZANZIBAR. 465 


ABTEA TRUNCATA Landsborough. (Pl. LXIV. fig. 3.) 


There are only small fragments from Prison Island, Zanzibar 
Channel. The creeping tube or stolon is dotted in just the same 
way as the erect tube. 

Loc. Arctic; British; Danish; Madeira, Naples, Rapallo, 
Cette. Prison Island, Zanzibar Channel (505), 8 fath., collected 
by Crossland. 

Fossil. Helvetian, Egypt (Canz). 


BRETTIA TROPICA, Sp.n. (Pl. UXIV. figs. 4, 5.) 


There are only the zocecia figured, and it will be seen that the 
species is closely allied to 5. australis Busk, but differs in the 
shape of the area, which in B. iropica is about the length of 
a zoecium. There are on the dorsal surface the two light disks * 
on each side as in B. longa Waters, but in &. tropica the distal 
ones are very minute with the proximal one much larger. The 
lower zocecium arises from a calcareous knob, from which stolons 
spread out. The zocecium is calcareous, and most of the species 
of Lrettia are more or less calcareous, but with such small 
fragments we cannot know much about its relationships. 

I found species of Grettia in the Arctic and one in the Antarctic, 
and though the differences are very small it does not seem that the 
present form can be placed with any of those already described. 
Brettia may have avicularia, but then it has been called Con "0 - 
porella Hincks. 

Loc. Wasin, Brit. East Africa, 10 fath. (501), collected by 
Crossland. 


SYNNOTUM AVICULARE Pieper. 
Waters, Rep. Sudanese Red Sea, p.-129. 


Loe. (additional). Wasin, Brit. EK. Africa, 10 fath, (501); Ras 
Osowamembe, 10 fath. (504); Meweni Bay, 6 fath. (510). 


SYNNOTUM PEMBAENSIS, sp.n. (Pl. LXIV. figs. 12-15.) 


Zoarium with a spreading ramifying stolon, from which several 
erect stems arise, just as in Stirparia ete. The stem is about the 
same sizeas the creeping stolon, and issmooth for about the length 
of two or three zocecia, then there is a short zocecium followed by 
the pairs of zocecia. 

The zocecia are shorter and stouter than those of Synnotwm 
aviculare P. At one side.at the distal end there is a round 
pedunculate avicularium, but no sessile avicularium as in JS. 
aviculare. The first zocecium of each branch is uniserial, 
is also the case in S. aviculare, Notamia bursaria L., Dimetopia, 
and Calwellia, whereas in Gemellaria loricata L. there is a pair 
of zocecia at each fresh bifurcation. The pairs of zoccia turn 
alternately slightly to right and left, and there are radicles from 
between the zocecia just as in Synnotwm aviculare. There seem 


* There are similar disks in Catenaria and Vittaticella. 


466 MR. A. W. WATERS ON 


to be several pores in the large distal rosette-plate. There are 
10 tentacles. 

Loc. Wasin, Brit. E. Africa, 10 fath. (501); Chuaka, Zanzibar, 
2 fath. (508); Chaki-Chaki Bay, Pemba Island, near Zanzibar 
(517), collected by Crossland. 


SYNNOTUM ConTORTA, sp.n. (Pl. LXIV. figs. 16-18.) 


Synnotum aviculare Robertson, ‘* Non-Incrusting Bryozoa,” 
Univ. of Calif. Publ., Zool. vol. 11. p. 286, pl. xiv. figs. 84, 85 
(1905). 

The zoarium is coiled up, especially at the end. The branches 
of the zoarium dichotomise, and consist of pairs of zocecia back to 
back directed alternately in the opposite directions at right 
angles. 

The zocecia are wide, subtruncate at the top, diminishing regu- 
larly to the base, with a sessile avicularium at one or both sides 
near the distal end, and there are a few large rounded, pedun- 
culated avicularia replacing one of the sessile avicularia, but none 
of these are found in the older zocecia; the area is large, occupying 
more than two-thirds of the front, and the calcareous layer is much 
more solid than in any other of the Gemellaridze examined. The 
first zocecia at the bifurcations are single, whereas the next ones 
are double, being back to back. There are long radicles from the 
side of the zocecia. In the lateral wall there are two rosette-plates 
near the distal end. 

There are 11 tentacles. 

We know Gemellaria loricata L. without any avicularia, S. 
pembaensis nov. with terminal pedunculated avicularia, the 
present species with sessile avicularia and a few short thick 
pedunculated avicularia, and S. aviculare also with both sessile 
and peduneulated avicularia. ‘he presence or absence of avi- 
cularia is constantly turning out an unsatisfactory generic 
character, and I have never felt quite satisfied that a new genus 
was required for Synnotwn. 

Miss Robertson’s figure shows the zocecia more attenuated 
below, but it certainly seeras that this is the species she 
described. 

Loc. Chuaka, Zanzibar, 2 fath. (508); Chaki-Chaki, Pemba 
Bay, near Zanzibar, low water (517); Wasin, Brit. East Africa, 
10 fath. (501). 


HucrAtTEA CHELATA Linn. 
See Miss Jelly’s Catalogue, and add :— 


Scruparia chelata WKirchenpauer, Bericht ther die Unter- 
suchungs-Fahrt der Pommerania, ‘‘ Bryozoa,” p. 181 (1875). 

Eucratea chelata Levinsen, Zool. Danica, p. 42, pl. 1. figs. 8-9 ; 
Calvet, Bry. de Cette, p. 12; Robertson, ‘“ Non-Incrusting 
Bryozoa,” Univ. of Calif. Publ., Zool. vol. ii. p. 248, pl. v. figs. 7-9 
(1905); Barrois, Emb, des Bry. p. 194, pl. xv. figs. 10-12 (1877) ; 


BRYOZOA FROM ZANZIBAR. A67 


Osburn, “ Bry. of Wood’s Hole Region” Bull. Bur. Fish. vol. xxx. 
p. 221, pl. xxi. fig. 15 (1912); Nordgaard, “‘ Die Bry. des West. 
Norwegens,” Die Meeresfauna von Bergen, p. 76. 

A tew zocecia were seen from Wasin. 

Loe. As far north as the Lofoten Islands; Atlantic; British ; 
Mediterranean ; California; Australia; S. Africa (4. W. I. coll.). 
Wasin, Brit. East Africa, 10 fath. (501), collected by Crossland. 


BEANIA SPINIGERA MacGillivray. 

Diachoris spirigera MacG, Trans. Roy. Soc. Vict. vol. - iii. 
p: 165, pl. uu. fig. 12 (1859); Prod. Zool. Vict. dec. v. p. 32, 
pl. xvi. fig. 3; Waters, Ann. Mag. Nat. Hist. ser. 5, vol. xx. 
p. 94 (1887). 

There are some small specimens from Wasin which correspond 
in most particulars, though the avicularia ave materially smaller 
than the type, more like those of B. intermedia Hincks. There 
are three terminal spines and usually 5-6 delicate lateral spines. 

Loe. Victoria (Australia); New South Wales. Wasin, Brit. 
Hast Africa, 10 fath. (501), collected by Crossland. 


BEANIA MIRABILIS Johnston. 

For synonyms see Miss Jelly’s Catalogue, and add :— 

Beania mirabilis Hincks, Ann. Mag. Nat. Hist. ser. 3, 
vol. vill. p. 36 (1862); op. cit. ser. 5, vol. xiii. p..357 (1884) ; 
op. cit., ser. 5, vol. xix. p. 215 (1887); Waters, Journ. Linn, Soc. 
Zool. vol. xxvi. p. 17, pl. u. fig. 1 (1896); Jullien & Calvet, 
‘ Bry. prov. des Camp. de l’Hirondelle,’ p. 38 (1903) ; Thornely, 
Ceylon Pearl-Oyster Fisheries, vol. iv. Suppl. Rep. xxvi. p. 109 
(1905): Robertson, “ Non-Incrust. Chil. Bry.” Univ. of Calif. 
Publ., Zool. vol. ii. p. 276, pl. xii. figs. 63, 64, & fig, in text (1905). 

Loc. Northern; British; French coasts; Atlantic (Jull. & 
Calv.); Mediterranean ; Ceylon (7/.); Burmah (H.); Australia ; 
Pacific coast of N. America (/od.). Meweni Bay, Zanzibar (510), 
collected by Crossland. 

Beania has been considered to belong to the Flustride by 
Busk, and to Bicellaride by Levinsen, but the large embryo 
found by me in &. magellanica* seems to indicate the proba- 
bility of the genus standing elsewhere. There are 20-26 
tentacles in Seania, whereas in Bicellaride and its allies there 
are usually fewer, 12-18. &. magellanica B. has 23-26; B. hirtis- 
sima Hell., 20-30; Bb. hyadesi Jull., 20; B. quadricornuta H., 
23 (W.); B. spinigera MacG., 20; 6. mirabilis Johnst., 20. 


BICELLARIA CHUAKENSIS, sp.n. (PI. LX VIII. figs. 7, 8.) 

The zoarium arises froin a long, erect primary with radicles ; 
it frequently anastomoses, forming a colony about 7-8 mm. high. 

The primary zoccium has an elongate area with nine short 


* Ann. & Mag. Nat. Hist, ser. 8, vol. ix. p. 493 (1912). 
Proc. Zoot. Soc.—1913, No. XXXII. 32 


468 MR. A. W. WATERS ON 


spines, the next zoccium has seven spines, while the younger 
zocecia have usually three spines, two outer ones and one inner, 
though there may be occasionally two or four spines instead of 
three. The area is less than half the length of a zocecium, and 
the pedunculate avicularia are placed at about half the height 
of the area, whereas in B. ciliata they are much below it. The 
avicularia are of moderate size and similar in shape to those of 
B. ciliata; however, avicularia do not occur on any of the lower 
zocecia, so that, counting from the primary, no avicularia will be 
found before about the 18th zocecium. 

The pedunculate ovicell, directed laterally as in B. ciliata, is 
situated on the inner side near the distal end. 

Loc. Taken in tow-net, Chuaka Bay, Zanzibar (515); Wasin, 
Brit. E. Africa, 10 fath. (500), on Steganoporella magnilabris ; 
Chuaka, 2-3 fath. (612), collected by Crossland. 


Srrrparta EXILIs MacGillivray. (Pl. LXVI. figs. 1-3.) 


Stirparia ewilis MacG. “ Desc. of New or Little-known Polyzoa,” 
pt. xiii. Proc. Roy. Soc. Vict. n.s. vol. 11. p. 107, pl. iv. figs. 1-16 
(1890). 

In the Wasin specimens the lower part of the stem is buried 
in sponge and cannot be completely examined ; the upper part is 
annulated for a short space; the rest is smooth, unjomted, and 
without any strengthening rods, but at irregular intervals there 
are contractions, or sometimes two or three together, with rosette- 
plates across the stem in places. Although these stems differ 
from those of the other two species found, yet they are divided 
up by these contractions into lengths often about equal to those 
of S. dendrograpta; however, the growth is much simpler, so 
that, perhaps, S. ewilis may ultimately have to be placed in 
another genus. Fresh branches are given off at right angles 
to the main stem and start from an expanded disk (fig. 3). 

The tuft is 5-6 mm. long, and the zocecia face to the outside 
of the tuft. There are about 12 tentacles. The area is about 
two-thirds of the length of a zoccium ; there are three spines, 
or, in parts two; and only a very few, almost globular, avicularia 
have been seen (about two in each tuft), and these are short with 
apparently a wide mandible rounded at the end. The avicularia 
are attached just below the area and there are no ovicells, only 
the commencement of one. MacGillivray found neither avicularia 
nor ovicells. 

The opercular opening is low down. At a bifurcation one 
zocecium extends up a short distance on the opposite side of the 
bifureation (fig. 1), and this is well shown in the British Museum 
specimens of S. exilis MacG., from Port Phillip Heads. 

The mounted specimen from the ‘ Challenger,’ named S. glabra, 
is S. annulata Mapl., though there are mounts of a stem of a 
species which are like those of 8. glabra and S. dendrograpta. 
Busk’s figure, however, seems to show S. glabra H., so that we 
have a puzzle; but we may be right in coucluding that S. glabra 


BRYOZOA FROM ZANZIBAR. 469 


and S. annulata were both obtained from the ‘ Challenger’ Station, 
off Bahia. In the British Museum ‘Challenger’ specimen there 
are six spines to the primary of the tuft, not a number as figured ; 
there are no avicularia or ovicells, but there is a central spine in 
the same position as the avicularium in S. glabra H. 

Loc. Port Phillip Heads (J/acG".). Wasin, Brit. East Africa, 
20 fath. (522), collected by Crossland. 


STIRPARIA ZANZIBARIENSIS, sp. n. (Pl. LXVIII. figs. 1, 2; 
Pl]. LXIX. fig. 14.) 


The stem throws out branches which may bear a tuft 6—7 mm. 
long, and the internodes of the stem are approximately equal. 

The zocecia are alternate and turn partly away from each other, 
that is, the central line of the branch is raised so that the zocecia 
slope laterally downwards. The zocecium is much wider at the 
distal end than below, and the area is rather more than half 
the length of a zocecium. The first zocecium of a tuft has 9-11 
long spines, usually six on one side and four on the other 
(or dorsal side) with the central spine long; then the second, 
third, and sometimes fourth zocecium have several spines, whereas 
the normal zocecia have usually only one stout spine at the upper 
inner angle, though occasionally there is also one at the outer 
angle. 

There is sometimes an avicularium to the second zocecium, and 
this and the subsequent avicularia, which are long and narrow, 
are situated close to the base of the zocecium. 

There are about 14-15 tentacles. 

On the lower part of the stem or stolon the radicles are some- 
times replaced by capsules, similar to those described in my paper, 
“ Bryozoa from Rapallo,” * and they may be filled with bright 
yellow homogeneous contents. Levinsent refers to similar 
capsules as occurring in Bugila caliculata Lev. 

The ovicells are pedunculate, and there is a calcareous cover 
over a part only, not exceeding the half of a globe, so that the 
embryo is thereby but slightly protected, sometimes not at all. 
The calcareous wall of the ovicell is made of plates deposited 
from centres and looking like the shell of a turtle. 

The ovaria are central immediately proximal to the cecum, 
and the testes fill up the proximal part of the zowecium. No 
ovarium in my sections has more than one ovarian cell. 

The zoarial growth is similar to that of S. dendrograpta, sp. n., 
with the long nodes as depicted in Pl. LXVI. fig. 4; but the 
tufts are longer, and both the stout spinous processes and the 
long avicularia are distinguishing characters. There is no line of 
chitinous thickening as in S. dendrograpta. 

Loc. West Australia, some imperfect specimens in my col- 
lection. Chuaka, Zanzibar, 3 fath. (506), collected by Crossland, 


* Journ. Linn. Soc., Zool. vol. xxvi. p. 19, woodcut fig. 6 (1896). 
+ Morph. & Syst. Studies on Cheil. Bry. p. 102. 
32* 


ATO MR. A. W. WATERS ON 


STIRPARIA DENDROGRAPTA, Sp. n. (Pl. LXVI. figs. 4-9.) 


The stems grow from spreading stolons, and at frequent 
intervals branches occur which may bear tufts of zocecia or may 
produce other branches, and both the original stem and the 
branches are divided into segments approximately equal, though 
an internode below a tuft is frequently shorter than the others 
From the base of most of the internodes there is a radicle or a 
pair with frequently a cervicorn grapnel at the end. The colony 
may grow to at least 50 mm. long, and the tufts 3-5 mm. long 
originate from a zocwcium entirely different from the later zoecia, 
having more or less the character of a primary zoccium. When 
first described the stem of Stirparia was considered to be the 
equivalent of radicles, but this is not the case. 

The first zocecium (figs. 4, 6) has the area a little more than 
half the length of the zocecium and is surrounded by eight very 
long spines, often attaining about four times the length of a 
zocecium. ‘The spines of the first zooecium of a tuft are, however, 
not bilateral, but are five on one side and three on the other, the 
smaller number being on the side from which the next zocecium 
grows. The next zocecium has a somewhat similar area, with 
about five spines, and the avicularium is near the base of the area, 
while the following zocecium approximates to the later zocecia in 
having the avicularium somewhat higher than in zoocium no, 2, 
though still low down. In subsequent zomwcia they are placed 
still higher, their normal position in the older zoecia being at 
the distal end on the outside corner. Typically they may be 
terminal in the fourth pair of zocecia, or they may continue 
lateral until the eighth, and after the appearance of a terminal 
one subsequent zocecia are also generally at the corner of the 
distal end. 

The avicularia are short with a distinct beak. 

The branches of the tufts do not form a complete cup as in 
S. exilis MacG. and S. zanzibariensis, sp. n., and the zoccia are on 
the inside of the cup, whereas in the others they are on the 
outside. The branches of the tuft dichotomise, and the spread- 
out fan-shaped tuft is 3-5 mm. long, having often ten pairs in 
succession. 

The zocwcia are alternate and diagonal with the area a little 
more than half the length of a zoocium, and the full number of 
spines is three long ones at the distal edge, though many of the 
lower zowcia may have one and the younger zoecia two spines ; 
nor do the same number occur on both sides of a branch, the 
zocecia on the outer zoarial side having more spines than those 
on the inner; with three on the outer, there is often only one on 
the inner side. 

The ring-shaped oblique chitinous thickening, to which 
Levinsen * refers as occurring in Bicellaria ciliata L., is often 

* Morph. & Syst. Studies on Cheil. Bry. p. 101. Levinsen puts B. caliculata 


Ley. under Bugula, but his Pl. ui. fig. 1 shows the character of Bicellaria in 
having the long tubular proximal part. 


BRYOZOA FROM ZANZIBAR. 471 


quite distinct on the dorsal surface without being seen on the 
anterior surface; in other cases a mark is seen all round 
(fig. 4). When put into Kau de Javelle the zowcium often 
breaks off at this line, and it is seen that this proximal tubular 
part is connected at the base with the lower zocecium through a 
rosette-plate. A rosette-plate higher up connects with the next 
younger zocecium (fig. 7). 

The ovicell is lateral and pedunculate, and there are about 
12 tentacles. 

Radicles may occur in abundance on the lower zowcia and not 
merely from the normal position at the proximal end of the node. 

Immediately below the cecum there is a small globular body 
which, as it grows, is seen to be the ovarium, but in no case has 
more than one ovarian cell been seen. The same thing occurs in 
various Cellularide. 

In many respects S. dendrograpta resembles S. glabra Hincks, 
but that species has the stem internodes long and short alter- 
nately and also there is no avicularium at the distal end in 
S. glabra. From S. exilis MacG., it differs in the internodes 
being approximately equal and in the different character of the 
avicularia. 9. eailis has 6-7 spines on the primary zocecium. 

The graptolite, Dendrograptus serpens Hopkinson *, has similar 
colonies growing on a stout stalk, and the subcolonies and branches 
are about the same size as those of the Stirparia ; and some sub- 
colonies I collected in Llandrindod Wells of D. ser “pens H., or a 
closely allied species, have the branching quite similar to this 
Stirparia and might have been an impress of it, but as competent 
authorities have found it to be a graptolite, it slows how identical 
the growth may be in widely different classes. The name is 
given on account of the superficial likeness. 

Loc. Chuaka, Zanzibar, 2 fath. (508), collected by Crossland. 

There is a specimen of S. dendrograpta from Port Phillip in 
the British Museum. 


BUGULA NERITINA var. MINIMA Waters. 


Waters, “ Rep. Sudanese Red Sea,” p. 136, pl. xi. figs. 4-7, 
for syn., and add Thornely, “ Mar. Polyzoa of the Indian Ocean,” 
Trans. Linn. Soc., Zool. vol. xv. p. 141. 

Loc. N. S. Wales (W.); Red Sea (W.); Cargados Reef, 
Providence, 50-78 fath. (7h.). Prison Island, Zanzibar Channel, 
8 fath. (505); Ras Osowamembe, Zanzibar Channel, 10—20 fath. 
(514); Chuaka, Zanzibar, tow-net (515), collected by Crossland. 


Buevza rosusta MacGillivray. (Pl. LXIX. figs. ae aa. 
Bugula robusta MacG. Trans. R. Soc. Vict. vol. ix. p. 129 


(1868) ; Prod. Zool. Vict. dec. viii. p. 29, pl. Ixxviil. fig. “1 (1883). 
Bugula capensis Busk, MSS. 
* Hopkinson & C. Lapworth, “On the Graptolites of the Arenig and Llandeilo 


Rocks of St. Davids,’ Quart. Journ. Geol. Soc. vol. xxxi. p. 665, pl. xxxvii. fig. 3 
(1875). 


472 MR. A. W. WATERS ON 


There is a specimen from Wasin which has the zoecia a trifle 
smaller than the South African and Australian specimens, and 
the avicularia are somewhat smaller; however, in a specimen in 
my collection, determined by Busk a B. capensis *, there is one 
small avicularium, while the rest are large. ‘There is no real 
spinous process at the outer angle, but neither do I find more 
than a projection in any of my specimens from other localities. 

The distal rosette-plates are all close to the basal wall and very 
small, so that it is difficult to distinguish them. In some other 
Bugule they are similarly situated, though in others they are 
spread over the wall. There are no ovicells in this specimen, 
but the species has lateral ovicells like those of B. neritina L., 
and the brown colour suggests its belonging to the neritina group. 

The primary zoecium is very long and narrow, followed by a 
second long one, then a zoceciuin about the ordinary length, after 
which the growth is biserial. The first two zocecia remind us of 
the segments of the stem of various Stirparie. 

Bugula, asa rule, shows no articulation, but B. reticulata B. 
is distinctly articulated—that is, at the bifurcation there are 
distinct thick articular chitinous tubes. 

Other tropical species of Bugula are:-—B. dentata Lamx., 
B. mirabilis B., B. versicolor B., B. reticulata var. unicornis B., 
B. gracilis B., B. neritina var. rubra Thornely, B. neritina var. 
tenuata Th., B. neritina var. ramosa Th. These last two in 
many particulars resemble B. reticulata Busk. Most of the 
tropical species have a very wide distribution. 

Loc. Victoria (McG); South Africa (as capensis); Port 
Hlizabeth, 8. Africa (A. W. W. coll.). Wasin, Brit. EH. Africa 
(501), collected by Crossland. 


SCRUPOCELLARIDS. 


Levinsen 7 does not consider that Menipea can be divided up, 
as If jroneeedl by the character of the jointing, and on p. 133 
gives his account of the articulation of “all Bryozoa that oceur 
in jointed colonies,” but his account is not exhaustive. Dealing 
now with much more material than on the previous occasion, 
my suggestion is more fully tested. 

The jointing in the Scrupocellaridz varies considerably, giving 
useful specific characters, and it is most important that we should 
trace it from its simplest to its most complicated condition. 
As I have shown, in most articulated Bryozoa the branches 
are at first continuous, and the last two or three bifurcations 
may show no sign of rupture, which only takes place after 
the chitinous tubes have been formed; and in most articulated 
Bryozoa the chitinous tube is formed within the calcareous wall, 
though in some it may be formed merely within the membranous 


* See my remarks, Journ. Linn. Soc., Zool. vol. xxxi. p. 137 (1909). 
+ Morph. & Syst. Studies on the Cheil. Bryozoa, p. 183 (1909). 
t{ Journ. Linn. Soc., Zool. vol. xxvi. p. 2 (1896). 


BRYOZOA FROM ZANZIBAR. 473 


wall. Usually, in Scrupocellaride, the inner zocecium of the new 
pair is jointed close to the proximal end (Pl. LX VIII. fig. 14), 
but the position of the articulation of the outer zocecium varies 
considerably, having the part below the articulation much larger 
than the similar portion in the inner zowcium. There are three 
exceptions to this rule—WMenipea cirrata Hill. & Sol., MM. smitti* 
Norm., MW. flagellifera Busk—all three of which are without a 
scutum f. 


A.—Beginning with what seems to me the simplest form of 
articulation, namely, that found in Canda retiformis Pourt. 
(Pl. LXIX. fig. 6), though both genus and species have previously 
been described as non-articulated in consequence of the chitinous 
tube being often entirely covered by the calcareous wall, in which 
as yet there is no rupture, so that decalcification is necessary in 
order to study the articulation. As mentioned on p. 480, in the 
younger branches no articulation is found, but in the older ones 
there is a chitinous tube on the inside of one of the two branches 
and a similar tube on the opposite side in the next branch, and 
so on alternately (fig. 6). 


B.—From the simple form of Canda, we pass on to that of 
Scrupocellaria with two chitinous tubes, but with the outer 
zocecium havine the articular tube near the middle of the 


zocecium (Pl. LXITX. fig. 8). As an example, S. jolloisii Aud. 


C.—In the following group the chitinous tube in the outer 
zowcium 1s very much lower than in B, but is not, however, close 
to the proximal end of the zocecium. The articular tubes are 
here narrower, more distinct, and often separated as in Menipea 
patagonica Busk (Pl. LXIX. fig. 11). As examples, I/. ternata 
Ell. & Sol., Bugulopsis peachii Busk (Pl. LXIX. fig. 10), J. occt- 
dentalis Trask, MW. portert MacG. 


D.—We pass next to a group in which both articular chitinous 
tubes are close to the proximal end of the zowcia. I spoke of 
the proximal ends of these new zocecia as small chambers, but 
now consider that it would be better to compare them with the 
“basis rami” (Harmer) of Crisia, and think the designation may 
be used here, remembering that they are really the beginnings of 
new zocecia. As examples, Menipea buskit W. Th. (Pl. LXIX. 
fig. 12), MW. erystallina Gray, M. cervicornis MacG., MM. occi- 
dentalis Trask, M. cirrata Ell. & Sol. 


* Waters, “ Bryozoa from Franz Josef Land,” Journ. Linn. Soc., Zool. vol. xxviii. 
pl. vii. fig. 8 (1900). 

+ As I have always noticed the articulation when opportunities have presented 
themselves, several have been carefully figured. In my paper on “ Bryozoa from 
Rapallo,” Journ. Linn. Soc., Zool. vol. xxvi. pl. i. the position of the polypides in 
the zocecia of Scrupocellaria inermis is SONY figs. 11, 12, and op. cit. vol. xxviii., 
the joints of Menipea gracilis, pl. vi. fig. 1 2, Scrupocellaria scabra, fig. 14, and 
S. smittii Norm., fig. 8, are shown. In Te last the articulation only occurs 
beyond the distal end of the outer zocium. In the present paper the articulation 
is shown in Pl. LX VILLI. fig. 14. 


A474 MR. A. W. WATERS ON 


E.— Lastly, Menipea cyathus Wy. Thomp. has only one chitinous 
tube (Pl. LXIX. fig. 13). In Scrupocellaride there is, in some 
species near the distal end, a long body which may be folded 
back as in S. ferow Aud. (PL. LXVIII. fig. 14), or it may be 
very long, extending to the proximal end oF the zoceclum as 1n 
M. fla gells fera Busk! There is much to suggest that this functions 
as a testis, and that it should be compared with the organ in 
Flusira abyssicola Sars, and Cribrilina figularis Johust. 


The classificatory groups may now be considered :— 


1. Canna. Articulation simple as A, p. 473; ovicell smooth, 
imperforate; vibracula have the sete serrate. Two vibracula at 
a bifurcation. Levinsen considers that the ovicell is enclosed in 
the widened proximal half of the avicularium, whereas I should 
say that the avicularium is on the ovicell. 


CapereA. Articulation internal tubes, ovicell imperforate, 
vibracula with smooth sete. 


3. ScRUPOCELLARIA (div. 1). Articulation as B, p. 473 ; ovicell 
smooth, imperforate, usually with two vibracula at a bifurcation ; 
sete smooth. This includes S. delilii Aud., S. scruposa L., 
S. scabra Van Ben., S. cervicornis Busk, S. macandrei Busk, 
S. ornithorhynchus W. Th., S. serupea var. dongolensis Waters. 
(S. scabra has sometimes one vibraculum at a bifurcation.) 


A, ScrupoceLLARia (div. 2). Articulation as B, p. 473; ovicell 
perforated, usually one vibraculum at a bifurcation, sete smooth. 
This includes S. bertholettii Aud., S. jolloisii Aud., S. mansueta 
Waters, S. reptans L., S. ferox Busk, S. obtecta Haswell, and 
probably S. portert MacG., and S. occidentalis Trask. 


5. Bueutopsis. Articulation as C, p. 473: ovicell imperforate, 
no vibracula. Example, B. peachii Busk. I follow Levinsen in 
using this generic name for the present, but have not had the 
opportunity of fully studying the genus. 


6. Menipra (div. 1). Articulation as D, p. 473; ovicell truly 
endozoccial, showing no external difference, no vibracula; 
avicularia sometimes suboral and sometimes lateral, short inter- 
nodes. ‘This includes J. cirrata Ell. & Sol., M/. crystallina Gray, 
M. cervicornis MacG. 


7. Munrpza (div. 2). Articulation as H, p. 474. Two zoeecia 
in an internode, scutum directed downwards from the distal end 
of the zocecium *. Possibly a new genus will have to be made for 
this. This includes J. cyathus. 


FLABELLARIS. Levinsen leaves in MZenipea species that cannot 
be referred to any other of the genera, and puts under it what I 


* Specimens of Menipea fuegensis Busk and WM. aculeata Busk have cases of one 
zocecium of a new braneh growing from the distal end of the terminal zocecium and 
also one from the side of the lower zocecium. 


BRYOZOA FROM ZANZIBAR. 475 


placed in a new genus Flabellaris*, and I then showed that 
Craspedozoum MacGillivray must be united with one of the types 
of Menipea Lamx., namely W/. flahellum Hill. & Sol., but it seems 
better to keep the name JJenipea for the first species mentioned 
by Lamouroux, namely I. civrata Lamx. However, this group 
of flabellaris does not seem to belong to the Scrupocellaride 
at all, but to the Membraniporide, having a Membraniporidian 
oviceli much like that of JZ. lineata L., MW. craticula Alder, 
M. unicornis Flem. The species included are /. flabellata Ell. & 
Sol., #. (M.) cuspidata Busk (Pl. LXIX. fig. 9), /. trisertata MacG. 
(specimens i in my collection have ovicells), /. (‘‘ Craspedozoum ”) 
reborata Hincks, F. (C.) ligulatwm Mee F. multiseriata Busk. 
fF. roborata and F. ligulatum when broken through have at the 
articulation interior tubes like the radicles. 

The ovaria of Flabellaris roborata are distal with many ovarian 
cells, and one or more grow to a considerable size. Jullien 
considered that Menipea must be merged in Scrupocellaria in 
consequence of having found one vibraculum on Menipea clausa 
Jull.=Serupocellaria marsupiata Busk, and this conclusion he 
considered was upheld by the fact that some colonies of S. scabra 
Van Ben. have no vibracula, while others have a few or some- 
times many. This seemed quite reasonable, and since then the 
idea has received further support, as Levinsen has found one 
vibraculum on WM. ternata, and, also, he found vibracula on 
Menipea benemunita Busk of the ‘ Challenger,’ for which Levinsen 
proposed the genus Caberiella. Also, the form and position of 
the radicle chamber in Scrupocellaria serrata Waters} suggests 
that a recent ancestor had vibracula. 

Although these cases prove that the presence of vibracula does 
not give a Sharp divisional line between what has been under- 
stood as Serupocellaria and Menipea, yet all the species could 
scarcely remain in one genus, and separation can be made on 
other grounds. The presence of avicularia gives but very limited 
assistance in classification, although there are characters in the 
avicularia which are very useful; so it is, therefore, not sur- 
prising to find that in some species vibracula may be found as 
an exception. 

Most of what have been called Menipea have an anterior 
avicularium immediately below the area or slightly to one side, 
though there are some species without any, as S. inermis, nor 
are lateral avicularia universal. 

Levinsen § considered that Caberea and his Caberiella had the 
avicularium divided into two chambers, whereas I was unable to 
find two, for while there is a prolongation of the vibracular 
chamber, this only seems to be for the groove in which the 


* Waters, “On Membraniporide,’ Journ. Linn. Soe., Zool. vol. xxyi. p. 672, 
pl. xlvini. figs. 10, 11; xlix. figs. 7-10 (1898). 

+ ‘Cap Horn,’ p. 69 (1888), and Bull. Soc. Zool. de France, p. 507 (1882). 

~ Report on Red Sea Bryozoa, Journ. Linn. Soc., Zool. vol. xxxi. p. 133, pl. x. 
figs. 11-14 (1909). 

§ Morph. and Syst. Studies on the Cheil. Bryozoa, p. 134. 


476 MR. A. W. WATERS ON 


vibracular seta les. Exactly the same thing, though not so 
marked, is seen in Scrupocellaria, for in several species the 
vibraculum extends beyond the median line, as in S. macandret 
Busk, S. incurvata Waters, ete. 

As I told Dr. Levinsen that I did not find two chambers, 
he kindly sent me some vibracula, skilfully separated, which, 
however, only confirmed what I had seen in my own specimens. 

In the Scrupocellaride, so far as I have seen, the ovaria are 
large, situated near the distal end, and contain several ova which 
are developed into large ova before they pass into the ovicell. 
On the other hand, in Bugula, Bicellaria, etc., the ovaria are, at 
the proximal end, usually very near to the base of the ceecum ; 
they are very small with usually two small ova, and when still 
extremely small, an ovum passes into the ovicell. It is very 
interesting to nal these generic differences in the ovaria, and 
undoubtedly the form, size, and position of the ovaria will be 
found to furnish useful characters i in many species of Bryozoa. 

The direction of evolution of the Scrupocellaridee seems to be 
indicated in the articulation, and a comparison of the changes in 
this family may help us to understand the Catenicellide better. 


SCRUPOCELLARIA FEROX Busk, (Pl. LXVIII. figs. 11-15; 
Pl. LXIX. figs. 7, 20.) 


Scrupocellaria feroc Busk, B. M. Cat. Mar. Polyzoa, p. 25, 
pl. xxu. figs. 1) 2; & 5. 

The avicularia vary considerably in size, being largest just 
below a bifurcation, and smallest or wanting in the younger 
zocecia, The avicularian chamber has the lateral projection to 
which I referred and figured in S. mansweta Waters*, from the 
Red Sea, and the long dorsal opening of the vibracular chamber 
in the older zocecia has a calcareous band across dividing it in 
two (see fig. 14). There is one vibraculum to a bifurcation some- 
what directed towards the front, as in S. cyclostoma Busk, and 
the radiele, which is hooked at the end, is not ringed as in 
S. cyclostoma, but there are only a few complete radicles in the 
specimens. No ovicells occur on the Zanzibar specimens. 

There are about 24 tentacles. 

The rosette-plate into the vibracular chamber is at the base of 
the chamber and has many pores; as this rosette-plate is not 
always very distinctly marked off these pores might be looked 
upon as several plates. Hach zocecium has its own lateral wall, 
so that when prepared in Hau de Javelle they may separate. 
Stained preparations show a band near the distal end bent 
back upon itself (fig.-126). The contents are granular, with 
hollow places at intervals, and probably the function is the same 
as in the bodies? I mentioned in Bugula bicornis Busk ¢, and they 


* Journ. Linn. Soc., Zool. vol. xxxi. p. 134, pl. x. fig. 15 (1909). 

+ Those on page 474 are compared with somewhat similar structures in 
M. flagellifera B., Flustra abyssicola Sars, and Cribrilina figularis Johnst. 

~ Résult. du Voyage 8.Y. Belgica, “ Bryozoa,” p. 21, pl. 1. fig. 4 (1904). 


BRYOZOA FROM ZANZIBAR. ATT 


are probably testes, or connected with the testes, and in all the 
specimens prepared these bands are found in all the zocecia, but 
there are no traces of ovaria, so it may be that the male organs 
are on one colony and the female on another, 

In one mount with about 130 zocecia all contain fully developed 
polypides, showing that degeneration does not always take place 
at such short intervals as has been often stated. There are, 
however, large buds developing by the side of the active 
polypides. 

Loc. Louisiade Archipelago, Bass’s Straits (B.). Prison Island, 
Zanzibar Channel (505), 8 fath.; Ras Osowamembe, Zanzibar 
Channel, 10 fath. (504 & 514); Wasin, Brit. EK. Africa, 10 fath. 
(507), collected by Crossland. 

Other species of tropical Scrupocellaria, not however found in 
the Crossland collections, are S. delilii Aud. ; S. ornithorhynchus 
B.; S. ciliata Aud. ; S. annectans MacG.; S. diadema B.; S. minuta 
Kirkp.; S. clypeata Hasw.; S. obtecta Hasw. 


SCRUPOCELLARIA CERVICORNIS Busk. (Pl. LXIX. figs. 3, 4.) 


Waters, ‘‘ Rep. Sudanese Red Sea,” p. 166. 
Loc. add Ras Osowamembe, 10 fath. (504); Wasin, Brit. Kast 
Africa, 20 fath. (522), collected by Crossland. 


SCRUPOCELLARIA MACANDREI Busk. (Pl. LX VIII. figs. 5, 6.) 


Scrupocellaria macandrei Busk, B. M. Cat. Mar. Polyzoa, p. 24, 
pl. xxiv. figs. 1-3 ; and add to Miss Jelly’s synonyms :—Haswell, 
W.A., “ Polyzoa from the Queensland Coast,” Proc. Linn. Soc. 
N. S. Wales, vol. v. p. 37; Philipps, “ Rep. on the Polyzoa,” 
Willey’s Zool. Results, pt. iv. p. 442 (1899); Calvet, ‘ Exp. Scient. 
du Travailleur et du Talisman,’ p. 375 (1907); Thornely, ‘‘ Mar. 
Polyzoa of the Indian Ocean,” Trans. Linn. Soc., Zool. vol. xv. 
p-. 140 (1912). 

This belongs to the S. scrwpea group. There are three outer 
spines and one inner near the peduncle of the scutum. The 
outer spines are often very long. The groove of the vibraculum 
is continued beyond the vibracular chamber, passing the median 
line of the zocecium, in this respect somewhat resembling S. i- 
curvata Waters. There are two vibracula at a bifurcation. The 
ovicell is smooth, placed somewhat diagonally, and has a clear 
space on the front. The vibracular seta is about the length of a 
zocecium, smooth but flat in the middle, so that perhaps we may 
call it sickle-shaped. The oral aperture is placed diagonally 
back, as in Caberea darwin Busk. 

Loc. Cape Verde Island, 1070-1150 fath. (B.) & 110-180 met. 
(Calv.) & Crossland Expedition ; St. Paul’s Rocks, N. Atlantic 
(B.); Coast of Spain (B.) ; Adriatic (Heller) ; Lifu (Phil.) ; Queens- 
land (Haswell); Providence, 50-78 fath., Amirante, 29 fath., 
Farquhar Reef, Cargardos, 30 fath., and Seychelles, 34 fath. 


478 MR. A. W. WATERS ON 


(Ind. Ocean, Thornely). Prison Island, Zanzibar Channel (505), 
8 fath., coll. by Crossland. 


SCRUPOCELLARIA PILOSA Savigny & Audouin (now Busk). 
(Pl. LX VITI. figs. 3, 4.) 


Crisia pilose Aud. Description de ’Egypte, Hist. nat. p. 241; 
Savigny’s pl. xi. figs. 1,—1,. 

Cellularia spatulata dOrb. Pal. Frang., Terr. Cret. p. 50 
(1850-52). 

Some dried specimens from Wasin, B. E. Africa, seem to be 
the species figured by Savigny. The zocecia are narrow, pro- 
ducing a wavy appearance, as figured by Savigny. The opesium 
has a very narrow border, and the scutum, which varies in shape, 
is small and does not nearly cover the aperture. Both the distal 
and proximal ends of the seutum are rounded, with the distal end 
the larger, and the scutum is without cervicorn marks. There 
are two or three oral spines on the outer side and one or two on 
the inner, the outer ones, especially the lower one, which is 
stouter, are sometimes long, though in the specimens they are 
mostly broken off. The vibraculum is small, partly free at the 
outside of the zoarium, with the groove extending slightly beyond 
the vibracular chamber, but not as much so as in S. macandrei B. 
The vibracular sete are smooth and rather longer than a zocecium. 
The zoarium at a bifurcation has a medium spine and one vibra- 
culum. The lateral avicularium is placed somewhat diagonally, 
instead of standing straight out. The radicles are large and are 
serrate near the ends. 

There are no ovicells on the specimens. 

Busk in the ‘Challenger’ Report, p. 24, describes a species 
from the southern hemisphere as S. pilosa Sav., but this seems 
a doubtful determination, for the shape of the scutum is different 
and the vibracular chamber is very wide and large. 

Busk speaks of it as the species of Audouin, whereas Audouin 
ealls it the species of Lamouroux, and supported the identity on 
Lamouroux having presented some of the type to M. Bory de 
St. Vincent which was compared. However, it was never the 
species of Lamouroux but of Pallas, and while his description 
would tally with this species, 1t would equally well cover a large 
number of other Scrupocellarie. D’Orbigny gave the name 
spatulata to Savigny’s figure, and in the same way he named 
many of Savigny’s other figures, although already named by 
Audouin, Savigny having given recognisable figures, we may 
suppose that Pallas and Lamouroux were dealing with the same 
thing, although this can never be known with certainty. 

This is much like Scrupocellaria pusilla Smitt, which, however, 
has cervicorn markings on the scutum. 

The various species now considered might well be ranged round 
S. scruped as varieties. 

Loc. ¢ Mediterranean (Pallas, etc.). Wasin, Brit. Hast Africa, 
10 fath. (500), collected by Crossland. 


BRYOZOA FROM ZANZIBAR. 479 


SCRUPOCELLARIA WASINENSIS, sp. n. (Pl. LX VIII. figs. 9, 10; 
P}, LXIX. tigs. 17-19.) 


Zoarium usually with 5 or 7 zocecia im an internode. Zocecia 
wide, with the round area occupying more than half the length 
of the zoccium:; at the distal end there are three exterior spines 
and one interior; the anterior avicularia are medium-sized, 
raised, tubular, with a narrow triangular mandible; the lateral 
avicularia are very small; the vibracular, together with the radi- 
cular chamber, is about half the length of a zocecium, the delicate 
vibracular setee are smooth and shghtly longer than a zocecium ; 
the vibracular chamber is separated near the base from the 
radicular chamber, and the distal end is contracted. There is 
one vibraculum at a bifurcation. A few large lateral avicularia 
have been seen which are divided at the end, and the mandible 
is forked like that of S. serrata Waters *. 

The globular ovicell has numerous pores. 

The radicles sometimes pass from one branch to another, as in 
Caberea retiformis Sim. 

There are about 16 tentacles. 

The ovaries containing many ova are at the distal end, while 
testes in the same zocecia are at the proximal end. The ovaria 
in Scrupocellaria contain several ova, but the ovaria in this 
species are peculiar, as there are at first a number of large homo- 
geneous cells of which usually only one or two show any nucleus 
(PL LXIX. fig. 19). The ovaria at this stage show no follicular 
wall, but ata later stage, when the yolk-mass of an ovum has 
become very large, then the follicular wall is distinct. 

In this species, as in other Scrupocellarie, a large ovum passes 
into the ovicell, whereas in Bugula the ovaria instead of being 
distal are proximal, and a small ovum in a very early stage passes 
into the ovicellt. The ovarian cells, which are frequently far 
apart, are often surrounded by a protoplasmic network which 
passes up to the rosette-plate of the next zocecium (figs. 18, 19). 
This species is of about the same size as S. mansueta Waters, but 
has four distal spines and a much smaller vibraculum, with sete 
more delicate, and about half to one-third the length of the latter 
form. The large lateral avicularia are known in S. varians 
Hincks, S. serrata Waters, and S. obtecta Haswell, but apparently 
are only the ordinary avicularia more developed. 

Loc. Wasin, Brit. Hast Africa, 20 fath. (522), collected by 
Crossland. 


CANDA RETIFORMIS Pourtales. (Pl. LXIX. figs. 1, 2, 6.) 


Canda retiformis Pourtales, Bull. Mus. Comp. Zool. Harvard 
Coll., I., No. 6, p. 110 (1867); Philipps, “Rep. on Polyzoa,” Willey, 


* “ Bryozoa of Sudanese Red Sea,” Journ. Linn. Soc., Zool. vol. xxxi. p. 133, pl. x. 
fig. 11. 

+ In Bugula there are two small ova in each ovarium, occasionally three, or even 
four, small ones; however, in B. murrayana Johnst., now called Dendrobeania 
by Levinsen, the ovarium is distal, and the ova in the ovarium grow to a large size, 
so that the material differences in the ovaria would alone suggest that murrayang 
does not belong to the same genus as B. avicularis, ete. 


480 MR. A. W. WATERS ON 


Zool. Results, pt. iv. p. 441, pl. xl. fig. 1 (1899); Thornely, 
“Mar. Polyzoa of the Indian Ocean,” Trans. Linn. Soc., Zool. 
vol. xv. p. 141 (1912). 

Caberea retiformis Smitt, “ Floridan Broyzoa,” pt. i. p. 16, pl. v. 
figs. 43-46 (1872); Thornely, ‘‘ Ceylon Pearl-Oyster Fisheries,” 
vol. iv. Suppl. Rep. xxvi. p. 109 (1905). 

2 Canda fossilis Waters, Q. Journ. Geol. Soe. vol. xxxvii. p. 322, 
pl. xvi. figs. 51, 52 (1881); MacG. “Tert. Polyzoa of Victoria,” 
p- 25, pl. ili. figs. 12-14, Trams. R. Soc. Vict. vol. iv. (1895). 


The specimens from Zanzibar have the scutum very narrow, 
ointed distally, and rounded proximally, as described by Miss 
Philipps, and I follow her in considering them to be the form 
described by Smitt, and although allied to C. arachnoides they 
seem quite distinct. It occurs in the Atlantic, the Indian Ocean, 
and the Loyalty Islands, without any connecting links being 
known. Caberea crassimarginata B., of the ‘Challenger,’ and 
Scrupocellaria clypeata Haswell, seem to belong to this group. 

No anterior avicularium has been described, but sometimes 
there is one attached to the inner side of the zocecium, and then 
it is usually just below a bifurcation. The avicularian chamber 
is wide, and the mandible is triangular. There are two vibracula 
at a bifurcation.’ 

Both Smitt and Levinsen have considered this species as having 
no articulation, and as the chitinous tube is often entirely covered 
by the calcareous wall in which there is as yet no rupture, it 
requires decalcification to study the articulation, which is peculiar 
and seems to differ from that of any other species examined. 
In the younger branches there is no sign of articulation, but in 
the older ones there is one chitinous tube on the inside of one of 
the two branches, but not of both, and a point to be noticed is 
that when the chitinous tube is on one side, say to the right, it 
is in the next branch on the left, in the one following on the 
right, and so on alternately, though in a few cases in the older 
parts of the colony I have seen a chitinous tube to each branch. 
In (. arachnoides Lamx., as I have shown*, there are two 
chitinous tubes, one on the inside of each branch. The articula- 
tion of CO. tenwis MacG. is somewhat similar. 

When decalcified, a chitinous tube is shown in the peduncle of 
the scutum, just as is seen in the base of many spines. 

Near the base of the vibracular seta there is a projecting 
delicate free arch at right angles to the axis of the seta, and in 
other species of the Scrupocellaride there seems to be a similar 
structure, though not so pronounced. 

One of the cross radicle tubes usually connects the two new 
branches at a very short distance from the bifurcation, and this 
may partly account for rupture at the articulation so seldom 


taking place (see fig. 1). 


* Ann. Mag. Nat. Hist. ser. 5, vol. xx. p. 89, pl. iv. fig. 7. 


BRYOZOA FROM ZANZIBAR. 481 


There are about 16 tentacles. 

The vibraculum of Canda retifornis is of the same character as 
that of Serupocellaria, in which genus some of the vibracular 
chambers extend beyond the median line of the zoarium, though 
not quite as much so as in the two species of Canda; also the 
vibraculum of Caberea has the same general character. 

I have several times maintained* that there are material dif- 
ferences between avicularia and vibracula, but that the length of 
the mandible or seta is of no importance in indicating which they 
are. The distinctions are in the chamber and the basal part of 
the chitinous organs. While the mandibles of avicularia are 
symmetrical and have the closing muscles attached by Tone or 
two long tendons, the base of asymmetrical seta of the vibracula 
is very complicated with a large number of curiously shaped pro- 
tuberances, to some of which the muscles are attached by a 
fascia, but without any long tendon, so that instead of there 
being two main muscles, there are more attached by a short band 
to various parts of the base of the seta. The vibracular base is 
very small, so that it is difficult to follow this complicated 
mechanism ; the reason for this complication is found in the seta 
being movable in all directions, whereas the mandibles of the 
avicularia only move in one. The mandibles all have a straight 
proximal edge, but this is not the case in the seta; further, the 
avicularian mandible works from this straight base either against 
the calcareous bar, or, in case this is not complete, then from two 
teeth. The universal movement could not take place with a 
eross-bar, and none has been found, nor must we ever expect to 
find one in vibracula. The vibracular chambers of this group 
are different from those of Cupularias, ete., in which the vibracular 
chamber takes the place of a zocecium. 

Loc. Florida, 68 & 270 fath.; Loyalty Islands (Ph.); Galle, 
deep water (7%.); Amirante, 23-29 fath., Saya de Malha, 55 fath., 
Seychelles, 39 fath., Cargados, 30 fath. (Thornely). Wasin, Brit. 
East Africa, 10 fath. (501); Prison Island, Zanzibar Channel 
(505), 8fath., Ras Osowamembe, Zanzibar Channel, 10 fath. (504), 
collected by Crossland. 


CATENARIA LAFoNTIT Audouin. 
Waters, ‘“ Rep. Sudanese Red Sea,” p. 131. 


Near the distal end there are small ovaria with two ovarian 
cells. 


Loe. Wasin, Brit. Kast Africa, 10 fath. (501); Prison Island, 


8 fath.; Chuaka, 2 fath. (508), 3 fath. (626); Meweni Bay, 6 fath. 
(510); Zanzibar town, shore (527). 


* Résultats du Voyage du S.Y. Belgica, “ Bryozoa,”’ p. 27. . Zool. Chall. Exp. 
vol. lxxix. p. 22, pl. 1. fig. 12. 

+ See page 529. 

ft Very minute glands occur in the vibracula of Cupularia. 


482 MR. A. W. WATERS ON 


CATENARIA DIAPHANA Busk. (Pl. LXIV. figs. 6-11.) 


Scruparia diaphana Busk, Q. J. Micr. Se. vol. vii. p. 281, 
pl. xxxi. fig. 1 (1860). 

Catenaria diaphana Busk, “ Polyzoa,” Zool. Chall. Exp. vol. x. 
pt. xxx. p. 14, pl. 11. fig. 3 (1884). 

Halysis diaphana Norman, “ Polyzoa from Madeira,” Journ. 
Linn. Soc., Zool. vol. xxx. p. 296 (1909); Levinsen, Morph. & 
Syst. Studies on Cheil. Bryozoa, p. 274 (1909). 

The ovicell has not before been described, in fact Levinsen says 
itis wanting. A growth of the outer calcareous wall of this recum- 
bent ovicell projects forward from the distal end in the middle 
and also from the sides, forming at first one elongate space which 
is subsequently divided, making two vacant spaces or large pores. 
The node with an ovicell or ovicells is never a single zocecium, 
and there are often many zocecia with ovicells in a node (fig. 9) ; 
six together have been counted many times. We have seen 
many ovicelligerous zocecia in a node in Catenicellide. The node 
may, however, be only one ovicelligerous zocecium followed by an 
ordinary zowcium. Asa rule, from the distal end of the older 
zoceclum a new zocecium arises in the median line, and there 
may also be one growing laterally from very near the end, or 
occasionally one on each side. 

The front wall is but little raised and is perforated, the per- 
forated part being bounded on each side by a raised ridge, and on 
the dorsal surface there are two lines of pores. The parietal 
muscles start under the longitudinal ridge. The operculum has 
a dark mark in the proximal part (fig. 8), and there are 20-22 
tentacles. There are radicles attached to the side of the zowcium, 
with the attachment elongate in the direction of the long axis. 
None of the ovaria seen in sections are surrounded by the 
follicular cells occurring in nearly all species. The ovarium 
(fig. 10), with many ovarian cells, is found near the basal wall, 
a short distance below the ovicell. The ovum is seen in the 
upper part of the zocecium, where there are strong muscles ready 
to force it into the ovicell from below internally, and sometimes 
there is an embryo or ovum in the ovicell as well as a large ovum 
below in the zocecium. Levinsen * considers that in some species 
of Bryozoa the ovum comes out of the zoecium and then enters 
the ovicell, but until proof is brought forward we may hold a 
position of doubt, as the contrary is known in so many cases. 

All the characters given by Norman for his genus Halysis 
apply to Catenaria lafontii Aud., and it is not clear that a new 
genus 1s required. Levinsen makes the absence of the avicularia 
and of ovicells a reason for separating this species from his genus 
Savignyella=C, lafontii Aud., but in a large proportion of genera 
there is sometimes an avicularium, sometimes none. The genus - 
Catenaria was discussed in my Report on the Sudanese Bryozoa 7, 


* Loc. cit. p 67. 
+ Journ. Linn. Soe., Zool. vol. yxxi. pp. 130, 131 (1909). 


BRYOZOA FROM ZANZIBAR. 483 


Loc. Madeira (.); St. Paul’s Rocks, N. Atlantic (Challenger), 
shallow water. Ras Osowamembe, Zanzibar Channel, 10 fath, 
(504); Prison Island, Zanzibar, 8 fath. (505), collected by 
Crossland. 


CATENICELLIDA. 


In the Catenicellide the ovicells are of much more use in 
classification than has been generally recognised, but Levinsen 
has ignored them in his synopsis of the genera, nor has he used 
the shape of the opercula. 

Based upon the form of the ovicells, there are two main 
divisions :— 

First, those in which the ovicell is a terminal gonecium, as 
Scuticella Lev., Costaticella* Maplestone, Cribricella ¢ Lev., all 
three of which have an operculum with a straight or but slightly 
curved edge, and Calpidiwm, which has a sinus. In all these 
genera the ovicelligerous zocecium has a much wider operculum 
than the ordinary zocecium, but the proximal edge is straight, 
although in both the goneecium of all of these, and the ordinary 
zoceclum of the first three, there may be an apparent sinus in 
the calcareous wall. 

SECOND, those in which the ovicell occurs in a node with other 
zoccia ; divided into 

I. Species with the ovicell occurring between two zocecia in a 
straight line and the operculum straight or nearly so on the 
proximal edge; and here we have Vittaticellat Maplestone, in 
which the ovicell is partly imbedded in the superior zocecium and 
is surrounded by a beaded structure; also Catenicella delicatula 
Wilson and Claviporella pulchra MacG. have the ovicell in the same 
position but perforated all over. Perhaps a new genus is required 
for these two, and C. wmbonata$ B. may have to be included. 

IL. Species with the tuberculate imperforate ovicell at the end 
of a mother zoccium of a biglobulus, namely, C. perforata B., 
C. taurina B., C. cornuta B., but very few ovicells have been seen 
in this group, and perhaps it is a matter of secondary importance 
whether they are on a biglobulus or a triglobulus. 

IIL. Species with the ovicell belonging to the mother zocecium 
of a triglobulus, including Pterocella Lev., which has a double area 
to the ovicell and the ovicelligerous aperture different from those 
of the ordinary zocecia ; Strongylopora || Maplestone, with a per- 
forated ovicell and the operculum straight at the proximal edge 


* “Wurther Desc. of the Tert. Polyzoa of Victoria,” Proc. Roy. Soc. Vict. vol. xii. 
n.s. p. 9 (1899). Levinsen, in making the genus Costicella, evidently overlooked 
the fact that Maplestone had already made a genus Costaticella, of which the type 
was lineata, a species included by Levinsen in his Costicella. 

+ As indicated in the * Zoological Record,’ the name Cribricella has already been 
used by Canu for a fossil belonging to the family Adeonide. 

{ The surface of none of the Vittaticella seem to be perforate, but smooth or 
papillose, but some have wrongly been described as perforate. 

§ Described as fusca by MacGillivray. 

|| This Levinsen calls Hincksiella, but Maplestone has priority, as his gens was 
described in 1899 (Proc. Roy. Soc. Vict. vol. xii. p. 4). 


Proc. Zoou, Soc.—19138, No. XX XITT, 33 


AR4 MR. A. W. WATERS ON 


of both the ordinary zoecia and the ovicelligerous zoecia, although 
the notch in the caleareous wall has been taken for an oral sinus ; 
Claviporella Ley., with a perforated ovicell and triangular aperture 
to both forms of zocecia. : 

The characters relied upon by Levinsen are none of them now 
mentioned, as my object is to show the importance of the ovicell 
and of the operculum in elassification, though of course all 
available characters must be used. 

Levinsen (p. 254, pl. xiii.) mentions a closure: in Vittaticella 
and other genera which he calls an “occlusion.” J have not seen 
anything quite like Levinsen’s description, in which it is said to 
start from three processes, which, of course. is the youngest stage, 
but in a specimen of Catenicella cornuta 8. from Western Port, 
Victoria, there is in the older zocwcia a calcareous closure which 
is, however, under the operculum and quite Independent of it. 
There are two round openings near the distal end, that is, one 
at each side, and a rather larger one at the proximal end. 
Tn a few very old zowcia these two distal pores eoalesce and the 
proximal opening becomes much larger, as if it were being dis- 
solved away. I have only been able to find this closure in 
C. elegans, in var. zanzibariensis, nov., and in C. cornuta as men- 
tioned, although I have looked through the Catenicellidee in my 
collection and the British Museum, though Levinsen gives it as 
a character of Vittaticella (Catenaria Lev.). However, it only 
occurs in the older and empty zocecia, and now, knowing this, 
T might on re-examination find some which had been overlooked. 
On p. 505 it is suggested that certain closures of Melicerititide 
should be compared. 


VITTATICELLA ELEGANS Busk. (PI. LXV. figs. 1-7, 12.) 


Catenicella elegans Busk, Brit. Mus. Cat. Mar. Polyzoa, p. 10, 
pl. ix. (1852); Zool. Chall. Exp., Polyzoa, vol. x. pt. xxx. p. 12 
(pars); Ortmann, “ Die Japanische Bry.,” Arch. fiir Naturgesch. 
vol. i. p. 27 (1890); MacGillivray, Prod. Nat. Hist. Vict. 
dec. ili. p. 23, pl. xxiv. fig. 10 (1895); Thornely, Ceylon Pearl- 
Oyster Fisheries, Suppl. Rep. xxvi. p. 109 (1905). 

Vittaticella elegans Maplestone, “ On a new name— Vittaticella— 
for the Polyzoan genus Caloporella McG.,” Proc. Roy. Soc. Vict. 
vol. xiii. n. s. p. 203 (1900). 

Specimens from Zanzibar growing on seaweed seem to be this 
species. There is sometimes on one side of the zocecitum a very 
long, raised avicularium with the mandible directed distally 
(figs. 1, 2). A similar avicularium occurs in a specimen in the 
British Museum from Avafura Sea. On the dorsal surface there 
are the two small dots as figured by me in Vittaticella conter Aud. 
(also in Brettia), and the radicles are in the same position as in 
V. contei. In one case a new zocecium springs from the anterior 
surface of an older zocecium, and this I have seen in other species 
of Vittaticella. 

The operculum is larger than that of V. contec, and is more 


BRYOZOA FROM ZANZIBAR, 485. 


curved on the proximal edge; on the other hand, it is not so 
large as that of V. buskei W. Th. These are all nearly related, 
but the difference in the operculum and the number of tentacles, 
besides other characters, indicate that they must be separated. 

There are often many ovicelligerous zocecia in one node. In 
one specimen there are two cases of a node having six ovicelli- 
gerous zocecia in one continuous line, and in both cases a lateral 
zocecium grows from the side of the unjointed node. In the 
ovicelligerous zocecia the avicularia near the oral aperture are 
directed forwards instead of laterally, as in the other zowcia. 
In the Tertiary beds of Curdies Creek, Australia, there is a form 
with long biserial nodes which I described as Catenicella inter- 
nodia*, but for which MacGillivray has since made the genus 
Ditaxipora ; also from the North Italian tertiaries I have described 
two forms with long biserial nodes as C. septentrionalist and 
C. continua, of which continua is. probably Vittaticella, but 
septentrionalis will require a new genus, so far as we can see at 
present. 

Among recent forms no long nodes have previously been 
described, but in the specimen from Madeira, which Norman 
considered to be C. contei Aud., there are two zocecia with ovicells 
following one another in an internode. Long ovicelligerous nodes 
are also found in Catenaria diaphana B. (see p. 482). 

The ovaria commence at the proximal end at one side, whereas 
the testes are near the distal end to one side. There are ciliated 
embryos in the ovicells, and below the ovicells there are several 
fleshy bands or tubes by which, no doubt, material for growth is 
transferred to the ovicell. ' 

Loc. Bass’s Strait, 47 fath., Banks’ Peninsula, Algoa Bay, 
Port Dalrymple; Tasmania (b.), Victoria (MJacG'.); Sagamibai, 
Japan (Ort.); Gulf of Manaar, on floating oyster-cages (Zh.); 
Arafura Bay (Brit. Mus.). Prison Island, Zanzibar shore (503) ; 
Ras Osowamembe, Zanzibar Channel, 10 fath. (504), collected by 


Crossland. 


VITTATICELLA ELEGANS, var. ZANZIBARIENSIS, nov. (PI. LXV. 
figs. 8-11.) 

Zoarium about 40 mm. high, with the branches curved over. 
Zocecia elongate ovoid, surface smooth or slightly papillose ; large 
lateral avicularia with a large pore at the base, in this respect 
differing from the type. There are on the dorsal surface near 
the distal end, and often resting on the large avicularia, two 
minute oval avicularia or an oval raised tubular opening. The 
radicles arising from the dorsal surface form a thick bundle. 


No ovicells are known. 
It is very similar to V. elegans B., but the larger beaked 


_* “Foss. Chil. Bry. from S.W. Victoria,” Quart. Journ. Geol. Soc. vol. xxxvii, 
p. 318, pl. xvi. figs. 78, 79 (1881). 
+ “ North Italian Bryozoa,” Quart. Journ. Geol. Soe. vol. xlvii. p. 5, pl. i. figs. 1-8 


(1891). 
Bon 


486 MR. A. W. WATERS ON 


avicularium with the large pore at the base and the minute 
dorsal avicularia suggest its being separated as a variety. 

The structure ofl She vittee has not received much attention, 
though Harmer* has alluded to it; but in this species it has 
been possible to obtain some explanation. The vitte are sunken 
perforated grooves in the calcareous wall. and along each groove 
there is a cylindrical tube, and within this, from the pore-tubes 
(the perforations just mentioned), organic cords spread out and 
reach the upper tree surface at definite spots or pores (fig. 10). 
It thus seems that the vittze should be compared with pore- 
chambers of many Cheilostomata in so far as there is indirect 
communication from the interior to the water-surface, through 
the vittee. 

Loc. Prison Island, Zanzibar Channel, 8 fath. (505); Wasin, 
Brit. Kast Africa, 10 fath. (500), collected by Crossland. Algoa 
Bay and Natal (Brit. Mus.). 


MEMBRANIPORA SAVARTII Audouin. (Pl. LXXI. figs. 1-4.) 


In my Report on the Bryozoa from the Red Sea (Journ. Linn. 
Soc., Zool. vol. xxxi. p. 138), I refer to the astonishing amount 
of anastomosing protoplasmic threads in a specimen from Zanzi- 
bar, and as some from the Sudan are also very full, this seems to 
be a specific character. It certainly seems strange to find such 
an extraordinary quantity, for though in my collection there are 
preparations of a large number of species showing the threads 
exceedingly well, I have never seen anything approaching these, 
and further study of the funicular cords is desirable. 

These threads are very abundant in zocecia with active poly- 
pides having digestion in full activity. In these threads are 
included small granular patches, either round or filiform, and 
where the polypides have degenerated or are degenerating there 
are large masses of this granular substance also surrounded by 
and connected with the protoplasma (fig. 2). In earlier stages 
the protoplasmic threads are in some cases surrounding the 
granular cord (fig. 4); in others there are only one or two plasmic 
threads by the sides of the granular cord or mass. 

The collecting together of these masses naturally suggests that 
waste products. are thus brought bepeuney and afterwards got 
rid of. 

Loc. Zanzibar Channel from the under side of buoy (528); 
Ras Osowamembe, Zanzibar Channel, 10 fath. (504); Prison 
Island, Zanzibar Channel, 8 fath. (505), collected by Crossland. 


?MEMBRANIPORA ARMATA Haswell (2072 Koschinsky). (PI. LX VII. 
fig, 10, & Pl. LX.XI. figs. 5-10.) 


Biflustra armata Haswell, ‘On some Polyzoa from the Queens- 
land Coast,” Proc. Linn. Soc. N.S. Wales, vol. v. p. 38, pl. i. 
fig. 7 (1880). 


* “ Morph. Cheil.,” Quart. Journ. Micr. Sc. vol. xlvi. p. 306 (1902). 


BRYOZOA FROM ZANZIBAR. A487 


Membranipora panhoplites Ortmann, “ Die Japan. Bry.,” Arch, 
Naturgesch. vol. i. p. 28, pl. 11. fig. 4 (1890). 
Membranipora armata Waters, “On Membraniporide,” Journ. 


Linn. Soe., Zool. vol. xxvi. p. 687, pl. xlvii. fig. 3 (1898). 


The specimen from Wasin is in parts in the hemescharan stage, 
in others in the bilaminate, and one in my collection from Port 
Molle is also bilaminate. This specimen from Wasin starts from 
a unilaminate incrusting layer, but in places the zoarium is 
tubular. 

The distal wall of the zocecium is slightly prominent, somewhat 
reminding us of what Busk calls the penthouse projection in 
Aspidostoma giganteum B. There are no ovicells in any of the 
specimens examined, and on most of the zocecia an avicularium 
occurs on one side only, at the distal end, directed proximally, 
while on the other side, in a long chamber, there is a long gland 
with distinct secreting cells (figs. 8, 9), but usually without any 
lumen. These glands are irregular in shape, sometimes lobed, 
and there may be two elongate lobes side by side. In all the 
specimens seen there is a distinct calcareous bar or arch to the 
avicularium, and there are two openings on the front of the avicu- 
larium, though sometimes the lateral projections in the avicularium 
do not meet, when there is, consequently, only one opening 
(fig. 7). In the membrane covering the avicularian chamber 
there is, under the mandible, a chitinous ring where the peculiar 
body * ends (fig. 9, pb.), and there are other species of Cheilo- 
stomata with a chitinous ring or other thickening. As we have 
seen, there is at one side an extremely long avicularian chamber, 
at the proximal end of which there are stout muscles (fig, 9 a) 
attached to a very long tendon by which the mandible is closed ; 
further up there are muscles also on both sides of the chamber, 
but much more delicate than the last (fig, 9 6), aud attached to a 
shorter tendon fastened to the base of the mandible with a median 
attachment, whereas in some species of Cheilostomata this muscle 
is attached at each side. 

The chamber containing the glands (fig. 9, gc.) is also very long, 
and may be close to the avicularian chamber of the zocecinm next 
above or below, but no connection with the avicularia has been 
found after careful examination of many microtome sections. 
On the inside there ave pores like rosette-plates, and the proto- 
plasmic threads from these are sometimes seen passing to the 
gland, but no other internal opening has been found, Externally 
there are three or four minute pores along the line of junction of 
the zoecia, which, however, were only noticed when carefully 
searching for openings, There are similar small pores over the 
avicularian chamber. We have here another Bryozoan puzzle, 
for the gland is not the same as the oral gland, and is contained 
in a caleareous chamber without any openings except small pores. 


* This peculiar body is in many Cheilostomata contained in a sheath, homologous 
with the tentacular sheath, but in this species no sheath was found. 


488 MR. A. W. WATERS ON 


The operculum is membranous, with a diagonal band or sclerite 
on each side to which the muscle for closing the operculum is 
attached ; but also attached to the same sclerite there is a muscular 
band fastened to the tentacular sheath. I have not noticed a 
double attachment like this before, but examination may show 
that it occurs in other species. 

In a large proportion of zocecia there are two polypides of 
about the same size, so that 20-30 zocecia with two polypides 
may be seen adjoining one another, whereas other pieces may 
show a much more limited number of double polypides. Although 
budding polypides in the same zoccia as mature polypides are 
known to us all, and have been described by Haddon*, Ostromoff 7, 
and Harmer {, they are only in a limited number of zocecia ; nor 
have I been able to see that they are the same as the two zoecia 
described by Prouho § in Aleyonidium duplex P. The two tenta- 
cular sheaths are side by side, and are attached to the operculum 
and the neighbouring wall. No ovaria or ova have been seen, 
and only in a very few cases were testes found occurring in round 
masses near the lateral wall. 

From the lateral walls there are bundles of muscles (6-12) 
attached to the frontal membrane of the zoecia. 

In all the lateral walls there are pores at fairly regular 
intervals all over the walls, and besides there are in some cases 
disks with numerous pores near the opercular wall. Further, in 
a bilaminate piece of 17. armata, there are in several cases large 
perforated disks on the basal walls, like those described in 
Petralia for the radicle attachment, though, strangely, in the 
unilaminate parts no distinct perforated disks have been found. 
This form cannot remain under J/embranipora, though I am not 
suggesting that it is Petralia, but call attention to various similar 
oer dehors in forms placed far asunder. 

There are about 30 tentacles, which is a larger number than 
has been found in any true Membranipora. Membranipora nigrans 
Hincks and WM. marginella H., with avicularia similarly placed 
near the distal end, have also cur ious large vicarious avicularia. 

Loc. Port Danion, Holborn Island, oy 0 fath. (Haswell); Port 
Molle, Australia ; Sagamibai, Japan, 40 fath. (Ortmann). Wasin, 
Brit. East Africa, 10 fath. (500); Zanzibar Channel, from under- 
side of buoy (528), collected by Crossland. 


MEMBRANIPORA CATENULARIA Jameson. 
For synonyms see Miss Jelly’s Catalogue. 
Although the Arctic W. monostachys Busk from Franz Josef 


Land has many points of similarity, the operculum in J. catenw- 
laria from Zanzibar is only about ‘half the width of that of the 


* Quart. Journ. Micr. Sc. vol. xxiii. p. 520 (1883). 

t Arch. Slaves de Biol. vol. 11. p. 341 (1886). 

apace Excretor, y Processes in Marine Polyzoa,” Quart. Journ. Micr. Sc. p. 189 (1891). 
( § “Cont. a Vhist. des Bry.,” Arch. de Zool. Expér. et Gen. 2nd ser. vol. x. p. 581 
1892). : 


BRYOZOA FROM ZANZIBAR. 489 


former species, and does not reach to the border of the opesium, 
so that they can be distinguished by this character. * 

Loc. Widely distributed, but there is much uncertainty about 
some of the determinations. Chuaka, Zanzibar, 3 fath. (506), 
2 fath. (508), on the dorsal surface of iseareponclia aan is b., 
collected by Crossland. 


Farcimia ocunata Busk. (Pl. LX VII. figs. 8, 9.) 


For synonyms see :— 


Farcimia oculata Waters, “Rep. on the Mar. Biol. of the 
Sudanese Red Sea, Bryozoa,” Journ. Linn. Soc., Zool. vol. xxxi. 
p. 167 (1909) ; and add Canu, “ Bry. des Terrains Tert. des Envy. 
de Paris,” Ann. de Paléont. vol. ii. p. 20, pl. 11. figs. 36, 37 (1907); 
pebsiye Helv. del Egypte,” Mem. 4, I Inst. Eeyptien, vol. Wie Dole 
pl. x. figs. 16-19 (1912). 

Nellia tenella Le svinsen, Morph. & Syst. Studies of the Cheil. 
Bry. p. 120, pl. i. figs. 13 a—13 e (1909). 

The growth is from a spreading stolon from which, at intervals, 
sub-colonies grow (figs. 8, 9), commencing with short calcareous 
nodes joined ‘by chitinous tubes. There are usually three or four 
nodes in the stalk, though there may be only one. Quite simi- 
larly sub-colonies on a stalk with internodes grow from delicate 
stolons in Chlidonia cordiert Aud., Diplodidymia complicata 
Reuss (Pl. LX VII. figs. 11, 12), Catenaria parasitica Busk *. 
I figured it some years ago for Chlidonia cordieri Aud. 7, and 
since then Calvet and Levinsen have dealt with the species, All 
these species have a somewhat similar operculum, about the same 
size, and the number of tentacles in all is 11-12. They are 
probably more nearly related than we have imagined. 

_- There are two very thick chitinous tubes connecting each new 
branch. 

The ovarium, with several ovarian cells, often in a row, is near 
to the inner wall, about equidistant from the distal and proximal 
ends. One ovum grows extremely large, and is pressed into all 
kinds of shapes thr ough want of room. The ovum passes into a 
sac near the basal “nal and the distal end, and before an ovum 
has passed into it muscle-threads are seen radiating over this sac, 

Levinsen has figured the ovicell, which is a ‘small cap-like 
growth, and this I have seen in a Basin cases, but often in zocecia 
with ovicelligerous zocecia no external difference is recognised. 
There is, however, often a vertical division separating the ovi- 
cellular wall from the rest of the zocecium, 

The triangular mandible of the avicularium is found with 
difticulty and is extremely minute, being about 0°006 mm., while 
a large number of the mandibles, such, for example, as in Retepora 
cellulosa, are 35 times as long; some, as for example in Lepralia 


* The Honourable Mary Palk informs me that Catenaria lafontii Aud., grows on 
a similar stalk. 
+ Journ. Linn. Soe., Zool. vol. xxvi. pl. 1. fig. 8 (1898). 


490 MR. A. W. WATERS ON 


occlusa B., are 60 times as long. In many cases there seems to 
be no mandible, only a disk, to which the peculiar body is attached, 
and this peculiar body is relatively very large. Smitt refers to 
the mandible of the Floridan specimens often being wanting, 
“presenting the opening closed only by a membrane.” 

There is no possibility of knowing what the Cellaria tenella of 
Lamarck was, and it certainly may have been Cellaria*, so that, 
as I have previously said, it is better to retain the name oculata. 
The genus Vellia was not satisfactorily described, and therefore 
various authors have adopted the genus Farcimia of Pourtales 
and Smitt. Fleming had made a genus Farcimia which might 
include Wellia, but as he made it for Cellaria, with C. fistulosa 
as type, it was always a superfluous genus, and Smitt considered 
it non-existent. Although Levinsen adopts WVellia, it seems 
better to adhere to Farcimia, the name used in my recent papers, 
and which has been used by most workers recently. 

Loc. See my paper referred to, and add Wasin, Brit. East 
Africa, 10 fath. (501); Prison Island, Zanzibar Channel, shore 
(503), 10 fath. (505); Ras Osowamembe, Zanzibar Channel, 
10 fath. (504); Meweni Bay, Zanzibar, 6 fath. (510); Chuaka, 
Zanzibar, 3 fath. (526), collected by Crossland. Texas and 
St. Thomas, W. Indies (Levinsen). 


DIPLODIDYMIA COMPLICATA Reuss. (Pl. LXVITI. figs. 11-15, & 
text-fig. 79.) 


Diplodidymia complicata Reuss, ‘‘ Foss. Fauna der Oligoc. von 
Gaas,” Sitzungsber. d. k. Ak. der Wissensch. Wien, math.-nat. Cl. 
vol. lix. Abth. i. p. 469, pl. ii. figs. 6-9 (1869). 

Micropora ratoniensis Waters, Ann. Mag. Nat. Hist. ser. 5, 
vol, xx. p. 185, pl. iv. fig. 5 (1887). 

Micropora articulata Waters, Quart. Journ. Geol. Soe. vol. xlvii. 
p. 14, pl. u. figs. 5, 6 (1891). é 

From Chuaka, Zanzibar, there are several pieces throwing 
light on this form, which has only been seen in fragments 
previously. 

The zoarium has a stalk consisting of long barren internodes, 
sometimes as many as eight, followed by long articulated inter- 
nodes with the zoeecia placed diagonally on the four sides. The 
contents of the barren internodes send out a branch to the 
surface, just as we have seen in Chlidonia cordierti Aud. From 
near the starting-point of each sub-colony a number of narrow 
radicles radiate, and sometimes from one of these radicles a 
fresh sub-colony grows, as is frequently the case in species with 
creeping stolons. 

The branches are dichotomous and articulated, having two 
chitinous tubes forming each articulation. Occasionally there 
are more than the four rows, but this will only be for a short 
distance near the articulation, and in one piece there is a median 


* Busk gives it with ?synonym of Cellaria gracilis Busk. 


BRYOZOA FROM ZANZIBAR. 491 


line on the dorsal surface with the zoccia on each side. The 
front wall has a minute pore on each side below the oral aperture 
(fig. 13), but these are only seen in Eau de Javelle preparations, 
and in the same way the Jong slit-like pore on the one side is 
scarcely seen so long as the membrane covers the walls. The 
muscles passing through this slit are wide, and are attached to a 


Text-fig. 79. 


Diplodidymia complicata Reuss. 


A. Showing small sa¢és hanging from the opercular region, and in each an ovum 
grows (ov.). At y. there seems as yet no ovuin in the small sac; m., the 
muscles of the opesium. X 50. 

B. Section showing an ovum in the small sac. X 250. 


C. Section in which the ovum has segmented and a blasluta is formed. X 250. 


chitinous thickening or sclerite on the front membrane, and are 
partly protected by a calcareous wall. There are two rosette- 
plates in the entire lateral wall. On the avicularian chamber 
there is a concave pit with a central perforation, no doubt indi- 
cating that radicles can be given off from this pit, and thereby 


492, MR. A. W. WATERS ON 


the two pores described in the fossil Wieropora articulata Waters 
are explained, the one being the avicularian and the other the 
radicular opening. ‘The avicularium is by the side of the aperture, 
and the mandible is triangular. 

There are 11—12 tentacles. 

The testes extend all down the lateral wall. A very few small 
ovaria with one to three small ovarian cells were found. 

A small ovum is found in a small sac hanging from the 
opercular region (text-fig. 79). The sac and ovum grow until 
ultimately the embryo nearly fills the zocecium, and now there 
are small muscles from the opercular wall to the ovicell and 
a strong lateral band. There is no external indication of any 
ovicell nor of which cells contain an embryo. The embryo is 
relatively large, with the couronne very large and distinct, and 
the way in which the embryo grows in the pendant sac may 
throw light upon the development of the ovum and embryo in 
Adeonella, but no stage has been found with a sac hanging from 
the opercular region in Adeonella. 

The present species, as [ have previously indicated, is closely 
allied to the fossil Cellularia diplodidymoides Meun. & Pergens 
from the Chalk, and both belong to the same genus *, Canu has 
described four fossil species from the Paris Basin, but, unfor- 
tunately, he has not given figures as well as photographs. In 
the work there are a number of magnificent photographs, 
showing the characters most beautifully, but some of the Paris 
authorities are making too hard and fast a line that everything 
must be photographed. The specimens in question (pl. v. figs. 
6-10) do not lend themselves to photography, and require either 
figures or full description for elucidation. 

These Zanzibar specimens are so different in the younger and 
older joints that with fossils several species might be made from 
one colony. 

Canu 7 says that the genus Diplodidymia is the Poricellaria of 
dOrbigny, but the description of this latter leaves recognition 
impossible without direct comparison, and therefore Canu is quite 
right in retaining the name Diplodidymia Reuss. 

Loc. Off Katow, New Guinea, 7 fath.; Singapore (fide White- 
legge, in lit.). Chuaka, Zanzibar, 3 fath. (506), collected by 
Crossland. 

Fossil. Gass, near Dax, 8. France, Oligocene; Montecchio 
Maggiore, N. Italy, Bartonian. 


CHLIDONIA CORDIERI Audouin. (Pl. LXV. figs. 15, 16.) 


EHucratea cordiert Aud. Descrip. de ’Egypte, Hist. nat. p. 242, 
2nd ed. p. 74; Savigny’s pl. xiii. fig. 3. 
Chlidonia cordiert Waters, Journ. Linn. Soc., Zool. vol. xxvi. 


* “ Bryozoaires du Systtme Montien,” Louvain, p. 8, pl. ii. fig. 3 (1886). 
+ “ Bryozoaires des Terrains Tertiaires des Environs de Paris,” Ana. de Paléont. 
t. liv. p. 39 (1907). 


BRYOZOA FROM ZANZIBAR, 493 


p. 18, pl. i. figs. 8, 9 (1896), which see for synonyms, and add: 
Calvet, ‘Bry. Mar. de Cette,” Trav. de I’Inst. de Zool. de Univ. 
de Montpellier, ser. 2, mem. 11, p. 13, pl. 1. figs. 1, 2 (1902) ; 
‘“‘Bry. Mar. des Cotes de Corse,” op. cit. mem. 12, p. 6 (1902) ; 
Levinsen, Morph. & Syst. Studies on the Cheil. Bry. p. 197, 
pl. viii. figs. 6 a-6 y (1909). 

From the front wall to the zocecial wall, through what has been 
called a second chamber, but is the equivalent of a compensation 
sac, there is a bundle of three, four, or more muscles (see fig. 15). 
The attachment of these on the front has been mistaken for a 
suboral pore, and in dried specimens there is frequently an 
opening here. 

No ova or ovaria have been found in any of my sections, 
whereas some show an embryo about half filling the zocecium, 
though no external difference has been noticed. 

The operculum is interesting, as it has at each side a projection 
or wing at right angles to the operculum (figs. 15, 16); also at 
each distal corner there is a slight projection. The wing reminds 
us somewhat of the thin membranous growth of many Mem- 
branipore and some Micropore. 

There are 9-11 tentacles. 

Loc. Red Sea (Aud.); Naples, Trieste, Rapallo, Nice, Cette, 
Corsica, Algiers, Tunis, Tyre, Calvados, Egypt, Victoria (Aus- 
tralia), Cape York, New Zealand, Atlantic (fide Carus); Canaries 
(dOrb.). Wasin, Brit. Hast Africa, 10 fath. (500), on calcareous 
seaweed, collected by Crossland. 


CELLARIA. 


Levinsen, following Norman, uses Cellularia for what we 
understand as Cellaria, but as these names have long been used 
for widely distinct genera I must certainly, in the most definite 
manner, refuse to use the name Cellularia for what we have for 
many years understood as Cellaria. 

Cellularia of Pallas was a simply ridiculous jumble of forms for 
which a place had not been found elsewhere. The species men- 
tioned by him are now placed in nine genera, one of which is the 
Cyclostome Crisia, and as the description of the genus refers to 
the ovicell of Crisia, perhaps the least objectionable thing would 
have been to have retained the name Cellularia for Crisia. The 
real difficulty, however, is that the name Cellulavia has been 
retained for a quite different group, and to interchange and now 
use a name long understood in a different sense would cause the 
greatest confusion. 

Solander employed the name Cellaria for a group approxi- 
mately, but not absolutely, similar to the Cellularia of Pallas, 
giving a definition also not quite the same, and it has been con- 
sidered to be only a change of spelling, though curiously, Ellis & 
Solander never indicate that they considered it was the same as 
Cellularia of Pallas, nor throughout the description of the genus 


494 MR. A. W. WATERS ON 


do they ever refer to Pallas, although many of the species dealt 
with had been mentioned by that author. This is certainly 
difficult to understand, and perhaps would have been different 
if Solander had lived to complete the work himself. 

Hincks considered that the species in Solander’s genus had all 
found places elsewhere, and that the genus of Solander had lapsed. 
He therefore took Cellavia as the genus of Lamouroux, who under 
Cellaria put Cellaria (as now understood) with the type C. sali- 
cornia and also Tubucellaria. Lamouroux says of all genera none 
seems to contain as widely distinct species as this, and that 
it seems to have been formed to contain everything that could 
not be placed under Flustra or Sertularia. 

Instead of dropping Celluluria as hopeless, Busk unfortunately 
retained it for a small division, and this has been accepted. 

The first species mentioned by Pallas in his genus is tubu- 
cellaria, which also is included in Lamouroux’s Cellaria, though 
not as the type, which was Cellaria salicornia; and Stoliczka*, in 
a long discussion of the subject, took the view that as tabucellaria 
was first mentioned the genus must be called Cellaria, and 
Cellaria, as now understood, must be separated as Salicornia. 

We now see that it is unfortunate that Hincks should have 
retained Cellaria, though it then appeared that this would not be 
challenged, and it has been adopted generally, and no genus 
seemed more firmly established. However, it is now clear that if 
Hincks had continued to use Salicornaria there would have been 
no possibility of the name of a now long recognised genus being 
replaced by one used in most various ways, and now limited to 
another small group. 

Those who are at work upon the class know how often the 
descriptions of the earlier authors are now meaningless, for the 
characters then used are found to be useless; but this can hardly 
be appreciated by those who have not had occasion to consult 
such descriptions. It is as if some well known tree had genera- 
tions ago received a name and a few lines of description which 
would apply to a quarter of our phanerogams and some cryptogams. 

I have previously shown that we are brought into a perfectly 
ridiculous position by being asked to adopt such names at all 
costs, when we often have no idea what they meant. It is not 
science, and since Cellaria as modified by Hincks is well established 
I shall still use it f. 


* Foss. Bry. aus dem tert. Griinsand. der Orakei Bay bei Auckland, pp. 142-149 
(1864). 

+ Norman (“Polyzoa of Madeira,” Journ. Linn. Soc., Zool. vol. xxx. p. 293 
(1909)) challenges the correctness of considering that Tubucellaria opuntioides 
should have been considered the type of Cellularia, but I cannot agree with his 
conclusions as to what I say being contrary to the British Association Rules of 
Nomenclature. My edition is later than Norman’s, but apparently is only areprint, 
and it says “ When they omit doing so” (7. e. fixing a type), “it may still in many 
cases be correctly inferred that the first species mentioned on their list, if found 
accurately to agree with their definition, was regarded by them as the type.” This 
rule of course means that if the generic diagnosis is taken from some one species, 
and that an error has been made in including the first, then common sense may be 


BRYOZOA FROM ZANZIBAR. 495 


CELLARIA GRACILIS, var. TESSELLATA, nov. (PI. LXVII. fig. 7.) 

For synonyms of Cellaria gracilis see Miss Jelly’s Catalogue 
and add :— 

Meissner, ‘* Liste der von Herren Prof. Simon bei Amboina und 
Thursday Island ges. Bry.,” Jena Denkschr. vol. vill. p. 730. 


A specimen from Ras Osowamembe growing on Hydrozoa 
seems to be a variety of C. gracilis. In this specimen the trabe- 
cule mentioned by Busk enclose an area formed of large tessellated 
divisions, about 20 (fig. 7); and curiously, another species, which 1 
eall C. wasinensis, sp. n., has also a tessellated area, but the 
avicularium of that species is triangular with an acute mandible, 
and belongs to the C. tenwirostris group. These large tessellated 
arens are unusual, and have not been found in various species of 
Cellariain my collection ; nor in the British Museum ‘ Challenger’ 
collections are they found in C. bicornis B., C. dubia B., C. malvi- 
nensis B., C. variabilis B., C. divaricata B., C. australis Hincks, 
CU. rigida MacG. The significance both of the trabecule and of 
the divisions is at present obscure. 

The opercula are a trifle larger than the type from Holborn 
Island, but the mandibles are the same. 

Loc. Type: Cumberland Island ; Cape Capricorn ; Victoria, 
8 fath.; Torres Straits; Holborn Island (Queensland); Katow, 
New Guinea, 7 fath. Variety: Ras Osowamembe, Zanzibar 
Channel, 10 fath. (504), collected by Crossland. 


CELLARIA WASINENSIS, sp.n. (PI. LXVII. figs. 1-6.) 


Zoarium about 25mm. high, with small branches dichotomously 
jointed and connected by two or three stout straight chitinous 
tubes and a “knot.” The zoarium swells out where the ovicells 
occur, as is frequently the case in Cellaria, there being sometimes 
two such swellings in an internode ; and it will be seen on fig. 6 
how these are equidistant from the joint in the two branches. 

The zowcia are rather elongate, hexagonal, or rhombie, with the 
lateral walls of the zocecium much raised, surface finely granular ; 
oral orifice wide, with two teeth on the distal edge, which is 
slightly turned up, forming a lip. 

The ovicellular aperture is large, round, with a plate from the 
proximal border spreading out and partially closing the aperture. 


used in saying the first is not the type, but surely it never meant that any individual 
could subsequently pick and choose which was the type among those agreeing 
equally well. However, in my work, to which Norman refers, I and other specialists 
were under an obligation to follow the rules of the Zoological Congress, and the rule 
in question is “other things being equal the name is to be preferred which stands 
first in the book or article.” In anything I write now the same obligation does not 
exist, though the rule entirely commends itself, and it does not seem to clash with 
the rules previously mentioned, drawn up by Strickland. Both practically say that 
when there are adequate reasons to indicate which was the type intended, that may 
be considered the type although not first mentioned, but without good cause to the 
Say the first is the type, and independently this is what workers are constantly 
oing. 


496 MR, A. W. WATERS ON 


The oyvicellular opening of C. gracilis Busk, var., C. australis 
MacG., C. rigida MacG., and C. hirsuta MacG., is similar; but in 
the few cases where the ovicellular aperture 1s mentioned, it is 
often only the incompleted opening in an early stage which is 
described. ; 

The avicularia are triangular with mandibles like those of 
C. tenuirostris B. In the avicularium the calcareous sub- 
mandibular wall rises up to the proximal edge of the mandible 
with a notch on each side, perhaps for the muscles (fig. 5), so that 
the submandibular part of the avicularium is almost entirely 
closed; and this 1s an interesting point, for Levinsen considers 
that the Melicerititide differ from the living Cheilostomata in 
having the submandibular portion entirely calcified. 

As I have mentioned *, there is, however, a specimen in the 
Museum dhistoire naturelle in Paris from the Banecs des 
Aiguilles, 8. Africa, which is probably the J/acropora cribrilifera 
Maplestone t, fossil from Mitchell River, in which the large 
vicarious avicularia have the submandibular part entirely 
calcified. Maplestone mentions that three of the zocecia have a 
‘“‘caleareous closure.” I have previously stated that there seem 
to me to be some points of relationship between Cellaria and 
Melicerititide. 

Levinsen deals but very shortly with the avicularia of Cellaria, 
and I am not quite sure that I follow what he means about the 
avicularium of C. malvinensis. He, however, says that the sub- 
mandibular er yptocyst, reaches up to the operculum in C. fistulosa. 
I have not seen it rise as it does in C. wasinensis in any of my 
specimens of C. fistulosa L., but there is a similar wall rising to 
the base of the mandible in C. variabilis B., C. hirsuta MacG.,’ 
and C. gracilis B.; C. variabilis has two slits in the submandibular 
part. More frequently there is an open rounded submandibular 
space as in C. dennantt MacG.t, C. malvinensis B.t, C. australis 
H., C. fistulosa, C. tenuirostris Busk, and C. wandeli Calvet. In 
C. fistulosa this is not much more than a wide round sinus. 

There are two species which have been taken for C. malvinensis 
B.. The first, which I have from Wanganui, New Zealand, has a 
fairly large submandibular space with a distinct ridge where the 
proximal end of the mandible comes, in fact in a few cases this is 
continued, forming a bridge across. The mandible soon contracts, 
with the distal end lanceolate. The other form is slightly 
smaller, from Baie Orange, S. Africa, and mentioned by Jullien as 
C. malvinensis. It has the sides of the zoecia straight, the distal 
end rounded, and the submandibular part of the avicularium rises 
up to the mandible, having two diagonal slits. The mandible is 
shorter than in the other species, sloping gradually to the apex. 
Whenever this is figured it might be called C. jullient. The 


* Résultats du Voyage du S.Y. Belgica, “ Bry oz0a," ’ p. 85 (1904). 

+ ° Further Desc. of the Tert. Poly. ot Victoria,” Proc. Roy. Soc. Vict. vol. xiii. 
n. s., p. 204, pl. xxiv. fig. 2 (1901). 

4€ Rés. du Voy. du 8.Y. Belgica, pl. it. fig. 9a & pl. viii. figs. 4, 5. 


BRYOZOA FROM ZANZIBAR. 497 


forms from Curdies Creek which I considered to be malvinensis 
have since been separated by MacGillivray as C. contigua, but the 
fossils from Bairnsdale and New Zealand are C. malvinensis. 

The operculum of C. wasinensis is nearly semicircular with 
large hollows fitting on to the teeth, and is similar to the 
operculum of C. gracilis. The trabeculee (figs. 3, 4) enclose in the 
lower part large divisions, about 7 in number, and the divisional 
walls are very thick, whereas in C. gracilis var. tessellata, nov. 
(fig. 7), they are linear with numerous divisions. 

There are about 13 tentacles. In Cellaria we find * the number 
of tentacles is approximately the same throughout the genus, only 
C. dennanti MacG., a species showing other differences, has 20. 
Now Cellaria is a well marked genus hhaving opercula of a special 
form, with a hollow cup fitting on to the teeth in the oral 
aperture, and the ovicell has a characteristic chamber, and also a 
characteristic ovicellular aperture. 

Levinsen 7 states that a zocecium does not correspond with an 
area, but with this I cannot agree, as I find the superficial 
divisions approximately mark off the zocecia, even though they 
may in parts extend somewhat under the divisional line, and this 
is the case In many genera. 

The ovarium usually has two ova, though there may be one or 
three, and the ovum is large when it enters the ovicell. 

Loc. Wasin, Brit. Hast Africa (501) (507), 10 fath.; Ras 
Osowamembe, 10 fath. (504); Prison Island, Zanzibar Channel 
(505), 8 fath. 

Cellaria tenwirostris B., C. salicornioides Aud., C. magnifica B., 
O. malvinensis B., C. gracilis B., have been previously described 
from tropical regions. 


THAIROPORA MAMILLARIS Lamouroux. 


Flustra mamillaris Lamouroux, ‘“ Polyp. corall. flexibles,” p. 110 
(1816) and add to Miss Jelly’s synonyms :— 

Thairopora mamillaris MacG. Prod. Zool. Vict. dec. xx. p. 351, 
pl. 196, fig. 2 (1890). 

Membranipora mamillaris Hincks, Ann. Mag. Nat. Hist. ser. 6, 
wolevaltenp.. 9 (USO): 

Thalamoporella mamillaris Levinsen, Morph. & Syst. Studies 
on Cheil. Bry. p. 194, pl. via. figs. 5 a—5 e (1909). 

There is a small piece from Chuaka, Zanzibar, which was de- 
calcified when received. It was preserved in HgCl,, but no 
doubt there was acid as well. The main poimts could, however, 
be seen, and the opercula and mandibles were separated, and 
although the mandibles are smaller than in the specimen received 
from the Red Sea, to which reference is made below, yet they are 
quite characteristic of 7. mamillaris. Levinsen has shown that 
there are calcareous spicules in all the Thalamoporellide, affording 


* Résultats du Voyage du S.Y. Belgica, “ Bryozoa,” p. 37 (1904). 
7 Morph. & Syst. Studies on the Cheil. Bryozoa, p. 211. 


498 MR. A. W. WATERS ON 


good specific characters, and the way in which he has dealt with 
them is one of the most important features of his book. 

Loc. Victoria (MacG’.); South Australia (my coll.); a weed, 
brought up on the sounding-line in the Red Sea, covered with this 
species, was given to me by a passenger. We may take it that 
there is every probability of its being from the Red Sea, though 
we cannot record it as such without pointing out the possibility 
of its having remained on the line from some previous locality. 


Chuaka Bay, Zanzibar, 2 fath. (509), collected by Crossland. 


SrEGANOPORELLA MAGNILABRIS Busk. (Pl. LX XIT. figs. 12-20.) 


Membranipora magnilabris Busk, Brit. Mus. Cat. Mar. Polyzoa, 
p. 62, pl. Ixv. fig. 4, in the explanation of the plate called 
M. grandis. 

Steganoporella magnilabris Harmer, “‘ Rev. of Gen. Stegano- 
porella,” Quart. Journ. Mier. Sei. vol. xli. p. 279, pl. xii. fig. 10, 
pi. xiii. figs. 31, 44-46, which see for synonyms. Add Thornely, 
‘“‘ Mar. Poly. of the Indian Ocean,” Trans. Linn. Soc., Zool. vol. xv. 
p. 145. 

Specimens from Wasin agree in most respects with the descrip- 
tion of this species, though the mandibles of the A zocecia having 
no teeth are more like those of S. buskit Harm. However, a 
comparison with the British Museum specimens has led to my 
leaving them under magnilabris, though some workers might * 
separate them asa variety. Thezoaria are bilaminate, irregularly 
contorted, and evidently attain to a very considerable size, as 
some pieces, evidently only fragments of larger ones, measure 
two or three inches across. 

There are very few B zoecia, in fact two mounts, each con- 
taining about 300 zocecia, had no B zoecia, On examining all the 
remaining material a few scattered B zocecia were found, and it is 
interesting to find that from these zocecia, which are somewhat 
larger than most A zowcia, two new zocecia usually arise, that 
is the row here bifurcates, and to this but few exceptions were 
found in all the material examined. 

I called attention to this duplication from the B zocecia occurring 
in some fossils, but Harmer ft has shown that it is by no means 
universally the case. At the time I wrote very little material 
was available for comparative study. It, however, is a fact that 
in many species two zoccia usually, or at any rate frequently, 
grow from the distal end of B the zoccia. In this species, when 
the operculum is removed, it is often not possible to be sure which 
are B zocecia, as there are large A zocecia, with a large shelf, in 
all respects similar excepting the operculum, so that with fossils 
there will often be uncertainty. In most other species there is a 
greater difference between the A and B zoecia. The A opercula 
vary considerably in size. In a specimen from (508) the B zowcia 
are much more common than in those from (500). 


* “Bryozoa from Aldinga,” Quart. Journ. Geol. Soe. vol. xli. p. 292. 
+ Revision of the genus Steganoporella, p. 266. 


BRYOZOA FROM ZANZIBAR, 499 


According to Harmer the A zocecia are in most species of 
Steganopor -ella more abundant than the B, and he says there are 
no B in 8S. simplex H., and goes on to say that they are rare 
in S. suleata H., S. lateralis H., and S. magnilabris B. In 
S. truncata H. the ratio of A to B is 4-5 to 1; on the other hand, 
in S. alveolata H., there are few A zoecia, and S. connexa H. has 
only B zoecia. 

As Harmer™* has shown, there is considerable difference in the 
teeth of A opercula of magnilabris from various localities, and we 
have opercula of this Wasin specimen without any, also the B 
opercula may be with or without lateral teeth. The A opercula of 
a ‘Challenger’ specimen (208-90°4.16.13) has the teeth so minute 
that they would be overlooked with a low power, but are seen 
with a quarter-inch objective. 

The operculum of Steganoporella can scarcely be compared with 
the opercula of other Bryozoa, as it closes the whole of the distal 
chamber, and in a zoarium there is an amount of variation in size 
of the opercula which is unknown in other genera of Bryozoa. 
In a British Museum specimen of S. truncata Harm. from Port 
Dalrymple, there are a few cases where the operculum is thrown 
back, and then the whole of the opercular opening (that is over 
the distal chamber) is covered by a membrane with a large round 
opening, thus furnishing a most interesting form of closure. 

The wall, dividing the two chambers, passes vertically from the 
opercular wall to the basal wall, and in the middle there is a 
round opening, sometimes with a slight tubular projection through 
which the polypide passes. The polypide rests partly in the two 
chambers, that is the tentacles are not entirely withdrawn into 
the proximal chamber. The embryos develop at the base of the 
distal chamber, but this chamber can by no means be spoken of 
as an ovicell. 

The Wasin specimens show some blind zoccia either entirely 
closed or with a central opening, and the same occurs in a speci- 
men of S. tubulosa H., where two zocecia are about the normal 
size, one with a large round opening, the other with an oval one, 
and in these the whole of the frontal wall is granular. On the 
other hand, a British Museum specimen of S. solace H. has two 
zocecia entirely closed by the perforated eryptocyst. 

The S. magnilabris from Wasin has floating in the proximal 
chamber many small oval bodies, surrounded by a membrane 
containing diatoms and other detritus. These are the excre- 
ment pellets, and they are evidently frequently ejected within 
the zocecium. 

Study of the growing ends is instructive. At first there is 
an absolutely empty oblong zocecium covered with a plain mem- 
brane, and in this either A or B opercula may be formed, but at 
first the operculum has no basal sclerite (fig. 20), as this is 
formed subsequently. Next the proximal part of the eryptocyst 


* Toc. cié. p. 234. 


Proc. Zoou. Soe.—1913, No. XXXIV. j4 


500 MR. A. W. WATERS ON 

is formed (fig. 20, z 2-2 3), then the tube or opening between the 
two chambers (fig. 20, 24) which is at a much lower level than 
the operculum, next the lip is formed, which at last reaches up to 
the operculum. The muscles are formed after the operculum, 
and the polypide not until after the tube and lip. 

It has not been mentioned that the part between the main 
sclerite of the operculum is covered by a membrane, thus enclosing 
a space (fig. 17), and the tentacular sheath is fastened * between 
the two muscular attachments of the operculum, having a tissue 
across from one muscular attachment to the other (fig. 19, 
from r-r). As mentioned, the large distal muscle is attached to 
the operculum, but the large muscle just proximal to it in the 
A zocecia is fastened by a tendon to the frontal membrane (fig. 19 d) 
close up to the operculum, where there is a slight thickening or 
sclerite. Harmer f speaks of this muscle as perhaps inserted into 
the frontal membrane immediately adjacent to the basal sclerite. 

Further back (proximally) there is on each side a diagonal 
sclerite (fig. 14, sc.), to which the tendons of a smaller muscle are 
attached, which draws down the frontal membrane {. These are 
called depressor muscles by Harmer $}). 

In the B zocecia the retractor muscles are much larger than in 
A, and are in two groups instead of one, but in the present species 
I am unable to find any very material difference between A and 
B zocecia. 

The embryos develop at the base of the distal chamber, but 
this chamber cannot be spoken of as an ovicell, for the polypides 
are, even when retracted, partly in this chamber. It seems that 
the embryo may exceptionally be in the proximal part of the 
zocecium surrounded by a membrane, really an internal ovicell. 
This requires investigation. 

The first polypide buds in the growing terminations are in the 
proximal end of the young zoccia as usual, and are nearly always 
in one of the corners, consequently the polypide grows diagonally 
across the zoccium, causing a slight amount of asymmetry in the 
zoecium. The buds of the Bryozoa usually grow from near to a 
rosette-plate, the position of bud and polypide being thus affected 
by the position and number of these plates. In S. magnilabris, 
and probably generally in the genus, there are two large distal 
rosette-plates. In decalcified preparations a curious large fleshy 


* We must keep in mind that in the Cheilostomata the tentacular sheath is 
fastened to the operculum and to the zowcial wall. 

+ “ Morph. of the Cheilostomata,”’ Quart. Journ. Micr. Sc. n. s., vol. xlvi. p. 318 
(1902). 

¢ This was not readily made out at first, but some thick sections stained in 
Chlorazol blue enabled me to see it all clearly. This is a stain that will be found 
useful for staining muscles and some other tissue, but it is not a good nuclear stain. 
Material can be stained and decalcitied at the same time, as acid does not affect the 
colour and it may be used for intra vitam staining. This stain was given to me by 
my friend Mr. Waddington, who has been unable to find out particulars as to its 
constitution. 

§ Loe. cit. p. 320. 

|| It is interesting to find that the large “pores” in the cryptocyst of Cupularia 
oweni Busk are only for the passage of depressor muscles. 


BRYOZOA FROM ZANZIBAR. 501 


structure is found in connection with them (fig. 16). On the 
inner side (towards the older zocecium) there is a large saucer- 
like portion with a fleshy club-like projection on the other. ‘The 
wall of the zocecium and the rosette-plate passes between the two. 

From the tissue round one of these rosette-plates the polypides 
of the growing parts start, but many of the polypides have the 
retractor muscles always attached to the lateral wall. The ten- 
tacular part of the bud seems to be growing in one distal corner 
while the gut parts are growing in the other, and in the mature 
polypide a long tube connects the two (fig. 12). This narrow tube 
connecting the two parts is a character of the mature polypide, 
and is somewhat like that of the cesophageal tube of the 
Ctenostomata. 

There are very few secondary buds to be seen (as the zocecia 
nearly all contain perfect polypides) ; they, however, arise from 
tissue on the membrane crossing the operculum. 

There are about 25 tentacles, four of which near the base are 
larger and broader than the others. 

In other species of Steganoporella there is also considerable 
variation in the teeth of the opercula ; for example, in a specimen 
of S. buskii H. from Algoa Bay, South Africa, in the British 
Museum, there is a B operculum without any teeth whatever, 
whereas the other opercula have small teeth. 

Steganoporella is not yet known before late Tertiary, whereas 
Thalamoporella is much older. 

Loc. Brazil; Jamaica; St. Vincent; Florida; China Seas ; 
Singapore ; Philippine Islands; Trincomalee ; Amirante Isl., 20- 
25 fath. (Thornely). Wasin, Brit. East Africa, 10 fath. (500) 
(520); Chuaka, Zanzibar Channel, 2 fath. (508), collected by 
Crossland. 

Fossil. Australian Tertiaries. 


CRIBRILINA RADIATA Moll. 

Loc. Cosmopolitan. Ras Osowamembe, Zanzibar Channel, 
10 fath., on Adeonella platalea, collected by Crossland. 

HippotTHoa DIVARICATA Lamouroux. 

Loc. Wasin, Brit. Hast Africa, 10 fath. (520), on shell, collected 
by Crossland. 

Hipporuoa pistans MacGillivray. 

For synonyms see Waters, ‘“‘ Bryozoa,” Résultats du Voyage du 
S.Y. Belgica, p. 54 & pl. i. fig. 8 (1904). 

Loc. Wasin, Brit. East Africa, 10 fath. (520), on shell, collected 
by Crossland. 

SCHIZOPORELLA UNICORNIS Johnston. 

Waters, “ Bryozoa of the Sudanese Red Sea,” Journ. Linn. 
Soc., Zool. vol. xxxi. p. 143, pl. xu. figs. 12, 13 (1909). 


Some pieces are either one or two layered, and here again I 
Sie 
otk 


502 MR, A. W. WATERS ON 


have been able to see the zoccia superimposed, so that both lateral 
and distal walls of the upper layer are above the similar walls of 
the lower layer *. Reuss has shown the same thing in Cumul- 
pora angulata v. M., and I have in previous papers referred to it 
in Melicerititide. 

In some other species of Schizoporella the superimposed layers 
grow quite differently (see p. 504). 

Loc. Add: “ From bottom of s.s. ‘ Juba,’ which always remains 
in Zanzibar waters” (511), collected by Crossland. 


SCHIZOPORELLA PERTUSA Hsper. 


See Miss Jelly’s Catalogue, and add :— 

Lepralia pertusa Calvet, “‘ Bry. Mar. de Cette,” Trav. Inst. de 
Zool. de l'Université de Montpellier, ser. 2, vol. 1. p. 515; ‘* Bry. 
Mar. de Corse,” op. cit. vol. ii. p. 26; Jullien & Calvet, “ Bry. 
prov. des Camp. de l’Hirondelle,” pp. 69, 134. 

Schizoporella pertwsa Calvet, ‘‘ Exp. Sc. du Travailleur et du 
Talisman,” p. 416. 

Loe. British; Labrador; Atlantic; Florida; Mediterranean ; 
Mazatlan; Australia; Samoa; China Seas. Wasin, Brit. Hast 
Africa, 10 fath. (500) (520), collected by Crossland. 


SCHIZOPORELLA NIVEA Busk. (Pl. LXX. figs. 1-3, 7-9, & 
P]. LXXIII. fig. 16; and text-fig. 80.) 

Schizoporella nivea Busk, Zoo]. Chall. Exp. vol. x. pt. xxx. 
p- 163, pl. xvii. fig. 1 (1884); Philipps, in Willey’s Zool. Results, 
pt. iv. p. 440 (1889); Thornely, Ceylon Pearl-Oyster Fisheries, 
vol. iv. p. 114(1905); Rec. Indian Mus. vol. i. pt. 3, no. 13, p. 189 
(1907); Waters, Rep. Mar. Biol. of the Sudanese Red Sea, 
‘“‘ Bryozoa,” Journ. Linn. Soc., Zool. vol. xxxi. p. 168, pl. xvii. 
figs. 2-4 (1909); Thornely, ‘“‘ Mar. Poly. of the Indian Ocean,” 
Trans. Linn. Soc., Zool. vol. xv. p. 148 (1912). 


Zoarium in Hemescharan form. Zocecia quadrate, distinctly 
separated, fairly large pores over the surface, an avicularium 
near each upper corner with broad triangular to nearly semi- 
circular mandibles, sometimes a small avicularium at one of the 
lower corners, Below the aperture there is frequently, in the 
older zocecia from (501), a mucro which may be long spreading 
out at the ends, or there may be a thin lamina radiating in four 
directions the whole length of the mucro. ‘These mucros are 
more frequent on the ovicelligerous zocwcia. The oral aperture 
is nearly circular, has a wide poster, and there are fine lines 
across the operculum (fig. 2). 

The ovicell is large, raised, globular, with a ridge from the 
middle of the proximal edge, shortly bifurcating to each side, at 
any rate in the older zocecia. ‘The pores over the surface of the 


* Reuss, ‘“‘ Zur Fauna des deutschen Oberoligocans,” pt. i3.,Sitzb. d. k. Akad. der 
Wissensch., Wien, math.-nat. Cl., vol. 1. Abth. 1, p. 644 (81), pl. ix. fig. 1 (1864) ; 
“Die Foram., Anth. und Bryozoen des deutschen Septarrenthones,” Denk. s. d. 
k. Akad. der Wissensch., Wien, math.-nat. Cl., vol. xxv. p. 179 (63), pl. vin. fig. 12 
(1865). 


BRYOZOA FROM ZANZIBAR. 503 


ovicell are fairly large, though slightly smaller than those 
of the zocecia. 

The small avicularium at the distal corner (figs. 7-9) has short 
and wide glands which at the lower part are joined together, 
and no other avicularian glands yet seen quite resemble these 
(text-fig. 80). Avicularian glands occur in Lepralia foliacea Hl. 
& Sol., ZL. clivosa Waters, L. margaritifera, Smittia trispinosa 
Johnst., Porella plana Hincks, P. acutirostris Smitt, Retepora 
cellulosa L., and other Retepore. In all these cases there are also 
oral glands. The oral glands of S. nivea are but small with the 


end cells the larger and darker (Pl. LX_XIII. fig. 16, g/.). 


Text-fig. 80. 


Avicularian glands of the small avicularia of Schizoporella nivea. 


A, the two glands separated, X 320. B, the same, X 100. 
C, glands united, xX 320. D, gland, X 320. 


There are 16 tentacles, no pore-chambers, but on the lateral 
walls there are about eight rosette-plates near the basal wall, also 
on the distal wall there are about six situated near the base. 

The internal membrane of the zocecium projects in a sac-like 
manner into the ovicell (fig. 16). This sac is about the width of 
the zocecium, and is provided with a large number of muscles 
(fig. 16, sc.) radiating from near the operculum. From a whole 
preparation it seems that the ovum passes into this sac, which 
is then ruptured (fig. 16, 7) to allow of the passage of the large 
ovum into the ovicell. I think there is no doubt of my inter- 
pretation, although a ruptured membrane has only been seen in 
one ease. ‘The ovarium consists of a number of large ovarian 
cells. 

Some of the older zocecia are closed by a calcareous layer over 
the operculum, with a calcareous bar or lamina, extending from 
side to side (Pl. LXX. fig. 3) curving round to the sinus, a position 
which is fairly constant, not accidental. In older zoccia there is 
sometimes a calcareous bar straight across the aperture, and there 


504 MR, A. W. WATERS ON 


may be no operculum remaining underneath it. Frequently a 
tubular projection occurs in the middle of the supra-opercular 
calcareous layer, and there is a closure of the same kind in Schizo- 
porella unicornis and other Schizoporellide and various other 
genera besides some Tertiary fossils; and a similar structure has 
been described in ‘“ Lepralia’’* syringopora Rss. and is found in 
various Adeonide 7. It can be seen in SN. nivea that these raised 
ledges across the operculum are the basal lines for a superimposed 
layer even though the layer is seldom completed; and a South 
African species which, though a larger and distinct form, is very 
similar to S. nivea, explains the growth across the operculum 
more fully. In it we have the commencement of a superimposed 
new layer of zocecia, and in one specimen there are about fifty 
zocecia together on which the walls for another layer are all 
mapped out, and the wall passes over the operculum, nearly 
always with the same curved line as in S. nivea. 

I described and figured a similar growth:in Meliceritites $ and 
apparently it occurs in various other cases. On the other hand, I 
have specimens of Schizoporella unicornis Johnst. from Cape Verde 
Islands and Zanzibar, in which there are many layers, and each 
following layer is formed by the new walls, nearly always 
growing exactly above the walls of the older layer. Nevertheless, 
there is, in some cases, a curved calcareous ridge over the oper- 
culum for which there seems no object. In S. wiicornis there is 
often a tubule on the calcareous closure. 

Some stained sections of Adeonella contorta Mich. in which 
there are superimposed layers, show this tubule as an inverted 
funnel with a long tube (over the operculum) attached to the 
tissues below the operculum. These closures can only be compared, 
in a limited sense, with the tubules of the closures of the Cyclo- 
stomata, as the operculum is unaltered and there is no perforation. 
There are also membranous closures, and the subject deserves 
further study. The Schizoporellide and the Adeonide are 
apparently the two families most likely to throw light on the 
closures of the Cheilostomata. Of course we do not find the 
closures in the younger zocecia, only in the older ones. In both 
these families blind cells are very frequent §. 

The Zanzibar forms are not separated as varieties or species, 
although the one with the tall mucro might perhaps be called var. 
wasinensis (Pl. LXX. fig. 1)on this account. It may be the Schizo- 
porella linearis of Hincks, Ann. Mag. Nat. Hist. ser. 5, vol. vii. 
p- 159, pl. ix, fig. 2, to which there is no description or locality. 


* Waters, “ North Italian Bryozoa,’ Q. Journ. Geol. Soc. vol. xlvii. p. 20, pl. iii. 
figs. 3, 4 (1891). 

+ Waters, “ A Structure in Adeonella contorta Mich.,” Ann. Mag. Nat. Hist. 
ser. 8, vol. ix. p. 498 (1912). 

+ **On Cheilostomatous characters in fossil Bryozoa,’” Ann. Mag. Nat. Hist. 
ser. 6, vol. vill. p. 52, pl. vi. fig. 4 (1891). 

§ Sincethe above was written Levinsen has published an important work, “ Studies 
on the Cyclostomata Operculata,” D. Kgl. Danske Vidensk. Selsk. Skr. 7 R., Nat. og 
Math. Afd. vol. x. pt. 1, 1912, dealing with Melicerititidee. 

What he calls the opercula of these fossils I should speak of as the closures, for 


BRYOZOA FROM ZANZIBAR. 505 


Frequently a hydroid is growing over the surface. ‘This seems 
to be a species of Clava and may be the same species as one 
growing in Holoporella columnaris B., and H. pigmentaria Waters, 
in which it may pass through several layers of zocecia. Sections 
showed that the stolons were spread under the H. columnaris, 
indicating that the growth of the Holoporella occurred over 
established colonies of Clava round which the Holoporella formed 
calcareous tubular walls *. 

(1) S. nivea belongs to a group in which all the species 
have a wide sinus or arc, the surface is perforated as is 
also the ovicell, there are quite small oral glands or but very 
moderate sized ones. The opercula have the muscular attach- 
ment close to the border and fairly near to the distal end. This 
group I referred to} as including the types of Schizoporella, 
and belonging to it are S. sanguinea Norm., S. linearis Hass., 
S. harmsworthii Waters, S. awriculata Hass., S. galeata B.% ete., 
anda new species from 8. Africa. Levinsen includes several other 
species under Schizoporella, but I am by no means sure that all 
his species will remain in the same genus, for in (2) S. wnicornis 
Johnst. and var. errata Waters, S. longirostris Hincks, S. spon- 
gites, S. biaperta Mich., the sinus is much narrower, and what 1s 
of most importance, the muscular attachments are some distance 
from the border of the operculum. In 8S. biaperta the ovicell 
has a flat area with perforations round the border. 

(3) In the group S. vulgaris Moll., S. viridis Thorn. , there is a 
similar operculum, and it is in part the Escharina of Levinsen. 

In the same place I showed that there was a group which 
might be placed in a modified Buffonella Jullien, in which the 
surface of both the zoarium and the ovicell is imperforate, also 
there is a small suboral avicularium, and the opercula have the 
muscular attachments some distance from the border as in the 
last group. It is represented by S. ridleyi MacG., S. simplex 


I consider that the chitinous operculum was under the closure, and I should like to 
see the name operculum confined to the movable chitinous appendage. Levinsen 
on Plate i. figs. 15, 16, shows Meliceritites vielbanci d’Orb. with tubules to the 
closures, such as I have seen in recent Cheilostomata, and a number of similar 
closures are figured. Without there being perhaps any wide divergence of view I 
should not describe these as being regenerated zoccia, for while Levinsen has de- 
scribed regenerated zocecia, I have not seen anything to suggest their being common, 
and further, although we know that the polypides are regenerated i in the closured 
zoccia. this by no means always takes place, as, for example, in the older parts of 
stems and when there is more than one layer. It therefore seems unadvisable to 
speak of them as regenerated zocecia when they are frequently permanently closed. 

I have suggested to Professor Levinsen that the sunk walls with few openings as in 
Meliceritites magnifica (op. cit. Pl. i. fig. 7), may be comparable with the perforated 
wall or closure UNDER THE OPERCULUM in Vittaticella (see page 484). ‘These are 
also to be compared with the partial closure I described in MW. royana Waters, 
Ann. Mag. Nat. Hist. ser. 6, vol. viii. pl. vi. figs. 2,6 (1891). Levinsen in his most 
valuable work has given full figures and descriptions of the structure of Melicerititidee, 
and as I believe that many analogous structures are found in the Cheilostomata, a 
thorough examination of the closures of livmg forms is much to be desired. 

* Waters, Report on Sudanese Bryozoa, p. 254. 

t Waters, Résultats du Voyage du S8.Y. Belgica, “ Bryozon,” p. 42 (1904). 


506 MR. A. W. WATERS ON 


VOrb., S. rimosa Jull., S. marsupifera Busk, S. twmida H., 
S. levata H., S. levigata Waters. 

It is quite clear that there are several groups that can be 
separated off from the old Schizoporella, but until more living 
and spirit specimens have been examined important characters 
remain unconsidered. 

Loc. Since I gave the localities of S. mivea Miss Thornely has 
reported it from Amirante, 29 fath. ; Seychelles, 39 fath.; Provi- 
dence, 50-78 fath. It was procured from Wasin, Brit. Hast 
Africa, 10 fath. (501); Ras Osowamembe, Zanzibar Channel, 
10 fath. (504); Prison Island, Zanzibar Channel, 10 fath. (505), 
and with the large mucro from Wasin, 10 fath,, which may have 
to be called var. wasinensis : collected by Crossland. 


SCHIZOPORELLA MONTFERRANDI Audouin. 


Lepralia montferrandi Waters, Rep. Mar. Biol. of the Sudanese 
Red Sea, “‘ Bryozoa,” Journ. Linn. Soc., Zool. vol. xxxi. p. 171, 
pl. xvii. figs. 15-18.(1909). To which add :— 

Schizoporella pachnoides MacG. Trans. Roy. Soc. Vict. vol, xxiii. 
p. 180 (1886); Prod. Zool. Victoria, dec. xix. p. 314, pl. 186, 
fig. 6 (1889). 

This must go to Schizoporella as now limited (see p. 505). 
It is closely related to S. galeata, but as I have said they must 
probably be separated. 

Add:—Loc. Victoria (MucG@.). Wasin, Brit. East Africa, 10 
fath. (520), collected by Crossland. 


GEMELLIPORA PROTUSA Thornely. (Pl. LXX. figs. 4-6.) 


Gemellipora protusa Thornely, Ceylon Pearl-Oyster Fisheries, 
vol. iv. Polyzoa, p. 119, pl. iv. fig. 7 (1905). 

Zoarium incrusting. Zocecia rhomboidal, raised, with pores 
round the border; surface with few pores and granular. Oral 
aperture clithridiate, very long, with large lateral contractions, 
and the peristome frequently much raised, especially at the 
side. At one side of the zocecitum directed downwards a short 
wide triangular avicularium with a stout bar, and sometimes a 
second roundish avicularium near to the side of the aperture. 
Ovicell small, raised, globose, with very thick walls, perforated, 
much open in front and not closed by the operculum. 

Loc. Gulf of Manaar (7/.). Wasin, Brit. Hast Africa, 10 fath., 
received dry (520), collected by Crossland. 


TRYPOSTEGA VENUSTA Norman. 


Lepralia venusta Norman, Ann. Mag. Nat. Hist. ser. 3, 
vol. xiii. p. 84, pl. x. figs. 2, 3 (1864). 

Schizoporella venusta Hincks, Brit. Mar. Poly. p. 276, pl. xxx. 
figs. 6,7 (1880): Kirkpatrick, Ann. Mag. Nat. Hist. ser. 6, vol.i. 
p. 76 (1888); ‘‘ Hyd. & Polyzoa from the China Sea,” Ann. Mag. 


BRYOZOA FROM ZANZIBAR. 507 


Nat. Hist. ser. 6, vol. v. p. 17 (1890); Proc. Roy. Dublin Soe. n. s. 
vol. vi. p. 612 (1890) : Calvet, Expéd. Scient. du Travailleur et du 
Talisman, vol. viii. p. 416 (1907). 

Trypostega venusta Levinsen, “Studies on Bryozoa,” Vid. Medd. 
f. d. Naturh. Foren. i Kjgbenhayn, p. 23 (1902); Morph. & Syst. 
Studies on Chil. Bry. p. 281, pl. xix. figs. 1a-ld, pl. xxii. 
figs. 13 a-13d (1909); Norman, “ Polyzoa from Madeira,” Journ. 
Linn. Soc., Zool. vol. xxx. p. 299 (1909). 

Gemellipora glabra, form striatula Smitt, ‘ Floridan Bryozoa,’ 
pt. 1. p. 37, pl. xi. p. 207 (1873) ; Thornely, “‘ Mar. Poly. Indian 
Ocean,” Trans. Linn. Soc., Zool. vol. xv. p. 149 (1912). 

Gemellipora striatula MacG. Prod. Zool. Vict. dec. xiv. p, 150, 
pl. 138. fig. 10 (1887). 

Lepralia striatula MacG. Tr. Roy. Soc. Vict. p. 134, pl. iu. fig. 17 

1882). 

ene striatula Waters, Q. Journ. Geol. Soc. vol. xi. 
p. 301 (1885) ; Philipps, ‘‘ Poly. Loyalty Isles, &c.,” Willey’s Zool. 
Results, pt. iv. p. 440 (1899). 

Mollia tuberculata VOrb. Paléont. Franc. p. 388; see Waters, 
Ann. Mag. Nat. Hist. ser. 7, vol. xv. p. 6. 

Lepralia inornata Gabb & Horn belongs to this group. 

I should have hesitated, at present, to put this in a new genus 
merely on account of the small chamber above the zowcium, 
which no doubt must be considered as a vestigial avicularium, 
but as a genus may be required it will now cause less confusion 
to use Levinsen’s name. Jepralia turgescens Reuss, ‘“ Foss. Bry. 
Oest.-Ung. Miociins,” Denk. K. Akad. der Wissensch. Wien, 
math.-naturwiss. Class., vol. xxxiii. p. 36, pl. viii. fig. 7, has an 
avicularium, which, judging from the figure, corresponds to 
the small chamber above the zoccium in 7’. venusta, and also 
the avicularium of Chorizopora brongniartti Aud. is often 
sunilarly situated. 

Sections of the specimen from Wasin show that there are no 
bands of muscles in the chamber above the zocecium, but there 
are protoplasmic bands to the frontal disk, starting from two 
rosette-plates at the proximal end of the small chamber, and 
near the distal end there is a small group of cells at each side, 
which must probably be considered as glands. There are a few 
irregular large chambers almost as large as the ordinary zocecia 
closed in the same way as these small supra-zocecial chambers. 

There are but few ovarian cells in the ovarium, usually two, 
sometimes three or four, and one may be seen developed to a 
considerable size. The embryos develop fully in the ovicell. 
No suboral glands have been found. 

Loc. British; Guernsey; Calvados; Florida; Madeira (WVor- 
man); Azores; Cape Verde Islands (C.); Lifu, Loyalty Isl. (Ph.); 
China Seas (Tizard Bank), 27 fath.; Torres Straits (A’.); Amirante, 
22-85 fath., Saya de Malha, 29-125 fath. (Zhornely); Mauritius. 
Wasin, Brit. Hast Africa, 10 fath. (500), collected by Crossland. 

Fossil. River Murray Clifts, South Australia. 


508 MR. A. W. WATERS ON 


ARTHROPOMA CECILIZ Audouin. 

Flustra cecilia Aud, ‘ Zool. Egypte,’ p. 66 (239), pl. viii. fig. 3 ; 
see Miss Jelly’s catalogue. 

Arthropoma cecilii Levinsen, Morph & Syst. Studies on the 
Cheil. Bry. p. 332 (1909). 

Schizoporella cecilit Thornely, ‘* Mar. Poly of the Indian Ocean,” 
Trans. Linn. Soe., Zool. vol. xv. p. 147 (1910). 


In my Report of the Antarctic Bryozoa from the Voyage du 
S.Y. Belgica, p. 50, I called attention to this group and now 
Levinsen has given ita name. As I previously said Phonicosia 
Jullien may belong here, but Jullien’s specimens are not suf- 
ficiently complete to settle the question. 

Loc. Distribution general in the north temperate zone, the 
tropics, and Australia. Prison Island, Zanzibar Channel, on the 
shore (513), collected by Crossland. 

fossil. Kuropean and Australian Tertiaries. 


OSTHIMOSIA ZANZIBARIENSIS, sp. n. (PI. LX XIII. figs. 9, 15.) 


Zoarium incrusting seaweed, small, irregular. Zocecia sub- 
globose, smooth. Oral aperture with distinct sinus, otherwise 
nearly circular, a small semicircular avicularium at each side of 
the aperture, and when there is an ovicell usually a second pair 
of avicularia. Surface of the zocecia imperforate with longi- 
tudinal ribs in the older zocecia, vicarious avicularia among the 
zocecia having expanded spatulate mandibles. Ovicell large, wide, 
globose, perforated all over. 

In having the perforated ovicells this is like Cellepora avicularis 
Hincks, to which it is closely allied, but the median avicularium 
is absent. A perforated ovicell occurs in C. avicularis H., 
C’. coroncpus B., C. megasoma MacG., C. conica Busk, C. re- 
douti* Aud. In the early stages this form resembles Schizo- 
porella biaperta Mich., but vicarious spatulate avicularia are 
unknown in SS. biaperta. The operculum is thinner than that 
of C. conica, and the walls also are thinner. I find that C. conica 
is the young form of a species determined by Busk as (’. simo- 
nensis ~ B.,an erect cylindrical species. The oral aperture of both 
C. conica and C. avicularis is slightly larger than that of this 
species from Prison Island. 

After removing from Cellepora the genera Holoporella and 
Lagenipora there is the present group with a triangular sinus, 
the ovicell perforated and the surface of the zocecium imper- 
forate except near the border, and the group includes C. avicularis 
H., C. conica Busk, C. coronopus S. Woods, leaving still another 
group with imperforate ovicells, but a small semilunar area or 
mark near the proximal part, and a marked sinus in the oral 
aperture, as, for example, C. evewa Jull., C. eatonensis B. 


* This is only a variety of C. avicularis H. I have a specimen from a sounding- 
line in the Red Sea. 
+ Waters, Zoo]. Chall. Exp. vol. xxxi. pt. Ixxix. p. 85 (1889). 


BRYOZOA FROM ZANZIBAR, 509 


Hineksin his ‘ Brit. Mar. Polyzoa’ says that the typeof Cellepora 
was pumicosa, but this is not the case, for as Leyinsen points out in 
his large work, Linnzeus refers to C. ramulosa as the first species 
of Cellepora. Hincks incorrectly considered it the genus of 
Fabricius. Levinsen is, however, mistaken in supposing that 
C. ramulosa is schizostomous, as it belongs to the holostomatous 
group, though not to Holoporella*. I have not as yet had any 
opportunity of cutting sections of 0. ramulosa, but the operculum 
has the proximal border but slightly curved, and the small 
muscular attachments are close to the thicker part of the border. 
In most Holoporelle the attachment is a little nearer to the 
edge, on the other hand it is much closer to the distal edge than 
is usual in the Schizostomous groups. 

The ovicell of ramulosa is cap-like as in Holoporella ete., and 
although Hincks says perforate or imperforate, none of the speci- 
mens in my collection, nor any in the British Museum general 
collections, including Busk’s and Hincks’s,’ have a perforate 
ovicell ; also in the Norman collection a few from each locality 
were examined without finding any perforate. There are two or 
three species externally corresponding with C’.. ramulosa, so that 
a mistake is easily made, and in three cases friends have sent me 
specimens so marked, of which only a part were ramulosa. 

By taking (even if provisionally) C. ramulosa as definitely 
described by Hincks, and perhaps by some before him, as the 
type of Cellepora, we get out of a difficulty, for when Schismopora 
was created by MacGillivray Cellepora remained for the holo- 
stomous division. C.ramulosa is the first of Linnzeus’ species, 
the others being spongites, punvicosa, ciliata, hyalina. Linneeus’ 
description of femmes would do for several species of branching 
forms, and we are doubtful what the other species weve meant 
for. In Linneus’ copy of the 12th edition of Syst. Nature, 
there is in his small writing, under C. ramulosa, a reference 
SOLOS © lemme One which was hieroglyphic to me until 
Dr. Daydon Jackson kindly explained that it referred to Det 
Kongelige Norske Videnskabets Selskabs Skriften, 4th part, 
1768-1774, in which there is a paper by Gunnerus, who supplied 
Linneus with both C. ramulosa and C. pumicosa. In this work 
(pl. i. fig. 6) is a figure of a Cellepora, which, judging from the 
locality, “Oceano Norvegico,” is probably t C. incrassata Sm., 
although C. coronopus 8. Woods, an entirely different species 
from the Mediterranean, corresponds equally well with this 
figure, which shows nothing but zoarial shape. It is, however, 
what Linneus described as C. ramulosa. On the same plate 
the figure 7 shows a_ similar growth, though with smaller 
branches, and to this under C. pumicosa Linneus refers by a 


* There are various specics of holostomous Bryozoa which do not belong to 
Holoporella Waters, as, for instance, C. sardonica Waters, which will fall into 
Holoporellidee. 

+ Waters, “ Bry. from Franz-Josef Land,” Journ. Linn. Soc., Zool. vol. xxviii. 
p. 94 (1900). 


510 MR. A. W. WATERS ON 


similar note. A long time ago I showed* that the C. pumi- 
cosa as we have understood it had nothing whatever to do with 

pumicosa I., as it does not in any way correspond with 
Linneus’ description, besides which he refers to a figure by 
Marsigli which probably is a figure of C. coronopus. I then 
referred to it as C. pumicosa Busk (non L.) and have continued 
to do so. Fig. 7 may be a figure of C. ramulosa Hincks, ete. 
though called pumicosa by Linnzeus; there are, however, several 
other species that it might represent. After this record of mis- 
takes about two well known species, showing how little we can 
know what the earlier writers meant, we should be allowed to 
return to our senses, and use these long established specific 
names for thoroughly described and well recognised species, 
namely C. pumicosa Busk, C. ramulosa Hincks, but if we retain 
Cellepora for ramulosa the C. pumicosa Busk becomes Osthimosia 
Jullien. This group was divided off in the same year by Jullien 
as Osthimosia, and as Schismopora by MacGillivray, but it seems 
that Juljien’s name was published a few months the earlier. The 
opercula of this group are all of the same type, with the opercular 
attachment some distance from the border of the operculum. 
The group is left by Levinsen under Cellepora, through mis- 
understanding C. ramulosa. The ovicell of Osthimosia is, at any 
rate, nearly always punctured. 

This leaves the Cellepora of Hincks divided into Holoporellidee 
(holostomatous), and the schizostomatous forms into Osthimosia 
and Lagenipora. 

Loc. Prison Island, Zanzibar Channel, 8 fath. (505); Ras 
Osowamembe, Zanzibar Channel, 10 fath. (504), collected by 
Crossland. 


LAGENIPORA ROTA ee eis 


C ellepor a rota MacG. Trans. Roy. Soc. Vict. vol. xxi. p. 116 
(11) pl. i. fig. 6 (1885) ; Prod. Zool. Vict. dec. xv. p. 184, pl. 148. 
fig. 3 (1887). 

Levinsen would call this Siniopelta, but it is what I have 
previously put under Lagenipora, as the position and character of 
the ovicells of L. socialis Hincks seem to me to be the same as in 
the group which Levinsen calls Siniopelta. As Levinsen has not 
agreed with me, I have also, besides again examining the British 
Museum specimens, through the kindness of Professor Hickson, 
examined the specimens from Miss Jelly’s collection in the 
Victoria University Museum. Miss Jelly first found ZL. socialis 
in Hastings, and presumably all known Hastings specimens were 
collected by her. The Hastings specimens in the Victoria 
Museum did not show the ovicell, ‘but one sonamed by Miss Jelly, 
from Guernsey, has some ovicells and also shows the spinous 
processes well. 

The ovicell of the Guernsey specimen is situated on the wall 


* Waters, “ Bryozoa of the Bay of Naples,’ Ann. Mag. Nat. Hist. ser. 5, vol. iii. 
p. 198 (1879). 


BRYOZOA FROM ZANZIBAR, Bt I 


of the peristome, which frequently extends beyond it as a com- 
plete tube, and the ovicell has an area which is flat or slightly 
rounded and is surrounded by a ridge, while at each corner 
inside the ridge there isa pit. It seems justifiable to call the 
part surrounded by the ridge an area, but if there is any 
objection to this it might be called a tabula. In this case the 
‘“‘ calcareous base” is very slightly developed and might be over- 
looked, and except that the Guernsey specimen is slightly 
granular or rather nodulated there is but little difference 
between it and Lagenipora lucida Hincks from Madeira, except 
that Z. Jucida has pores round the area as figured by me*, but 
I have not the opportunity of now comparing any ZL. lucida 
having ovicells. 

Now in Lagenipora nitens MacG. from Port Phillip Heads, 
which I considered only a variety of lucida, the ovicell is just 
the same shape and in the same position, while close to the ridge 
of the area there is a row of pores; also L. boryii Aud. from the 
Mediterranean has a row of pores in the same place, whereas in 
L. rota MacG. and some other species there are radiating grooves. 
An Australian species of Lagenipora has the area evenly perforated 
all over. The pores or pits of ZL. socialis, though difficult to 
trace, are no doubt small and close to the edge. When JI first 
thought I saw pores it was puzzling not to find a complete row, 
and there was a doubt whether they might only result from 
accidental damage, but no doubt the explanation is that there is 
only one at each corner, . There must be something wrong if 
Lagenipora socialis and L. lucida ave placed in different genera. 

Loc. Victoria (MacG’.). Wasin, Brit. East Africa, 10 fath. 
(507), collected by Crossland. 


HASWELLIA AUSTRALIENSIS Haswell. 


Myriozoum australiense Haswell, Proc. Linn. Soc. N.S. Wales, 
vol. v. pt. 1, p. 43, pl. iii. figs. 9-11 (1880). 

Haswellia australiensis Busk, “ Polyzoa,” Zool. Chall. Exp. vol. x. 
pt. xxx. p. 172, pl. xxiv. fig. 9 (1884); Kirkpatrick, Proc. Roy: 
Dublin Soe. vol. vi. p. 612 (1890); Meissner, M., aus Semon 
“Zool. Forsch, Reisen in Australien und Malay,” Jena. Denkschr. 
vol. vill. p. 731 (1902); Levinsen, Morph. & Syst. Studies of the 
Cheil. Bryozoa, p. 297, pl. xvi. figs. 2 a—2 6 (1909). 

Porina coronata var. labrosa Waters, Zool. Chall. Exp. vol. xxxi. 
pt. lxxix. p. 32 (1889). 

Levinsen places this in Myriozoide t, but it seems somewhat 
doubtful whether the genera grouped together by Levinsen will 
remain in the same family (see p. 520). 

The series of zocecia in H. australiensis are all on the same 
horizon, and usually at the beginning of each branch there are 
six zocecia in a whorl, there is then another whorl with six, then 
one with eight when a bifurcation takes place, and then again 


* Journ. R. Mier. Soc. 1899, p. 18, pl. ili. figs. 25, 27, 30 (1899). 
+ Loe. cit., see page 296. 


512 MR. A. W. WATERS ON 


each of the new series commences with six. H. australiensis has 
19-21 tentacles. Unfortunately none of the specimens now 
examined have ovicells. The //aswellia group is well represented 
in the North Italian Tertiaries, and although some appear to be 
almost identical with living species, it seems better to keep them 
distinct, as all the structures cannot be compared. 

There is over the surface an outer membrane, and then under 
this there are membranes of the two walls of the shell showing 
when decalcified the position of the pore-tubes, Some specimens 
from Wasin have a pink colour. 

The operculum of the species and of the H. coronata H. or 
H. gracilis, whichever we may have to call it, is identical, having 
the opercular attachment very high up and elongate (see my 
figure in Ann. Mag. Nat. Hist. ser. 5, WO, so TO, Wik)s Ine 
operculum of H. auriculata B. has the muscular attachments much 
lower down and not so near the edge ; further, the ovicell which is 
but slightly raised, has a semicircular area, and the surface of 
the zocecium has not poresall over. It seems exceedingly doubt- 
ful whether H. auriculata should remain in Haswellia, but I have 
never had sufficient material in my hands for a complete 
examination. H. grandipora Waters is Haswellia, and I now 
think must stand as a species. 

Loc. Holborn Island, Queensland (asw.); near Torres Straits 
(Chall.); Thursday Island, Torres Straits (Jeissner); Formosa 
Channel (Levinsen). Wasin, Brit. Hast Africa, 10 fath. (500, 
501, 507, 520), collected by Crossland. 


T'UBUCELLARIA CEREOIDES, var. CHUAKENSIS Waters. 

Tubucellaria cereoides var. chuakensis Waters, ‘ Tubucellaria,” 
Journ. Linn. Soc., Zool. vol. xxx. p. 130, pl. xv. figs. 10-13, 18, 
19, pl. xvi. figs. 20-25 (1907); Thornely, “Mar. Poly. Indian 
Ocean,” Trans. Linn. Soc., Zool. vol. xv. p. 146 (1910). 

Tubucellaria fusiformis Busk (non d’Orb.), Zool. Chall. Exp. 
vol. x. p. 100 (1884). 

Loc. Torres Straits (Busk); Grahamstown, 8. Africa; Provi- 
dence, 50-78 faths. Chuaka, Zanzibar, 3 fath. (512, 524); 
Wasin, Brit. E. Africa, 10 fath. (501); Chaki-Chaki, Pemba 
Island (517), low water, collected by Crossland. 


TUBUCELLARIA FUSIFORMIS d’Orbigny. 

Waters, loc. cié. p. 131, pl. xv. figs. 1, 2, 3, 14. 

Loc. Malacca, Ainirante Is., Chuaka, Zanzibar, 3 fath. (528) ; 
Wasin, Brit. East Africa, 10 fath. (520), collected by Crossland. 


TUBUCELLARIA ZANZIBARIENSIS Waters. 

Waters, loc. cit. p. 131, pl. xv. figs. 4-7; Thornely, “ Mar, Poly. 
Indian Ocean,” Trans. Linn. Soe., Zool. vol. xv. p. 146 (1912). 

Loc. Saya de Malha, 145-150 faths.; Farquahar Reef ; 


BRYOZOA FROM ZANZIBAR. ils 


Cargados, 28 fath. Wasin, Brit. East Africa, 10 fath. (501) ; 
Ras Osowamembe, Zanzibar Channel, 10 fath. (504, 514); Prison 
Island, Zanzibar Channel (505); Chuaka Bay: collected by 
Crossland. 


SMITTINA TRISPINOSA, var. PROTECTA Thornely. 
Smittia trispinosa, var. protecta Vhornely, Ceylon Pearl-Oyster 
_ Fisheries, vol. iv. Polyzoa, p. 123 (1905); Waters, “ Mar. Biol. 
of the Sudanese Red Sea,” Journ. Linn. Soec., Zool. vol. xxxi. 
p. 173, pl. xvii. figs. 5, 6 (1908). 

Smittia nitida Hincks, Ann. Mag. Nat. Hist. ser. 5, vol. vii. 
p. 159, pl. ix. fig. 5 (1881). 

There are two specimens with the large avicularia situated 
diagonally distal to the oral aperture. The ovicells of this variety 
and of typical nitida are similar, and there are some specimens 
with a large, almost spinous, process at the proximal part of the 
peristome, with a similar process on the ovicell just distal to the 
area of pores. An identically similar form occurs off the Cape 
Verde Islands, and Osburn * menticns an umbo in some forms of 
nitida “behind the orifice,” and he shows how WS. trispinosa var. 
nitida is subject to great variation with regard to the avicularia 
and the peristome. 

Loc. Gulf of Manaar (7h.); “ Africa” (J.); Red Sea (W.). 
Wasin, Brit. E. Africa, 10 fath. (520), collected by Crossland. 


SMITTINA TRISPINOSA, var. SPATHULATA MacGillivray. 


Waters, “ Mar. Biol. Sudanese Red Sea,” Journ. Linn. Soc., 
Zool. vol. xxxi. p. 156. 


The ovicell has a distinct area, the border of which sometimes 
rises aS an umbo, or is divided into two sharp, erect processes. 
S. trispinosa var. bimucronata Hincks belongs to this group, but 
it is doubtful whether it should be separated asa variety, as there 
is normally such considerable variation in the zoecia. From 

(504) there are very large vicarious avicularia, directed either 
distally or proximally. 

Smittina is used instead of Smittia, though not including all 
that Levinsen refers to it, for I consider it a group in which the 
operculum is usually very thin, almost membranous, with the 
lower edge straight. In the species so far examined, the oral 
glands are quite small and usually more or less attached to the 
tentacular sheath. In the aperture a lyrula is usually found with 
the operculum under the cardelle but over the lyrula. Psewdo- 
flustra solida Stimp., and Lepralia pallasiana, together with its 
allies, show many differences and do not seem to belong here. 

Loc. Bass's Straits; Torres Straits; Red Sea, Wasin, Brit. E. 
Africa, 10 fath. (520); Ras Osowamembe, Zanzibar Channel, 
10 fath. (504), collected by Crossland. 


* “The Bryozoa of the Woods Hole Region,” Bull. Bur. of Fisheries, yol. xxx. 
p. 246 (1912). 


514 MR. A. W. WATERS ON 


SMITTINA TROPICA Waters. t 

Smittia tropica Waters, “ Mar. Biol. of the Sudanese Red 
Sea,” Journ. Linn. Soc., Zool. vol. xxxi. p. 174, pl. xvii. figs. 10-14 
(1909). 

Loc. Red Sea. _ Wasin, 20 fath. (520), collected by Crossland. 


SMITTINA Sp. 


There is a small piece of Smittina encrusting ddeonella platalea, — 
which has the peristome very much raised, especially at the distal 
part, and there is apparently a triangular avicularium in the lip. 
There are perforations round the border of the zocecium, and the 
small ovicell is not much raised and is at the base of the peristome, 
and at each side of the ovicell close to the edge there is a small 
clear spot. There are two openings in the ovicell of S. oculata 
MacG., and sometimes others; and I find in a specimen sent me 
by Jullien as 8. longirostris J., there is a similar spot at the side 
of an ovicell. Jullien’s species has large frontal avicularia, 
whereas none are found on the small fragment. The lyrula is at 
first very narrow but spreads out widely on each side. I hesitate 
to give it a name until better specimens are found. 

Loc. Ras Osowamembe, Zanzibar Channel, 10 fath. (504). 


LEPRALIA FEEGENSIS Busk. (Pl. LXX. figs. 21, 22.) 


Lepralia feegensis Busk, Zool. Chall. Exp. vol. x. pt. xxx. 
p. 144, pl. xxii. fig. 9 (1884); Philipps, in Willey’s Zool. Results, 
pt. iv. p. 446, pl. xl. fig. 7 (1889); MacGillivray, Proce. Roy. 
SocwVaets mass) vole iis p: Sl, plo ox. shlosh a(S oi) seh onmelive 
Ceylon Pearl-Oyster Fisheries, vol. iv. p. 121 (1905); Rec. of 
Indian Mus. vol. i. pt. 3, no. 13, p. 190 (1907); “ Mar. Polyzoa 
Tndian Ocean,” Trans. Linn. Soc., Zool. vol. xv. p. 150 (1912). 

Hippopodina feegensis, Levinsen, Morph. & Syst. Studies on 
the Cheil. Bry. p. 353, pl. xxiv. figs. 3 a—3 f (1909). 

Miss Philipps described and figured the ovicell as with opaque 
granulations, globular and depressed ; and Miss Thornely, not 
aware that the ovicell had been seen, described it as with large 
circular pitted areas punctured in their centres. In a specimen 
in my collection, from “ Singapore or Philippines,” the ovicell has 
the pores fairly similar to, and about the same size as, the pores 
of the zocecia. MacGillivray had previously described the ovicell 
of his LZ. feegensis, but as the avicularium, which is directed down- 
wards, is in quite a different position, I doubt whether it is the 
feegensis of Busk. He described the ovicell as large, rounded, 
prominent, and marked similarly to the zocecia, In the specimens 
from Zanzibar (501) there are no ovicells, glands, ovaria, or 
testes, but in those from (511) there are many ovicells which are 
perforated, though when the ovicell contains an ovum or embryo 
these perforations look dark and opaque. 

There are about 26 tentacles, and the operculum closes the 
ovicell. There ave several ovarian cells in an ovarium. 


BRYOZOA FROM ZANZIBAR. 515 


In the submandibular part of the avicularium there is a 
hammer-shaped thick chitinous piece for attachments of muscle. 
The proximal muscles are in several bundles and are attached to 
the base of the tentacles. 

Lepralia is a genus which we have long wanted.to see brought 
into order, but certainly further work is required. Hincks made 
a group Lepralia, and, no doubt, on the whole the species of his 
group are closely related, but he unquestionably placed some 
there which should have gone in his Schizoporella and other 
genera. Neviani, taking species with a horse-shoe aperture, 
made a genus Hippoporina, but he still retained some which 
must be separated, and very little was gained by his new 
name. In Lepralia the operculum is usually thick and has a 
strengthening band at each side with the muscles attached near 
the distal end of the band. ‘The sides of the operculum are either 
straight or much contracted where the lateral teeth of the 
aperture occur. This will remain a fairly large group even when 
extraneous species have been removed, but we hardly know where 
Levinsen would place them, as hardly any of them are mentioned. 
L. adpressa, L. hippopus, L. rectilineata Hincks, find no place. 
Lepralia feegensis is made the sole. representative of a new genus 
Hippopodina, but the characters given seem insufficient for 
separation, as there are a considerable number with the same 
general characters. 

Some Lepralia Levinsen places under Smittina, a new name for 
Smittia, but the true Smittia to which he alludes have a very 
thin membranous operculum with straight proximal edge hardly 
separated from the frontal membrane, and there is usually a lyrula, 
which is perhaps a structural correlation with the thin mem- 
branous operculum; further, there are very small oral glands 
often partly attached to the tentacular sheath. I find it quite 
impossible to place Lepralia pallasiana and its allies side by side 
with Schizoporella auriculata Hass., S. linearis Hass., S. triangula 
Hincks, ete., as the group seems very unnatural. 

Levinsen’s Cheilopora* contains species some of which have the 
ovicelligerous zocecia with a quite differently shaped aperture from 
that of the ordinary zowcia, as, for instance, Lepralia cirewmeinetu 
Neviani, a species found fossil, but also living, from Naples, 
Capri, and Oran (Algiers) ; whereas LL. longipora MacG.= L. pre- 
longa Wincks, and L. prelucida H. have large raised perforated 
ovicells. This dimorphism occurs in several Lepralide, but in 
Schizoporellide I only remember it in S. swbimmersa MacG. We 
hardly know yet how far these characters are reliable for generic 
divisions. 

Such forms as LZ. adpressa Busk, LZ. hippopus Sm., L. recti- 
lineata H., and the bulk of what we have known as /#pralia 
should remain there until ample living and spirit material of 
numerous specimens has been examined. 


* Haime made a genus Chilopora and Michelotti called one Cheilopora- 


Proc. Zoou. Soc.—1913, No. XXXV. 35 


516 MR. A. W. WATERS ON 


Loc. Philippines, 18 fath. (Chall.) ; Singapore or Philippines 
(A. W. W. coll.); Hong Kong; Lifu (Ph.); Manaar (7h.); 
Andamans (7h.); Cargados (Ind. Ocean) (Zh.); (¢) Nichol Bay, 
N. W. Australia (ZacG.). Wasin, Brit. East Africa, on Adeonella, 
10 fath., from bottom of s.s. ‘Juba,’ which always remains in 
Zanzibar waters (511), collected by Crossland. 


LEPRALIA TURRITA Smitt. (Pl. LX XIII. fig. 10.) 


Lepralia turrita Smitt, Floridan Bryozoa, pt. 1. p. 65, pl. xi. 
figs. 226-228 (1873); Kirkpatrick, Ann. Mag. Nat. Hist. ser. 6, 
vol. v. p. 16 (1890); Thornely, “Mar Poly. of the Indian Ocean,” 
Trans. Linn. Soc., Zool, vol. xv. p. 150 (1912). 

The small specimen from Wasin has four long calcareous 
spinous processes, irregularly placed round the aperture. One 
or more of these processes may bear an avicularium at one side 
near the base, as in the spinous process of H. albirostris Sm. 
Besides these avicularia there are on the surface of the zoarium 
numerous small semicircular avicularia, irregularly scattered. 
The ovicell is not much raised and has an elongate elliptical 
opening in the front. A specimen of Lepralia turrita in my 
collection has short blunt processes with similar semicircular 
avicularia, and the ovicell, which is more raised, has a similar wide 
opening in the front. Perhaps, on account of the different 
character of the ovicell, it should at least be made a variety, but 
until more material is available it is allowed to stand. 

The operculum has the sides nearly straight with the proximal 
edge curved ; there are two muscular dots a little distance from 
the edge and two articular thickenings ; the width is about 0°2 mm. 
This operculum does not correspond with any with which I am 
acquainted, for though in many respects it is like those of what I 
should call, ina restricted sense, Lepralia, there is no lateral ridge 
for the muscular attachment, which is not close to the border. 

In the British Museum there is a specimen marked turrita, 
which is Holoporella, and it seems as if there were more than one 
with a series of calcareous processes round the oral aperture. 
What Ridley and what I called Z. twrrita may each have to be 
placed elsewhere. 

Loc. Ascension Island ; Florida (Sm.) ; China Seas; Amirante, 
29 fath.; Cargados, 30 fath. Wasin, Brit. Hast Africa, 20 fath. 
(522), collected by Crossland. 


LEPRALIA WASINENSIS, sp.n. (Text-fig. 81.) 


A small piece with only seven zocecia was found on re-examin- 
ing some material, when the paper was almost completed. Only 
the caleareous part remains, which was probably bilaminate, but 
this cannot be stated with certainty. The zoccia have the sides 
straight and the distal end somewhat rounded, with thick borders 
to the zoecia. The oral aperture is subrotund with a wide 
curve on the distal end surrounded bya thick band. The frontal 


BRYOZOA FROM ZANZIBAR. 517 


surface is closely pitted, and at each side of the oral aperture, 
somewhat below it, there is a large subtriangular chamber which 
is probably avicularian. In the band surrounding the oral 
aperture there is an opening at the proximal end which appears 


to be avicularian. 


Text-fig. 81. 


Lepralia wasinensis, sp.n. X 25 


The species seems most nearly related to Lepralia (Mucronella) 
prelucida Hincks, which has a large avicularium by the side of 
the oral aperture as seen in my specimens, and as Professor R. 
Osburn * mentions in his specimens from Labrador. 

Loc. Wasin, Brit. East Africa, 20 fath. (520), collected by 
Crossland. 


LEPRALIA CLEIDOSTOMA, var. INERMIS Ortmann. 


Lepralia cleidostoma, var. inermis Ortmann, ‘“ Die Japanische 
Bry.,” Archiv fiir Naturgesch. vol. i. p. 49 (1890). 

Lepralia cleidostoma, var., Hincks, Ann. Mag. Nat. Hist. ser. 5, 
vol. xii. p. (41) (1884). 

This form has the surface smooth and silvery with frequently 
an umbo below the aperture and often one on the ovicell. The 
ovicell is scarcely at all raised and is also smooth and imperforate, 
and has a small semilunate area. There is sometimes a small 
knob on each side of the aperture. Both the aperture and the 
operculum are about the same size and form as that of LZ. cleido- 
stoma Smitt from Madeira 7, which also occurs at Florida (Sm.), 
Japan (Ort.), and Bermuda (my coll.) ; but L. imermis var. differs 
in having no avicularia, and it is a question whether it is advisable 
to consider it a species or a variety. 

Norman says ¥¢ that L. cleidostoma is the L. porcellana Busk, but 


* “Bryozoa from Labrador, etc.” Proc. U.S. Nat. Mus. vol. xliii. p. 283, 
pl. xxxiv. figs. 3, 3a, 3c (1912). 

+ Waters, “ Bryozoa from Madeira,’ Journ. R. Micr. Soc. 1899, p. 10, pl. iii. 
fig. 16. 

2; “Polyzoa of Madeira,” Journ. Linn. Soc., Zool. vol. xxx. p. 305, pl. xl. figs. 1, 2 


(1909). 
35* 


518 MR. A. W. WATERS ON 


as it has been impossible to recognise it from Busk’s figures the 
name cleidostoma must stand. 

Loc. Queen Charlotte Island (Hincks); Japan, 100 fath. 
(Ortmann). Wasin, Brit. Kast Africa, 10 fath. (520), collected by 
Crossland. 


PETRALIA JAPONICA Busk, 


Lepralia japonica, Waters, Mar. Biol. of the Sudanese Red Sea, 
Journ. Linn. Soc., Zool. vol. xxxi. p. 149, pl. xiii. figs. 10-12. 


Loc. Add (Natal, as Hemeschara gigantea Busk MSS. in the 
Brit. Mus.) Wasin, Brit. Hast Africa, 10 fath. (501), collected by 
Crossland. 


PETRALIA CHUAKENSIS, sp. n. (Pl. LXX. figs. 10-14.) 


Specimens from Chuaka correspond in most particulars with 
Busk’s description of Lepralia dorsiporosa *, but the notch at the 
distal end of the zoccium is wanting, and in chuakensis the 
proximal edge of the aperture is serrate. The avicularium 
at the side is curved and is larger than those figured by Busk. 
The Zanzibar specimens have ovicells which are unknown in 
dorsiporosa, and the wall, which is not much raised, is finely 
perforated. 

The perforated area in the dorsal surface is for a broad radicle, 
which, however, is not always developed ; a similar area occurs in 
Petralia vultur var. armata, and also in Petralia japonica there 
is often a similar large radicle. Levinsen refers to the radicle 
tube in P. dorsiporosa. There is a complete bar to the avicularia 
often disappearing in Eau de Javelle preparations. There are 
numerous uni- or few-pored rosette-plates scattered over the 
distal wall. On the lateral wall, about halfway between the frontal 
and basal walls, there is a row of about 5 few-pored rosette-plates. 

There are about 28 tentacles. In the ovarium there are but 
few ovarian cells. 

It has been thought best to give a specific name, although there 
is much in common with P. dorsiporosa from the Torres Straits. 

Loc. Chuaka, Zanzibar, 3 fath. (502), collected by Crossland. 


PETRALIA vuLTUR Hincks, var. ARMATA, nov. (Pl. LXX. 
figs. 15-20). 

Type, Mucronelta vultur Hincks, Ann. Mag. Nat. Hist. ser. 5, 
vol. x. p. 98, pl. viii. fig. 2 (1882). 

From Chuaka, Zanzibar, there are unilaminate specimens 
differing from the Mucronella vultur as described by Miss 
Thornely 7, in having a triangular mandible to the avicularium 
attached to the mucro. ‘There are also a large number of raised 
round avicularia, with semicircular mandibles, and a distinct 
cross-bar. These avicularia are sometimes on the top of a raised 


* Zool. Chall. Exp. vol. x. p. 148, pl. xviii. fig. 4. 
+ Ceylon Pearl-Oyster Fisheries, vol. iv. p. 124 (1905). 


BRYOZOA FROM ZANZIBAR. 519 


tube. There are a few large spatulate avicularia at one of the 
distal corners, and these also have a distinct cross-bar. Speci- 
mens of JZ. vultur in Miss Thornely’s collection show the mandible 
serrate. 

There is a fairly broad lyrula and two cardelle, and the lyrula 
is formed before the front wall is complete. Only one ovicell has 
been seen, and that is partly broken, but it is broad and minutely 
perforated. 

On the dorsal surface there are perforated spaces, no doubt for 
radicles, but there are usually several small ones instead of one 
large one as in Lepralia dorsiporosa B. 

Similar ovicells with minute perforations occur in Petralia wi- 
data MacG., P. japonica Busk, P. vultur Hincks, P. ellerii MacG., 
P. (Mucronella) magnifica Busk, P. (Lscharella) bisinuata Smitt 
(non Busk), P. thenardii Aud., P. (JL) porosa Hincks, P. (JZ) 
castanea Busk, L. crassa Thornely; and these Levinsen would 
piace in his family Petraliide, and thus it would seem as though 
a genus can be separated based largely on the ovicell. In the 
species of this group which it has been possible to examine, there 
are a large number of tentacles, viz., Petralia undata MacG., 
about 26; P. japonica B., about 25; P. castanea B., about 23 ; 
P. vultur var. armata, nov., about 25; P. chuakensis, nov., 
about 25. 

There seem to be two tentacles larger than the rest (figs. 15-17), 
and these, instead of being triangular, have the inner surface 
nearly straight, with a number of long nuclei. On this inner 
surface there are cilia, but unfortunately the condition does not 
admit of exact study of this point. These two larger tentacles 
occur in all the species examined of the group, and larger tentacles 
occur in other groups to which I have previously alluded. These 
large tentacles are most marked near to the base of the tentacles 
when they are commencing to divide, whereas nearer to the ends 
there is but little difference. 

Loc. Chuaka, Zanzibar, 3 fath., 29.38.1901 (506), collected by 
Crossland. 


? EsCHAROIDES OCCLUSA Busk. 


See ¢Lepralia occlusa Waters, ‘‘ Mar. Biol. of the Sudanese Red 
Sea,” Journ, Linn. Soc., Zool. vol. xxxi. p. 152, pl. xiii. fig. 15, 
pl. xiv. figs. 1-9, 13 (1909). 

The distal and lateral walls have numerous scattered uniporous 
rosette-plates, especially near the inner borders. At the side of 
the zoarium there are often two large spatulate opposite avicu- 
laria, whereas in Adeonella the avicularia are usually on a median 
line. 

Levinsen would put this under Myriozoidea, but I hardly think 
he can have had sufficient material before him when writing about 
the family, and that is often our difficulty in considering classi- 
fication. 


520 MR, A. W. WATERS ON 


The type of Myriozowm must, of course, be I. truncatwm, and 
Tam not sure that any other living species can be put in the 
same genus. It has a large number of tentacles, 26-27, which 
should lead us to consider carefully whether the species with about 
16 tentacles belong to the same genus, though we cannot conclude 
from that alone that they do not. 

Levinsen makes one of the characters of Myriozoidea, ‘“ de- 
pendent avicularia seem always to be present,” but in the type 
M. truncatum no avicularium has been found; another of his 
characters of Myriozoum and Myriozoella is no transverse bar in 
the avicularium, but Schizoporelia bituwrrita H. and what I have 
called S. fuberosa Rss. and Hscharoides occlusa have a distinct bar. 
In WM. truncatum the ovicell is a very large swelling with the 
covering wall resembling the zowcial wall. S. crwstacea has a 
prominent round ovicell with radiating ornamentation. S. biter- 
rita H. and S. tuberosa have very large wide raised ovicells. In 
M. simplex Busk no ovicell is externally apparent, though cal- 
careous sections reveal a concealed ovicell. In S. polymon phe the 
ovicell is also under the calcareous wall, giving, as a rule, no 
external sign (see my suppl. ‘Challenger’ Rep. vol. xxxi. pl. i. 
figs. 22, 23). In S. biturrita and S. polymorpha B. there is an 
avicularium at each side of the oral aperture, and in J/. mario- 
nensis B., at each side well within the peristome, there is a 
small avicularium with a mandible rounded at the end. 

Now, in ? Escharoides occlusa there is an avicularium on the 
lip of the peristome at one side*, the ovicell is but moderately 
raised, with a few large openings, the avicularia have a bar and 
there are very large glands, at first paired, but often ultimately 
uniting to form a large multilobular gland. So far as my sections 
go, namely, S. crustacea, M. coarctum, M. subgracile, and M. trunca- 
twm, there are no glands in the species considered as Myriozoidea 
except in H. occlusa, and these are quite different from any others 
known. Also the general absence of glands indicates that Myrio- 
zoidea of Levinsen are not, as considered by him, closely allied to 
the Reteporid, in which glands are so well developed, and which 
usually have a sublabial pore and a lamina to the ovicells, also 
vibices indicating two distinct layers. ‘The operculum of Z. occlusa 
is quite different from that of any other species placed by Levinsen 
under Myriozoidea. It has a thick border all round, and near 
the distal edge are two knobs for the attachment of the muscles. 
Myriozoum truncatum has a very large operculum of a shape 
quite different from that of the others mentioned, the proximal 
portion of the operculum being somewhat triangular with the 
attachments near to the proximalend. M/. subgracile, M. coarctum, 
and MM. crustacea have the sides of the moderate-sized operculum 
nearly straight, with the muscle near the side. Haswellia 
australiensis has the muscular attachment near the border of the 


* The position of the avicularium is similar in Rhamphostomella, etc., but we do 
not yet know the systematic value of this character. 


BRYOZOA FROM ZANZIBAR. SPAM 


operculum. J/yriozowm marionensis would seem to be Cellarinella 
or related to it. Gephyrophora polymorpha has a large operculum 
nearly round, with a wide vanna fitting into the poster, and the 
muscular dot is some distance from the edge, in fact it is of a 
Schizoporellidan character. 

It seems probable that H. occlusa must be made the type of a 
new genus, and I expect that the examination of material suit- 
able for cutting will result in a re-arrangement of classification 
of the Myriozoidea of Levinsen, as the ovicells present such great 
differences. I am loth to make new genera until we can see from 
sufficient material what is to be grouped together ; at any rate, I 
fail to see good reason for placing Hscharoides occlusa * under 
Myriozoum. 

Loc. See Waters, ‘“‘Red Sea Bryozoa” p. 156. Wasin, Brit. 
East Africa, 10 fath. (501) (520); Ras Osowamembe, Zanzibar 
Channel, 10 fath. (504), collected by Crossland. 


HoLoPoRELLA COLUMNARIS Busk. 


Cellepora columnaris Busk, Zool. Chall. Exp. vol. x. pt. xxx. 
p. 194, pl. xxix. fig. 11, pl. xxxv. fig. 16 (1884); Waters, Ann. 
Mag. Nat. Hist. ser. 6, vol. iv. pl. il. figs. 1-6 (1889); Ortmann, 
“Die Japanische Bryozoenfauna,” Archiv f. Naturgesch. vol. i. 
p- 55 (1890). 

Cellepora cidaris MacG. Prod. Zool. Vict. dec. xvii. p. 243, 
pl. 165. fig. 4 (1888) ; Thornely, ‘‘Ceylon Pearl-Oyster Fisheries,” 
vol. iv. Poly. p. 126 (1905); Records of Indian Museum, p. 195 
(1907). 

The oral aperture has a denticle at each side. As shown in my 
paper mentioned above, the columns are often very thick, ex- 
tending through several layers of the zoarium. A specimen 
of Holoporella in my possession, named in manuscript celosia by 
Busk, has similar columns passing through several rows of zocecia, 
and the avicularian mandible has a double columella, while the 
shape of the operculum also shows that it is a distinct species. 
This is not, however, the species so named in the Busk collection 
in the British Museum, where there are two other species named 
celosia, MSS. 

I have had the opportunity of comparing Busk’s types and 
MacGillivray’s co-types (7. e., duplicates) in the British Museum, 
and there can be no doubt as to the identity of columnaris and 
cidaris. 

In a specimen of colwmnaris from Wasin there are numerous 
small hydroids extending far into the zoarium within, protecting 
internal walls, formed by the WHoloporella, prolonged slightly 


* Hincks considered H’scharoides, a name given by Milne-Edwards for a subgenus 
of Cellepora, as being now the genus of Smitt, who used it for species with an 
avicularium in the lip on one or both sides, but Smitt never fully diagnosed it. As 
Escharoides, a name originally meaningless, has been used in a quite different sense 
from Levinsen’s, it would be much better if he recalled his genus Peristomella and 
used it instead of E’scharoides. 


He 


Oo, MR. A. W. WATERS ON 


beyond the zoarium. Sometimes they start from the base, and 
the poiypides stand out from the upper surface of the zoarium. 
A similar case from the Red Sea is mentioned in the addenda to 
my Report*. Probably the hydroids in both are Clava. 

There are about 18 tentacles. There are a great number of 
ovaria with two small ovarian cells, and in only one or two cases 
has a moderate-sized ovarian cell been seen, and the ovaria are of 
the Bugula type. The ovaria may be surrounded by testes. 

Loc. Bass’s Straits, 38 fath. (Chall.); Port Phillip Heads 
(MacG’.); Port Jackson, N.S.W. (Waters); Sagamibai, Japan 
(Ort.); Manaar, 34 fath.; Ceylon Coast, 32-34 fath.; off Port 
Blair, 100 fath.; St. 59, 32 fath., St. 77, 35 fath. (Zhornely). 
Wasin, Brit. Hast Africa, 10 fath. (501), collected by Crossland. 


HoLoporeEuuA APERTA Hincks. 


Waters, ‘Mar. Biol. of the Sudanese Red Sea,” ‘“ Bryozoa,” 
Journ. Linn. Soc., Zool. vol. xxxi. p. 161, pl. xvii. figs. 20-23 
(1909). 

Loc. Additional. Wasin, Brit. Kast Africa, 10 fath. (500); Ras 
Osowamembe, Zanzibar Channel, 10 fath. (504), collected by 
Crossland. 


HoLoporELLA ALBIROSTRIS Smitt. (Pl. LX XIII. fig. 11.) 


Cellepora albirostris, forma typica Smitt, ‘ Floridan Bryozoa,” 
pt. ii. p. 70, pl. xii. figs. 234-239 (1873). 

Cellepora albirostris Busk, Journ. Linn. Soc., Zool. vol. xv. p. 347, 
pl. xxvi. fig. 2 (non fig. 1) (1881); Zool. Chall. Exp. vol. x. pt. xxx. 
p. 193, pl. xxxiv. fig. 7, pl. xxxv. fig. 3 (1884); Waters, Quart. 
Journ. Geol. Soc. vol. xli. p. 304 (1885); op. cit. vol. xlii. p. 68 
(1887); Ann. Mag. Nat. Hist. ser. 5, vol. xx. p: 197 (1887); 
MacGillivray, Prod. Zool. Vict. dec. xvii. p. 249, pl. 167. fig. 1 
(1888) ; Mon. Tert. Poly. p. 109, pl. xiv. fig. 11 (1895); Thornely, 
“Mar. Poly. Indian Ocean,” Trans. Linn. Soc., Zool. ser. 2,-vol. xv. 
Oise hs fos Ltaysy (ILS) : 

In HZ. albirostris there is great variation in the colour, some 
being very dark with the ends of the rostra white, others may be 
nearly white. 

There are vicarious duck-bill avicularia and the border of the 
rostral avicularia is denticulate ; the oral glands (fig. 11) are very 
long, and there are 16-17 tentacles. 

A specimen is partly adnate, from which bilaminate branches 
arise. The unilaminate portions, when seen from below, are quite 
regular with the sides of the zocecia parallel, as in so many 
Schizoporellide. 

Both Smitt and Busk expressed doubt as to whether //. albi- 
rostris and H, bispinata were synonyms, but this is not the case, 
for albirostris has a long spine below the oral aperture, while 
bispinata has a short avicularium ; further, im 7. albirostris there 


* Journ. Linn. Soc., Zool. vol. xxxi. p. 254, p. 168, pl. xv. fig. 16 (1910). 


BRYOZOA FROM ZANZIBAR. De 


isa pair of the minute avicularia at the side of the zocecium 
to which I referred in my Report of the Sudanese Bryozoa, but 
none were found in H. bispinata. Further, the operculum of 
H, albirosiris has a muscular ridge turning sharply inwards and 
ending off short, whereas the operculum of H. bispinata has a 
ridge, very thick at the base, extending as a thin band nearly to 
the apex. H. albirostris has a very long oral gland nearly the 
length of the polypide, while the gland of H. lispinata, though 
also cylindrical, is much smaller. 

Loc. Florida, 25-35 fath. (Sim.); Heard Island, 35 fath. (Chall.); 
New South Wales (Waters); Victoria (ZacG'.) ; Amirante, 12-18 
fath. (Zhornely). Wasin, B. East Africa, 10 fath. (501); Chuaka, 
Zanzibar, 3 fath. (525), collected by Crossland. 

Fossil. Australia, N. Zealand (WV); Victoria (JlacG.). 


MiICROPORELLA CILIATA Pallas. 


Loe. Northern Seas; British; Atlantic; Mediterranean ; Red 
Sea; Ceylon; Australasia and varieties in Japan, Australasia ; 
Zanzibar (H.), the N. Indian Ocean. Wasin, Brit. KE. Africa, 10 
fath. (500), collected by Crossland. 

Fossil. Pliocene and Miocene of Europe, and Tertiary of 
Australia and New Zealand. 


RHYNCHOZOON PROFUNDUM, var. LAMINATUM, nov. (PI. LX XII. 
Hees) 

The specimens from Wasin may be the 2. profundum as first 
described by MacGillivray, but it is not the profundwm of the 
Prodromus, which MacGillivray considers is the same as 2. longi- 
rostre Hincks. 

The species of Rhynchozoon are difficult to distinguish, but the 
ovicell has a lamina in the front like the lamina of Retepora, to 
which it is related, and the shape of the lamina furnishes specific 
characters of value. In this Zanzibar variety the lamina is long 
and rounded at the end, whereas in 2. profundum it is shorter, 
very broad, and spreads out at each side. In &. bispinosum Johns. 
it is quite short. 

In this variety as in &. profundum the denticle is broad and 
uncinate, and the ovicell is but little raised above the general 
surface. ‘There are narrow pointed avicularia over the surface, 
as usual in Lynchozoon. 

Loc. Wasin, Brit. Hast Africa, 10 fath. (520), collected by 
Crossland. Noumea (in A. IW. W. coll.). 


RETEPORA HIRSUTA Busk. 


Retepora hirsuta Busk, Zool. Chall. Exp. vol. x. p. 119, pl. xxvi. 
fig. 4 (1884); Waters, ‘‘ Mar. Biol. of the Sudanese Red Sea,” 
Journ. Linn. Soc., Zool. vol. xxxi. p. 175, pl. xviii. figs. 24-26 
(1909). 


Specimens from (500) and (520), which are undoubtedly the 


524 MR. A. W. WATERS ON 


same as Busk’s ‘ Challenger’ 2. hirsuta, have besides minute cir- 
cular avicularia round the fenestra, also on the inner side of each 
of the zocecia bordering a fenestra, an elongate avicularium with 
the beak bifurcate but with an acute mandible. This was de- 
scribed by Busk for F. harsuta and is one of the most important 
characters. Large pieces were examined without finding any of 
the long antenniform spines described by Busk, and then in 
some places they were found in abundance. This is much like 
Rk. monilifera var. umbonata MacG., but in that there are only 
occasionally the long bifid avicularia on the zocecia bordering on 
the fenestra, and they must be separated because, among other 
reasons, the branches have more zoccia in A. wmbonata; the 
more delicate hirsuta is a tropical form, whereas wmbonata is 
Australian. 

The monilifera group of Retepora seems to be well defined, and 
perhaps should have generic value given to it. All have the 
well known “ trifoliate stigma” on the ovicell, and in most there 
is a minute avicularium on the lip to one side; the opercula 
generally are fairly similar with a nearly straight proximal edge, 
and in shape rather wider than long, with the muscular attach- 
ments rather high up and near the border. In &. formosa 
MacG. and R. contortuplicata B., the operculum is more curved 
on the proximal edge, and no labial avicularia have been found 
in these; apparently all have the labial pore, which is often the 
end of a long tube opening into the zocecium * proximally to the 
operculum, and I agree with Jullien and Calvet in speaking of 
this as the Reteporidan pore, as it is characteristic of a large 
number of Retepore. 

There is another group with widely open ovicells, and in these 
none have a labial avicularium, unless 2. incrustata Calv. belongs 
here; the proximal edge of the operculum is not straight, and in 
some it is contracted in the middle like the opercula of some 
Lepralia (for example L. hippopus Sm.). In this group are 
R. imperati B., R. elongata Smitt, LR. tessellata Hincks, R. solan- 
deria Risso, R. inordinata Calv., having a broad thickened border ; 
while &. sinwosa Kirkp., R. novezelandie Waters, R. hippocrepis 
Waters, R. gelida Waters, 2. lepralicides Waters, have long bands 
at the sides of the operculum, for the muscular attachments, and 
lateral contraction of the oral aperture. The Reteporidan pore 
has only been observed in this group in &. solanderia Risso. 

Another group has a fissure in the ovicell as in #. cellulosa L. 
and the proximal edge of the operculum is nearly straight, and 
very similar throughout this group, labial avicularia occur in 
some but not in all. There are a few Retepore with imperforate 
ovicells. Throughout the genus the oral glands are very well 
developed. 

The minute round or oral avicularia with thin membranous 


* Waters, “ Medit. & New Zeal. Reteporide,’’ Journ. Linn. Soc., Zool. vol. xxv. 
pp. 255-6; Expéd. Antarct. Belge, pl. vi. fig. 6 a (1904). 


BRYOZOA FROM ZANZIBAR, 525 8 


mandibles occur very frequently in the genus and may be called 
Reteporidan avicularia. There are other small avicularia with 
solid chitinous semicircular mandibles like those near the oral 
aperture. These have two muscular tendons, whereas the large 
triangular ones have a single tendon, and this muscular distine- 
tion between the round and triangular mandibles is very frequent 
in the Cheilostomata. 

In the monilifera group the growth of the ovicell can be well 
followed. Looking down the peristome the opening of a large 
pore is seen in the distal wall (Pl. LX_XJI. fig. 1) until the ovicell 
commences to grow (figs. 1 & 2), when the pore is more or less 
hidden by a disk growing on a small stalk (fig. 2). This disk 
becomes concave as it grows, showing this to be the commence- 
ment of the inner wall of the ovicell (fig. 3), then in a later 
stage part of the front is open (fig. 5), next the slit of the upper 
part of the trifoliate stigma is seen (fig. 6), and soon the ovicell 
is complete (fig. 7). 

More or less similar commencements of the ovicell occur in 
other genera, and sometimes specimens show these disks when 
no ovicells are developed. 

There is one large rosette-plate at the distal end of the 
zocecium, and the muscles for withdrawing the polypide are 
arranged on each side of this with a chitinous thickening to 
which they are attached. 

The avicularia are very variable in etepora, and we must not 
trust to them too much in determining species. 

Loc. Off Cape York, 8 fath. (Busk): Red Sea (Waters). Wasin, 
Brit. East Africa, 10 fath. (500), 20 fath. (520), collected by 
Crossland. 


Rerepora PropuctaA Busk. (PI. LX XII. figs. 9, 10.) 


tetepora producta Busk, Zool. Chall. Exp. vol. x. pt. xxx. 
p. 108, pl. xxv. fig. 7 (1884); MacGillivray, ‘‘ Tert. Polyzoa of 
Victoria,” Trans. Roy. Soc. of Victoria, vol. iv. p. 115 (1895) ; 
Thornely, Records of Indian Museum, vol. i. pt. 3, no. 13, p..193 
1907). 

Retepora porcellana, var. laxa MacGillivray, Prod. Nat. Hist. 
Victoria, dec. x. p. 15, pl. 95. fig. 6 (1885). 

The small semicircular avicularia are very much raised, which 
is not usualin Retepora, and are crenulate, as is frequently the 
case in the round avicularia of the genus. These avicularia do 
not occur on the younger zocecia but are common on the others. 
The zocecia are much smaller than those of 2. porcellana MacG. 
and the spines are ridged, but Iam not sure that it is right to 
speak of them as articulated. In the zowcia near the border 
there are often triangular avicularia with a single beak. The 
ovicells in the Wasin specimens are not very much raised and 
have a distinct median slit. On the dorsal surface there are 
long narrow avicularia; the meshes are large, but “ papillary 
eminences ” are not found as a frequent character. 


326 MR. A. W. WATERS ON 


Loc. Tongatabu (Polynesia), 18 fath. (Chall.); Samboangan, 
10 fath. (Chall.); Port Phillip Heads, Victoria (J/acG.); San Pedro 
Shoal, 25 fath.; Daros Island, Amirante Group (in my collection, 
sent to me under another name). Wasin, Brit. East Africa, 10 
fath. (507), 20 fath. (520), collected by Crossland. 

Fossil. Victoria, Tertiaries (MacG.). 


RETEPORA DENTICULATA Busk. (Pl. LX XII. fig. 8.) 


Retepora denticulata Busk, Zool. Chall. Exp. vol. x. p. 109, 
pl. xxvi. fig. 1 (1884); Philipps, “Rep. on Polyzoa,” Willey’s 
Zool. Results, p. iv. p. 449, pl. xliii. fig. 13 (1899); Calvet, “ Bry. 
@Amboine,” Rev. Suisse de Zool. vol. xiv. p. 620, pl. xxi. 
figs. 2-9 (1906); Thornely, ‘“‘ Mar. Poly. of the Indian Ocean,” 
Trans. Linn. Soc., Zool. vol. xv. p. 147 (1912). 


The lamina of the ovicell is not quite so long as that figured 
by Calvet, nor has it the orifice mentioned by him. Only a 
few ovicells have been found. The trabecule are not barren. 
There are large bifid avicularia on many of the zocecia near to 
the distal end, which are probably only the small labial avicularia 
modified ; also there are small semicircular or oval avicularia in 
the older zocecia, near the middle of the front surface, but in the 
younger zoccia they are wanting. Similar avicularia also occur 
in the ‘ Challenger ’ specimens. 

There are about 11 tentacles. 

Internally, at the proximal end on each side of the zocecium, 
there is a chitinous thickening (fig. 8) for the attachment of the 
muscles which withdraw the polypide, and between these two 
sets of retractors there is a thick funicular bundle passing to 
the next older zoecium. The interior wall of the zocecium is 
lined with very large square cells, and very similar cells line the 
zoecial and vibracular chambers and the tube-pores of some 
Cupularie, recent and fossil. 

Reteporella dendroides Ortmann is similar in many respects. 

Loc. Sandwich Is., 20-40 fath. (Calvet); Lifu, Loyalty Is. 
(Ph.); Amboina (Calvet); Amirante, 15-35 faths., Seychelles, 34 
fath. (7h.). Wasin, Brit. East Africa, 10 fath. (501), collected by 
Crossland. 


RETEPORA JERMANENSIS Waters. 

Waters, “ Mar. Biol. of the Sudanese Red Sea,” Journ. Linn. 
Soc., Zool. vol. xxxi. p. 176, pl. xviii. figs. 11-16. 

Loc. Red Sea (Waters). Prison Island, Zanzibar Channel (513), 
the shore, collected by Crossland. 


RErEPoRA TUBULATA Busk, var. 

Ketepora tubulata, type, Busk, Zool. Chall. Rep. vol. x. pt. xxx. 
p. 121, pl. xxviii. fig. 2 (1884); Thornely, “‘ Ceylon Pearl-Oyster 
Fisheries,” vol. iv. p. 125 (1905); Records of Indian Museum, 
vol. 1. p. 193 (1907). 


BRYOZOA FROM ZANZIBAR. 527 


The fureate avicularium at the proximal end of the fenestra 
is not so large or as much divided as in the ‘Challenger’ specimen, 
but this divided avicularium is a character of much use in dis- 
tinguishing the species from F. hirsuta B. and some others about 
the same size with various zoarial characters similar. There are 
no other large avicularia on the fenestral border, and there is a 
delicate spine at each side of the oralaperture. As Miss Thornely 
found in the specimen from the Indian Ocean, the growth some- 
times brings the celluliferous surface to the outside as in 
R. philippensis B. 

Ortmann has described a species as FR. axillaris which has 
an avicularium at the proximal end of the fenestra, but the 
shape of the avicularium and mandibles differs from the species 
from Zanzibar in which the avicularia are similar in shape to 
those of 2. hirsuta. 

There are 11 tentacles, and the oral glands are moderately 
large but not much elongated. 

Loc. Cape York, 8 fath. (Busk); Gulf of Manaar (7.); Gaspar 
Straits, Malacca Straits (7h.), var. Wasin, Brit. East Africa, 10 
fath. (501), Ras Osowamembe, Zanzibar Channel, 10 fath. (501), 
10-20 fath. (504, 514), collected by Crossland. 


? BIFAXARIA VAGANS Thornely. (PI. LXV. figs. 13, 14.) 


Thornely, ‘‘ Mar. Polyzoa of the Indian Ocean,” Trans. Linn. 
Soe., Zool. vol. xv. p. 145, pl. viii. fig. 7 (1912). 


Zoarium erect, cylindrical, about 0-3-0-4 mm. in diameter, 
branches dividing dichotomously at nearly right-angles, continuous 
in the younger portions, but in the older the junctions are broken 
across and there is a chitinous tube inside the articulation. 
Zocecia face opposite all round the stem in four directions; the 
surface of the zocecia has longitudinal ridges and large pits. The 
secondary orifice is round with a small mucro bearing a small 
avicularium, with an acute triangular mandible extending into 
it, while the primary orifice, which is some distance down the 
peristome, has the proximal edge straight. There are no ovicells 
on the specimens examined, which were growing on Retepora, 
nor were any ovaria or testes found in the sections made. There 
does not appear to be any bar to the avicularium. There are 
about 16 tentacles. 

This is much like the ‘ Challenger’ Lifaxaria reticulata * Busk, 
but in the British Museum specimens there are no avicularia, 
and the surface is reticulate and apparently not perforate. 

Bifaxaria of the ‘Challenger’ is not a satisfactory genus, and 
will be broken up. It might be retained for B. submaucronata B. 
and £. corrugata B., which are closely allied, and have the 
avicularia at the proximal end of the zocecia, not at the distal 
end as supposed by Busk. JB. denticulata B. has been removed 


* Busk, Zool. Chall. Exp. vol. x. pt. xxx. p. 82 (1884). 


528 MR. A. W. WATERS ON 


to Sclerodomus by Levinsen; B. papillata B. should perhaps be 
put to Urceolipora; B. abyssicola B. probably belongs to the 
Cellarinella group; B. levis B. must be placed elsewhere; B. 
minuta B. perhaps belongs to the B. reticulata B., B. vagans 
Thornely, group. 

Loc. Seychelles, 34-39 fath., Amirante, 34 fath. (Thornely). 
Wasin, Brit. East Africa (500); Ras Osowamembe, Zanzibar 
Channel (504), collected by Crossland. 


ADEONID4. 


In a short paper in the Annals and Mag. of Nat. Hist. for May 
1912*, I have given grounds for believing that the Adeonide 
will be found to be a much more important group than has 
generally been supposed, and this in spite of none of the characters 
upon which it was founded being of the importance then supposed. 
A large number of the species are pigmented, most have long 
pore-tubes, there are triangular avicularia on the front, and 
often vicarious avicularia on the zoarial border, and both kinds 
are without any cross-bar. There are no external ovicells, 
the embryos being developed in an ovicellular sac which about 
half, or, in some cases, nearly wholly fills up the zocecium ; and in 
a considerable number the zocecium in which the embryo is 
developed is larger than the others, with a differently shaped 
aperture, together with a difference in the frontal pores, and these 
larger zoccia have been called goneecia. The number of species, 
however, in which there is no difference in the ovicelligerous 
zocecia 1s very large, perhaps as many are indistinguishable as are 
externally recognizable. There are no oral glands in any species 
examined. I had previously shown that in the Adeonella of 
Busk’s ‘ Challenger’ Report there were species in which the pore 
entered into the zocecial chamber, while in others the pore is 
above the operculum, also that the opercula in some are nearly 
straight on the proximal border, others have a broad curve. At 
the time there was not sufficient material for complete generalisa- 
tion, but it is now clear that I was right in separating from 
Adeonella species with a pore entering into the zocecial chamber, 
and it is now seen that they also have a more or less straight edge 
to the operculum. This group is now known as Adeonellopsis of 
MacGillivray ; and Adeonella, much the same as I restricted it, is 
accepted. 

The wall of the sac (Pl. LX XIII. figs. 3, 5, s.) containing the 
embryo (figs. 3, 5, emb.) is much thicker than that of the sac in 
such species as Lepralia cucculata B., which also has no external 
ovicells. The very earliest stages of this sac have not been seen, 
and while it does not ever seem to hang from the opercular 
region as a small pendant, yet a comparison with the sacs of 


* Ser. 8, vol. ix. p. 493 (1912). 


BRYOZOA FROM ZANZIBAR. 529 


Diplodidymia complicata Rss. (text-fig. 79, p. 491) may throw 
some light upon the growth. 

Blind cells, that is cells without an oral aperture, are very 
frequent throughout the family, and sometimes the central or 
stalk portion is composed of such cells. A few have a closure 
somewhat like those of the Cyclostomata, but not entirely so, as 
a membrane from below the operculum spreads to the front to 
make the funnel-like closure. Membranous closures are also 
occasionally found, and these may be perforated. These closures 
with tubuli occur in Adeonella contorta Mich., and I have found 
them in Lepralia syringopora Rss., one of the Adeonide. 

The gonecia in Adeonide, when distinguishable, frequently 
are situated near the border, but in one species they are all 
situated on the median line, while in another they are in groups 
in various places, distinguishable by the naked eye. 

In my recent paper dealing with Adeonide I alluded (p. 496) 
to the occlusor muscles being, as a rule, single in triangular 
mandibles, whereas the semicircular mandibles usually have two 
tendons. A slightly different statement would have been better, 
as I was aware of some exceptions, and since writing I have come 
upon more—in fact in Cellepora and Holoporella many of the 
triangular mandibles have two tendons, but in Flustra, Membrani- 
pora, Retepora, and Cribrilina, etc., and in most genera the rule 
seems to be, only one tendon to the triangular mandibles. Some 
of the exceptions may be in semicircular avicularia modified 
into triangular ones. I have not got a record on this point of all 
the mandibles prepared, and must make further examination. 


ADEONELLA PLATALEA Busk. (Pl. LUXXIII. figs. 3-5, 8 & 
text-fig. 82.) 

Eschara platalea Busk, Brit. Mus. Cat. Mar. Pol. p. 90, pl. ev. 
figs. 1, 2 3, pl. eviii. fig. 4 (fide Busk) (1854); MacGillivray, 
Prod. Zool. Vict. dec. v. p. 41, pl. xlviil. fig. 4 (1880). 

Eschara hexagonalis Haswell, Proc. Linn. Soc. N.S. Wales, vol. v. 
p. 41, pl. ili. figs. 1, 2 (1881). 

Adeonella platalea Busk, Zool. Chall. Rep. vol. x. pt. xxx. 
p. 184, pl. xxi. figs. 4, 4a & fig. 50 (1884); Meissner, M., “ Liste 
der von Herrn Prof. Semon bei Amboina und Thursday Island 
gesammelten Bryozoen,” Jenaische Denk. vol. viii. p. 731 (1902) ; 
Waters, “ A Structure in Adeonella (Laminopora) contorta Mich.,” 
Ann. Mag. Nat. Hist. ser. 8, vol. ix. p. 489 (1912). 

Escharinella cecilleana VOrb. Pal. Frang. vol. v. p. 207 (1850) ; 
and see Waters, Ann. Mag. Nat. Hist. ser. 7, vol. xv. p. 5 
(1905). 

I have not had the opportunity of re-examining the Z. cecilleana 
d’Orb. since this collection was taken in hand, but the name 
cecilleana cannot be retained, as the description was insufficient. 

The ovicellular sac (fig. 3) is formed near the distal end 
and ultimately fills up nearly the whole of the zoccium, but 


530. MR. A. W. WATERS ON 


the very earliest stage has not been seen. The wall of the 
sac is formed of short cells and at one part touches the embryo. 
The embryo in early stages (fig. 5) is surrounded by large and 
long cells, and in some eases is filled up by formative tissue. In 
later sta ges, when the embryo is nearly ready for a free existence 
the large cells of the calotte are readily followed (fig. 6). 


Text-fig. 82. 


Fi iota 


ise 


Dy 
Ve ee 2s 


Lateral surface of Adeonella platalea. X 25 


When the ovicellular sac has grown to a moderate size there 
may be, near to the opercular wall, an ordinary polypide, but 
often there is only a very small one looking like a simple bud, 
being sometimes not more than a straight tube. There are strong 
muscles at the distal end on one side for contracting the walls of 
the ovicellular sac, to which they are attached, as well as to the 
distal wall. The ovieells of A, polystomella Rss., are smaller than 
those of the other species, but there is the same thick-walled 
ovicellular sac. Also, in a specimen. of Beania magellanica B., 
from Chatham Islands, there is a large embryo in a thick-walled 
cellular sac nearly filling the zoecium. This I do not find in any 
of my specimens from | Naples, but there is a pair of sac-like 
bodies near the distal end, and in the Antarctic specimens these 
bodies are large and were thought to be for the formation of 
testes, although there are ordinary testes near the distal end in 
which the spermatozoa can easily be seen. 

It has been supposed that all the large zocecia of the Adeonida, 
namely the goneecia, are merely for embryos, but this is by no 
means certain, as some may be for the testes, with which some are 
nearly filled, but the material available does not permit of my 
saying much on this point. However, it is only in a limited 
number of species that there are large zocecia, and the same changes 
may take place in these as in ordinary zocecia ; for as the gonecia 


BRYOZOA FROM ZANZIBAR. 53h 


contain polypides and buds, the embryos are only a stage in the 
life of the goneecia. 

In the present species the two outer rows of zocecia are, in parts 
of the colony, larger than the rest, forming goneecia with a larger 
and wider oral aperture, also the pore below it is wider, and a 
short tongue from the distal end sometimes divides the pore into 
two. In: many species of the Adeonidee where there are goncecia 
they occur at the border of the zoarium or near a Rigarestions but 
apparently in no species are they continuous all along the border. 
When the zoccia are blind or closed there is often a dise with a 
central tubule in about the position of the oral aperture. 

In A. platalea there are at the border of the zoarium, besides 
the large avicularia, very minute ones between the two zocecia, 
and sometimes the small chamber has a round opening and no 
mandible. 

Loc. Bass’s Straits; Cape Capricorn; Victoria, Queensland 
(MacG.) ; Port Darwin (Brit. Mus. Coll.); China Seas, 82-102 
fath. (Chall.); Thursday Island (Meissner); Ye, Burmah Coast 
(Thornely); Basilan, Indian Ocean (dOrb.). Wasin, Brit. E. 
Africa, 10 fath. (501,520); Ras Osowamembe, Zanzibar Channel, 
10 fath. (504); Prison Island, Zanzibar Channel, 8 fath. (505): 
collected by Crossland. 


ADEONELLOPSIS CROSSLANDI, sp. n. (Pl. LX XIII. figs. 1, 2, 6, 7, 
12-14.) 

This is, in many respects, like 4. swbsuleata Smitt, but there 
seems sufficient reason for separation, for the ridge surrounding 
the area in swbsulcata is here wanting and the suboral avicularium 
is directed laterally. 

The zoarium consists of flat branches (a little over 1 mm. wide 
and less than | mm. thick) frequently bifurcating and sometimes 
anastomosing. 

All the zocecia are about the same size, no difference being 
discoverable externally in the ovicelligerous zocecia, although in 
sections zocecia steele filled by the embryo are found occurring 
indifferently in any position except in the outer zocecia. The 
proximal border of the aperture is crenulate and this, as mentioned 
by Miss Thornely, is the case in subsulcata ; also the broad median 
pore is denticulate ; however, in some zocecia, especially those next 
to the outer row, there is no apparent pore. 

There is a small triangular avicularium immediately below or 
within the aperture, and often a few of the zoccia near a bifurea- 
tion, and some of the bordering zocecia have also an avicularium 
at the proximal edge. 

The zocecia at the sides of the zoarium are opposite as, in fact, 
are all the zocecia, so that the oral apertures on both sides rise to 
the same height. On the side of the zoarium there are no 
vicarious avicularia, whereas they occur in A. distoma, A. sub- 
sulcata, and A. imperforata. 


Proc. Zoou. Soc.—1913, No. XXXVI. 36 


532 MR. A. W. WATERS ON 


The tentacular sheath is attached to the operculum close to its 
border, and on the inner side of the opercular region there is a 
small globular body (fig. 2), immediately below which the new 
bud grows. We see the same thing in various species, among 
others, in ? Bifaxaria vagans Thorn., A. polystomella Rss., ete., 
and it is very large in Adeona foliacea var. fascialis Kirchenpauer, 
being about 0-4 mm. in diameter, which is about double the 
diameter of the globe in the present species. The wall is thick 
and it may give support to the new bud. 

There are about 13-14 tentacles. 4. distoma has 14-16. 

Tt would seem that the ovicellular zocecia very frequently occur 
near to a bifurcation, and there may be a group of three or four 
ovicellular zocecia close together. The few ovaria seen consist of 
two very small ovarian cells, at first situated at the distal end of 
the zocecium, but others, no doubt older, occur by the proximal 
end of the ovicellular sac. The ovaria are similar to those of 
Bugula, and are of about the same size in Adeonella platalea, 
A. polystomella*, and A. crosslandi. The early stages of develop- 
ment have not been made out. 

The ordinary zoccia may have the polypide entirely surrounded 
by a testis, in fact, may be practically filled with it. 

Loc. Wasin, Brit. East Africa, 10 fath. (500, 501), collected by 


Crossland. 
EXPLANATION OF THE PLATES. 


Prats LXIV. 


Fig. 1. Aetea anguina L. Showing the ovicell (ov.) at the end of the tubular pro- 
jection; the retractor muscles (7m.) attached just 
below the tentacles, and also attached at the other 
end near the commencement of the basal expansion, 
where the ovaria (oa.) also occur. X85. Fron 
Wasin (500). 

2. Do. do. Front view, showing the operculum and the tentacular 
sheath (¢s.) to which the muscles are attached. 250. 


3. Aetea truncata Landsborough. X25. From Prison Island (505). 

4. Brettia tropica, sp.. X85. Dorsal surface. From Wasin (501). 

5. Do. do. < 25. Anterior surface. 

6. Catenaria diaphana Busk. X25. From Ras Osowamembe (504). 

al Do. do. Lateral view, X 25. 

8 Do. do. Operculum, X 85. 

9 Do. do. X 25. Section showing embryos (/.) in the ovicell; 


also an ovum (ov.) in the zocecium, and ovaria below 
the ovicell (oa.). 

10. Do. do. Ovarium, X 250. 

11. Do. do. < 85. Section of zocecium showing ovicell (ov.), 
diaphragm (d.), compensation-sac (es.). 

12. Synnotum pembaensis, sp. n. X25. From Chaki-Chaki Bay, Pemba 
Island (517). 


118} Do. do. Mandible, X 250. 

14, Do. do. Avicularium, X 85. 

15. Do. do. x 3. 

16. Synnotum contorta, sp.n. X 25. From Chaki-Chaki (617). 
Wife Do. do. Avicularium, < 150. 

18. Do. do. Mandible, X 250. 


* A. polystomella occurs in the Red Sea, as I find from the examination of a 
specimen in the British Museum, since my Red Sea paper was written. There is 
also a specimen from Brishane (65.5.29.4). 


BRYOZOA FROM ZANZIBAR. 533 


PLATE LXV. 


Wig. 1. Vittaticella elegans Busk. XX 25. Showing large avicularium. From 
Prison Island, Zanzibar Channel (503). 


2. Do. do. Mandible of large avicularium, X 85. 

3), Do. do. Small avicularium, X 150. 

4. Do. do. Distal end, showing the avicularium, X 86. 

5: Do. do. Showing internodes composed of many zocecia with 
ovicells, X 15. (503). 

6. Do. do. Section of embryo im ovicell, X 400. 

7 Do. do. Section showing zowcia with polypides and ovicells 
(ov.) in position, X 85. 

8. Vittaticella elegans, var. zanzibariensis, nov. Distal end, showing the 


avicularium, X 85. From Wasin, Brit. KE. Africa 
(500). a, mandible, X 250. 


9. Do. do. do. x 50. Showing small avicularium on the 
dorsal surface near the distal end. 
10. Do. do. do. Upper half of the contents of the vitte, « 250. 
From Prison Island, Zanzibar Channel (505). 
ik Do. do. do. < 85. Showing internal zocecial muscles. 
(505.) 
12. Do. do. Semidiagrammatic transverse section through the 


vitte (v.), X about 200. 

13. 2 Bifaxaria vagans Thornely. X 25. From Wasin (500.) 

14, Do. do. Operculum, X 85. 

15. Chlidonia cordieri Aud. Section showing muscles (m.) from the parietal 
wall to the base of the chamber, X 250. From 
Wasin, Brit. E. Africa (500). 

16. Do. do. Operculum, X 250. a, mandible, X 85. 


Prats LXVI. 


Hig. 1. Stirparia exilis MacG. 25. Only the lower part of the tuft is shown. 
From Wasin, Brit. E. Africa (522), 20 fath. 


2. Do. do. x 2. 

3. Do. do. X 85. Stem showing disc from which a fresh branch 
grows. 

4, Stirparia dendrograpta, sp.n. X 25. Only the lower part of the tuft 


is shown, and for clearness ouly a few of the spines 
are drawn full length, but are shown as broken off, 
which is frequently the case. A pair of radicles are 
shown growing from the base of a node. From 
Chuaka, Zanzibar (508), 2 fath. 


5. Do. do. < 250. Avicularium. 

6. Do. do. x 25. First zowcium of a tuft, showing long spines. 

ie Do. do. < 85. Dorsal surface, showing the polypide in the 
zocecium with ovaria (ovar.). 

8. Do. do. < 250. Ovarium from below the polypide, with one 
ovarian cell. 

9. ote dos xX 2. On the left a few zocecia are shown in a natural 
position; the others are spread out as in a mounted 
slide. This piece is entirely unattached. 

Prats LXVII. 

Fig. 1. Cellaria wasinensis, sp. n. X25. Specimen prepared with Eau de 
Javelle, showing the openings of the ovicell in an 
enlarged part of the zoarium. From Wasin. 

2 Do. do. Mandible, & 85. 

3 Do. do. Showing the frontal membrane, with the trabecule and 
the tessellated area below the operculum, X 86. 

4. Do. do. Tessellated area, X 250. 

5 Do. do. Avicularium prepared with Eau de Javelle, showing 
the plate im the submandibular part. XX 8d. 

6 Do. do. x 3. 


7. Cellaria gracilis, var. tessellata, nov. ‘Tessellated area, X 250. From 
Ras Osowamembe. 
Figs. 8,9. Farcimia oculata Busk. Stalk growing from stolon, X 265. 
36% 


Fig. 


Fig. 


MR. A. W. WATERS ON 


Membranipora armata Haswell. Opercular region, seen from the interior. 
The opercular muscle is attached to a linear sclerite (sc.) on the mem- 
branous operculum, and from the sclerite there is also a muscular band 
to the tentacular sheath (¢.s.). Two protoplasmic bands (0.) pass to the 
distal rosette-plate. XX 50. 


11. Diplodidymia complicata Rss. X 6. Colony from Chuaka (506). 
12. Do. do. X 25. Lower part of colony. 
13. Do. do. x 85. 
14. Do. do. < 250. Operculum. 
15. Do. do. X 250. Mandible. 
Pratt LXVIII. 
1. Stirparia zanzibariensis, sp. n. X 85. From Chuaka, Zanzibar, 8 fath. 


(508). 
Do. do. <2: 
Serupocellaria pilosa Aud. X85. Dorsal surface. From Wasin, Brit. 
KE. Africa, 10 fath. (500). 
Do. do. X25. Anterior surface. 
. Scrupocellaria macandrei Busk. X85. Dorsal surface, showing the 
groove extending beyond the vibracular chamber. 
From Prison Island, Zanzibar Channel (505). 
Do. do. < 85. Anterior surface. 
. Bicellaria chuakensis, sp.n. Avicularium, « 250. 
Do. do. < 25. From Chuaka Bay, Zanzibar Channel, (515). 
. Scrupocellaria wasinensis, sp.n. X25. Dorsal surface. From Wasin,. 
Brit. E. Africa, 10 fath. (522). 
Do. do. X 26. Anterior surface. 
. Scrupocellaria ferov Busk. X25. Anterior surface. From Prison 
Island, Zanzibar Channel, 8 fath. (805). 


12. Do. do. Mandibles—(a) anterior, (4) lateral, X 85. 
13. Do. do. Base of vibracular seta, * 250. 
14. Do. do. Decalcified piece, showing the articulation with the 
chitinous tube already formed to the inside zoecia 
of the new branch, X 25. 
16 Do. do. Dorsal surface, & 25. 
PratE LXIX. 
1. Canda retiformis Pourt. X25. From Ras Osowamembe, (504). 
2. Do. Do. Dorsal surface, X 28. 
3. Serupocellaria cervicornis Busk. X 25. From Wasin (522). 
4, Do. do. Dorsal surface, showing the articulation, < 25. 
5. Caberea ellisii Flem. Vibracular chamber seen from the inside: a, },. 


vibracular chamber; 4, ¢, continuation of the groove beyond the 


chamber. 
6. Canda retiformis Pourt. Dorsal surface, to show the articulation. X about 6. 
7. Scrupocellaria ferox B. Do. do. _X about 6. 
8. Scrupocellaria jolloisii Aud. Do. do. X about 6. 
9. Flabellaris (Menipea) cuspidata B. Do. do. X about 6. 
10. Bugulopsis peachii Busk. Do. do. X about 12. 
11. Menipea patagonica B. Do. do. X about 12. 
12. Menipea buskii W. Th. Do. do. X about 25. 
13. ? Menipea cyathus W. Th. Do do X about 25. 


. Stirparia zanzibariensis, sp.n. Showing basal stalk from which there are 
delicate rootlets, and also capsules nearly filled with a bright yellow 
substance. XX 12. 

. Bugula robusta MacG. Showing the earlier zocecia, X 25; a, natural 

size. From Wasin (501). 


16. do. do. Avicularium, X 8d. 

17. Serupocellaria wasinensis, sp. n. Zocecia showing the position of the 
ovaria (ov.), testes (¢.), rosette-plates (7.). > 85. 

18. Do. do. Ovarian cells with protoplasmic threads from the 
rosette-plates, K 250. 

19. Do. do. Ovaria showing two nucleated ovarian cells, X 250. 

20. Serupocellaria ferox Busk. Band which starts near the distal end and 


passes down the side of the zocecium; also small bundle of protoplasmic: 
threads running parallel with the granular band. 250. 


BRYOZOA FROM ZANZIBAR. 535 


PLATE LXX. 


Fig. 1. Schizoporella nivea Busk. X25. From Wasin (501). This may perhaps 
be called var. wasinensis. 


2. Do. do. x 85. Operculum. 
3. Do. do. X 85. Oral aperture closed by calcareous wall. 
4. Gemellipora protusa Thornely. X 25. From Wasin (520). 
5. Do. do. X< 85. Operculum. 
6. Do. do. < 85. Interior of the zoccium showing the oral 
aperture. 
7. Schizoporella nivea Busk. X 25. From Wasin (501). 
8. Do. do. X 150. Basal avicularium. 
9: Do. do. X< 150. Lateral avicularium. 
10.8 Petralia chuakensis Busk. > 25. From Chuaka, Zanzibar. 
Make Do. do. x 85. Operculum. 
12. Do. do. x 85. Mandible. 
13. Do. do. < 250. Mandible showing the crenulated border. 
14, Do. do. X< 25. Dorsal surface showing spaces for the attach- 


ment of radicle tubes. 
15. Petralia vultur, var. armata, nov. X 85. Section through the zocecium, 
showing the tentacles, with two larger than the 


others. 
16. Do. do. do. < 85. Operculum. 
AWE Do. do. do. < 550. Section of tentacles near the base 


showing the two large fan-shaped tentacles with 
long nuclei near the edge. 


18. Do. do. do. x 25. Eau de Javelle preparation. From 
Chuaka. 

19. Do. do. do. > 85. Mandible of oral avicularium. 

20. Do. do. do. > 85. Mandible of round avicularium. 


21. Lepralia feegensis Busk. X 85. Operculum. 
22. Do. do. X85. Mandible. 


Prate LXXI. 


Fig. 1. Membranipora savartii Aud. Zoccia showing the network of protoplasmic 
threads. In the upper zocecium histolysis of the 
polypide has commenced, and in the right hand 
zocecium (7) histolysis has progressed further, and 
now considerable granular masses are formed. In 
the lower zocecium, of which only the distai end is 
shown, the polypide is in full vigour with digestion 
active and the tentacles unaltered. Drawn trom a 
decalcified mount, < 85. From Zanzibar Channel 


(528). 

2 Do. do. Section showing plasma threads by the side of finely 
granular cord and parts of narrower cords, X 1000. 

3 Do. do. The granular masses shown in the right zocectum in 
fig. 1, X 150. 

4. Do. do. A similar mass shown in transverse sections sur- 
rounded by protoplasmic threads, x 1000. 

5. ? Membranipora armata Haswell. Mandible, X 85. 

6 Do. do. Decalcified avicularium showing retractor muscle (a), 


divaricator (6), with tendon attached to the mandible 
in the middle of the base, and the “peculiar body” 


(ec), X 85. 
Uke Do. do. Avicularium, X 85. 
8. Do do. Lower portion of lateral gland, X 320. 
9. Do. do. Decalcified zocecium showing the avicularian chamber 


on the left with muscles as in fig. 6, and the glan- 
dular chamber (ge.) on the right, X 86. 
10. Do. do. X 25. From Wasin, Brit. East Africa (500). 


536 


ON BRYOZOA FROM ZANZIBAR. 


Puate LXXII. 


Fig. 1. Retepora monilifera, var. umbonata MacG. X 25. Showing in the lower 


9 
10. 


1 


Do. 
Do. 


Do. 
Do. 


Do. 
Do. 


do. 
do. 


do. 
do. 


do. 
do. 


zoccia the large pore within the oral aperture, and 
in the upper zoccia the stalked dise which grows in 
front of this pore. From Victoria. 

x 85. Aperture showing the stalked disc. 

x 50. The disc is shown larger and can now be 
recognised as the commencement of the wall of the 
ovicell. 

X 50. The same rather larger. 

x 50. The sides of the ovicell are commencing to. 
grow, but the ovicell is entirely open in front: 

x 50. The ovicell showing a large cruciform opening.. 

x 50. The ovicell shows the complete trifoliate- 
stigma. 


. Retepora denticulata Busk. X 250. Section of the proximal end of the- 


zocecium showing the muscles (m.) attached to the base of the polypide 
and to the wall of the zocecium, which has a chitinous thickening where 
the muscle is attached. Between the two sets of muscles a thick funi- 
cular cord (f.) passes through a large rosette-plate. 


Do. 


. Retepora producta Busk. X 25. From Wasin (520). 
do. 


xX 50. Ovicell. 


. Rhynchozoon profundum, var. laminatum nov. Diagrammatic figure of 
the aperture and ovicell. 
. Steganoporella magnilabris Busk. X about 50. Showing the tentacular 


8}, Do. 
14. Do. 
15. Do. 
16. Do. 
Y/, Do. 
18. Do. 
19. Do. 
20. Do. 


do. 
do. 


do. 


do. 


do. 


do. 
do. 


do. 


parts attached to the side wall of the proximal 
corner, while the stomach, etc., is in the other corner 
connected by a long narrow cesophageal tube. 

x 50. Operculum of B zoccium. 

x 50. Decalcified distal end of the zocecium as seen. 
from the front with the polypide seen through the. 
wall and (se.) sclerites to which the frontal muscles: 
are attached. 

X 250. Section of the edge of the operculum. 

X 85. Section of the distal wall through two rosette- 
plates. Probably the fleshy parts belong in part to: 
the distal and in part to the proximal wall. 

X 50. Section cut through an operculum showing a 
membrane (m.) between the main sclerites. 

x 50. Operculum of A zoccium. 

< 85. Decalcified distal end seen from inside with 
the basal wall removed. The tentacular sheath (é.s.); 
is held by four bands (6.) attached to the distal wall, 
also there are muscles (¢.s.m.) from the distal end of 
the tentacular sheath to the wall (a.) which divides 
the zowcium. The large retractor muscles (7.) are 
seen attached to a tendon, while below there is a 
smaller pair of muscles (d.) not attached direct to, 
the operculum, but to the frontal wall quite close 
to the operculum. 

X 25. Growing end with the membrane covering the 
surface. The end zocecia are as yet quite empty 
and the operculum of a B zocecium (B.) is growing, 
but no basal sclerite is yet formed. In the second 
row the frontal calcareous wall only extends a slight 
way into the zoccium and here the zoccium is still 
quite empty. Next the short tube (¢.) connecting 
the two parts of the zocecium grows and the lip 
commences, but is still some distance from the 
frontal membrane, though later it rises right up to: 
the proximal edge of the operculum. The frontal 
sclerite (sc.) to which the opesial muscle is attached 
does not occur in the younger zocecia, but is seen im 
the mature zocecia. 


Fig. 1 


Or 


a 
) 


~ 


<6 OO NI 


ON TWO AFRICAN ANTELOPES. DOT 


Prare LXXIII. 


. Adeonellopsis crosslandi, sp. n. X 250. Showing an ovarium in the 


distal end of the zocecium. From Wasin. 
X 250. Showing round body under the operculum 
with the protoplasmic threads to the buds below. 


. Adeonella platalea Busk. X 150. Showing the early stage of the ovicel- 


lular sac (s.) near the distal end containing the 
embryo (emb.). From Wasin (501). 

X 85. Section showing the ovicellular zowecium nearly 
filled up by the embryo, with an ovarium (ovar.) 
now proximal to the embryo. From Wasin (501). 

< 150. Section of embryo showing the zocecial wall 
(z.w.), the ovicellular sac (s.), and the embryo 
(emb.). The muscles are seen at the distal end on 
the right in the zocwecial chamber. From Wasin (501). 


. Adeonellopsis crosslandi, sp. n. X 250. Thick section of embryo 


showing the aboral end. Fig.6a, 250. Diagram- 
matic outline of vertical section. 
< 85. Opereulum. 


. Adeonella platalea Busk. X85. Operculum. 
. Osthimosia zanzibariensis, sp.n. X 85. Operculum. 


10. Lepralia turrita Sm. X 85. Operculum. 


TL 
12 


14, 


. Holoporella albirostris Sm. X 85. Polypide showing the long oral 


glands. 


. Adeonellopsis crosslandi, sp.n. X 25. 
13. 


x 50. Transverse decalcified section showing the 
opposite zocecia at about the same stage of develop- 
ment. 

Natural size. 


15. Osthimosia zanzibariensis, sp.n. X 50. 
16. Schizoporella nivea Busk. X50. The two zoccia figured were near 


together but not in the same row, and are placed 
together to save space. A sac-like structure (sc.) is 
shown at the base of each ovicell (oc.), but of the 
numerous muscles in this sac only about half of 
those in focus are drawn. It seems that the ovum 
(ov.) passes into this sac which is then ruptured (7.), 
and the ovum is pressed forward into the ovicell 
where it segments. The small oral glands are 
shown (g/.). 


35. Notes on Albinism in the Common Reedbuck (Cerw- 
capra arundinum), and on the Habits and Geographical 
Distribution of Sharpe’s Steenbuck (Raphiceros sharpez). 
By Major J. StevENson-Haminton, C.M.Z.S., Game- 
Warden of the Transvaal. 


[Received April 8, 1913: Read April 22, 1913.] 


INDEX. Page 
(CAP UORLY DTH GHROKDCIODTID, "53 oon soutien ves 2o0ane oob oan esdnsoson SEH 
IMO YOOGUROS SUCH VED ogi aroaae neddossanoadaxscousuee soxdu0%90 bon AY) 


Albinism in the Common Reedbuck (Cervicapra arundinum). 


During the early weeks of 1911 a rumour reached me of three 
white Reedbucks having been seen near the south-western border 
of the Sabi Game Reserve. I gave orders that they should be 


538 MAJOR J. STEVENSON-HAMILTON ON 


kept under observation so far as possible, though the very broken 
and hilly nature of that area rendered any close following of 
their movements impracticable. 

A month or two later a troop of hunting dogs (Lycaon pictus) 
passed through that part of the country and, it is believed, killed 
one of the animals, as only two were afterwards seen. I then 
made preparations for an expedition to capture the survivors, if 
possible, for the Pretoria Zoological Gardens, but almost imme- 
diately afterwards heard that they had wandered out of the 
Game Reserve and that the male had been shot by some unknown 
person. Towards the end of 1911, the last survivor, a female, 
came back to the Reserve. The Museum authorities having 
expressed a desire to obtain the skin, failing success in capturing 
the animal alive, I proceeded to the ground. Under the circum- 
stances it seemed that the capture would be very difficult, and 
that failure would almost certainly drive the animal out of the 
Reserve, where it would be killed at once by natives or white 
men, and so be irretrievably lost. I therefore shot it, and 
sent the skin, bones, and measurements to the Pretoria Museum, 
where it is now set up. 

The animal proved to be a female about two years old, and of 
a perfectly pure white colour throughout. The muzzle and inside 
of the ears, membrane round the eyes, ete., were pinkish or flesh- 
colour. 

The skin on the back of the ears was scabby and unhealthy- 
looking. Eyelashes white. Pupils reddish black. Inner irides 
grey blue. Outer irides pearly with darker rays. White of eyes 
normal. Hoofs and lateral hoofs pale horn colour, the former 
nearly white at bases. 

Measurements for a two year old animal about normal. 

Head and body 52 inches; tail 73 ; fore girth 32. 

When found she was running with an ordinary ram, and 
contained a perfectly formed male feetus, which was normal in 
all respects. I should be inclined to think, in view of the time 
which had elapsed since the disappearance of the male albino, 
that she had been impregnated by the normal ram with which she 
was found ; but this isa point I cannot speak on with any certainty. 

It is worth recording that within a mile of the spot where I 
saw this animal, and in the midst of the comparatively small 
area where these Reedbucks had been born and bred, was a 
village in which lived an albino male native, his hair nearly 
white, and his eyes of almost the same curious light grey-blue 
shade as those of the Reedbuck. Native information states that 
albino children have been born before in the same village, and 
that white Reedbucks have also been seen in former years in the 
district. ; 

Some two months later (January 1912) a report came to hand 
of another albino Reedbuck at a point 60 or 70 miles north of 
where the female was secured. ‘This animal was always noticed 
alone, and was repeatedly seen by passengers by the Selati train. 


TWO AFRICAN AN'LELOPES. 539 


T sent a ranger up to endeavour to capture it alive, but attempts 
were unsuccessful, owing to the mesh of our net being too large. 
It was seen close enough, however, to be recognized as a female, 
nearly pure white and about one year old. After the unsuc- 
cessful attempts to capture her alive, she was noticed at intervals 
for another month ; but before another attempt could be made, 
she disappeared, probably killed by wild dogs. 

These animals, which may have been born in 1909 in the first 
ease and 1910 in thesecond, were found at a considerable distance 
north and south of one another, and the incidents almost certainly 
had no connection whatever, but both occurred within a few miles 
of or among the foothills of the Drakensberg Mountains, on the 
western border of the Reserve. In nearly eleven years’ caretul 
observation of Reedbucks in the Reserve, these are the only 
eases of albinism that have been brought to my notice. 


Sharpe's Steenbuck (Raphiceros sharpei). 


Spiti-pite or Pitsi-pitsi of the Thonga tribes of the North- 
east Transvaal and neighbouring Portuguese East Africa. 
Isigulane of the Swazis. 


(reographical Distribution. —The most northerly habitat of the 
species appears to be British Nyasaland, where it was discovered 
and recorded by Sir A. Sharpe. Thence it extends down through 
Mashonaland, possibly hugging the vicinity of the eastern hilly 
country, to the North-eastern Transvaal. It is there found all 
along the course of the Lebombo Hills (but never in the Drakens- 
berg Mountains, sixty miles further west) as far as the Crocodile 
River at Komati Poort. It becomes very numerous between the 
Limpopo and Letaba Rivers, and is there spread through broken 
ground to as much as thirty miles from the Lebombo. South of 
the Olifants it becomes progressively scarcer, and is very rare 
indeed between the Sabi and Crocodile Rivers. It reappears, 
however, in Swaziland (Mr. R.'T. Coryndon), and is found all 
along the border of that country and Portuguese East Africa on 
both sides of the Lebombo. Mr. Coryndon is of the opinion that 
it occurs also in Northern Zululand—that is to say, as far as the 
end of the Lebombo Hills. 

Whatever be the case in Nyasaland and Mashonaland, in the 
Transvaal and Swaziland it occurs only in one narrow strip com- 
prising the Lebombo Hills and their immediate surroundings. 

No antelope answering to the description seems to be reported 
from Natal or Northern ‘Pondoland, and in the south of the latter 
we arrive at the extreme known northern limit of the Grysbuck 
(2. melanotis), at Port St. John’s. 

The geographical distribution of Sharpe’s Steenbuck, therefore, 
seems to be between 14° and 28° south latitude, and between 
about 312° and 332° east longitude, the line of distribution 
following more or less the south- -westerly trend of the coast, 
though at some distance from it. 


540 MAJOR J. SLEVENSON-HAMILION ON 


There seems to be a gap of 4° of latitude in which no small 
grizzled antelopes are found between the most southerly habitat 
of Sharpe’s Steenbuck and the most northerly of the Grysbuck 
(32° south). 

Description.—The animal found in the North-eastern Trans- 
vaal bas been described from specimens obtained by Mr. Grant 
(collector for Mr. Rudd). Females and immature males are, 
generally speaking, of a brighter red colour than old males. 
The distribution and profuseness of the white in the coats tends 
to vary in different specimens, and some show more than others. 
Individuals display small patches of white hairs here and there 
on back or sides. 

In a half-grown female the dark horseshoe mark on the crown 
was found to continue backwards in the form of a narrowing dark 
brown line as far as the centre of the back, where it suddenly 
broadened and then disappeared. In the same animal the face- 
markings were much more highly contrasted than in older 
specimens. 

There are four mamme present in the females. 

I fancied that I detected a very slight thickening of the tissues 
at the points where the lateral hoofs are found in other species. 

The horns of the males are very short and rather conical in 
shape. They seldom exceed an inch and a half in length, and 
I should imagine two inches to be about the maximum length. 

Mr. Selous has recorded shooting certain small grizzled animals 
in the Mashonaland Hills which possessed horns over three inches 
long, and this seems to point, perhaps, to an interesting variation 
of the type in that country. 

Habits, etc—Sharpe’s Steenbuck is found either in patches of 
thick bush, which may be at a considerable distance from any 
hilly country, or among the stones and boulders of the lesser 
ridges of the Lebombo Hills and the thick covert sometimes 
growing at their bases (North-eastern Transvaal). In no case has 
it been noticed at any very great distance from water. In the hill- 
country it especially favours the rough boulders interspersed with 
rank grass and bushes, crowning the crests of long stony ridges 
which gradually merge into grassy slopes on either side. Its 
colour almost exactly matches the red tint of the Lebombo rocks, 
and consequently, when it chooses to lie close, it is a most difficult 
creature to see. One or more individuals have a custom of 
returning to the same spot to deposit their droppings, but the 
habit is not a constant one. It feeds at night or in the very late 
evenings and early mornings, and, except on dull days, always 
lies up for the daylight hours among boulders or long grass. 
In cloudy weather, especially in spring, when the grass is young 
and fresh, it may be observed quite late in the day feeding close 
to one of its refuges, moving slowly about, and lying down at 
frequent intervals. In the stomachs examined, I have found 
about half the contents to consist of young grass and the rest of 
the small leaves of ground-shrubs. 


TWO AFRICAN ANTELOPES. 541 


When disturbed while lying up for the day, Sharpe’s Steenbuck 
usually springs up 30 or 40 yards away, and makes off at once. 
Occasionally, however, when well concealed, it lies very close, 
and only gets up whenapproached within a few yards. In no case 
that I have observed did it ever pause before going, resembling 
many other small buck and hares in this respect. It always goes 
at best pace for a considerable distance, squatting again suddenly 
when it has found suitable covert. Its gait isa scuttling run, and 
it never bounds like a Steenbuck or a Duiker. Nevertheless, its 
speed is considerable, and, in the rough country affected, it takes 
a very good dog to run down an adult of either sex. 

It is very solitary in habit, and even when a pair are put out. 
of the same patch of bush, they seem generally to have been lying 
in different parts of it. Bush and rocks seem to be regarded 
equally as natural refuges, and in following up individuals I 
could not discover any preference for one over the other. Even 
in the heat of the day the shade of a large stone on a hillside, 
destitute of a blade of grass or a scrap of bush, is sometimes 
sufficient for the animal’s requirements. 

Most of the females appear to be in lamb(North-eastern Trans- 
vaal) in October and November, and I imagine the bulk of the 
young to be born in the early or mid summer months, though I 
should hesitate to affirm that they do not, like many other small 
buck, breed more or less all the year round. 

From observation of appearance and habits in the field it would 
be impossible to recognize its kinship with the true Steenbuck. 
In habit it approximates much more closely to the Grysbuck, for 
which it used often to be mistaken by hunters. The native 
(Thonga) name is Spiti-pite or Pitsi-pitsi for Sharpe’s Steenbuck, 
while they call the Steenbuck Jnginana and the Livingstone 
Antelope Jnhlengana. 

Mr. R. T. Coryndon, the Resident Commissioner of Swaziland, 
confirms these observations of the animal’s habits in the North- 
eastern Transvaal as being applicable to what he has himself 
observed in Swaziland, and gives the Swazi name for it as 
Isigulane, for the Steenbuck /ngcina, and for the Livingstone 
Antelope Znhlengana. The Zulus call the common Steenbuck 
Iqina. The natives therefore recognize, judging as they do 
merely from habits and outward appearance, no affinity between 
the two species of Steenbuck. 

Sharpe’s Steenbuck is generally spoken of by Colonists in South 
Africa as “Grysbuck” or “ Grys Steenbuck,” and in this Province 
it received its proper title for the first time in the Game Laws. 
of 1912. 


542 MR. H. J. ELWES ON THE 


EXHIBITIONS AND NOTICES. 
April 8, 1913. 
Sir Jouy Ross Braprorp, K.C.M.G., M.D., D.Sc., F.B.S., 


Vice-President, in the Chair. 


THe Secrerary read the following report on the additions that 
had been made to the Society’s Menagerie during the month of 
Wlevela JUSS} 3 

The number of registered additions to the Society’s Menagerie 
during the month of March last was 133. Of these 45 were 
acquired by presentation, 58 by purchase, 11 were received on 
deposit, 6 in exchange, and 13 were born in the Gardens. 

The number of departures during the same period, by deaths 
and removals, was 167. 

Amongst the additions special attention may be directed to :— 

1 Kandt’s Cereopitheque (Cercepithecus kandti), from Lake 
Kivu, new to the Collection, deposited on March 7th. 

2 Canadian Poreupines (Hrithizon dersatus), from North 
America. presented by W. O. Danckwerts, Esq., K.C., F.Z.S., 
on March 19th. 

1 Kordofan Giraffe (Giraffa camelopardalis antiquorum) 3, 
born in the Menagerie on March 13th. 

1 Greenland Falcon (Hierofalco candicans), captured in Mid- 
Atlantic, and presented by Lt.-Col. F. B. Drage, R.H.G., and 
the Hon. John Hubert Ward, C.V.O., on March 30th. 

2 Purple Kaleege Pheasants (Genneus horsfieldi), from the 
Himalayas, presented by the Marquess of Tavistock, F.Z.S., on 
March 3rd. 

2 Keysser’s Cassowaries (Casuarius keysseri), from German 
New Guinea, received on deposit in immature plumage last 
September, but identified as new to the Collection on March 
12th. 

4 Banded Trichogaster (7'richogaster fasciatus), from India, new 
to the Collection, purchased on March 14th. 

1 Central-African Mud-fish (Protopterus cthiopicus), from 
Uganda, new to the Collection, presented by C. W. Woodhouse, 
Esq., on March Ist. 

2 Gibbous Scorpions (Buthus gibbosus), from Sardis, Asia Minor, 
new to the Collection, presented by Miss Flora Russell on 
March 11th. 


Mr. H. J. Exwes, F.R.S., F.Z.S., exhibited the head of an 
Asiatic Wapiti (text-figs. 83 & 84), remarkable for the outward 
extension of the bay or bez tines, and made the following 
remarks :— 

“The antlers I exhibit were bought in Moscow on my return 
from Formosa in May 1912 of Mr. Lorenz, who informed me that 


ASIATIC WAPITT. 943 


they had come direct from the Sayansk Mountains, east of the 
Upper Yenesei River. The only original account I know of the 
Deer of this region, though no doubt more recent information 
exists in Russian, is by Radde in ‘ Reisen im Siiden yon Ost- 
Sibirien,’ St. Petersburg 1862, vol. i. p. 284. Radde considered 
the species to be C. elaphus, and says that it was found up to 


Text-fig. 83. 


Front view of antlers of Asiatic Wapiti from the Sayansk Mountains. 


and above tree-level on the Sayansk, Baikal, Apfel, and Chingan 
Mountains, frequenting the thickest forests and islands on the 
Amur, but wandering in summer as high up as the glacier 
of Munku-Sardik and over the bare peaks of the Sayansk 
Mountains. 


544 ON THE ASTATIC WAPITT. 


es 


Semi-profile view of antlers of Asiatic Wapiti. 


MR. R. H. BURNE ON MALFORMED ANTLERS. 545 


“He gives the rutting cry, as imitated by the native hunters 
with a hollow stem of Calisace daurica, as follows, which seems to 
me to have some resemblance to that of the American Wapiti :— 


“In a paper read by me at the Linnean Society on Dee. 15, 
1898*, I spoke of some horns from the Yenesei Valley in the 
St. Petersburg Museum, which [ thought had more resemblance 
to those of Cervus elaphus, having a distinct cup or crown 
of 6 or 7 tines branching from the same point on the beam, as 
sometimes seen in old Kuropean specimens of C. elaphus, but 
never, so far as I know, in any other Asiatic form of Wapiti. 
But I have not been able to examine them recently; and the 
horns shown to-night belong, without doubt, to a race of the 
Asiatic Wapiti known as C’.. eustephanus Blanford, which name I 
think preferable to C. canadensis, var. siberica Severtzoft, though 
the latter has priority. But until the races of Asiatic Wapiti 
are better known, I think it is premature to name them definitely, 
and it cannot be done without careful examination of specimens 
in the St. Petersburg Museum. 

“The bez or bay antlers in my specimen (text-fig. 83) are 
clearly abnormal in their position. The development of the back 
tines, though clearly of Wapiti type, is also poor.” 


April 22, 1913. 
EH. T. Newron, Esq., F.R.S., in the Chair. 


Mr. R. H. Burne, M.A., F.Z.8., exhibited two pairs of 
malformed antlers of an Axis Deer (Cervus axis), lately presented 
to the Museum of the Royal College of Surgeons by Capt. Stanley 
S. Flower, F.Z.8. The deer was born in the Giza Zoological Gardens 
in 1899, the malformed antlers being shed in 1905 and 1906. The 
second pair showed a common malformation, i. é. duplicity of the 
brow tine, but the first pair suggested rather an injury during 
growth than a congenital malformation, Ata similar point in 
each antler the beam was abruptly bent inwards upon itself at a 
very acute angle. The apical tines also were stunted, probably 
owing to an interference with the blood-supply brought about by 
the sharp bend in the beam. 


* Journ. Linn. Soc., Zool. xxvii. p. 23. 


546 DR. R. BROOM ON 


Mr. E. G. Boutencer, F.Z.S., Curator of Reptiles, exhibited a 
number of living specimens of the Leaf-Insect (Phyllium cruri- 
folium), presented to the Society by Dr. Alfred Russell, which 
had been reared from eggs laid in captivity, and which showed 

various stages of development. 


May 6, 1913. 


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


Mr. E. G. Boutencer, F.Z.5S., Curator of Reptiles, exhibited a 
living melanistic specimen of the Green Lizard (Lacerta viridis) 
recently received from Dalmatia. 


A new Species of Golden Mole.* 


Dr. R. Broom, C.M.Z.S., exhibited an example of a new species 
of Golden Mole from the Transvaal, of which he gave the following 
account :— 

‘“‘ For a couple of years I have known that a species of Golden 
Mole occurred near Johannesburg in the Transvaal, but until a 
few weeks ago the only specimen I had seen was a badly stuffed 
skin without the skull. Recently I found that two specimens 
had been ploughed out on the farm Vischkuil, near Springs, and 
had been skinned by a native. Fortunately the complete carcase 
of one was still to be had, and most of the skin. On examining 
the skull I found to my great surprise that the Mole is nota 
typical Chrysochloris, but belongs to the subgenus Bematiscus, 
hitherto only known from Natal and EKastern Cape Colony. The 
species is a near ally of Lematiscus villosus Smith, one of the 
least known South African forms. 

‘““So few specimens of BL. villosus are known that we do not at 
present know the degree of variability of the species, and thus 
cannot be quite sure whether the few specimens at present placed 
under that specific name really all belong to one species. The 
type specimen described by Smith is in the British Museum. It 
is nearly full grown, and is stated to have come from Natal. The 
underfur is of ver y fine texture, and of a slaty-grey colour. 
The long hairs, which are compar atively few in number, have the 
outer, flattened portion 10 to 15 mm. in length and of a pale 
greyish-brown tint, rather darker towards the tips. The fewness 
of the bristly hairs and their length give the fur a rough harsh 
feeling. 


* [The complete account of this new species appears here; but since the name 
and a preliminary diagnosis were published in the ‘ Abstract,’ ‘it is distinguished by 
being underlined.—Ep1ror. | 


A NEW GOLDEN MOLE, DAT 


“The skull of Smith’s type is well figured by Dobson, and 
though barely mature is sufficiently near full size for purposes of 
comparison, 

** Dobson described and figured a second specimen which he 
believed to be Chrysochloris villosa, but he recognised differences 
which he thought might be specific. When working at the 
Golden Moles some years ago, I thought it well to keep the 
varieties of 6. villosus under the one species, though I called 
attention to the fact of one of the specimens in the Maritzburg 
Museum having a dark slaty, almost black fur. The specimens 
are so few in number—there being, so far as I am aware, only 
five skins in the museums of the world—that one hesitates to split 
up the Natal supposed B. villosus specimens into two species, but 
it seems likely that when more specimens are obtained at least 
two subspecies and possibly even species will be recognised. 

““ Whatever be the case with the Natal specimens, there is no 
doubt that the present Transvaal specimen is not Smith’s Chryso- 
chloris villosa, and almost as certainly it is not the same as 
Dobson’s type. 

“The Transvaal specimen, for which I propose the name 


BEMATISCUS TRANSVAALENSIS, 


Abstract P. Z.S. 1913, p. 25 (May 13), 


has a thick soft fur very unlike that of Smith’s B. villosus. The 
underfur is much shorter and less woolly, and the terminal 
flattened portion of the hair is less bristly and very much shorter, 
being only about 7 or 8 mm. in length, instead of 10 to 15 mm. as 
in B. villosus. The terminal portion of the hair isa rich reddish- 
brown, becoming very dark towards the tip. The whole dorsal 
side of the animal has thus a reddish-brown tint. On the 
abdominal surface the reddish tinge has almost disappeared and 
the fur has a slaty tint. The fur from the sides of the head 
and nose has been destroyed. 

“The claws of the manus of &. transvaalensis measure: 
Ist, 3°7 mm.; 2nd, 16°5; 3rd, 10; 4th, 1:5. The hind foot 
measures 16 mm., and the length of the body is about 155 mm. 
The type is a female. 

“The skull differs from that of b. villosus in a number of 
characters. The huge crest which rises up from the zygomatic 
arch is even better developed than in B. trevelyani, and a plane laid 
across the tops of the crests is 2 mm. above the cranial wall. In 
B. villosus the crests pass forwards and outwards, and ata distance 
of 7 mm. in front of the top of the occiput are 10°5 mm. apart 
in Smith’s type. In Dobson’s type they are 10 mm. apart. In 
B. transvaalensis the crest runs forward for some distance nearly 
parallel, and at a point 7 mm. in front of the occiput are only 
6-5 mm. apart. Another important point in the’skull is the very 
much larger temporal bulla. In J. villosus the bulla is about 
7 mm. in diameter, in B. transvaalensis it is 10 mm. in diameter. 


Proc. Zoou. Soc.—1913, No, XX XVII. 37 


548 ON A NEW GOLDEN MOLE AND YOUNG LIZARDS. 


‘¢he followine skull measurements (in millimetres) indicate 
oO 


the slight differences in size :— 
Length. Breadth. Height. Dental Series. 
ae 


Ga SSOUMIEUS UNOS go soc0oeosa0 c55000 33 20°5 15:5 13°5 
Dobson’s specimen............ 34 22 17 1B 
B. transvaalensis ............ Bylo, D355 es 14:5 


“The teeth in the specimen of B. transvaalensis are in front 
partly the milk and partly the permanent set. The Ist upper 
premolar has no internal cusp. The molars are appreciably larger 
than in B. villosus. In B. villosus the 2nd molar measures 2 mm. 
in width; in B. transvadlensis it measures 2°5 mm.; and there 
is an even greater difference in the size of the last molar. 

“The type has been deposited in the British Museum.” 


Dr. Broom also exhibited an adult female of the large 8. African 
Lizard, Zonurus giganteus, with two newly-born young (PI. 
LXXTIV.), and three adult male specimens of the allied species 
Pseudocordylus microlepidotus. 


He gave the following particulars of Zonuwrus giganteus :— 


“The mother was given to me by Mr. F. W. Fitzsimons, F.Z.8., 
of the Port Elizabeth Museum. On opening the box containing 
the specimen on arrival in London it was found that two young 
had been born on the voyage. The young differ in appearance 
very considerably from the mother, being much more brightly 
coloured. Young specimens in the British Museum collection 
are described by Mr. G. A. Boulenger as “light yellowish, 
marbled and cross-barred with blackish brown ; the spines not at 
all developed, and all the scales more strongly imbricate.” 

‘These young also show the marked imbrication of the scales 
which is largely due to their having no supporting dermal ossi- 
fication. This gives the tail especially avery different appearance 
from that of the adult. The colour may be described as yellowish, 
with, on the back, irregular cross-bands of black. The top of the 
head is blackish, but in most of the antorbital region the yellowish 
colour predominates. The scales round the eye are mostly bright 
yellow, but there is a black spot on the upper eyelid and a less 
distinct one on the lower. The 4th lower labial has a large black 
mark, and a black mark extends above this on the upper jaw to 
the eye. The scales above the tympanum are yellow. On the 
back the light bands, especially towards the lower half, become 
very distinctly reddish, and on the tail about half the scales are a 
light brick-red. The legs are irregularly banded yellow and 
black, and the front of the body is pale straw-coloured except the 
neck, which has a number of large black spots. The young 
measure about 5 inches in length. Mr. HE. G. Boulenger assures 
me that the young are feeding satisfactorily and look lively.” 


“SNALNVOID SNYNNOZ 
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ON MAMMALIAN CESTODES. 549 


PAPERS. 


36. Contributions to the Anatomy and Systematic Arrange- 
ment of the Cestoidea. By Frank HE. Bepparp, M.A., 
D.Sc., F.R.S., F.Z.8., Prosector to the Society. 


[Received April 8, 1912: Read May 6, 1913.] 
(Text-figures 85-94.) 
X. On Two Spectres oF TAPEWORMS FROM GENETT.A DONGOLANA. 


INDEX. 


Page 
Dipylidiim dongolense; Sp. Ne ...-----2--never se cee 2-2 549 
Diplopyliduuni Sen NOVe seers eee eseeee ee eeske ead OOO 
DDG CWECEE ASPs, Men det rir siasisteateceee meee TIT eeiee 559 


An examination of the small intestine of a Dongolan Genet 
(Genetta dongolana) yielded about a dozen small tapeworms and 
the hinder part of the body of a larger worm. ‘The latter I have 
not studied ; the former belong to two species which are described 
in the present paper. Inasmuch as the Genet, which died in 
November of last year, had been about four years in the Gardens, 
it is uncertain whether the parasites are to be regarded as indi- 
genous to Africa or had been acquired in captivity. From this 
particular variety of the Genet I believe that no tapeworms have | 
been recorded; but from allied forms species of Cestodes are 
known.  v. Linstow, in his ‘Compendium der Helminthologie,’ * 
mentions three from “ Viverra genetta,” viz. Mesocestoides ambiguus 
Vaillant 7, Tenia platydera and 7’. geneite, both of Gervaist. The 
latter, as I point out later, may be identical with my Dipylidiwm. 
Tenia platydera seems$ to have no rostellar hooks, and may 
quite possibly be an Anoplocephalid. I refer later to other species 
from Genets in my description of Dipylidiwm dongolense. 


(1) Dipylidium dongolense, sp. n. 


There were altogether five specimens of this species, all of 
which I have carefully examined either entire or cut into series 
of transverse and longitudinal sections. The largest specimen, 
after preservation in alcohol, measures just 5 mm. in length 
and 1 mm. in breadth at the broadest part. During life 
individuals reached a length of 6 mm. The accompanying 
illustration (text-fig. 85) represents the individual referred to as 
the largest. The scolex is not sharply marked off from the 
ensuing neck, which is very short and soon passes into the 


* Hannover, 1878, p. 37. 
+ Comptes Rend. Soc. Biol. Paris, 1863, p. 48. 
{ Mém. Ac. Sci. Montpellier, 1847. 
§ Cf. Diesing, Syst. Helminthum, i. p. 519. 
Sit 


550 DR. F, E. BEDDARD ON 


strobila, The neck is at once wider than the scolex. The pro- 
truded rostellum has several rows of the usual minute thorn-like 
spines characteristic of the genus Dipylidium. The rostellum has 
a three-pointed appearance, which will be obvious from the 
drawing, the narrowest part being the apex. 


Text-fig. 85 


Dipylidium dongolense. 
View of entire worm, magnified. 
p. Extruded cirrus. 7. Rostellum. 


T believe that there are four rows of spines upon the rostellum. 
The four suckers are not very large. The shape of the terminal 
segment of the body seems to me to prove that the specimen 
which I here figure is a complete individual with no posterior 
segments shed. It will be noticed that the proglottids are 
nowhere longer than broad, and that they overlap laterally. The 
segmentation of this species is thus very different from that of 


MAMMALIAN CESTODES. 551 


the type species of the genus, Dipylidiwm caninum. An in- 
spection of the figure will render a detailed description of the 
proglottids of the present species unnecessary ; it will be noticed 
that their number is small, not more than 28. In another 
specimen I only found 22. In this example (or in one very like 
it) the last nine segments became suddenly very much more 
elongate, and were three to four times as long as broad. In 
transverse sections the body shows an elliptical form pointed at 
the two ends laterally and nearly as deep as wide. 

The general structure of the body shows certain differences in 
the less mature and more mature segments, which do not appear 
to me to be altogether due to the greater tension in the latter. 
The specimen which I selected for examination by means of 
transverse sections was much like that of which I give an entire 
view in text-fig. 85. But in the posterior segments, though 
short, the sexual organs were well developed, and there were 
ripe ova lying in cavities (of which a full description will 
be given later). In these transverse sections the cuticle is 
very thick and the layer of subcuticular cells very conspicuous 
and deeply stained; they have the usual flask-shaped form 
and lie in a dense layer, being closely adpressed. Their thinner 
outer ends are in contact with the cuticle above them. Upon 
this layer follows a layer of lax tissue, and then a strong 
longitudinal layer of muscular fibres, which are themselves 
separated from the medulla by a thinner layer of transverse 
muscular fibres. These fibres are associated together in bundles 
of three or four fibres, which are very stout ; there appear to be 
also a few fibres to the outside, which are not associated in 
bundles but implanted singly. In longitudinal sections it is 
rather easier to count the number of fibres which lie in a single 
radial row of this longitudinal layer, and I find that there are not 
more than five or six. Here and there between the bundles are 
parenchymal cells whose deep staining as contrasted with that 
of the muscular fibres emphasizes their existence. In sagittal 
sections the subcuticular layer is also obvious in the more anterior 
proglottids. I have not been able to get any transverse sections 
of the more elongated posterior proglottids (owing to the limita- 
tion of my material); but in longitudinal (sagittal) sections I 
could find no trace whatever of elongated flask-shaped subcuticular 
cells. The general cortical parenchyma reaches absolutely up to 
the (here) very thin cuticle. There are, I think, only two possible 
explanations of this appearance of vanished subcuticular cells. 
First, that they have altered their form owing to the pulling 
out of the segment, and have become broader and round like the 
cells which secrete the calcareous bodies; or secondly, that they 
are really absent from this region of the body. It is clear from 
the observations of Lonnberg that the subcuticula varies among 
tapeworms. It is, for example, in Tetrarhynchus tetrabothrius* ~ 


* “ Anatomische Studien tiber Skandinavische Cestoden,” K. Svensk. Ak. Handl. 
xxiv. 1891, pl. i. figs. 6, 11. 


552, DR. F. E. BEDDARD ON 


a layer two or three cells deep, of which the outer ones are close 
to the cuticle. On the other hand, in Ptychobothrium belones the 
more scanty subcuticula lies in the parenchyma some way below 
the cuticle. I find on a re-examination of sections of Hyracotenia 
hyracis* that the subeuticula occurs in patches on the mature 
segments, and seems, therefore, to be in course of disappearance. 
It is, moreover, mentioned by Gough? that in Séilesia the sub- 
cuticular layer is not to be found in ‘older portions of the strobila 
in segments where the paruterine organ is fully developed.” 
Gough holds that this absence of the subeuticula in mature 
segments is a secondary character. The state of affairs seems to- 
bear out the suggestion that in my Dipylidiwm the subcuticula 
has actually disappeared ; in the mature segments. 

The testes occupy all the available space in the proglottids, and 
are thus mainly between and posterior to the ovaries. In the 
more elongated and riper proglottids the testes are only one row 
deep and not more than 6—9 (according to the length of the 
segment) in a single row; they are thus evidently not so 
numerous as In some species. 

The two generative orifices lie one on each side of the body not 
far from the anterior boundary of the proglottids. The ewrrus-sac 
is in front of the vagina, and, as is shown in sagittal sections, in 
the same straight line with it, being neither dorsal nor ventral. 
The cirrus-sac has very definite, rather thick muscular walls, 
which are even quite obvious when the sac is dilated in mature 
segments, though not so thick in appearance. The cirrus is long 
and much coiled within the cirrus-sac. The sperm-duct is also 
much coiled, and the two sperm-duct coils wey nearly come into. 
contact in the middle of the segment. 

The ovaries of this Dipylidiwm are, as a rule, clearly two in each 
proglottid lying rather laterally and anteriorly. They are, in fact, 
like those of other species, such as D. zschokkei of Hungerbihler, 
as to their position in the proglottid. But I do not find in cases 
which I have particularly examined that there is an absolutely 
complete separation between the two ovaries of a given proglottid. 
In this matter I refer to rather anterior proglottids which are 
not elongated and stuffed with ripe eggs containing embryos. 
However, in other proglottids I found a distinct separation between 
the two ovaries, the fact being that they approach very near to 
each other in the middle line of the segment. There is, however, 
no doubt about the entire separation of the vitelline clands, each 
of which lies behind the ovary of which it is an adjunct. The 
ovaries occupy a good deal of the space in the short proglottids 
where they occur when full sized. They are posterior to the coils 
of the vas deferens, but reach very nearly to the posterior end, 
where the testes occur ; the testes also occupy the available space 
between them. 

The receptaculum seminis penetrates the middle of the ovary 


* For species, see P. Z.S. 1912, p. 593. 
+ “A Monograph of the.... Avitelline,”’ Q. J. M.S. lvi. 1911, p. 345. 


MAMMALIAN CESTODES. 553 


and is greatly swollen so as to be almost spherical in this region. 
It is several times the diameter of the vagina. 

The wterws in the genus Dipylidium is thus described by Ran- 
som in his definition of the genus * :—‘“ Uterus at first reticular, 
later breaking up into egg capsules, each containing one or more 


Text-fig. 86. 


Dipylidiun dongolense. 
A completely mature proglottid seen in sagittal section. 


m. Longitudinal muscles. o. Ripe embryos, each in a separate cavity. 
¢. Remains of testes. 


eggs.” Nearly the same definition, but not quite so explicit, is 
given by Fuhrmann t, who writes :—‘‘ Der Uterus lést sich in 


* “Cestodes of N. American Birds,” Bull. U.S. Nat. Mus. no, 69, 1909. 
+ “Cestoden der Végel,” Zool. Jahrb. Suppl. vol. x. 1908. 


554 DR. F. E. BEDDARD ON 


einzelne ein oder mehrere Hier einschliessende Siickchen auf.” 
These definitions are, as I presume, chiefly based upon the many 
observations made upon the best-known species of the genus, 
viz. Dipylidium caninum (=Tenia cucumerina, mult. auct.), 
of which they are a correct restatement. But Fuhrmann has 
himself described in an Avian Dipylidium* a series of sinuous 
uterine tubes preceding a series of ege-capsules each containing a 
single egg. Furthermore, although he gives no details, Diamare f 
asserts that the development of the uterus in other species is as 
in D. caninum. 


Text-fig. 87. 


Dipylidium dongolense. 


A portion of the section represented in text-fig. 86, more highly magnified 
to illustrate the absence of a special lining membrane to the egg-holding spaces. 


Lettering as in text-fig. 86. 


In the species of which the present paper is an account I can 
find no confirmation of Diamare’s statement concerning the de- 
velopment of the uterus, though his figures of the completely : 
mature uterus of D. trinchesti and D. pasqualii agree very closely 
with what I have seen in mature proglottids of D. dongolense (see 
text-fig. 86). In the largest specimen which I have been able to 
examine, in which the last nine proglottids were elongated, it was 
only the last two which agreed really closely with the figures given 
by Diamare and just referred to. In these alone the embryos were 
contained each in its own separate compartment, which it did not 
by any means fill. I lay particular stress upon the fact that they 


* “Taenien der Raubvoégel,” CB. Bakt. u. Paras. xli. 1906. 
+ “Tl genere Dipylidium,” Atti R. Acc. Napoli (2) vi. 1894. 


MAMMALIAN CESTODES. 55D 


were embryos, since in Diamare’s figures they are clearly eges 
which have not yet segmented to form emb1 ‘yos. In Dip alidlinm 
caninum (as 1 suppose), an example of which I have studied from 
the Asiatic Jackal, the separate chambers of the mature uterus 
were very plainly lined with a cellular layer. J may observe of 
this Dipylidium incidentally that the number of embryos in a 
given chamber was very much greater than what is figured in 
that species by Diamare, who only represents 3-7. I found quite 
30 or more. 


Text-fig. 88. 


Dipylidium caninwn. 
A portion of uterus in a mature proglottid. 


ep. Epithelial lining of uterine cavities. O. Embryos. 


In Dipylidium dongolense there was not, so far as I could see, 
any special lining membrane to the small chambers which lodged, 
each of them, a developing embryo. It will be observed in the 
text-figure (text-fig. 87) which illustrates the mature “ uterus” 
of the present species, that the appearances presented are con- 
sistent with the view that the chambers which contain the eggs 
are merely interstices in the general medullary network. Their 
walls are perfectly continuous with the medullary tissue lying to 
the inside of the longitudinal muscles and forming the peripheral 
layer of the medulla. Though the walls of the cavities often 
contain imbedded in them crowded nuclei, it must be borne in 
mind that the peripheral layer of the medulla is also crowded 


556 DR. F. E. BEDDARD ON 


with quite similar nuclei. Furthermore, the nuclei (see text-fig. 
87, p. 554) do not give the impression that they line the cavities ; 
they belong rather to the tissue of the walls: there are, moreover, 
often considerable tracts without any nuclei at all, as is often 
the case with the medullary tissue of tapeworms. This is, of 
course, opposed to the idea of a lining membrane. 


Text-fig. 89. 


Dipylidium dongolense. 


A portion of a section through a proglottid less mature than that represented 
in text-figs. 86 & 87. 


m. Longitudinal muscles. o. Egg-holding spaces. 


All this contrasts greatly with the uterine spaces of D. caninwm 
(see text-fig. 88, p. 555), where there is invariably a close row 
of nuclei with accompanying protoplasm actually lining the 
cavity and in which the nuclei of the medullary tissue are 
not numerous and, as a rule, different from those of the lining 
membrane. ‘The thedulla, in fact, forms a reticulum in éhe 
meshes of which lie the developing embryos; there is not, 
at any rate in the fully mature stage, any uterus at all. A 


t 


Oo 
1) | 


MAMMALIAN CESTODES. DDT 
consideration of the disposition of the eggs in proglottids anterior 
to the two that have just been described bears out this view. 
In the two proglottids anterior to the two terminal ones, the 
arrangement of the eggs, or rather developing embryos, in the 
medullary parenchyma was as follows :—The medullary region is 
not so entirely given up to these eggs as it isin the case of the 
last two segments that have been described above. In the two 
latter but slight traces are left of the testes; in the less mature 
proglottids more testes are found. They have, however, com- 
menced to disappear, for in the proglottids antecedent to these the 
testes are more abundant and therefore take up more room. The 
space apparently thus left by the degeneration of the testes is 
occupied by the developing eggs. It is not, however, a continuous 
cavity running from end to end of the proglottid (see text-fig. 89). 
It is divided here and there by delicate trabecule into compart- 
ments, each of which contains a considerable and varying number 
of developing embryos*. Further forward still, the embryo-con- 
taining spaces are less conspicuous owing to the full development 
of the testes, which thus occupy more room. JI can draw no line 
of demarcation between these cavities in which lie the developing 
embryos on the one hand, and spaces which lodge ova and form 
actually a part of the ovary on the other hand, in the more 
mature proglottids which have begun to become elongated. 
As already mentioned in describing the ovary of this species 
of Dipylidium, the ovaries are mature or, at any rate, 
possess plenty of full-sized ova in proglottids which are not 
at all elongated and are, indeed, still broader than long. In 
these proglottids, however, there is no trace that I could find 
of a definite uterus, and nothing at all like the obvious retiform 
uterus of not fully mature proglottids of D. caninum. However 
we may interpret the embryo-holding spaces of D. dongolense, it 
is clear that they cannot arise in the same way that the egg- 
sacs of D. caninum arise, i.e., by a breaking up of a continuous 
retiform uterus into a number of discontinuous sacs. 

With what, then, can we compare the spaces which surround 
the developing ova in this and other species of Dipylidiwm—in 
fact, possibly in all with the exception of D. caninum? I do 
not think that any genus has been described as possessing an 
exactly similar mode of lodgment of the developing eggs. But 
there are nevertheless some comparisons to be made with other 
forms. 

The final stage shown in Dipylidiwm dongolense, where the 
eggs are uniformly scattered through the medullary region, each in 
its own separate compartment, is suggestive of Oochoristica, with 
which genus Dipylidiwm is usually considered to be properly 
associated. The comparison, however, can hardly be exact, since, 
as we have seen, this final stage is preceded by larger spaces con- 
taining each of them many developing embryos. Jt seems to 


* Tn one specimen I have also seen ova enclosed by threes and fours in cavities in 
the cortical layer. But this seems exceptional. 


558 DR. F. E. BEDDARD ON 


me that we must come to the conclusion that a growth of the 
medullary parenchyma between the individual eggs produces the 
final result seen in the terminal and penultimate proglottid of 
the specimen which I have studied. It might be argued that 
this process is really comparable to the formation of paruterine 
organs which are growths of medullary tissue encapsuling the 
eges though with histological change. In Dipylidiwm we have 
no histological change, but all the same encapsuling growths of 
parenchyma. 

Having given an account of the structure of this Dipylidium, 
it remains to be decided whether it be a new species or identical 
with one of those already described. It is, as I think, hardly 
necessary to say much of D. caninwm, with which there can be no 
possibility of confusing the present species. The much greater 
size of that species and the form of its uterus at once distinguish 
it from that which is the subject of our present communication. 
It seems indeed to me that the genus Dipylidiwm requires really 
division into two genera—J/. caninum belonging to one, and all 
the other species that are adequately known to the other. It is 
from the latter that it is not so easy to distinguish D. dongolense 
of the present paper. If D. genette of Gervais * and D. gervaisi 
of Setti were not much larger species than mine, I should have 
been inclined to identify it with one or possibly both of them. 

D. echinorhynchoides (from Megalotis cerda) is much larger than 
my species, has a much longer rostellum and a longer neck. 
D. trinchesti is a smaller species, but still considerably larger 
than D. dongolense, measuring 2°5 cm.; it has, however, a 
short rostellum, like that of my species; but it is to be 
distinguished by the greater length of the mature segments, 
which do not overlap, and the invariable inclusion of each ripe 
ovum in a separate compartment. Though a receptaculum 
seminis 1s said to be present, it is hardly represented in the 
figure. D. pasquali, on the other hand, has an obvious recep- 
taculum {, but it is a much larger species, 7.e. 200 mm. The 
uterus is as in the last, and therefore differs from that of D. don- 
golense; but it is to be noted that the ripe ova extend into the 
cortical layer, as do the egg-chambers of D. dongolense. Occurring 
as 1t does in the intestine of a closely allied species, it might be 
expected that D. gervaisic of Setti $ would be more closely allied 
to D. dongolense than the other species of Dinylidiwm referred to. 
It is not, however, as it would appear, any nearer in structure. 
For in this species, as in most others, the ripe ova are figured as 
embedded separately in the parenchyma, without any previous 
inclusion in larger cavities. It is also rather larger, being from 
1-4 em. in length. 

Nor can the present species be confused with D. zschokkei ||. 

* Mém. Ac. Sci. Montpellier, 1847. 

+ Diamare, loc. cit. pl. 1. fig. 9. { Loe. cit. pl. i. fig. 8. 

§ “ Dipylidium gervaisii,’ Atti Soc. Lig. Sci. Nat. vi. 1895. 

|| Hungerbiihler, “Studien au Gyrocotyle und Cestoden,”’ Denkschr. Ges. Jena, 
xvi. 1910, p. 516. 


MAMMALIAN GESTODES. 559 


For the latter reaches a length of 120 mm., and the ripe pro- 
glottids may be four times as long as broad. Moreover, the neck 
of D. zschokket “muss als sehr lang bezeichnet werden,” It 
appears also that my species has a much more pronounced 
receptaculum, which is mentioned by Hungerbihler as existing 
in D. zschokkei, but is hardly indicated in his figure*. This 
species occurs in Cynictis penicillata. Dipylidium triseriale of 
Lihe also appears to odin a species which occurs in the African 
Civet cat. For Liihe says of that species that the ripe eggs are 
imbedded singly in the parenchyma, and makes no mention of 
previously existing spaces, a state of affairs which I have described 
in my species. Moreover the genital pores of D. triseriale are 
further back than in my species, and it is larger in size, In 
D. triseriale, moreover, the proglottids are described by Liihe 
as being “ungefihr quadratisch,” which I presume means that 
they do not overlap as In my species. 

A second species, Dipylidium monoophorum, 1s also described by 
Liithe from the same host, which only measures 10 em. in length 
and therefore more nearly approaches the dimensions of ). dongo- 
lense of the present paper. But in this species, as in D. triseriale, 
the length of the ripe proglottids is much greater; and further- 
more, the two separate small round ovaries (with the vagina to 
the outside of each) are not what I have seen in D. dongolense. 

Diamare = has described, besides the well known JD. caninum, 
D. echinorhynchoides Sonsino, two species named by himself, 
viz. D. trinchesti and D. pasqualii (both from the common cat), 
and two doubtful species, which are D. (?) genette Gervais and 
D, monticellir, 


Diplopylidium genettz, gen. et sp. n. 


I refer what I believe to be a new species of Tapeworm to the 
new genus Diplopylidiwm for reasons which I shall discuss after 
setting forth the structure of this worm. The intestine of the 
Dongolan Genet contained fewer specimens of this worm than of 
the Dipylidiwm which has been described above. They were, 
however, of about the same size, measuring up to 6 mm. or so 
in length, and consisting of about 28 segments at most. 

The accompanying drawing (text-fig. 90) represents an average 
sized example to which the above statements apply. The scolex 
is as broad as the body which follows, and there is only a slight 
increase up to the end of the body. The worm has thus a sturdy 
form, and is so far much like the Dipylidiwm which has just been 
described. It agrees with it, moreover, in thickness, the trans- 
verse sections being oval in outline and not much flattened from 
above downwards. The scolex is furnished with an armed ros- 
tellum, which is large in contrast to the comparatively small 

* Toc. cit. pl. xix. fig. 18. 

+ “Beitrage zur Helminthenfauna des Berberei,’ SB. Ak. wiss. Berlin, 1598 


p- 626. 
ft “Il Genere Dipylidium,” Atti R. Acc. Napoli (2) vi. 1894. 


560 DR. F. E. BEDDARD ON 


suckers. These latter are unarmed and show no peculiarities of 
moment; they face outwards and slightly forwards. The hooks 
of the rostellum are large and arranged in two rows, 17 to each 
row. Their shape and relative size can be understood from an in- 
spection of text-figure 91, which represents a longitudinal section 


Text-fig. 90. 


8 


Diplopylidium genette. 


View of entire worm, magnified. 


g. Extruded cirri, 7. Rostellum. 


through the retracted rostellum of this species. The rostellum is 
a muscular one, but with fewer external layers of muscles than is 
sometimes met with, which act as protractors of the rostellar bulb. 
The stout retractors (see text-fig. 91) are of course continuous 


MAMMALIAN CESLODES. 561 


with the longitudinal muscular layer of the body, which is poste- 
viorly not thick but quite evident. In comparing the present 
species with Dipylidium dongolense in the living state, I noted a 
difference in the dispersion of the calcareous corpuscles. In both 
species they were abundant throughout the body generally; but 
in the Diplopylidium alone did I succeed in detecting any in 
the scolex. 


Text-fig. 91. 


Diplopylidium genette. 
Longitudinal section through anterior end of body. 


FH. Hooks on rostellum. S. Sucker. 


The éestes of this species are numerous in proportion to the 
size of the proglottid. They are ripe in the earlier seg- 
ments, where the ova are not mature, but still attached to the 
ovaries. In such segments the testes are closely crowded 
together and fill all the availabie space of the proglottid from 
one end to the other. They lie, however, within the ovaries, 
thus occupying the middle of the proglottid. I counted up to 
twelve or so in a single longitudinal row. The cirrus-sac of 


562 DR. F, E. BEDDARD ON 


the worm is large and has an obvious and thick muscular coat. 
A peculiarity about this organ is that it does not he in a straight 
line extending inwards from the point of opening on to the 
exterior, and thus at right angles to the long diameter of the 
proglottid, as is so usually the case. The sac, on the other hand, 
often bends at right angles near to its external pore and passes 
backwards ; at other times it is more irregular in its folding. 
Its large size is responsible for this folding, as the two cirrus-sacs 
of a given segment would meet in the middle line did they run 
straight inwards from the external pore. The cirrus itself is a 
very fine tube upon the end of which I could detect no spines ; 
it is coiled within the cirrus. The sperm-duct after emerging 
from the cirrus-sac forms a coil which is anterior to the sac and 
near to the front of the segment. 

The vagina of this Cestode together with adjacent parts of the 
genital system is shown in text-fig. 92, The most remarkable 
fact is that that tube opens anteriorly to the cirrus, a position 
which is made plain in the text-figure referred to. It would 
also appear from a study of a series of sagittal sections—which 
are the best in which to study these particular relationships— 
that the vagina has not a separate orifice on to the exterior of 
the body. But the truth of this conclusion will be doubted after 
an inspection of text-fig. 92. For in that figure the vagina 
seems to open separately, and of course anteriorly to the cirrus- 
sac. It may be that this is at times the case; but it is also 
clear that in tracing the vagina in a series of sagittal sections the 
tube is lost sight of in the walls of the male duct before the latter 
reaches the exterior. I believe that among the Tetracotylea the 
genus Tetrabothriwum is the only genus in which the female 
openings are in front of the male. It is true that in the Ichthyo- 
teniide generally, if not universally, there is an irregular alter- 
nation in these conditions, the female duct lying in one segment 
in front of and in another behind the male pore. But it is 
doubtful whether this family is to be safely referred to the 
Teeniade. The figure referred to (text-fig. 92 C) shows the narrow 
vagina lying in front of and parallel with the stouter cirrus-sac, 
the two forming in this region a perfectly straight line, so that 
their relative positions is a matter of ease to ascertain. I found 
the same relations in other series of sections besides that from 
which the text-figure referred to has been taken. Thus I am 
able to state that there is not an alternation in the positions of 
the male and female openings, as in Jchthyotenia. The vagina 
is darkly stained and of small calibre in this tapeworm, the small 
bore of the tube bearing a relation to the fineness of the cirrus. 
Another remarkable circumstance is the nature of the recep- 
taculum seminis and less mature and more mature proglottids. 
In text-figure 92C the slender vagina is seen to open into a minute 
spherical chamber or dilatation and to issue from the opposite side 
as an equally slender tube. This chamber is, as I suppose, to be 
compared to a receptaculum seminis; but it has in this section 


MAMMALIAN GESTODES. 563 


the appearance of the mere beginning of such a chamber. There 
is no trace of sperm to be found in it, and it is so small that I 
have not been able to detect it in other sections of neighbouring 
segments. The particular section to which attention has been 


Text-fig. 92. 


(E Wipics 
Three sections through the generative region of Diplopylidiwm genette. 


-A. Section to show relations of vagina (@ ) and cirrus-sac (¢), the latter lying 
behind the former. 


B. A transverse section through the receptaculum seminis and adjacent parts 
of the generative system. 


ce. Cirrus-sac. o. Vitelline gland. 7.s. Receptaculum seminis. 


v. Vagina. 
vit. Ovary. 


C. Transverse section showing immature receptaculum seminis (7.s,) and 
anterior position of vagina (v.). 


c. Cirrus-sac. 


N.B.—There is an appearance of a diverticulum of the cirrus probably due to 
irregular staining. 


Proc. Zoot. Soc.—1913, No. XXX VIII. 38 


564 DR. F. E. BEDDARD ON 


called is of a proglottid filled with completely mature eggs, 
enclosed in their shell, and which have developed into embryos. 
It is rather remarkable to find that the receptaculum is so small 
and immature ina section which is otherwise quite ripe. For 
the receptaculum does not suggest a reservoir from which the 
contents have been recently expelled; it distinctly suggests an 
incompletely developed receptaculum. On the other hand, text- 
figure 92.B is from a section of which the maturity was less 
advanced. The testes were fully ripe, and the ovaries quite 
developed ; but there were no ova scattered through the paren- 
chyma at all. And if I have missed any in the examination of 
the section, they must at most have been few. Yet in this 
section, as will be seen from the drawing cited, the receptaculum 
seminis is very large and quite distended with abundant sperm. 
Nor can this difficulty be explained away on the assumption that 
one of the two vagine in the fully-ripe proglottid was more 
mature than the other, and may have been the storehouse of the 
sperm after copulation. For I ascertained that the vagine on 
both sides of the body were in an identical state of maturity. 

The ovaries in this species are distinctly double, and lie one on 
each side of the proglottids a little way behind the point of 
opening of the generative ducts. The ovary is in each case 
inmediately Follow ed by the vitelline gland. There seem to be no 
features of special interest about either the ovaries or the vitelline 
glands. On the other hand, the cavities lodging the ripe eqys are 
remarkable, and like those of but few other tapeworms that have 
neon described. I do not give to these cavities the name of 

“uterus,” for it does not seem to me to be proved that they 
actually ‘correspond to the uterus of such genera as Choanotenia, 
Hymenolepis, etc. ‘The spaces indeed to which I here refer are more 
comparable, as it appears to me, to those of the Dipylidiwm, which 
have already been deseribed in the present paper, and to be rather 
lacune in the medullary network than deliberate pre-existing 
cavities at first united to form a single cavity. I believe, imoleeal 
that there is no such fraementation| here of a pre-existing uterus 
as, for example, I have lately described * in my genus Otiditenia. 

For it is hard to believe that a uterus like that of so many 
tapeworms can exist in the present species. As already men- 
tioned, in the more anterior proglottids, where the testes are 
fully ripe and the ovaries fully formed, there is no trace of any 
sac which might be considered to be a uterus. Nor, indeed, 
among the closely adpressed testes which occupy so much of the 
interior of these proglottids does it seem that room exists for 
the development of a uterus. In any case nothing obvious of 
the kind is visible. It is in riper proglottids which are more 
elongated in their form that the “ uterine” spaces referred to are 


first visible. They le (text-fig. 93) scattered among the testes, 


* “Ona New Genus of Tapeworms (Otiditenia) from the Bustard (Hupodotis 
kori), P.Z.S. 1912, p. 194. 


MAMMALIAN CESTODES. 565 


and are developed in the posterior region of the proglottid 
only, but all the same occupy the greater part of such proglottids. 
There is no regularity in the dispersal of these cavities, so far as 
I have been able to make out. They lie here and there, and are 
never crowded like the testes ; among them lie the remains of the 
testes in the most posterior segments, where the testes are most 
fully reduced. The cavities increase in size pari passu with the 
development of the embryos contained within them. In the more 
anterior part of the body the spaces are small and contain but a 
single egg or embryo just commencing to divide, which does not 
nearly fill the cavity. 


Text-fig. 93. 


Diplopylidium genette. 


A portion of a nearly ripe proglottid showing ova (e.) lying im spaces in 
the medulla. 


In the centre of the field is one ovum which has no cavity round it. 


Further back the cavities are larger to accommodate the large 
embryo with its wide egg-shell. Nowhere is there anything of 
the nature of a lining membrane to these cavities. No nuclei 
are distinguishable as forming a layer immediately surrounding 
them, though the nuclei of the medullary parenchyma are, of 
course, to be found frequently in the neighbourhood of the 


limiting layer of the cavities; but in these cases there is no 
38* 


566 DR. F. E. BEDDARD ON 


relation to be observed between these cells and the egg-holding 
spaces. I have never found more than a single embryo in one 
cavity ; if there are occasionally more, such instances must be 
rare. The accompanying drawing (text-fig. 94) shows a number 
of the embryo-holding spaces in a fully ripe proglottid. It will 
be noticed that between the embryo itself and the wall of the sac 
in which it lies is a tumbled mass of a membranous appearance. 


Text-fig. 94. 


Po Oi 
/ a ids 


do i 
™ 


Diplopylidium genette. 


A portion of a more fully mature proglottid. The embryos are now surrounded by 
a delicate often crumpled membrane, and the spaces in which they lie are 
larger. 


e. Embryo. m. Egg-membrane. w. Cavity surrounding embryo. 


This is not all stamed by the hematoxylin, which has amply 
stained the embryo itself and the surrounding tissues of the 
medulla. The inference, therefore, appears to be that this 
membranous material is in reality the egg-shell and is of 
a chitinous nature. In some cases it has a more regular 


MAMMALIAN CESTODES. 567 


appearance, and seems to lie in concentric layers surrounding the 
embryo; in others this is not apparent, and it even simulates a 
retiform tissue in the laxity of its arrangement. 

The ultimate arrangement of the uterine spaces in this worm 
is, therefore, the same as we find in the species of Dipylidiwm 
that has just been described. But the two genera differ in the 
fact that this is preceded in Dipylidium by another developmental 
stage not represented in Diplopylidiwm. In the latter we find 
the scattered cavities lying among the remains of the discharged 
testes, and as both a final and an initial stage. There is no inter- 
mediate condition that I have found where the ova are contained 
in larger cavities, which subsequently become reduced to chambers 
each containing a single egg such as is found in Dipylidium. 
The cavities in the two cases are, therefore, not exactly equiva- 
lent; otherwise, as has been said, there is a close agreement 
between the two worms. In neither can any lining membrane 
be detected forming a wall to the ege-cavities. The actual size 
of the individual spaces is much the same, but in Dipylidiam 
they are more closely packed together. In formulating this 
difference between the two genera I rely upon what I have seen 
and not seen in my sections. It would be unwise—because of 
the negativity of my conclusions—to assert this fact in a more 
positive way by using it as part of the generic definition of 
Diplopylidium. It is not only in Dipylidiwm that we find a 
series of egg-holding spaces like those of Diplopylidium. The 
same kind of arrangement occurs in Oochoristica, so named 
on account of the scattering of the mature ova through the 
parenchyma. But in this genus there is an obvious uterus which 
exists before the final scattering of the eggs. Moreover, when 
imbedded in the parenchyma, the appearance is as if the eggs 
were firmly imbedded in the homogeneous ground material of the 
medullary tissue and not received into cavities. The eggs, in 
fact, are immediately surrounded by solid tissue and not by the 
rest of a cavity. More like the disposition met with in Dipy 
lidiwm and Diplopylidium is that which is to be seen in J/ono- 
pylidium, a genus to which the present genus, as I point out 
later, is probably related. But it should be remarked, first 
of all, that the absence or presence of a cavity round the singly 
imbedded ova is not a matter of great importance. For, as will 
be seen by an inspection of text-fig. 93, it happens occasionally 
in Diplopylidiwm that imbedded ova have no cavity round them. 
This may well be a preliminary state of affairs, the subsequently 
appearing cavities being due to the growth of the egg into the 
embryo. In this case there will be no doubt whatever that the 
egg-holding cavities of Diplopylidiwm are totally different from 
those of Dipylidiwm dongolense, since the latter are formed as 
subdivisions of an earlier larger cavity. 

In Monopylidium the ova are in the same way scattered 
through the parenchyma in cayities which contain one, or in 
some cases more than one, ovum. It is evident, however, from 


568 DR. F. E. BEDDARD ON 


the definition of this genus by Ransom *, that he regards these 
spaces as disjected portions of a pre-existing uterus. It seems 
clear, from the illustrations given by Mola‘, that the egg-holding 
cavities are not lined by cells in Zenia marchali (a species which 
Fuhrmann refers to the genus J/onopylidiuwm), and in this fact 
we find an agreement with the two tapeworms described in the 
present paper. A very remarkable condition is described and 
figured by Fuhrmann? in Monopylidium rostellatum. In this 
species the eggs are scattered singly through the medullary region, 
and each egg fits closely into a hexagonal mass of delicate tissue, 
which itself fits equally closely into the medullary parenchyma, 
leaving no gaps or spaces anywhere. It seems to me to be just 
possible that this “‘ parenchyma vacuolaire”” which immediately 
surrounds the egg may be simply the outer egg shell, which 
I have myself observed in Diplopylidiwm, when crumpled, to 
present quite the appearance of a retiform tissue. It is finally 
to be noted that the disposition of the mature ova in Diplo- 
pylidium is perhaps also to be compared to the scattering of the 
eges in Inermicapsifer capensis § before the inclusion of these in 
the parenchymal organs. But it must always be borne in mind 
that in the case of Diplopylidium, Oochoristica, and Monopylidium, 
completely mature proglottids may not yet have been seen. A 
final stage may be the formation of paruterine organs, though 
this does not seem to be very likely. 

We may characterise this new form as follows :—Size small, up 
to 6 mm.; number of segments up to 28. Scolex with retractile 
muscular rostellum armed with two rows of hooks, 17 in each 
row. Suckers unarmed. Neck short; posterior proglottids two 
or three times as long as broad. They do not project posteriorly 
or overlap. Generative orifices lateral and paired in each pro- 
glottid, opening at or rather in front of the middle of the 
proglottid. The ducts pass between the longitudinal water- 
vascular tubes. The female orifice is situated in front of the 
male, The water-vascular tubes are two on each side, the dorsal 
being of narrower calibre; they are placed one above the other. 
A longitudinal muscular layer, only a few fibres thick, separates 
the cortical and medullary regions. The testes are numerous and 
large, filling up all the proglottid not occupied by other parts of 
the generative system ; testes disappear in posterior proglottids 
when egg-chambers appear. The cirrus-sac is large, with muscular 
walls, not lying in one straight line, but bent backwards in rela- 
tion to the fact that there are two in each proglottid. The cirrus 
is very slender and coiled within its cirrus-sac. The sperm-duet 
is coiled and lies on each side in front of cirrus-sac. Ovaries 
distinctly paired in each proglottid, lying laterally behind the 


* “The Teenioid Cestodes of N. American Birds,” Bull. U.S. Nat. Mus. no. 69, 
1909, p. 76: “ Uterus breaks down into egg-capsules.” 
+ Bull. R. Ac. Belg. 1907, p. 806. 
+ “ Nouveaux tenias d’Oiseaux,” Rev. Suisse Zool. xvi. 1908. 
P. Z.S. 1912, p. 586, text-fig. 76. 


MAMMALIAN CESTODES. 569 


generative pores and at the middle of the segment ; each ovary 
lies i in front of the corresponding vitelline gland. The vagina is 
narrow and passes forward, running in front of and parallel to 
terminal section of cirrus- pouch ; ‘poster iorly it dilates into a 
receptaculum seminis lying close to and in front of ovary and 
behind dilated region of cirrus-sac. Uterus as a simple cavity is 
absent (2). The developing ova are lodged singly in cavities of 
the medullary parenchyma, which they nearly fill, and are without 
any lining epithelium. Eggs with very wide but delicate shell. 

From this general résumé we may now endeavour to form a 
generic definition of Diplop ylidium by eliminating from the above 
characters those which are in all probability to be regarded as 
only specific in value. 


Diplopylidium, gen. nov. 


Retractile muscular rostellum armed with two circles of hooks. 
A double set of reproductive organs in each proglottid. Genital 
canals pass between dorsal and ventral excretory tubes. Vagina 
lying in front of cirrus-sac. Cirrus-sacs large and muscular, bent 
upon themselves ; cirrus coiled, very slender, and unarmed ; sperm- 
duct forms a coil. Testes numerous, filling wp available space in 
progiottid. Ovaries two, in front of vitelline glands. Vagina 
narrow, with receptaculum seminis. Uterus represented by nume- 
rous cavities, each containing one egg only. 


We have now to consider the affinities of this tapeworm and 
the family within which it should be placed. 

There is not a very large number of genera in which the 
reproductive organs are double in each segment. We find, how- 
ever, such forms in nearly all of the families into which the 
Tetracotylean Cestodes are divided. We may at once place on 
one side those genera, such as Cittotenia and Monieza, which 
belong to the Anoplocephalide ; for in the genus which forms the 
subject of the present communication there are not, as in these 
forms, persistent uteri; and, moreover, the worms of the family 
Anoplocephalide are not provided with a hooked rostellum. Nor 
can we place the present genus, which I propose to term Dzplo- 
pylidium, with either Diplophallus* or Diploposthe, two genera 
which are nearly allied to each other, though placed by Ransom + 
in separate families, viz. the Acoleide and Teeniade. It must be 
noted, however, that Fuhrmann {, in his well-known revision of 
the tapeworms of birds (those belonging to the Cyclophyllidea), 
regards Diploposthe as not strictly referable to the Tzniade, but 
as forming an intermediate type between this family and that of 
the Acoleide. 


* For anatomy of Diplophallus, see Cohn, Zeitschr. wiss. Zool. ]xvii. 1897, p. 277, 
‘and Wolfhiigel, “‘ Beitr. z. Kenntniss d. Vogelhelminthen,” Inaug.-Diss. Freiburg- 
im-Br. 1898. 

+ Bull. U.S. Nat. Mus. 1909, no. 69. 

t Zool. Jahrb., Suppl. vol. x. 1908. 


570 DR. F. E. BEDDARD ON 


Both of these genera have a rostellum which is armed with 
ten hooks. The ‘preater number of hooks in Diplopylidiwm 
might be regarded as a difference of merely specific value. But 
while in Diploposthe * and Diplophallus the uterus is persistent, 
and in mature segments occupies an enormous amount of space 
in each ripe proglottid, Diplop, ylidium has probably not a per- 
sistent uterus at all; or, if the spaces containing eggs which have 
been described above are to be looked upoh as remnants of a 
uterus or of uteri, the conditions are obviously very different. 
This important diacronce: as it appears to me, renders any con- 
fusion between these forms quite impossible. I have referred 
above to certain other comparisons between my genus and the 
two that have just been compared with it. 

My genus presents certain points of likeness to the genus 
Cotugnia, founded by Diamare in 18937. The chief point. 
of likeness is that in Diplopylidium the ripe eggs are imbedded 
singly in the medullary parenchyma, as is stated to be the case 
with Cotugnia. But in the latter genus, Fuhrmann {, though he 
gives no figures showing detail, speaks of a ‘“ parenchymkapsel ” 
as surrounding the eggs. This is probably to be compared to 
that of the allied Davainea, and, therefore, presumably is not 
like that which I describe in the present paper. The vagina, too, 
of Cotugnia has a dilated receptaculum seminis. Otherwise I do 
not think that the two genera can be confused. The worms are 
rather large species, and the rostellar hooks are numerous and 
minute, and have the typical Davaineid form which is unlike 
that which I find in Diplopylidium. But Cotugnia appears to: 
differ from other Davaineids in having no hooks upon the suckers 
[‘“‘ Ventose grandi e inerme” (Diamare)]|, and thus to approach 
Diplopylidium, to which its double generative pores and possibly 
the nature of the egg-cavities affine it. 

It may be furthermore pointed out that the genus Co otugnics 
seems to be characterised by its short and broad proglottids, in 
which it clearly differs from the worm upon which I report im 
the present communication. DiamareyT, in his paper upon 
Cotugnia (and other tapeworms), assigns to his new genus 
a species described by Monticelli§$ from material named by 
von Siebold. This species (‘‘ Tenia bifaria”) has, as it appears 
to me, rather more claims to be allied to my genus Diplopylidium 
than to Cotugnia, if Diamare is right in supposing Monticelli to- 
have overlooked the rostellar hooks. For in his figure || of the 
generative organs (which is repeated in Bronn’s ‘ Thierreichs’ @ ), 


* For the anatomy of Diploposthe, see Jacobi in Zool. Jahrb. x. Anat. Abth- 
1897, p. 287; Kowalevsky, Bull. Ac. Cracow, 1903, p. 518, for a brief account of 
Diploposthe sui-generis n. sp.(?); and Fuhrmann, Centralbl. Bakt. xl. p. 218, for 
a general résumé of this genus. 

+ Boll. Soc. Nat. Napoli (1) vii. 1893, p. 11. 

t “Neue Davaineen,” Centralbl. f. Bakt. u. Paras. xlix. p. 115; and Zool. Anz. 
xxiv. 1901, p. 273. 

§ Boll. Soc. Nat. Napoli (1) v. 1891, p. 151. || Loe. cit. pl. vill. fig. 12. 

¥ Pl. lvi. fig. 9. 


MAMMALIAN CESTODES. 57} 


Monticelli places the vagina in front of the cirrus-sac as I have 
found it in Diplopylidium. But this tapeworm has a cirrus 
which is armed with spines which I have not found, and which 
also does not seem to occur in Cotugnia. Nor does Monticelli 
say anything of the eggs: which makes any comparison more 
difficult. The segments, however, are more elongated than in 
Cotugnia and thus like those of Diplopylidiun. 

Passing by the family Amabilide, of which the type-genus 
Amabilia has double reproductive organs, and to which Diplo- 
pylidium does not, as it appears to me, show any affinities, we 
come to the Hymenolepide. To this family, and especially to the 
subfamily Dipylidiine (in the sense of Ransom), Diplopylidiwm 
shows points of likeness. In this subfamily there are two genera, 
viz. Dipylidium and Pancerina, which have reduplicated gene- 
rative organs. After the description of a Dipylidium, which I 
have given in the present paper, I do not think it necessary to 
emphasize further the distinctness of Dipylidiwm and Diplo- 
pylidium. As to Pancerina, that genus has no hooks upon the 
vostellum ; it may be merely an Oochoristica with double gene- 
rative organs, or, as I have ouegesied elsewhere * an Ichthyo- 
teniid. Our ‘knowledge of its anatomy is at present insufficient 
to place it with accuracy. 

But although Diplopylidiwm is certainly not identical with 
Dipylid ium, it agrees with that genus and some other Dipylidiinze 
in the characters of the ese holding spaces (which I do not term 
uterus for reasons given ¢ above )r. It is obviously quite near in 
this respect to the species of Dipylidiwm which I describe in the 
present paper. Oochoristica also has the ripe ova scattered at 
random through the medullary parenchyma ~. Monopylidium has 
also the same “type of egg-holding cavities van I have referred 
to more at length above in describing this part of the generative 
system of Diplopylidium. Furthermore, in Monopylidiwm there 
are species (e. g. Jf. macracanthum $) where the hooks are not 
very numerous (22) and arranged in two circles. On the whole, 
Tam inclined to place my genus in the near neighbourhood of 
Dipylidium and Monopylidium. 


P. Z.S. 1913, p. 6. 

Supra, p. 567 ete. 

Cf. e. g. Beddard, P. Z.S. 1911, p. 633, text-fig, 150 e. 
Fuhrmann, Centralbl. f. Bakt. u. Paras. xlv. 


tt % 


572 MR. J. A. MILNE ON THE 


37. Pacific Salmon: An Attempt to evolve something of 
their History from an Hxamination of their Scales, 
By Joun ApvAm Mitne, of Ardmiddle, Turriff, 
Aberdeenshire *. 
[Received April 9, 1918 ; Read May 6, 1913]. 
(Text-figures 95-118.) 


SALMONIDA: SeRuctuRE, DevetorpmeEnt, EvHonoey. 


The above title indicates what is perhaps a somewhat bold 
venture on the part of one who lives on the east side of the 
Atlantic, and who has only once seen a freshly killed Pacific 
salmon during a short visit to Vancouver in 1893. The 
reasons for my undertaking it are that, so far as I have been 
able to ascertain, the study of salmon scales has not yet made 
much progress in America, that undoubtedly much may be dis- 
covered from them, and that even such inadequate observations 
as I have been able to make may afford most valuable hints to 
others better able to procure materials for the study of the habits 
of the so-called salmon of the Pacific. I say so-called, because with 
one exception, Salmo gairdneri, the Steelhead Trout, the Pacific 
salmon do not belong to the same division of the genus Salmo as. 
the salmon of the Atlantic Ocean. Excepting the Steelhead, 
they belong to the subgenus Oncorhynchus, while our salmon and 
trout belong to the subgenus Salmo. 

I shall have to notice five species of Oncorhynchus which 
breed in the rivers and streams of Western North America. 
They are O. quinnat, or O. tschawytscha, the Quinnat, King, Tyee, 
or Spring Salmon ; O. nerka, generally known as the Sockeye, 
from the sunken appearance of the eyes, and also called the 
Blueback and the Red Salmon ; O. kisutch, the Cohoe, Silver, 
White, or Fall Salmon; O. gorbuscha, the Humpback, so called 
from a peculiar hump which appears on the backs of the males 
at spawning time; and O. keta, the Dog or Chum Salmon, A 
sixth species, O. masu, is found on the Siberian coast and in 
Japan, but I shall not deal with it here, as I intend to confine 
my remarks to the salmon of the Pacific coast of North America. 
It is there that full knowledge of the habits of the Pacific salmon 
is of the greatest importance on account of the magnitude of the 
canning industry, to which every one of the native species 
now contributes its quota. 

When the canning industry was first started in the West, the 
Quinnat only was cared for, but soon the Sockeye was recognized 
asa fish of much greater importance. The value of the remaining 
species has only recently been appreciated. 

It is generally believed that none of the fish of the genus Onco- 
rhynchus that go to the rivers ever return and that all die after 


* Communicated by the SECRETARY. 


HISTORY OF THE PACIFIC SALMON. Diliey 


spawning. But although the upper tributaries of the Fraser River 
are almost unapproachable for some time after the spawning 
season, on account of the numbers of dead and putrefying fish 
which they contain, I do not think it is absolutely proved that 
all the fish do die after spawning. Later on I shall produce 
something approaching a proof that they do not. Most of them 
undoubtedly do die then. Many of our Atlantic salmon are so 
exhausted after spawning that they promptly die. Much more 
so must that be the case with these Pacific salmon, which ascend 
the rivers not for a few tens of miles as our fish do, but for some 
hundreds, or even for many hundreds of miles. The probability 
certainly is that when they go very far from the sea, none of them 
return. But they do not all ascend to extreme distances; and 1 
can see no reason why some of those which have not had far to 
go, or great difficulties to surmount on the journey, should not 
have sufficient strength to recover and to spawn again. I well 
remember the manager of one of the largest canneries on the 
Fraser saying to me that the idea that all the fish died was based 
on the statements of Indians only; that no one else knew any- 
thing about it (I speak of twenty years ago), and that millions of 
kelts might come down the middle of the Fraser with the stream, 
and not a soul be any the wiser. To my mind the fact that no 
kelts are ever seen does not prove their absence. No netting is 
going on when the kelts would come down, so, as no Pacific 
salmon can jump, and as no Pacific salmon has ever been known 
to take a bait in fresh water, it is most improbable that they 
would be seen. 

It was in the hope that some definite evidence might be forth- 
coming upon this point that I first took up the study of the 
scales of Pacific salmon. One certain spawning mark would 
disprove the idea that every fish dies. But even on our Atlantic 
Coast the percentage of fish that are recaptured after having 
spawned is very small; and on the Pacific Coast the percentage 
must, for the reasons mentioned, be much smaller. Supposing 
it to be as high as one in a hundred, which it probably is not, it 
might be necessary to examine the scales of some thousands of 
fish before that one happened to be among them. I have not 
been able to examine the scales of more than a few dozens, but I 
think that I have been so lucky as to find a spawning mark 
on the scales of a large Quinnat. One of its scales is shown 
in text-fig. 95 (p. 574), and I believe that the mark about a 
quarter of an inch from the edge of the photograph is a spawning 
mark, I shall have more to say about it later on when I come to 
deal with the Quinnat salmon in detail. 

As against the general belief that Pacific salmon never survive 
spawning, I have heard the argument that the largest Quinnats, 
which weigh from 50 to 100 lbs., must be very old fish and that 
they must almost certainly have spawned. In text-fig. 96 (p. 575) 
is shown the scale of one which weighed 623 Ibs. It has certainly 
not spawned, and it is also certainly not old. It appears to be 


574 MR. J. A. MILNE ON THE 


only in its fifth year but to have grown very regularly and fas 
both in summer and winter throughout its life. 

As I have to reason largely from analogy, I must now digress 
for a time from my immediate subject to say a little about 
what has been discovered from the scales of our salmon, Salmo 
salar, which is also the salmon of the east side of the American 
continent. 


Scale of a Quinnat (Oncorhynchus tschawytscha) 33+ lbs. Length 415 ins.; girth 
252 ins. Captured at New Westminster, B.C., in 1912. Supposed spawning 
mark shown at S. 


It appears to have occurred to Leuwenhoeck so long ago as 
the year 1696, and to Réaumur in 1716, that the concentric lines 
which are to be found upon the outer surface of the scales of most 
fishes are formed with some relation to the age of the scale, and 
therefore must give an indication of the age of the fish to which 


HISTORY OF THE PACIFIC SALMON. 575 


the scale belongs. But no very close or reliable investigations as 
to how the ages of salmon might be read from their scales 
were made until Mr. H. W. Johnston took up the study 
a few years ago. Although I am not actually quoting from 
anything he has written, I think his authority for every- 
thing I am going to say, unless I may quote someone else, will 
be found in one or other of his contributions to the Reports of 
the Scottish Fishery Board. (See Parts II. of the Reports for 
1904, 1906, and 1907.) 


Text-fig. 96. 


Scale of a Quinnat (O. tschawytscha) 623 lbs., 6. Length 502 ins.; girth 31 ins. 
Captured at New Westminster, B.C., in 1912. 


A salmon scale is divided into two areas, one, the anterior and 
larger part, being enclosed in a pocket in the skin, to which it is 
loosely attached; the other, the posterior area, being the only 
part of the scale which we see while it is still attached to the 


576 MR. J. A. MILNE ON THE 


fish. The anterior part is covered by a mass of concentric lines ° 
or ridges, from the relative positions of which much may be 
learned. ‘The posterior part is almost without lines. This 
formation is common to the scales of all members of the 
genera Salmo and Oncorhynchus. The whole scale is covered 
by a membrane and grows with the fish. When the fish—and 
with the fish, the scales—grows slowly the concentric lines, which 
seem to be produced at a fairly uniform rate as to number, are 
situated closely together ; when they grow quickly the lines are 
placed further apart. When the fish does not grow at all, the 
scales also cease to grow, and no lines are added to their surfaces. 
The scales first appear upon young salmon as minute bony plates 
under the skin. They may first be noticed when the fish is 
about three months old, and about #? inch in length (Vogt, 
‘Embryologie des Salmones,’ 1842, Klaatsch 1890, and Dahl, 
‘Age and Growth of Salmon and Trout in Norway,’ 1912). 
When about four months old and 1; imches long the little fish 
may have from 2 to 5 rings round their scales ; and when about 
eight months old, that is just before the winter comes on, there 
may be, roughly speaking, from 8 to 20 lines already visible on 
each of their scales. Within small limits the number of lines 
varies in different scales even from the same fish. We know that 
the yearlings when kept in ponds feed well and grow rapidly 
during the summer months, and that as winter approaches they 
take less nourishment, finally at times fasting completely for 
three or four days on end. We also know that their growth 
progresses in proportion. This mode of life is clearly depicted 
on their scales. Surrounding the nucleus, the lines formed in 
summer are at some small distance from each other, and each 
line can usually be traced right round the scale. The lines 
formed in winter are much closer together and are usually more 
numerous on the anterior portion of the scale. 

When the growth of the next year begins in the spring, the 
lines first formed are again wider apart and generally continuous 
all round the scale, and, later on, the winter formation of the 
first year isrepeated. Thus there is formed an area of well-spaced 
lines followed by a band composed of lines very close together. 
This band is usually most noticeable in front, and is followed by 
another area of open lines and asecond band which is probably not 

uite so well defined as the first. The accompanying photograph 
(text-fig. 97) of the central part of the scale of a Steelhead Trout 
(S. gairdnert) well illustrates the formation. The line A points 
to the completion of the first winter band, B of the second winter 
band, © of the third, and the end of the line D shows the point 
in the next year when the smolt left the river and commenced to 
grow rapidly in the sea. 

Tn the sea growth proceeds as before, but at a much increased 
rate, and the difference between fast summer and slower winter 
growth continues to be apparent on the scales. 

Fish that are born, and spend all their lives in the sea, show 


HISTORY OF THE PACIFIC SALMON. Bite 


annual winter bands like the others ; but, except for these bands, 
the spacing between the lines shows no sudden increase due to 
any abrupt change in the mode of life. Compare text-fig. 98, 
the photograph of a scale of a large haddock, with the other 
photographs illustrating this paper, and the difference in the 


Text-fig. 97. 


tis) 
WU 


ag 


is 
: Aye a ge PS ; 
ae. The amie a ae 4 ted atte ae is ee 


Centre of scale of a Steelhead Trout (S. gairdneri) from the Fraser River, much 
magnified, showing three winter bands before migration to the sea. (For references 
see the text, p. 576.) 


i. ed » nee a et 
eae ry 
: . Hi 


centres will be at once apparent. I wish to emphasize 
this difference because the Pacific salmon scales, if we except 
those of O. keta, the Dog Salmon, show a sudden change 
of growth just as do those of the true salmon; and this, to my 


578 MR. J. A. MILNE ON THE 

mind, at once disposes of the idea that these fish go down to the 
sea almost as soon as they are hatched and commence feeding 
only when they get there. 


Text-fig. 98. 


iis 

uy 
neal 
i : 
i ei 
ANE 
aes 


Scale of a large Haddock (Gadus eglefinus). XX about 40. 


Mr. Johnston kindly gave me some of the scales taken from 
salmon which had been marked as smolts in the estuary of the 
Tay in the early summer of 1905. The smolts were then on 
their way to the sea, and the marks were still attached to the 
fish when subsequently recaptured as salmon. ‘Thus the exact 
times spent by them in the sea were known, and it was found, 
as had been expected, that the summer and winter bands on the 


HISTORY OF THE PACIFIC SALMON. 579 


scales agreed with the known times. Text-fig. 99 is a scale from 
one of these fish which was captured in the Tay nets on May 7th, 
1907. It had been marked as a smolt when the scale had grown 
to the point C, and when the fish was just over two years old. 
The winter band formed in the winter 1905-6 is apparent 
between the points marked D, and the band formed in the winter 


Text-fig. 99. 


Scale of 124 Ib. salmon (Salmo salar) caught in the Tay nets on May 7th, 1907. 
Previously marked as a smolt in May 1905. (For references see the text, supra.) 


1906-7 between the points E. Outside the last band is seen 
the quick growth made in the summer of 1907 previous to 
May 7th, when the fish was netted. Summer growth is the term 
used in distinction to winter growth. It does not mean growth 
taking place only in June, July, and August, our summer months. 
Tt evidently starts and finishes at varying times, the probable 


Proc. Zoou. Soc.—1913, No. XX XIX. 39 


580 MR. J. A. MILNE ON THE 


limits being about the middle of March and the middle or end 
of September. 

The foregoing remarks may suffice to explain the regular 
markings on salmon scales, and how they show the age and time 
of migration of the salmon. Scales, however, may show much 
more. 

Mr. Johnston noticed that the scales of the older fish some- 
times seem to have stopped growing, and become jagged and 
broken at the edges, and then to have gone on growing again. 
When this has happened the lnes of the new growth do not 
exactly follow the contours of the lines of the old growth, and 
thus a clear mark is formed round the anterior portion of the 
scale, often also apparent as a thickening round the posterior, 
the unlined, portion. Further, it was noticed that when such a 
mark occurred the number of lines between the winter bands on 
each side of it was frequently far from normal. ~The normal 
number differs somewhat in different fish, and also within certain 
limits on scales from different parts of the same fish; but never- 
theless it is fairly constant, and any wide departure from the 
usual state of things required explanation. A reason for the 
mark suggested itself to My. Johnston from the known fact that 
salmon, whether or not they take food in fresh water, do not 
take it in sufficient quantity, after the smolt stage is passed, to 
nourish them. Therefore neither salmon nor their scales can 
grow in fresh water. It was further noticed that most scales 
taken from kelts---that is to say from salmon which, having 
spawned, have not yet returned to the sea—were broken and 
torn at the edges. From these facts Mr. Johnston argued that. 
the sort of marks illustrated at C and D on the accompanying 
photographs (text-figs. 100 & 101) showed that the fish bearing 
them on their scales had entered fresh water and stopped growing 
there; that they had spawned and become shrunken after 
spawning, so that, from mechanical reasons, the scales being im- 
bricated, 7.¢e. overlapping each other like tiles on a roof, had become 
frayed at the edges; and, lastly, that on the salmon’s return to 
the sea the new growth had started again evenly round the scale 
and thus left the mark. Spawning operations mean a winter 
at least spent in fresh water and, therefore, if the fish enters 
a rviver early in the year there will be fewer than the normal 
number of lines between the winter bands formed on each side 
of the spawning mark. If it comes in early in the spring there 
will be none in that year, and in the year following there will 
be fewer than usual on account of the fish taking some time 
after its return to the sea to make up lost condition before it 
starts again to imerease in size. Of course, the later the 
salmon came into the river the more lines would have been 
formed behind the mark, and the later it returned to the sea the 
fewer would there be in front of it. 

Besides the irregular arrangement of the lines on the scales 
when a fish has spawned, there is a thickening round what was 


HISTORY OF THE PACIFIC SALMON, 581 


the edge of the scales at the time the salmon left the sea, and 
this thickening is apparent both on the anterior and posterior 
parts of the scales if the edges have not become too much 
worn during the stay in fresh water. JI account for it by 
supposing that the materials which go to form the scales 
are still being secreted in fresh water, and that as the skin 


Text-fig. 100, 


Scale of 33 lb. Salmon (Salmo salar) caught in the Tay in August 1903. 


This fish had spawned twice, and the marks are shown at C and D. 


pockets have ceased to grow they can be deposited only on the 

scale edges already formed. The cells from which the concentric 

lines on the upper surface are evolved are already dead, except 

close to the periphery of the scale (Klaatsch, ‘ Zur Morphologie 

der Fischschuppen,’ 1890, and Stuart Thompson, ‘ Journal of the 

Marine Biological Assoe.’ vol. vii. no. 1, 1904). This in itself 
3o™ 


582 MR. J. A. MILNE ON THE 


would account for the marginal thickening if the cells still remain 
active in fresh water and the scale cannot expand, 


Text-fig. 101. 


eee 
WI 
Wee 


iy 


A, 7. 


Hix 


GALT 
i 


Scale of a 334 lb. Salmon (Salmo salar) caught at Port Gordon, Aberdenshire, on 
August 19th, 1908. Previously marked as a kelt on the Deveron, March 23rd, 
1908. This fish had also spawned in the previous year, and the two marks 
are shown at Cand D. The original scale has been lost at the end of the first 
summer in the sea, and its position filled by a new scale without lines. 


On both text-figs. 100 & 101 two spawning marks are to be 
seen, but the figures illustrate what I have just said as well as if 
they had but one mark. Scales which like these show two marks 
are far from common. The central part of text-fig. 101 is 
without lines, so that the parr scale does not show at all. ‘This 
is because the original scale was lost by the fish towards the end 
of its first summer in the sea, and the place occupied by it in the. 
skin pocket was filled by a new scale which, the line producing 


HISTORY OF THE PACIFIC SALMON. 583 


cells having died, is devoid of concentric lines. In both text- 
figs. 100 & 101 the first winter band formed in the sea is shown 
between the ends of the lines AA. The second winter band is 
between the lines BB. The first spawning mark in each case 
is at the end of the line marked C and the second spawning mark 
at D. Text-fig. 100 is a scale from a 33 ]b. salmon caught in 
the Tay in August 1903. It can be seen, from the few widely 
spaced lines between the band BB and the spawning mark C, 
that when it first spawned it was a summer fish about 43 years 
of age. After spawning the scale had become much worn, so 
much so that, except in ” front, not only the summer growth but 
the band BB had disappeared, and the new growth formed after 
the fish returned to the sea can easily be distinguished, its lines 
being again continuous round the scale. In foxt- fis. 101 the 
winter band of the 5th year had begun to form before the 
spawning mark C, so it was evidently a late autumn salmon. 
In text-fig. 100 the second spawning mark is indicated by the 
line D. The fish must have returned to the sea after the first 
spawning early in the year, and the lines between the two 
spawning marks show it to have come in to spawn the second 
time late in the summer of the same year. The following summer 
it was coming back to spawn a third time when it was captur ed, 
The thickening on the posterior part of this scale, due to both 
spawning periods, is clearly visible in the form of dark lines. 
In text-fig. 101 the second mark D, like the first mark C, denotes 
an autumn fish. It is not a very clear spawning mark, but I 
have chosen it for that very reason, because it is so like the mark 
on the scale of the Quinnat which I believe to have spawned 
(see text-fig. 95, p. 574) and because I have undoubted evidence 
that mark D on text-fig. 101 is a true spawning mark. The fish 
was caught as a kelt, weighing 222 lbs., and marked by my own 
samekeeper, Peter Bowie, at “Netherdale on the River Deveron, 
on March 23rd, 1908. Tale put in its fin one of the Seotbish 
Fishery Board labels numbered 3613 B. The scales then showed 
only the first spawning mark C. The fish was recaptured as a 
clean salmon, weighing 333 lbs., in a bag-net at Port Gordon on 
the Aberdeenshire coast on August 19th, 1908, after only five 
months’ interval. The mark D thus certainly represents the 
second spawning period during which my keeper caught and 
marked the fish. 

Many other instances have occurred in which fish that have 
been marked with distinctive numbers as kelts have been caught 
again, and in every case they have proved the truth of Mr. John- 
ston’s theory by showing a spawning mark on their scales. On 
the other hand, no fish that one knows from its age could not 
have spawned has ever been found to bear quite the same sort 
of mark, although various marks indicating sudden checks in 
feeding may appear. 

I have entered somewhat fully into the above description of 
spawning marks, because I hope that now many scales of Pacific 


584 MR. J. A. MILNE ON THE 


salmon will be examined upon the chance of finding one. Much 
can be done with an ordinary high-powered pocket- -lens, and for 
close observation only a very lor powered microscope is either 
necessary or desirable. I find a 14-inch objective powerful 
enough for all purposes. One wants to be able to see as much as 
possible of the scale at the same moment. 

Mr. Knut Dahl (‘ Age and Growth of Salmon and Trout in 
Norway, London, 1912) has shown that the size of a fish at 
any period of its life may be deduced from its scales. 

A fish does not change its seales. From the time they form 
they are retained through life unless removed accidentally. If a 
fish does by chance lose one it is replaced by another of the same 
size and shape but without the concentric lines—see text-fig. 101. 
It follows that as the fish grows, but remains covered by the same 
number of scales occupying the same relative positions, the scales 
must grow with the fish, and the growth of each scale be pro- 
portionate to the growth of the fish. If, then, the protected 
part of the scale be measured along its length from the centre to 
the anterior edge and again from the centre (by which I mean 
the nucleus) to, say, the first winter band, the lengths of the 
scale at these two points will be proportionate to the lengths of 
the fish at corresponding ages. An example will make this more 
clear. Suppose we have a salmon 75 cm. long and that one of 
its scales, when magnified, measures 55 mm. from the centre to 
the anterior edge, and 11 mm. from the centre to the point at 
which the rapid growth, consequent on migration to salt water, 
is seen to begin. Then as 55 is to 11 so is 75 cm. to the length 
of the fish when it entered the sea. The smolt was therefore 
15 cm. (or just under 6 inches) long. Another scale from the 
same fish magnified to the same degree, might be only 45 mm. 
long, but then the other measure would be found to be 9, and the 
sum would work out just the same. 

Dahl has proved his theory by measuring the scales of 
hundreds of salmon and trout from different rivers. He has 
always found that the actual average lengths of the fish in 
different rivers at various ages, agree almost exactly with the 
lengths calculated by him from the scales of the older fish. 

I have been trying to apply this method of measurement to 
the scales of Pacific salmon, and I will give the results when I 
come to deal with each species separately. First, however, 
I must make some criticisms on what I have set out above as 
my understanding of the claims made by Dahl. I tried to 
check his theory on the scales of two salmon which had been 
marked as kelts and subsequently captured as clean salmon. 
Photographs of both of these scales form illustrations to 
Mr. Johnston’s second paper on salmon scales in the 25th 
Annual Report of the Fishery Board for Scotland. The lengths, 
on the occasion of each capture, are given elsewhere in the 
24th and 25th Reports, so it is known that the salmon marked 
No. 9194 was 27 inches long when taken as a kelt, and 31 inches 


HISTORY OF THE PACIFIC SALMON. 585 


long when recaptured. The other fish, marked 1180, was 
264 inches long as a kelt and 36 inches long when recaptured. 
The measurements on the photographs of the scales from the 
centre to the anterior edge, and from the centre to the spawning 
mark are 130 and 116 for No. 9194, and 198 and 175 for No. 1180, 
the unit of measurement being ~;th inch. This would give the 
kelt measures as 272 inches and 32 inches respectively. The 
former measure is only 2 inch wrong, but the latter is nearly 
6 inches wrong, and shows either that the scale is abnormal or 
that Dahl’s system of measurement is not applicable to a fish 
that has spawned. 

Another criticism is that measurements from several different 
scales of the same fish seldom all agree exactly, and I have there- 
fore come to the conclusion that it is very unsafe to rely on the 
measurement of one scale in estimating the size of the fish at 
various ages. The reason may be that it is not easy to recognize 
either the exact centre of growth, or the exact limits of the 


Text-fig. 102. 


JU 


Quinnat (O. tschawytscha). 25 bs. 13th October, 1911. Shuswap, 
South Thomson River. 


The black lines show the variations in the long axis of the scale. 


various bands, but Ido not think that the scales grow quite 
equally. Still the idea seems so well founded in theory, and to 
have worked out so exactly in practice on a large scale in Norway, 
that I believe I may consider myself justified in drawing con- 
clusions from the average measurements of a considerable number 
of scales taken from the same fish, and whenever possible, I have 
measured 20 scales. A smaller number might suftice in the case 
of a true salmon, but the scales of the Pacific salmon are more 
difficult to read. As a rule, the limits of the bands are less 


586 MR. J. A. MILNE ON THE 


clearly defined, and the change of growth, which I can only 
suppose to be due, as in the case of the true salmon, to the 
commencement of sea life, is more gradual and not so well marked. 
In addition to this, the long axis of the scale frequently shows 
more than one change of direction, the scale apparently being 
liable to get turned round in the skin-pocket. The accompanying 
photograph (text-fig. 102) will show my meaning. I have not 
measured such scales as these. 

To come now to what is known, and to what I think I may claim 
to have found out about the various species of Pacific Salmon. 


THE SockEYE. (Oncorhynchus nerka.) 


First I take the Sockeye, because it is the mainstay of the 
canning industry of the West. The number of Sockeyes canned 
on the Pacific coast in 1909 must have attained to the enormous 
total of about 50 millions. The exact weight was214,980,448 lbs., 
or nearly 100,000 tons. 


Text-fig. 103. 


Sockeyes (Oncorhynchus nerka) running up Scotch Creek, a tributary of the Fraser 
River, British Columbia. (Photograph reproduced by kind permission of 
My. Frank Parry.) 


The Sockeye is not much in evidence south of the Columbia 
River, in which it is known as the Blueback, but is plentiful from 
there the whole way north to Bering Sea. In Alaska it is known 
as the Red Salmon. The chief Sockeye stream is the Fraser 
River, which these fish ascend to spawn in countless myriads. 
The feeding-ground of the Sockeyes is somewhere far out in the 
Pacific, and the fish seem to cease feeding before they approach 
the coast, for even when caught in the sea near the coast their 


HISTORY OF THE PACIFIC SALMON. 587 


stomachs are invariably empty. This, however, may be from 
lack of means rather than from want of will. When one hears 
of a shoal of fish, seven miles broad and of unknown length, 
heading for the land through the Straits of San Juan, one can 
well imagine difficulties in the commissariat. Part only of this 
big shoal enters the Fraser River, the remainder moves on up the 
coast of British Columbia. The photograph (text-fig. 103), which 
I give by kind permission of Mr. Parry, late of Granite Creek 
Hatchery, shows a detachment on its way up Scotch Creek, a 
subtributary of the Fraser River, about 300 miles from the sea. 

The big run of which I have been speaking strikes the south- 
west coast of Vancouver Island in J uly and August, coming from 
the north-west, but a few Sockeyes run as early as April. In 
the Fraser River itself the main run is in August, and some con- 
tinue to come in until October. In the far north of Alaska the 
main run is as early as June, which goes some way towards 
showing that the feeding-grounds are in the north, unless a 
natural instinct to get spawning over before the winter sets in has 
determined the habits of the fish fr equenting the most northerly 
rivers. In the Fraser district spawning begins in August, and 
may go on until November; spawning takes place only in streams 
running into or out of lakes. In this district, from which 
most of my information and all my specimens have come, several 
hatcheries have been established, which in 1910 liberated 
134,639,200 Sockeyesin British Columbian waters. Besides these 
4,544,825 were liberated from U.S. hatcheries in Puget Sound, 
and 257,021,790 in Alaska. 

Tam indebted to Mr. W. J. Sim, who was employed at Granite 
Creek for some years, for the information that the fry are liberated 
from the Canadian hatcheries as soon as the yolk-sac has been 
absorbed, at a period of the year which varies from January to 
April, according to the date of spawning. The main liberation is 
about the first week in February. In that district the fry may 
remain in the creek for two months or less; then they move into 
Shuswap Lake. The exodus from the lake takes place from June 
to September. The fry are supposed to be under one year old at 
the time of their exodus from Lake Shuswap, but no one really 
knows how long they may have remained in this, or in any other 
lake, and it is admitted by some observers that they may be in 
their second year. Dr. Greene, writing in the Bulletin of the U.S. 
Bureau of Fisheries, vol. xxix. 1909, on the Migration of Salmon 
in the Columbia River, and quoting Evermann, says the fry 
begin their seaward journey not sooner than September of their 
first, and not later than July of their second year. That means, 
I presume, that they leave the lakes when from nine to eighteen 
months old. But others, e. g. Rutter, Bulletin U.S. Bureau of 
Fisheries, vol. xxii. 1902, p. 102, say that they begin to descend the 
rivers as soon as they can swim, and reach the sea in about three 
months. My. Sim says he is ori that there are no fry in the 
rivers after September, and that they could not possibly avoid 


588 MR. J, A. MILNE ON THE 


being swept down with the current as soon as they leave the 


lake. 
The question as to the age at which the fry enter the sea is 


one which scale-readings should easily settle. So far, I have been 


Text-fig. 104. 


3 


ALSk. 
Sari 
nn 


Nv 


RA 


Scale of Sockeye (O. nerka), $, 35 lbs. Length 202 inches; girth 103 inches. 


A=end of first year’s growth. B=nmigration to sea. 
C=end of second winter, D=end of third winter. 


unsuccessful in my attempts to procure any specimens of fry of 
known ages for the purpose of seeing how their scales do actually 
grow, but I give some illustrations of Sockeye scales from full- 
grown fish (text-figs. 104-106). I have no doubt but that each 


HISTORY OF THE PACIFIC SALMON. 589 


one of these Sockeyes spent a year at least in fresh water, and 
the first of them pr obably not less than 15 months. I can 
conceive of no other way in which scales with centres similar to 
these could have been formed. They are the only Sockeye scales 
which I have from measured fish, but I have others from six fish 
of which I know only the weights, and a great many more from 
Sockeyes that unfortunately were neither weighed nor measured. 
All have similar centres. 


ary 
ni a Py A 
Scale of Sockeye (O. nerka) 2. 5; lbs. Length 24+ inches; girth 121 inches. 
A=end of first winter. B=migration to sea. 
C=end of second winter. D=end of third winter. 


The Sockeye from which the scale, shown in text-fig. 104, was 
removed, was caught last August at New Westminster, near the 
mouth of the Fraser River. It weighed 3} lbs. and measured 
202 inches in length and 104 renee in birth when captured. 
I have measured 20) scales from this fish, the aver age length of 
the enlarged images to which [applied the measure being 33 mm. 
from the centre to the anterior edge. The other average 
measurements to the points denoted by the lines drawn on the 


590 MR. J. A. MILNE ON THE 


figure were 5 mm., 6°5 mm., 10 mm., and 20 mm., and the lengths 
of the fish at the various ages indicated work out as follows :— 
At the end of the first year 2? inches; when it entered the sea 
34 inches, at the end of the second year 6 inches, at the end of 
the third year 124 inches, and when caught it was as stated 
202 inches. The points indicated by the lines drawn on text- 
fig. 104 are calculated from the averages of the 20 scales, and 
they appear to coincide absolutely with the points I should have 
marked from examination of this scale alone. 


Text-fig. 106. 


B 
aie ice a A 


Scale of Sockeye (O. nerka), $.7 lbs.. 26% inches long; girth 144 inches. 


e 5 , 
A=migration to sea at end of Ist year. Bend of 2nd year. C=end of 3rd year. 


The next scale, text-fig. 105, is from a Sockeye caught at the 
same place and time. In the illustration the parr or fingerling 
scale is not very clear, but the examination of 20 scales enables 
me to put the lengths of the fish at the various points and times 
indicated by the lines drawn on the picture as follows :—1st year 
2% inches, entered the sea during the second year when 3 inches 
long, was already 9 inches long at the end of the 2nd year, 
193 inches long at the end of the 3rd year, and, as was known, 


HISTORY OF THE PAGIFIC SALMON. 591 


243 inches at the time of capture near the end of the fourth 
summer, 

The scale of a Sockeye, also taken at New Westminster in 
August, is shown in text-fig. 106. In the case of this fish, the 
entry into the sea seems to have coincided with the end of the 
first year’s growth. That is to say, it was some time during the 
winter ee 10. The readings from 20 scales show it to have 
been then 2? inches long. At the end of its second year it was 
113 inches long, at the end of its third year 217 inches, and when 
coq ed 264 inches. 

The great difference in growth of the three fish in the second 
year 1s very noticeable, but it seems quite natural, and goes far 
to confirm the accuracy of my reading of the parr scale, when it 
is observed that the fish which appears to have spent the whole 
of its second year in the sea has grown most, and that the one 
which spent the longest part of it in fresh water has made the 
least progress. 

It is further to be noticed that these fish all returned to spawn 
when of the same age, namely at the end of their fourth year. I 
have the scales of nine Sockeyes from the Fraser River. They 
varied in weight from 37 to 84 lbs., but all are in their fourth 
year. In British Co imeealaken the Sockeyes weigh from 3 to 10 lbs. 
Fish up to 17 lbs. have been caught, but over 10 lbs, they are very 
rare. I therefore seem to have got specimens of all the average 
weights. That they are all of the same age may be an accident, 
but much more probably it is not, and if it is not an accident it 
is a fact of the utmost importance to the Fraser River District. 

In the Fraser River the run of Sockeyes in every fourth year, 
the year after leap year, is almost six times as large as in any of 
the intervening periods of three years, and this has been the case 
as long as records go back. Whether the fourth years have shot 
ahead of the others, or whether all years were once equal to them 
will never be known, but most likely adverse breeding seasons in 
the intermediate years have gradually diminished the stock. If 
it proves upon further examination of scales that every Sockeye 
returns to breed in its fourth year im this district, it might well 
be centuries before the breeding stock, once diminished, increased 
to its original numbers. British Columbia has now only one 
season out of four up to what might be the mark; and, if I am 
right, the inference is obvious that the efforts of the hatcheries 
should be devoted chiefly to the collection of ova in the lean years 
even if they have to import them from other districts. In the 
springs of 1906 and 1910, the years following the last two bi 
years, 100,479,000 and 105,312,500 Sockeyes were liberated from 
British Columbian hatcheries. In the intermediate years the 
numbers were only 36,965,900; 51,855,200; and 41,909,500. 
My contention is, that if it is humanly possible the numbers 
hatched out in the seasons following the lean years should be 
made even greater than after the fat years. Some people may 
say, “ But how do we know that if we turn out fry in the Fraser 


592 MR. J. A. MILNE ON THE 


River or its tributaries, they will return there when they grow 
up?” The very fact that it is in that district alone that three 
years out of every four are lean years shows that the fish return 
to their own river. If it were not so, the shortage in these years 
would have gradually spread itself over a wider area. It seems all 
the more wonderful that it has not done so, when it is remembered 
that the big shoal which comes in in the summer is by no means 
solely composed of fish making for the Fraser. The facts further 
appear to show that not only do the fish return to the Fraser, 


Text-fig. 107. 


Scale of Sockeye (O. nerka) removed after spawning. 


but to the very creek in which they were hatched, for how 
otherwise can one account for many of the creeks which are full 
of spawning fish in the big years being always absolutely un- 
tenanted in the years between them. Salmon marking in our 
own country has now shown almost conclusively that when the 
access to a river is unimpeded the salmon born in it invariably 
return to it. The fish that have been marked in one river and 
subsequently recaptured in another have all been-from some 


HISTORY OF THE PA@IFIC SALMON. 593 


comparatively small stream, or from some river with a bad mouth 
which they might well find difficulty in entering again when 
they wished to do so. 

The value of hatcheries is a much-disputed point on which I 
will not express any opinion. For the moment I am content 
to assume that they are of some value on the Fraser, and that 
being so, to point out how they can best be utilized. But I will 
say that if the falling off in the catches which has been apparent 
since about 1900, when each year is compared with the fourth year 
before it, proves to be more than a mere temporary fluctuation, 
hatcheries alone will certainly not remedy the evil, and it will 
be necessary for a time at any rate to restrict the number of 
fish that are permitted to be caught. 


Text-fig. 108. 


Piece of skin taken from a Sockeye (O. nerka) after spawning. 


I should mention that I have scales from Sockeyes in their 
fifth year, but they came from the State of Washington, not 
from the Fraser. 

In text-fig. 107 is shown the scale of a Sockeye which weighed 
6; lbs. and was caught at Morris Creek near Lake Shuswap, 
about 300 miles from the coast. So far as I can judge it is quite 
typical, and its interest is in showing, from the worn condition 
of its edge, that if a Sockeye ever returned to the sea after 
spawning and was caught again, a clear spawning mark should be 
apparent on its scales. The scales le so far apart, and are so 


594. MR. J. A. MILNE ON THE 


flexible, that unless one saw how much they may become worn 
one would hardly believe it possible. 

It will, J think, be news to most of those employed on the 
hatcheries that the spawning Sockeye possesses scales at all. 
Last year when I asked for scales from a hatchery, I was told 
that it was well known that the Sockeyes absorbed all their 
scales as food in the course of their run up the river and never 
had any when they reached the spawning beds. IJ argued the 
point, and the hatchery people ultimately sent me the piece of 
skin shown in text-fig. 108 taken from a Sockeye that had 
spawned, to convince me that I was wrong. The scales were 
there, but are by no means so clear in reality as they are in 
the photograph. They are very deeply imbedded in the skin, 
and, in order to remove some of them for examination, it was 
necessary first to soak it well and then to pull and stretch it so as 
to open the skin-pockets. After that the scales were still invisible 
and adhered closely to the skin which formed the top of the 
pockets. It was possible, however, to remove them with a pair 
of foreeps without much further difficulty. So even those who 
have handled spawning Sockeyes for years may be excused for 
thinking them to be without scales.—How many fishermen in 
this country know that eels are covered with scales @ 


THe Quiynat (Oncorhynchus tschawytscha). 


The known range of the Quinnat on the American Coast is from 
the Ventura River in California to Norton Sound, Alaska. It 
probably really extends into Aretic regions. ‘The Quinnat is also 
known as the Chinook Salmon, the King Salmon, the Tyee, the 
Red or the White Spring Salmon, and the Black Salmon. The 
flesh is generally of a deep salmon-red colour, but in the south of 
Alaska and down to Puget Sound, sometimes as many as one- 
third of these fish have white flesh. Sometimes one half of the 
body is red and the other white, and sometimes the flesh is 
mottled. The white-fleshed fish are of little use. All the scales 
sent me and marked as from either Red or White Spring Salmon 
are from fish which have spent three winters, and a considerable 
part of the feeding season following the last winter, in the sea. 
Analogy from the true salmon would lead one to suppose that 
the white and mottled fleshed Quinnats might be fish that had 
spawned, but I can find no trace of a spawning mark on any of 
their scales. It has been suggested that the Red and White 
Spring Salmon are distinct varieties and that the specimens 
with mottled flesh are crosses, but upon this point I can offer no 
opinion. Differences in food might account for much, as in the 
case of the Brown Trout (Salmo fario). 

The name Black Salmon arises from the colour assumed by the 
Quinnat at spawning time. 

In 1909 the weight of Quinnats canned on the Pacific Coast 
was 12,640,344 Ibs. 


HISTORY OF THE PACIFIC SALMON, 595 


As a rule Quinnats weigh from 18 to 30 lbs. in British 
Columbia, but much larger specimens are taken, and in Alaskan 
waters they have been caught over 100 lbs. in weight, and average 
23 lbs. in some seasons. 

Quinnats are very powerful swimmers and ascend fast-running 
streams in preference to others, making for the head waters, and 
sometimes running up for many hundreds of miles. 


Text-fig. 109. 


Seale of Quinnat (Oncorhynchus tschawytscha). 15 lbs. Length 314 inches ; 
girth 193 inches. New Westminster, August 1912. (For references see text, p. 596.) 


Most of the principal rivers have spring and autumn runs, 
and some of them a summer run also, so Quinnats may be said 
to run from January until October. 

The scales I have examined show the running Quinnats to be 
in their fourth and fifth years, those over about 30 lbs. in weight 
being a year older than the others. Numbers of them are 


Proc. Zoou. Soc.—1913, No. XL. 40 


596 MR. J. A. MILNE ON THE 


spawned in the hatcheries, from which it was estimated that 
90,740,472 were liberated in 1910. 

I have already shown photographs from scales of Quinnats 
which eee 333 lbs. and 623 lbs. (see text-figs. 95 and 96, 
pp. 574 & 575) and 1 now append another’ (text-fig. 109) from a 
fish which weighed 15 lbs. The parr or fingerling ; stage of each 
scale shows that the fish spent a year (A) and the greater part of 
a second year (B) in fresh water. 

I have examined scales from eighteen Quinnats varying in 
weight from 10 to 623 lbs., and all show about the same duration 
of the freshwater stage. 

When the little fish enter the sea they seem to be already 
larger than the Sockeyes, and ae measure from 4 to 6 inches in 
length. The lengths of the 623 Ib. fish at various ages, deduced 
from the qneAS SMAI: of olen seales, were as follows :—At the 
end of the first year 22 inches, when it entered the sea in its 
second year 44 inches, at the end of the second year 133 inches, 
at the end of the third year 25 inches, at the end of the fourth 
year 39 inches, and when captured it measured 503 inches. The 
scale illustrated (text-fig. 96) agrees exactly with these averages 
up to the end of the third year, but the position of the fourth 
winter band on it would show the fish to have been about 
40 inches long at that time instead of 39 inches, if that scale 
alone were to be depended upon. 

The lengths of the 15 1b. fish,as shown from 22 scales, were, at 
the end of its first year 34 inches, when it entered the sea 
5 inches, at the end of its second year 13 ; inches, at the end of its 
third year 26 inches, and when captured it measured 312 inches. 
Text-fig. 109 shows the points from which the measurements were 
taken, the scale being exactly an average one. 

The measurements of the 337 Ib. fish, averaged from 16 scales, 
were, at one year 37 inches, when it entered alae sea 43 inches, at 
the end of the second year 127 inches, at 3 years 213 inches, at 
4 years 31 inches, and when captured 412 } inches. 

The above seem to be about the average lengths of Quinnats at 
the ages given. None of the twelve specimens I have measured 
depart far from them.* 


The Spawning Mark. 


On the scale of the 33} lb. Quinnat (text-fig. 95, p. 574), 
check in growth (S) may be observed beyond the fourth winter 
band (KE). It is equally clear on every scale of this fish, and 
IT think it must be a spawning mark. One might expect 


* Note:—Since this paper was written I have ascertained that in the Natural 
History Museum, South Kensington, there are specimens of Quinnat parr, about 
4 inches in leneth, which came from a lake near the head weters of the Fraser River. 
I have had the priv elege of examining the scales of one of these specimens which 
measured about 44 inches. The first winter band is quite clear, and five lines 
formed in the second summer are apparent. 


HISTORY OF THE PACIFIC SALMON, 597 


a better defined scar, but Quinnat scales are very thin and 
flexible, and besides are well protected by a great thickening of 
the outer skin at spawning time, so they may never become much 
worn at the edges. ‘That they may not have become much worn 
up to the actual time of spawning is shown by text-fig. 110, a 
photograph of a scale removed from a 12 1b. Quinnat after it had 
spawned in the South Thomson River, about 300 miles from the 
sea. That scale not only sustained the wear from the shrinkage 
of the fish owing to lost condition between the time at which it 
left off feeding, which the lines following the last winter band 


Text-fig. 110. 


7, 


1g” naar . WAX 
Liat pak 
Midis GC 2) 


SP 4 


<x heey 
SS e 
A rt Sl 


Scale of 12 1b. Quinnat (O. tschawytscha) from South Thomson River. 


Fish speared after spawning, 13th October, 1911. 


show to have been quite early in the year, until it was speared 
for me under the supervision of Mr. John Brown of Celista, B.C., 
on the night of 13th October, 1911; but the skin was sent to me 
dried and folded, and I removed the scales from it myself. As this 
scale shows so little sign of wear at the edge, it seems to me quite 
conceivable that the mark S on text-fig. 95 is a spawning mark. 
Tf it is not, Tam unable to suggest any other explanation of it. 
Compare text-fig. 95 with text-fig. 109. These scales were 
taken from fish caught at the same place at the same time. There 
are more lines after the last winter ring on the former than on 


40% 


598 MR. J. A. MILNE ON THE 


the latter, and that in spite of what is a very obvious check in 
growth. ‘The check clearly lasted long enough for the edges of the 
scales to become somewhat worn, for the lines following it do not 
at all points exactly follow the contours of those immediately 
preceding them. ‘This fish cannot in these circumstances have 
grown faster than its fellow, which was feeding all the time, 
and yet there are 20 lines as against 12. If, however, it 
spawned it added the twelve lines between the winter band and 
the scar in the early part of the year 1911, and the eight lines 
outside the scar in 1912. It would have spent the early part of 
1911 in the sea, then come in to fresh water to spawn, and 
remained there after spawning for some little time. It would 
then have returned to the sea in poor condition and possibly not 
very early in 1912, so that by the time it had made up condition 
and started to grow again it would add but another 8 lines or so 
to its scales before the spawning instinct again induced it to seek 
fresh water, and thus brought it to the place of its capture. 


Text-fig. 111. 


Quinnat Skin 
10 lbs. Nat. Size 


Piece of skin taken from a Quinnat (O. tschawytscha) after spawning. 


Compare this mark also with the outer spawning mark on 
text-fig. 101, and note how like they are. 

A Cutemet of about 30 lbs. is just the size on which I think a 
spawning mark is most likely to be found. The larger fish, like 
the 623 pounder, a scale of which is shown in text- fig. 96, Tere 
most probably attained their size on account of ane feeding 


HISTORY OF THE PACIFIC SALMON, 599 


instinct being more pronounced than the sexual. Scale reading, 
so far as it has gone, has shown that our own very large salmon 
are all maiden fish. 

Text-fig. 111 shows a piece of Quinnat skin taken from a fish 
that had lately spawned. 


Tur Conor (Oncorhynchus kisutch). 


The salmon with which J shall next deal is the Cohoe, also 
known as the Silver Salmon and the Fall Salmon. The photo- 
graph of its skin (text-fig. 112) shows that the scales overlap 
much more than those of either of the species already described. 
A spawning mark should therefore be better defined if one were 
to be found. 


Text-fig. 112. 


Cohoe Salmon | 


Piece of skin of the Cohoe (Oncorhynchus kisutch). 


The Cohoe Salmon gets the name of ‘ silver” salmon from its 
appearance when it first comes in from the sea. It is then a 
most brilliant silver with a greenish tint on the back. As 
spawning time approaches it becomes a dirty red. The Cohoe is 
found in almost all the Pacific Coast streams from Monterey Bay 
northwards. The run in the Fraser River is in September and 
October, but the Cohoes are on the coast from July to November. 
The weight of Cohoes canned in 1909 was 17,789,890 lbs., but 
large numbers, 1,152,452 lbs. in United States waters alone, were 
dealt with in other ways. The Cohoes in British Columbia 
- weigh usually from 3 to 8 lbs., but larger specimens are by no 


600 MR. J. A. MILNE ON THE 


means uncommon and may weigh as much as 30 lbs. More 
Cohoes would probably be canned if the run did not occur so late 
in the year, at a time when most of the canning stations are 
closed for the winter. 

The number liberated from the hatcheries in 1910 was 
50,424,586, but in some years the numbers have been much 
greater. 


Text-fig. 113. 


Scale of Cohoe (O. kisutch), 2. 4 lbs. 60z. 17th November, 1911. 


I find the scales of the Cohoes the most difficult of all to read. 
I have specimens of scales from eighteen fish from Puget Sound 
and from Shuswap, on the Fraser River. These fish varied in 
weight from 3 to 18 lbs. What strikes one as most curious is 
that, with the possible exception of one fish of 5 Ibs. 6 oz. 
(text-fig. 113) in which a few close lines between B and C, which 
I think due to a slight check in feeding, may really be the second 
winter band, their scales show them all to have been of the same 
age, namely nearing the end of their fourth year. 

I am satisfied that the Cohoes spend the whole of their first 
year in fresh water. At the end of it they measure, according 
to their scales, from 23 to 3 inches. One rapid grower, which 
weighed 15 lbs. near the end of its fourth year, measured as 
much as 4 inches at the time of migration. The growth in the 


HISTORY OF THE PACIFIC SALMON. 601 


second year is very difficult to make out. I feel convinced that 
the greater part of it (I mean as to time) has taken place in fresh 
water in nearly every instance, and in no instance am I quite 
convinced that the whole of it did not doso. If the whole of the 
second year’s growth took place in fresh water, the fish are a year 
older than I have stated above. In all cases (I speak, of course, 
only of the fish I have examined) the first entry into the sea 
appears to have taken place between 18 months and 2 years 
after hatching. Further certainty is added to this statement 
by the measurements of the scales, which show the second year’s 
growth to have been comparatively small. In three instances in 
which, from the arrangement of the rings, I think the fingerlings 
entered the sex when about 18 months old, the little fish were 
three times as long at the end of their second year as at the end 
of their first. None of the others even quite doubled their 
length. That is what one would expect at that stage from fresh- 
water feeding, but not from seafood. In the third year growth is 
very rapid, and hardly diminishes even as winter comes on. 


Text-fig. 114. 


Scale of Cohoe (O. kisutch), g. 14 1bs. Puget Sound, 18th October, 1911. 


Text-figs. 114 and 115 are typical Cohoe scales. 

All the scales from Puget Sound are from fish captured on 
18th October, 1911, and all show at their edges the lines coming 
clese together to form the next winter band, see text-fig. 114. 
A similar formation may be noted on text-fig. 113, which reached 


602 MR. J. A. MILNE ON THE 


its spawning ground a month later. But compare these scales 
with text-fig. 115. This fish was forwarded to me from Toronto 
on October 17th, so that it must have been procured at Lake 
Shuswap some days earlier. 


Text-fig. 115. 


acvAN 


Scale of Cohoe (O. kisutch), 9. 34 lbs. Shuswap Lake. 
Weighed after spawning. ° 


According to Dr. Greene (Migration of Salmon in Columbia 
River, Bulletin of U.S. Bureau of Fisheries, vol. xxix. 1909), a 
Silver Salmon travels about 7 miles per diem. This fish, then, 
to reach Shuswap by, say, October 12th, must have ceased feeding 
in the sea not later than 23rd August. It will be noticed that 
the outer lines of the scale are still widely spaced, showing that 
summer feeding and growth were still in progress up to the time 
at which the fish left the sea for the river. 


THe Houmppack Saumon (Oncorhynchus gorbuscha). 


I have but little to say at present about the Humpback, the 
Dog Salmon, and the Steelhead Trout. 

The Humpback has flesh of a pale pink colour. It was not 
used for canning purposes until quite lately, but now there is 
a good trade in it with China and Japan, where it is much 
esteemed. The Chinese, not being accustomed to red-fleshed 
fish, were very shy of the canned Quinnats and Sockeyes when I 
was in China in 1893, and this may account for their partiality 
to the Humpback. 

The total weight of Humpbacks canned on the Pacific Coast in 
1909 was 100,326,144 Ibs., and over another 3,000,000 lbs. weight. 


was used in other ways. 


HISTORY OF THE PACIFIC SALMON. 603 


Humpbacks were first dealt with in the hatcheries in 1904, and 
since then have been turned out from them intermittently, but 
never in very large numbers, comparatively speaking. 

They take their name from a peculiar hump which appears 
behind the heads of the males at spawning time. They are from 
3 to 11 lbs. in weight, and have very small scales. 

The main run is in Alaska. South of Puget Sound the 
Humpback is almost unknown. In the Fraser River the runs 
in the big Sockeye years and in the years next but one after them 
are much larger than in the other years. The fish, however, 


Text-fig. 116. 


Scale of Humpback Salmon (O. gorbuscha), from Puget Sound. 
(For reference see text, p. 604.) 


seem to come in at various ages, so this state of affairs] will 
probably gradually right itself again if the netting is not too 
severe. J am judging from possibly but a few specimens. A 
large number of Humpback scales have been sent me, but, 
unfortunately, with no further information than that they were 
taken from Humpbacks in Puget Sound on their way to the 
Fraser River. I append a photograph (text-fig. 116) of one of 
these scales. The centre is only moderately well defined, but in 
all the others it is even less clear, so that I have not been able to 


604 MR. J. A. MILNE ON THE 


form any definite opinion as to the time spent in fresh water. 
This scale shows clearly one winter band formed in the sea (AA). 
I presume that it represents the growth of the second winter. 
All the scales show a similar hema and many of them another 
evidently formed a year later. ’ 

I can give no measurements as I do not know the length, or 
even the weight, of any of the fish when caught. 


THe Doe Saumon (Oncorhynchus keta). 


The Dog Salmon frequents the Pacific Coast all the way from 
San Francisco to the Arctic Circle, but is most plentiful between 
Puget Sound and South-East Alaska. ‘This fish is also called the 
Chum Salmon, and is known in Japan as Sake, and in Siberia as 
Kita. 

I have not before referred to the prevalence of any of the 
species on the other side of the Pacific, and only do so now 
because the Dog Salmon has long been a staple food of Japan, 
and it is because this fish is white-fleshed, or almost white, that 
the red colour of the other varieties has caused them to be 
viewed with suspicion when exported to the East. This fact 
was ascertained by me when I was in China and Japan in 1893. 

The Dog Salmon gets that name from the distorted appearance 
of the mouth of the males at spawning time. They then look 
not unlike snarling dogs. The average weight of the Dog Salmon 
is about 8 lbs. In British Colombia it is higher, from 10 to 
12 lbs. 16 lbs. would be about the maximum w eight. 

The weight canned in 1909 was 25,660.845 lbs., and over 
4,000,000 Ibs. weight was otherwise dealt with for export. 

It is only in Washington State that these fish have been 
spawned in the hatcheries. There an average of about 7,000,000 
fry have been turned out in recent years beginning with the 
year 1900. 

Asa general rule the Dog Salmon comes in late in the year, 
September to November. But in Alaska the run begins in June, 
and in the Fraser River in the middle of August. 

In British Columbia they spawn close to the sea. This I had 
gathered from the appearance of the scales before I found out 
that it was known to be the case. I have received scales from 
two specimens weighing 114 and 12 lbs. respectively, both caught 
in Puget Sound on their w ay to the Fraser River. They show 
no indication of fresh water feeding at all. The fry evidently 
descend to the sea as soon as hatched out. The measurements of 
10 scales of the 114 lbs. fish, one of which is shown in text- 
fig. 117, enable me to give the rate of growth as follows :—At 
the end of the first year 71 inches, at two years 13 inches, at 
three years 22 inches, at four years 274 inches, and when caught 
in its fifth year it measured 31 inches. Its capture was evidently 


HISTORY OF THE PACIFIC SALMON. 605 


an early one, as the summer growth is only just complete. Un- 
fortunately I have not got the date. The 12 lbs. fish was caught 
at the same time and place. It was shorter, 302 inches, but 
thicker, 17 inches. Its length at the end of each winter was 
54, 11, 203, and 263 inches. From a smaller beginning it was 
steadily overtaking the other fish in length, and had already done 
so in weight when caught. 


Text-fig. 117. 


Scale of Dog Salmon (Oncorhynchus keta). 9. 11% lbs. 
Length 31 inches; girth 16? inches. 


THE STEELHEAD Trout (Salmo gairdneri). 


The last of the Pacific Coast salmon with which I am dealing 
is the Steelhead, which is there called a trout, but is the only 
Pacific member of the sub-genus Salmo, to which the true salmon 
belongs. It is believed to be a migratory form of the Rainbow 


606 MR. J. A. MILNE ON THE 


Trout. The flesh is pale-coloured, and is not much used for 
canning. 

The Steelhead is found from Carmel River, California, north- 
wards to Central Alaska. In California it may be fished for only 
with hook and line. In 1909 4,229,704 lbs. weight was canned 
and 1,960,000 lbs. weight was otherwise used in the United 
States fisheries, without counting those frozen or salted in British 
Columbia. 


Text-fig. 118. 


Scale of Steelhead Trout (Salmo gairdneri), 8. 122 lbs. Length 33 inches ; 
girth 163 inches. New Westminster. Autumn,1912. (For references see text, p. 607.) 


In 1910 there were liberated by the U.S. hatcheries 12,023,646, 
but none was turned out in Canada. 
The spawning season is in the spring, but Steelheads are to be 


HISTORY OF THE PACIFIC SALMON. 607 


found in fresh water at most seasons of the year. They weigh, on 
an average, at different places from 8 to 15 Ibs., witha maximum 
of about AD lbs. 

The scales, of which I have specimens from several fish, but 
from only two that had been weighed and measured, show that 
life in fresh water before the first migration to the sea may be of 
considerable duration. In this species, as in the case of our own 
salmon, no doubt is possible as to the limits of the part of the 
scale formed in fresh water. From examination of a large 
number of scales from my two measured fish, I judge them to be 
about 7 inches long when they enter the sea. As spawning takes 
place in the spring, the number of lines on the scale at the end of 
the first winter is naturally very small. The scale illustrated in 
text-fig. 118 shows five up to the point A, and that seems the 
usual number, but the first band is not always visible on all the 
scales, which is not to be wondered at seeing how minute they 
are, =; inch long at the most, at that time. 

A second winter in fresh water is shown in text- fig. 118 by the 
band B, and a third by the band C. Outside of i are two or 
three more lines added to the parr scale in the fish’s fourth 
year, before it entered the sea. All the Steelhead scales which T 
possess show similar lines and bands, so I suppose one may take it 
that these fish remain in fresh water until well on in their fourth 
spring, counting that in which they were spawned as one. They 
would then be just three years old. 

When they get to the sea they grow very rapidly. The first 
band completed in the sea within a year of migration shows my 
two measured fish to have been then already from 194 to 202 inches 
long (see text-fig. 118, E), and the second winter band shows a 
length of from 29 to 301 inches (text-fig. 118, F), but I am not 
sure that I have identified this band cor rectly on the seale 
illustrated. These two fish were both caught near the mouth of 
the Fraser River in the autumn of 1912, when they measured 
312 and 33 inches respectiv ely and weighed 12? and 13 lbs., 
the shorter and thicker fish being a female. They Tad then done 
a considerable amount of feeding i in their sixth year. 


This completes for the present my review of the salmon of the 
Pacific Coast of North America. 

It must always be borne in mind that my observations have 
been made on the scales of fish from the Fraser River district 
alone, and that readings from the scales of salmon caught else- 
where might show considerable differences. Some dive gence is 
almost cer Seat in the duration of freshwater life, for observations 
of the scales of the true salmon (Salmo salar) hee shown that 
the further north one goes the longer does the young salmon 
remain in fresh water, Band mtn as “aeh at all unlikely that the 
Pacific salmon are influenced in the same way by climatic 
conditions. 


608 MR. J. A. MILNE ON THE 


The strongest argument which I have heard in favour of the 
prevailing idea that Pacific salmon migrate to the sea as soon as 
they can swim, is that many of the rivers which they frequent 
are small, and, at times, are either dried up or frozen hard. This 
argument was put before me by a gentleman who is largely 
interested in the Pacific Coast fisheries and generally recognized 
as an authority thereon. It therefore seems to demand an 
answer. My answer is that I have examined only the scales of 
fish hatched in the large watershed of the Fraser River, and that 
possibly the fry of the Pacific salmon can adapt themselves to 
varying conditions. It is known that our own salmon, which 
generally migrate to the sea at two years of age, may enter it at 
one year without hurt, or may remain in fresh water for three, 
four, or even five years where the circumstances, as in Norway, 
make it desirable for them to do so. 

But, even supposing that they can adapt themselves to a 
certain extent, I fail to see how any salmon can perpetuate its 
species in a river which is frozen absolutely solid in winter, or 
which is bone-dry in summer. In fact, I believe that the fish 
that run into small rivers where such conditions prevail have not 
been hatched in them at all, but are really natives of one or other 
of the large river systems, and that, when they get into one of 
these smail rivers, they become at once of no further use to their 
own species, and might as well all be promptly utilized as food 
for the good of ours. 

For a fact which supports this belief I have only to refer to 
what Mr. W. L. Calderwood says in a communication to the 
‘Salmon and Trout Magazine’ for December 1912, page 24. He 
there describes Capt. Callbreath’s hatchery in Alaska. Iam not 
now concerned with the hatchery itself, but with the fact that it 
is on a river, the Jadeska, only half a mile long, which runs out 
of a small lake. A dam has been thrown across the river, only 
100 yards from the mouth, so as to intercept all fish, and enable 
the operators to select the Sockeyes and refuse the other salmon. 
In addition to this dam a fence of racks has been erected having 
at one place a trap. Here the Sockeyes have been taken out and 
lifted over the dam, if not required for the hatchery, while the 
other species have been left below. This selection has gone on for 
eighteen years, and still continues, although the Sockeye hatchery 
was a failure and hatching operations were discontinued in 1906. 
Mr. Calderwood says :—‘ It is an interesting point that, although 
the natural and artificial propagation of the Humpback and Dog 
Salmon were entirely discouraged, the number of these fish 
continued apparently undiminished.” He adds, and this is my 
point, ‘‘ The moral may be that the fish in this river, like the 
fish in other small rivers, are largely drawn from other sources.” 
But, in this case, as none but Sockeyes have been admitted 
to breed, it would seem proper to substitute the word “ entirely ” 
for the word ‘“ largely,” and I submit that what applies to one 
small river may equally well apply to them all. 


HISTORY OF THE PACIFIC SALMON. 609 


If the perusal of what I have written induces others to pursue 
this fascinating study of scales upon the lines I have suggested, 
my work will have served its chief purpose. 

That the study is a most important one cannot be denied. The 
salmon fisheries of the Pacific Coast are a source of vast wealth to 
the countries so fortunate as to share in them. But that source 
of wealth, like our own salmon fisheries, already shows signs of 
having been too lavishly drawn upon. It is easy to kill the goose 
that lays the golden egg. By legislation much may be done to 
prolong its life, but legislation can effect its purpose only if based 
upon thorough knowledge. In this country, if salmon scales had 
been studied before the Acts which regulate our fisheries had been 
passed, these Acts would have been very different from what they 
are. The study of scales has so revolutionized our knowledge 
that, of the eleven facts in the life of the salmon, mentioned by 
Mr. Cholmondeley Pennell in 1886 as absolutely proved, one, 
the most important, has now been shown to be absolutely 
wrong, and two others, hardly less important, require much 
modification. 

I have not made myself familiar with the varicus fishery laws 
and regulations of the Pacific Coast, but I know that many 
experts think them insufficient. If the regulations as to netting 
are to be made more stringent, let it be done with as complete 
knowledge of the various salmon as possible ; and in order that 
that knowledge may be obtained let their scales be systematically 
studied. For months, or years, the fish are hidden from our eyes, 
but from their scales we can fill in the gaps while they are out 
of view, for it has been well said that every salmon carries a 
record of its life on each of its scales. Whether I have read the 
records written on the scales of the Fraser River salmon correctly 
or not, I can at least claim to have proved that the records are 
there. 


In conclusion, I have to thank Mr. W. J. Sim of Northville, 
Kings Co., Nova Scotia, for much valuable information, and for 
having procured me numerous specimens through his friends 
Mr. Frank Parry, now also of Northville and formerly of Granite 
Creek, Mr. J. Brown of Celista, B.C., and Mr, Josephson of 
Bellingham, Washington, U.S.A., and Mr. J. Kirkpatrick of 
Vancouver. The various statistics I have given have been 
compiled mainly from “The Salmon Fisheries of the Pacific 
Coast,” by J. N. Cobb, Bureau of Fisheries, Document 751, 
Washington, 1911, and partly also from the Report of the 
Commission of Conservation on ‘ Lands, Fisheries and Game, 
and Minerals,” Ottawa, 1911. 


The following table may y;rove of interest :— 


ON THE HISTORY OF THE PACIFIC SALMON. 


| 


| | 
sot S = josh 
i~ | . PS =| | O& 
. aS s as PS S -S 
Oe Geen a Hea bee iss 
ae Bs |} Sos aS | CaS ot 
jz 2 SS | 22 Bars | pe | oS 
os SS | os sl p} o's 
= Ss | Oo BS ey =a 
RO os | S =. iS) a) 
| ai We aes mlcesie 
| a 
it Pane Ee eee 
Ome | Number of gill-rakers) 
Ze) Ss | onthe anteriorarch ...) 32-40 | 20-28 | 20-25 | 28-36 | 20-25 
on | | 
BESS | Saar | * 
: a | d 
3 B sae | Number of well-deve-, 
I San | loped rays in anal fin.) 14-15 | 15-18 | 12-14 15-16 | 13-14 10-12 
3 out e —= ae | 
fs oF — | | | 
° ABS ‘Number of scales on 
fe »o Me | dJateral line............... 125-185 | 140-1655 | 125-135 150-170 135-145 
Aaa | 
| fa, “Se = 
io = Q2s). ||| 
| ear 4s : - 
sa» Number of tmes of scales | | 
S845 between lateral line’ | 
q |e antd dowsalefiney ee. 20 25 nO BO BS 
| | 
Number of pounds canned, pickled, 
frozen, or salted on the Pacific Coast 
in 1909, in millions of pounds—nearest 
snillion zie) 13 iQ | 1108 | a0). | ° 
F Daan 
Number of fish caught in 1909, calculated | 
from total weight as given above, and | | 
general average weight in millions, | | 
after allowing one-fifth for waste ...... 54 3 | 5 Bt z 
| 
eda De | | | u 
Number of fish liberated from the) | 
hatcheries in 1910 in millions—nearest | 
million 316 Oi | sO 2 " 12 
Average weight of fish, as caught, in) | 
pounds 3-10 18-30 3-8 A Sales alo 
Maximum weight attaimed, in pounds. | 
aboubie erence cana enbacce ce sautiea decisis 17 100 | 30 11 16 45 
| 
Approximate time spent in fresh water! 
before migration to the sea, in Fraser 
River district, in months 12-15 18 18-24 ? 3orless, 36 
ee | ary 
Approximate length at time of migration 
to the sea, in Fraser River district, 
in inches 3-33 4-5} 23-5 P jlkorless 7 
s a . diss 
Age at time of return to fresh water, m) 
Fraser River district, in years; as) 
33 33-42 33 25-38 43 55 


| shown by my specimens. .................. 


MISS KATHLEEN HADDON ON PERIPATOIDES WOODWARDI. 611 


38. Notes on Peripatoides woodwardt Bouvier. 
By Karaieen Happon *. 


[Received April 11, 1913: Read May 6, 1913.] 


This species has been described by Prof. Bouvier t. It may 
readily be distinguished from other Australian species by the 
fact that it has 16 pairs of legs; in this respect this species and 
P. sutert from New Zealand are unique among all the Australasian 
Peripatoides. . 

The material was obtained through Prof. J. Stanley Gardiner, 
from Western Australia. 

There are twenty specimens, well preserved and extended, but 
they tend to be rather bent, thus making exact measurement 


difficult. The approximate measurements (in millimetres) are as 
follows :— 


Females ... 46, 43, 43, 41, 37, 37, 33, 23, 21, 17. 
Males ...... 31, 30, 30, 29, 27, 24, 24, 24,23, 29. 


It will be seen, therefore, that these specimens are considerably 
larger than those described by Prof. Bouvier {, the greater 
number of which measured from 10°5 to 17 mm., only one 
reaching the length of 21 mm. 

Bouvier recognises the following types of coloration :— 


1. Uniform, with predominance of blue-green pigment, 
yellowish pigment being strewn in spots equally dis- 
tributed over the dorsal surface. 

2. Striped, with the yellow spots more numerous, sometimes 
becoming predominant, and forming a longitudinal 
band, with or without dark papille, on each flank above 
the base of the legs. 


A variation of the first type had the ground colour light 
reddish yellow, almost all the papillae being darker, at any rate 
at the summit; some were quite black and arranged in short 
transverse rows. Thus the general tone was a dingy grey-black 
with small spots of yellowish pigment round some papille. 

A variation of the second type showed a yellow-red ground 
colour, with stripes of dark papille in transverse rows in the 
dorsal region and no mixture of dark pigment in the lateral 
stripe. 

All his specimens showed two large light-coloured papille on 
the flanks between each pair of legs, arranged parallel to the 
long axis of the animal. 


These types are all represented in my specimens and also all 


* Communicated by Prof. J. Srantey Garpiner, M.A., F.RS., F.Z.S. 


+ ‘Die Fauna Siidwest-Australiens,’ Bd. ii. No. 18, Onychophora. M. E.-L. 
Bouvier. Jena, 1909. 


i Doc. cié. ps 315. 
Proc. Zoou. Soc.—1913, No. XLI, 4] 


612 MISS KATHLEEN HADDON ON PERIPATOIDES WOODWARDI. 


the intermediate stages between them. Those belonging to the 
first type show in addition a well-marked lateral stripe of a 
greenish-blue colour. The antenne are also of a greenish-blue, 
thus differmg from those of Prof. Bouvier’s specimens which he 
describes as always being black. Four of the animals are 
very dark in colour, one being practically black, but all have a 
brownish-green tinge : the ventral surface is lighter, but even 
then darker than the backs of the other specimens. The darkest 
specimen, which is a male, differs from all the others in having 
the crural papille black instead of white. All the animals show 
a patch of green pigment on the inner side of each leg which is 
extended on to the ventral surface of the body. In most cases 
the ventral and pre-ventral organs are not well-marked, but some 
of the smaller specimens show the ventral organs distinctly. 
The two lght coloured papille described by Prof. Bouvier as 
situated between each pair of legs are not apparent in these 
specimens. 

The coloration of these Peripatus agrees fairly closely with 
that of Peripatoides orientalis *, except that the dark variety is 
by far the most common in the last-named species (772 per cent. 
according to Steel t), whereas in P. woodwardi they are in the 
minority, as far as our present material goes. 

With regard to the external features, these specimens of 
P. woodwardi agree with those described by Prof. Bouvier ¢ in 
every respect except for the number of crural papille in the male. 
These are very prominent and capable of retraction within the 
leg, and he describes them as present on all the legs with 
the possible exception of the 15th and 16th, that is the last two 
pairs, while he found coxal glands in all the legs except possibly 
the 15th pair. 

I have examined all the males in my material, and although 
the papille show extremely clearly in most cases, I can find no 
traces of them on the 4th or 5th legs; they appear to be entirely 
absent from these legs, as well as from the 15th; the last pair 
of legs may show a small papilla. In the male that I dissected 
the coxal glands showed very clearly in the other legs, but no 
trace appeared in the 4th, 5th, or 15th pairs. The absence of 
these glands in the 4th and 5th pairs of legs seems to be correlated 
with the great development of their nephridia, which open by a 
urinary papilla on the proximal of the three spinous pads of the 
foot. 

As deseribed by Prof. Bouvier, the first pair of crural glands 
are extremely long and lie in the Jateral sinus with the nephridia ; 
the last pair are elongated and iridescent and lie in the lateral 
sinus or in the general cavity of the body, while the remaining 


* “Monographie des Onychophores.” E.-L. Bouvier. Ann. des Sc. Nat. vol. v. 
9th series, 1907, p. 233. 

+ “Observations on Per ipatus,” Proc. Linn. Soc. N. S. Wales (2) vol. xxi. 1896, 
p- 94. 

{ ‘Die Fauna Stidwest-Australiens,’ p. 320. 


ON THE ENEMIES OF BUTTERFLIES. 613 


ones are of the same type as those of P. orientalis, a vesicular 
gland with a narrow duct lying in the leg. 

The only other particular in which my specimens differ from 
those of Prof. Bouvier is in respect to the female generative 
organs. In the two examples that he dissected there was little 
or no trace of a receptaculum seminis, but he points out that the 
ovaries were very large, thus making the dissection difficult. 

In both the females that I dissected it was easy to trace the 
oviducts for the whole of their course, and shortly after their 
emergence from the ovarian sac, where they at once diverge from 
one another, there was a well-marked receptaculum seminis on 
each side, full of sperm. This is another point of resemblance 
between this species and P. orientalis, which seems on the whole 
to be the most nearly related form. 


39. Field-Observations on the Enemies of Butterflies in 
Ceylon. By J.C. F. Fryer, M.A., F.E.S.; Fellow of 
Gonville and Caius College, Balfour Student in the 
University of Cambridge. * , 


[Received April 22, 1913; Read May 6, 1913.] 


Ermonocy (Birds AND BUTTERFLIES). 


The subject of Birds and Butterflies in Ceylon has already been 
admirably dealt with by Col. N. Manders in his paper entitled 
“« An Investigation into the Validity of Mullerian and other forms 
of Mimicry, with special reference to the Islands of Bourbon, 
Mauritius, and Ceylon”. In spite of this, however, I venture 
to place on record a few further notes from the island, partly 
because they deal more fully with the butterfly-eating habits of a 
most interesting bird, the Wood-Swallow (Artamus fuscus Vieill.), 
but also because the subject, with its direct bearing on the various 
theories of mimicry, is essentially one which requires as many 
observations as possible before any fair generalisations can be 
mace. 

A residence of a year and nine months, largely devoted to 
entomological research, gave an opportunity for keeping a con- 
tinual watch on certain of the more common insectivorous birds ; 
and the following notes comprise every observation made during 
that period. The locality in each case, unless otherwise stated, 
is the celebrated Botanical Garden at Peradeniyat. Other 
localities mentioned are Vavonia-Valunkulam, Maha I]lupalama, 
and Trincomalee—all places in the hot dry northern country, 
which is characterised by the large area of jungle it contains, 


* Communicated by Prof. J. SranntEy GarpiIner, M.A., F.R.S., F.Z.S. 
+ Proc. Zool. Soc. 1911, p. 696. 
t Elevation about 1600 feet. 

Ase: 


614 MR. J. C. F. FRYER ON THE 


and also, in my experience, by a greater quantity of butterflies 
and other insect life than can usually be found in the hills. 

The notes are given in chronological order, and are extracted 
from my journal practically without alteration, as this method, 
though possibly wearisome, places a check on the personal pre- 
judices of the observer, the introduction of which it is difficult to 
avoid in a mere summary of results. 


PERADENIYA. 


1911. April 7th. A drongo* took a Papilio polytes Linn. 3, 
which could hardly fly, as it had only just emerged ; it then 
caught a Melanitis ismene Cram., also released soon after it had 
left the pupa. 

April 2nd-9th. General note that “drongos were only observed 
to take small butterflies, chiefly Yphthima ceylonica Hewit., 
Lyceenids, and occasionally Precis iphita Cram.” ‘ Of the various 
attempts quite one out of two missed their object, while in many 
cases the swoop at a butterfly seemed to be taken more for amuse- 
ment than to gain food.” ‘“ Paradise fly-catchers (Terpsephone 
paradisi Linn.) being more retiring, were harder to observe ; they 
certainly took Yphthima ceylonica and small Lycenids, but were 
not observed to attack anything large.” [This week, April 2nd— 
9th, was spent mainly in bird observations, and the above is the 
result. | 

April 15th. A drongo was seen to attack Catopsilia pyranthe 
Linn., but apparently without success. 

May 9th. A drongo was seen to take an Yphthima ceylonica in 
the garden. ... Three drongos chased a Papilio parinda Moore, 
but did not catch it.... Two bulbuls (A/olpastes hemorrhous Gm.) 
chased an orange-coloured moth, probably Argina sp., but lest it. 

June 25th. A Paradise fly-catcher swooped at a Limenitis 
calidasa Moore, which escaped. 

July 9th. A magpie-robin (Copsychus saularis Linn.) took a 
specimen of the Chalcosiid moth, Heterusia cingala Moore +, and 
flew away with it. 

October 16th. A drongo chased a Papilio parinda, but lost it. 

October 19th. A drongo caught a Papilio polytes 3, and ate it. 

October 21st. A scanty P.-polytes migration for the last three 
days... . A drongo swooped at a P. polytes 2 form polytest, but 
missed it. Many females of the romulus form § with the entire 
hinder portion of the hind wings missing. 

November \st-15th (at Vavonia-Valunkulam). A continual 
migration of Appias (A. paulina Cram. and A. albina Boisd.), 


* Throughout, the word ‘“‘drongo” is used for either Dicrurus leucopygialis 
Blyth, or D. cerulescens Linn. The former was more common; but the species 
in wost cases was not ascertained. 

+ This should undoubtedly be an “inedible” species, as it has all the Chalcosiid’s 
nauseous characters to a marked degree. 

“ + P. polytes 2 polytes resembles Papilio aristolochie Fabr. 

§ P. polytes 2 romulus resembles Papilio hector Linn. Both P. aristolochie 

and P. hector are common Ceylon insects. 


ENEMIES OF BUTTERFLIES. 615 


Catopsilia (C. crocale Cram. with a few C. pyranthe Linn.), 
Papilio polytes, P. jason Linn., P. nomius Esper, and P. hector 
Linn. Bee-eaters (Merops sp.?) were common, but were only 
observed to chase butterflies (P. jason, Appias sp.) on three 
occasions, each attempt being unsuccessful. Wings of all the 
above butterflies were found occasionally along the roads, but not 
in any number. 

November 25th (at Peradeniya). A drongo seen to take a 
Papilio polytes 2 romulus, which was flying badly. 

November 29th. A number of wings of Huplaa and Danais 
found on the ground in the garden. ‘The only birds seen near 
were wood-swallows (Artamus fuscus Vieill.). The wings iden- 
tified were :—Huploaa core Cram., fore wings 6; #. kollari Feld., 
fore wings 2; HL. coreta God., fore wing 1; Danais septentrionis 
Butl., fore wings 5, hind wings 3. 

November 30th. Spent an hour, 11 A.m.—12, watching the wood- 
swallows hawking butterflies, which appeared to be forming their 
sole food. The birds sat on the top branches of a leafless tree, 
and swooped out after their prey, usually effecting a capture. 
‘The successful attempts were estimated at two out of three. The 
butterfly was carried to the perch and the wings there discarded. 
Butterflies on migration were extremely abundant, and indi- 
viduals of the genus Catopsilia were perhaps most numerous, 
though Eupleas, Danaids, Papilio polytes and P. demoleus Linn. 
were all quite common. ‘The wood-swallows limited themselves 
almost entirely to Eupleas and Danaids, only three swoops at 
Catopsilia and one at Papilio demoleus being observed, and these 
were all failures. The majority of the discarded wings were 
blown into the river, but the following were collected, practically 
all of which must have fallen during the morning. 

Result :—Danais septentrionalis: fore wings 61, hind wings 9. 

Huplea core: fore wings 48. 

EL. coreta: ss BAe AS) 

E. kollari : bs , 6. 

Kupleine hind wings 9, fragments 4. 

Papilio demoleus: fragments 2. 

P. polytes 3 ov 2 cyrus-form*: fragments 7 
(2 fore wings). 

P. polytes 2 romulus: fragments 5 (3 fore wings). 

P. polytes 2 polytes: hind wing 1. 


The predominance of fore wings is explained by their greater 
weight, hind wings being blown further. 

A pair of bee-eeters were feeding near the wood-swallows. 
Most of their attacks were on small insects; but four attempts 
on Catopsilia and two on Papilio demoleus were noted, all 
of which failed. These bee-eaters were either unskilful or 
only amused themselves with butterflies. A young Paradise 


* Papilio polytes 2 eyrus-form resembles the g. 


616 MR. J. C. F. FRYER ON THE 


flycatcher and a drongo confined themselves to small insects, 
in spite of the swarms of butterflies passing. 

December \st. Again visited the wood-swallows, but they left off 
feeding early ; the butterfly migration was larger, with a higher 
percentage of Huploea. It was distinctly observed in one case that 
the bird carried the butterfly, a Danais septentrionis, to its perch 
and then held it with its feet while it pulled the wings off with its 
beak *. Watched the bee-eaters again, and saw one attack first 
a Papilio agamemnon Linn. and then a Huplaa, but it missed 
each time. They were continually hawking small insects, which 
they presumably caught. No count could be made of discarded 
wings, as all had been blown by a high wind into the river. 

December 2nd. Again watched the wood-swallows hunting. 
The butterfly was always carried back to the perch before the 
wings were removed, and it was noticed that portions of the wings 
were often left on onal eaten.... After failing in a stoop, the birds 
varely made another at the same insect. ... Butterflies at some 
distance were usually taken, perhaps to allow the bird to attain 
greater pace... . If a butterfly noticed that it was being pursued, 
it at once dodged and, flying rapidly downwards, frequently 
escaped. Wings were again collected :— 


Danais septentrionis: fore wings 35. 
D. Vimniace : s Kage 
Huplea core: é 
EF. coreta: one whole insect and fore wings 10. 
i. kollari: fore wings 4. 

Papilio polytes 2 romulus: fore wing I. 


December 7th. Wood-swallows hunting in the afternoon; they 
usually stop before midday. 

December 10th. A young Paradise flycatcher swooped at two 
Papilio polytes,a 3 anda 2 polytes form, which were flying round 
inside a large breeding-cage, but of course could not get at them. 

December 16th. A drongo was seen to attack a Papilio aga- 
memnon and snap off a portion of the hind wings. The butterfly 
escaped. 

1912. January \st-6th (at Maha Illupalama). Bee-eaters 
were very common and butterflies, notably of the genera Cato- 
psilia and Appias, were abundant. The only attack observed was 
that of a bee-eater on a Leptosia xiphia Fabr., which was caught 
and eaten. 

January 10th (at Peradeniya). A drongo caught a Papilio 
polytes Q eyrus-form, which had just emerged and had escaped 
from a breeding-cage. 

February 2 Ond. A Papilio polytes 2 cyrus escaped from a cage 
and was eaten by a drongo; it was flying Heay) weakly, as one 
wing was broken halfway down the costa. 


* Legge, ‘Birds of Ceylon,’ vol. 11. p. 668, states that the wood-swallow beats off 
the wings of its prey. 


ENEMIES OF BUTTERFLIES. 617 


March 6th. Noticed three wood-swallows sitting on the 
telephone-wires crossing the River Maha-Weliganga. They were 
hawking butterflies continually, and during the time they were 
watched selected species of Huplea only from a passing migration 
which consisted largely of Appias paulina. A search for wings 
failed, as the birds confined themselves entirely to the river. ‘The 
reason for this seemed to be that the butterflies were only caught 
with ease when away from cover; among vegetation they dodged 
their pursuers nine times out of ten. This fact was specially 
evident during this migration, as species of Huplea and Danais 
were uncommon, and the birds were often forced to swoop at 
butterflies travelling along the river-bank instead of waiting for 
individuals to come out into the open. 

March 8th. The migration of butterflies almost ended; but to 
obtain confirmation, by a fresh witness, of the fact that wood- 
swallows attack esgectlly members of the genera Huplea and 
Danais, Alexander * and I watched three wood-swallows for an 
hour, and we both plainly observed them eat over a dozen Hunlea 
and one Danais. We also saw one bird with a Papilio, either 
P. polyies 2 polytes or P. aristolochie, but were unable to 
ascertain which. As before, the birds usually carried the butter- 
fly in the beak, though occasionally it was transferred to the 
claws when the distance back to the perch was great. 

March 15th. Watched the wood-swallows for three-quarters of 
an hour; Appias spp. common, Luplea and Danais scarce. There 
were only two birds feeding, one probably young, for when the 
other caught a Danais aglea Cram. the young one opened its 
beak and fluttered its wings as if asking for food, though it 
gained nothing by this proceeding. The old bird then ceased 
feeding, but the other, possibly encouraged by the sight of the 
Dearne, worked steadily, soaring round in circles over the river. 
It ca ught five Eupleas- out of seven attempts. 

March 16th. Watched a wood-swallow for a quarter of an hour 
and saw it take two Kupleeas. 

March 17th. Watched a wood-swallow from 11.30-12. It took 
two Euploeas, but then seemed to have no room for more, as it sat 
for a long time with the abdomen of the last victim protr uding 
from its mouth like a cigar. 

March 18th. In five minutes saw two wood-swallows take three 
Eupleas and a Papilio, either P. polytes 2 polytes or P. aristo- 
lochie, almost certainly the former. 

March 22nd. Species of Huplea and Danais abundant on 
migration. A wood-swallow picked out two Danaids (D. limniace 
and D. septentrionis) and then a Huplaa, after which it stopped 
feeding. 

Mar ch 23rd. Watched the wood-swallows from 11.40 a.m.—12; 
two were under observation most of the time, occasionally four. 
They took six specimens of Huplaa and two D. septentrionis. 


* Mr. Alexander, Assistant Curator of the Perth Museum, W.A., to whom I am 
indebted for much critical assistance during his brief visit to ‘the isl ind, 


618 ON THE ENEMIES OF BUTYERFLIES. 


April 23rd. A drongo swooped at and probably took either a 
Papilio polytes 2 polytes or a P. aristolochic. 

April 24th. Mr. Pole of Maskeliya* wrote that he saw two 
large dragonflies take two Appias sp. ? and one P. polytes 3. 

July Tth (at Maha Illupalama). Noticed the remains of Danais 
septentrionis, Papilio jason, Appias paulina and A. albina, killed 
probably by Asilid flies (Scleropogon piceus and other species), as 
the bodies were intact though empty. Asilid flies were twice 
seen carrying Appias sp.? 3. 

August 8th (at Maha Illupalama). Caught an Asilid fly with a 
medium-sized dragonfly +, which it was sucking through the eye, 
and later saw another Asilid killing a large Cicada $ which was 
flying madly about, shrieking (stridulating) all the time. A 
slight bite from an Asilid was most painful, and felt as if 
poisonous matter had been injected. 

August 13th (at Maha Illupalama). Noticed Asilid flies kill 
Appias sp.? 3 and Papilio jason. 

August 28th (at Habarana, Maha Illupalama District). An 
Asilid attempted to kill the formidable Ceylon hornet, Vespa 
cincta, which, however, escaped. 

August 31st (at Trincomalee). A few Euplea wings were found 
scattered about Fort Ostenburg ; they were probably dropped by 
wood-swallows, which were hawking round the inner harbour. 

September 1st (Trincomalee). On the Nilavelli-Kuchivelli road 
bee-eaters were numerous. One was seen to catch an Appias sp., 
and wings of Huplea, P. jason, and Appias spp. were occasionally 
found on the ground. 

September 7th (at Peradeniya). Released an Actias selene 3. 
It flew fairly well, but was soon snapped up by a drongo, which, 
before eating it, first pulled off the “ tails” and then the rest of 
the wings. 


The observations having now been given at length, it is 
perhaps permissible to summarise the impressions gained in their 
making :— 


i. Butterflies do not form any large percentage of the food 
of the more common insectivorous birds in Ceylon. 

il. With the exception of the Wood-Swallow, birds are by no 
means clever in capturing butterflies. 

ili, The Wood-Swallow is the only bird which actually lives on 
butterflies, and it almost always chooses butterflies of the 
so-called nauseous genera Danais and Huplea; it seemed, 
however, that this preference was due to the difficulty of 
catching faster-flying butterflies, and not to the superior 
flavour of the Danais or Huplea. 


* Hill district, elevation about 4000 feet. 

+ This and one or two subsequent notes, though not dealing with butterflies, are 
included, as they seem worthy of record and yet are insufficient for a separate 
contribution. 


Pola ch Sie, Pl LQ, 


C.D. photo. 


EUPARKERIA CAPENSIS 


1913. Pl. LXXVL 


28) 


a 


University Press, Cambridge. 


R-Broom ,del. 


EUPARKERIA CAPENSIS. 


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R.Broom,del. University Press, Cambridge. 


9,10. ORNITHOSUCHUS WOODWARDI. 11. 0. TAYLORI. 


P.Z.S. 1913. Pl. LXXVIUL 


O 
G en Oeononome 


R. Broom ,del. 


University Press, Cambridge. 


MESOSUCHUS BROWNI AND OTHERS. 


P.Z.S. 1918. Pl. LXXIK. 


R.Broom,del. University Press, Cambridge. 


BROWNIELLA AFRICANA AND OTHERS. 


ON FOSSIL REPTILES. 619 


iv. The various species of Bee-eaters, when attacking butter- 
flies, usually choose members of the Papilionide and 
Pieridee. 

v. The Asilid diptera are formidable enemies to all other 
living insects. They are, however, present in large 
numbers only in the northern country, and then only for 
a limited period of the year. 

vi. In Ceylon a resemblance to the genera Danais and Huplea 
is doubtfully of value ; in fact, in the neighbourhood of 
Wood-Swallows it is a distinct danger. 

vii. The mimetic females of Papilio polytes are not obviously 
protected by their mimicry, and as a whole probably 
suffer about as much, or as little, from the attacks 
of birds as any other fast-flying butterfly in Ceylon. 


40. On the South-African Pseudosuchian Kuparkeria and 


Allied Genera. By R. Broom, M.D., D.Sec., C.M.Z.S. 
[Received and Read May 20, 1913.] 
(Plates LXXV.-LXXIX.*). 


InprEx. 4 Page 
JD OG DOTIRAPIGL CHYDCISOS VEXTIOIN, oo nao ococboone a. 5senad ade coe no8eo0K0qbC 619 
Ornithosuchus woodwardi Newton .........0.. cc. ccc cee vev eee eee eee 624, 
Onmthoswenusiianloniasp aeeeerere eae eee eee ee OLD) 
Herpetosuchus granti Newton .......2......:c0se000.se0ese00sss0200 626 
Browniellanafricand, See CbiSpsMe 5: assess -eeteasseeeaee eee OT 
TESOS WEDS WIRDUTOR NN ENBIOD aoe sao coo tosenseceansene sAseseoasosannosns (Ae 
Scleromochlus taylori Smith-Woodward ........................... 629 
Affinities of the Pseudosuchia.................20.c..csseeese0eeeeeeeeee 629 


KUPARKERIA CAPENSIS Broom. (Pls. LXXV., LXXVL., 
LXXVIIT., LX XTX. figs. 1-8, 17-20, 22, 24.) 


In the collection of Mr. Alfred Brown of Aliwal North are 
a number of imperfect skeletons of a small Rhynchocephaloid 
reptile of very great interest, which he discovered in Upper 
Triassic beds about four yearsago. Mr. D. M.S. Watson examined 
them a year ago, but unfortunately he mistakenly regarded them 
as belonging to the little acrodont-toothed reptile named by 
him Mesosuchus brown. Though the two forms were met with 
together and are of about the same size, they certainly belong to 
different families, if not to different suborders. In a short note 
commmunicated to the ‘ Records of the Albany Museum’ (6), I 
have called attention to the confusion and proposed the name 
Huparkeria capensis for the Thecodont form. 

As the type, I take a specimen showing a beautiful skull 
with most of the skeleton. Though this specimen was in the 


* For explanation of the Plates see p. 633, 


620 DR. R. BROOM ON 


collection as seen by Watson, it was scarcely at all developed, 
and he did not recognise that the block of stone contained an 
almost perfect skull which, had he seen it, would have prevented 
the confusion of the two animals. 

In the type specimen is preserved the skull almost complete 
but shghtly crushed, and with the bones in beautiful condition and 
showing nearly every detail of structure. A number of cervical 
and densa vertebre are shown, both coracoids, the interclavicle, 
the right clavicle and scapula, and the right marae on radius, and 
ulna. There are also seen in the specimen the whole of the 
abdominal ribs in perfect condition, the pelvic bones in good con- 
dition, but displaced, and most of the left hind limb. Other 
specimens show the pelvic bones in position and most of the tail. 
One specimen shows a dentary with a perfect right pes. 

The skull is very similar to that of Or “ailioenieaus woodward, 
and there can be little doubt but that Hauparkeria belongs to the 
same suborder and to the same family, though the two genera 
must be regarded as distinct. From the snout to the occipital 
condyle the skull measures about 83 mm. From the snout to 
the front of the orbit is about 45 mm. The orbit is almost round 
and has an antero-posterior diameter of 23 mm. There is a 
large antorbital vacuity about 20 mm. in length. The infra- 
temporal opening measures 18 mm. in height and its greatest 
antero-posterior length at its lower part is 17 mm. The supra- 
temporal fenestra is small, measuring 13 mm. by 10mm. The 
width across the frontals between the orbits is 14 mm., and the 
width across the squamosals where they form the temporal arch 
is 34 mm. 

The front part of the premaxilla is missing from the type, but 
it was probably somewhat similar to that in Ornithosuchus and 
other allied types. It forms the lower half of the posterior 
margin of the rather large nostril. It supports at least two and 
most probably three flattened pointed thecodont teeth. 

There is no evidence of a septo-maxillary on the face. 

The maxilla is a long slender bone, which forms the lower 
and anterior borders of the antorbital vacuity. The anterior 
ascending process passes up behind the premaxilla and the 
descending anterior portion of the nasal, and meets the anterior 
end of the large lacrymal. The posterior horizontal portion 
passes back to below the middle of the orbit and meets the jugal. 
It supports apparently 13 thecodont teeth, of which 5 are 
preserved in the specimen. These are pointed flattened teeth, 
very similar in general shape to those of carnivorous Dinosaurs. 
They are feebly serrated behind and probably also in front. 

The nasal is rather peculiar in shape. When viewed from 
above, it appears as a long narrow bone about twice as wide 
behind, where -it meets the frontal, as in front. In reality the 
front is as wide as the back part, as it forms a curious downward 
process behind the nostril to meet the premaxilla. The peculiar 
shape will best be understood from the figures. 


FOSSIL REP'LILES, §21 


The lacrymal is an unusually large bone. It forms most of the 
anterior orbital margin. Above, it has a long articulation with 
the prefrontal, and below, a short one with the jugal. Anteriorly, 
it has a large development which forms the upper margin of the 
antorbital vacuity meeting the ascending process of the maxilla. 
Of the anterior process much is below the level of the general 
surface of the face, suggestive of the antorbital vacuity having 
lodged a large gland. 

‘Lhe prefrontal is a small narrow bone which forms about half 
of the upper orbital margin. It is bounded above by the frontal 
and nasal and below by the lacirymal. 

The frontal is a long narrow bone. Behind, it articulates with 
the parietal and laterally with the postfrontal, and in front 
with the nasal and laterally with the prefrontal. It only forms 
a small part of the orbital margin. 

The postfrontal is a small triangular bone which articulates 
with the frontal, parietal, and postorbital, and forms a small part 
of the upper orbital margin. 

The postorbital is a tiivadiating bone. The upper process 
passes upwards behind the postfrontal and meets the parietal. 
The inferior process passes downwards and articulates with the 
jugal, partly lying in front of it and forming with it the post- 
orbital arch. The posterior process is shorf. It meets the 
squamosa! and forms with it the temporal arch. 

The jugal is also a triradiating bone. The anterior process is 
the strongest. It forms most of the infraorbital arch, forming a 
long suture with the maxilla and meeting the lacrymal. ‘The 
upper process meets the postorbital and forms with it the post- 
orbital arch. The posterior process is long and slender and 
forms the zygomatic arch meeting the quadrato-jugal. 

The quadrato-jugal is an angular bone which supports the 
quadrate and the jugal and binds them together. Above, it 
meets the squamosal. A large foramen is present between the 
quadrato-jugal and the quadrate near the lower part of the 
bone. 

The squamosal is small but fairly strong. It may be regaraed 
as a clasping bone which holds together the quadrate, quadrato- 
jugal, postorbital, parietal, and opisthotic. A small upper 
process lies in front of the lateral process of the parietal. The 
relations to the quadrato-jugal and quadrate will be better 
understood by the figures given. 

The quadrate is a long, well-developed bone. Its upper end 
is firmly articulated with the squamosal and possibly also with 
the opisthotic, and also meets the quadrato-jugal. The lower 
end forms the articulation. There is a large opening between 
the quadrate and quadrato-jugal. The shape of the bone is seen 
in the drawings given. 

The parietal is smaller than the frontal. It forms the back 
part of the upper cranial wall and has a postero-lateral process 
which forms much of the posterior wall of the upper temporal 


622 DR. R. BROOM ON 


fossa, and at its outer end meets the squamosal. There is no 
trace of a pineal foramen. 

Behind the parietals and partly wedged between them is a small 
interparietal, It forms the upper part of the occiput, and partly 
divides the parietal from the supraoccipital. 

The supraoccipital forms the middle part of the occiput. It 
articulates with the exoccipitals, the interparietal, and the 
parietals. 

The exoccipital forms part of the occipital condyle and passes 
outwards, fusing with the opisthotic. 

Only a small part of the basioccipital shows in the specimen 
as the middle part of the condyle. 

The lower jaw is well preserved. The dentary forms the 
anterior half, and the angular and surangular the greater part 
of the posterior half. A lar ge oval opening is seen on the outer 
side of the jaw between the angular and surangular. 

A pair of long rib-like bones represent portions of the hyoid 
apparatus. These are evidently the ceratobranchials, and indicate 
that Huparkeria had a birdlike tongue. 

There are well-developed sclerotic plates in the eye, which are 
curved as in the bird. 

Two slightly displaced bones are probably the proatlas and 
portion of the atlas. There are probably about 9 or 10 cervical 
vertebrae, of which the upper 3 or 4 are hidden by matrix. The 
lower cervicals have comparatively short centra. They are 
practically amphiplatyan or incipiently procelous. The ribs are 
double-headed and have small uncinates. There are 2 sacral 
vertebra, and apparently 26 presacral. The tail is very long and 
has powerful chevrons, 

The shoulder-girdle is well preserved. ‘There is a long slender 
interclavicle which is apparently narrow even at its anterior end. 
The clavicle is also long and slender. The coracoid* is large, 
measuring 24 mm. in antero-posterior diameter and 16 mm. in 
its transverse diameter. ‘There is a large oval foramen near the 
scapular articulation and a little in front of the median plane of 
the bone. The scapula is long and slender. It measures in 
greatest length 38 mm., and its lower end is 15 mm. across and 
the upper end 12 mm. in width. There is no distinct acromion 
process, and the clavicle has been probably rather loosely attached 
to the front of the lower third of the bone. 

The humerus is very slender. Its length is 36 mm. Only the 
outer aspect is displayed, and it cannot be seen whether there 
is an entepicondylar foramen. There is no indication of an 
ectepicondylar foramen. The deltopectoral ridge is very short. 

The radius and ulna are slender straight bones. The radius 
measures 32 mm. in length and the ula) is probably slightly 
longer. The manus is lost from the specimen. 


* Though to avoid any confusion I have used the universally accepted term 
“* coracoid,” as I have elsewhere recently shown the bone ought more properly to be 
called the“ precoracoid.” 


FOSSIL REPTILES. 623 


There is a large broad plastron of abdominal ribs which 
occupies the whole of the front of the abdomen, from a little 
behind the coracoid to the front of the pubis. The riblets are 
all slender and arranged in series of threes—a long outer riblet, 
a shorter middle one, and a still shorter inner one. In front the 
innermost series remain free, but on passing back they become 
anchylosed into united groups of two, three, and four. There 
are probably four series of riblets to each vertebra, Altogether 
there are 31 groups of ribs. 

The pelvis is preserved in two specimens besides the type, and 
every detail of its structure is known. The ilium is similar to 
the type found in Belodon and Hrythrosuchus. ‘The acetabulum 
is large, and closed, and the ilium has a well-marked supra- 
acetabular ridge. The iliac crest is developed much more back- 
wards than forwards, and the whole crest measures 28 mm. in 
length. The pubis is comparatively short and broad. It forms 
a complete symphysis with its neighbour, and the outer border, 
which is considerably thickened, passes almost directly down- 
wards. A very remarkable feature of the bone is the presence of 
two pubic foramina. The upper one, which is apparently the 
homologue of the normal pubic foramen, is the smaller of the two 
and is fairly close to the acetabulum. The other is close to the 
symphysis. In the type specimen both pubes are well preserved 
and the foramina are exactly similar on both sides. The ischium 
is a variety of the plate-like form, but long and slender. It has 
a complete symphysis with the other ischium. 

The femur measures in the type 58 mm.: in a second specimen 
55mm. It has a slight double curve as in most reptilian femora. 
The ends have been largely cartilaginous. On the inner and 
posterior side, near the union of the upper and middle third, there 
is a well-marked small trochanter. 

The tibia and fibula are not perfectly preserved in any of the 
specimens, but in one or other both upper and lower parts are 
shown. They are apparently a little shorter than the femur and 
considerably more slender, 

The tarsus consists of two proximal elements, and probably 
four distal tarsals.. The astragalus is irregularly cubical, and the 
caleaneum considerably broader than long. There appears to be 
little in the way of a heel. Of the distal tarsals the 4th is the 
only large one. 

The metatarsals are all well developed, and the 5th has the 
peculiar Rhynchocephalian development. Of the others the 3rd is 
the longest. The 4th is slightly longer than the 2nd, and the 2nd 
considerably longer than the Ist. The Ist, 2nd, and 5rd digits 
have well-developed claws, but the 4th has the phalanges rather 
weak and the claw, if present, was very small. The 5th toe 
also has a small claw, though larger than that of the 4th. The 
digital formula is 2, 3, 4, 5, 3. 

Dermal ossifications are present along the back from the head 
to at least well down the tail. These are arranged in pairs, one 


624 DR. R. BROOM ON 


on each side of each vertebral spine. All the best preserved 
scutes are about twice as long as broad and have the long axis 
lying antero-posteriorly, 

Some at least of the ribs of the middle region of the body 
appear to have uncinate processes. These are little ossifications 
about 3 or 4 mm. in length and about 1 mm. in width. They 
are firmly attached to the posterior side of the ribs but not 
anchylosed. Just possibly, though much less probably, they are 
smal] lateral dermal scutes. 

Before discussing the affinities of Huparkeria and the Pseudo- 
suchians generally, it will be well to consider some points in the 
structure of the allied genera from Elgin. These very interesting 
specimens, preserved in the British Museum, reveal a few points 
in the structure of the group not seen in the specimens of 
Euparkeria, and afford a very thorough knowledge of the sub- 
order. Aétosaurus ferratus, on which the subondies Pseudosuchia 
was founded, has never been very thoroughly described, but 
Mr. Watson assures me that there is no doubt that it has two 
temporal vacuities like those of Huparkeria and Ornithosuchus, 
and not one as described and figured by Fraas. Though there 
is thus httle doubt that Aétosaurus belongs to the same embondler, 
it differs in a good many points from the Elgin and South-African 
forms. 


ORNITHOSUCHUS woopwARDI Newton. (Pls. LXXVII.- 
NONE XG ois Oe OG 525).)) 

The type of this Pseudosuchian is a fairly complete skeleton from 
Elgin, described in 1894 by Mr. E. T. Newton. The specimen is 
preserved in the British Museum, and through the kindness of 
Dr. Smith Woodward I have been enabled to make a fresh study 
of it in the light of the new knowledge obtained from the allied 
South-African form. Newton’s study of the type has been so 
thoroughly and carefully done that there is very little in the 
specimen he has failed to observe, and the points where I incline 
to differ from him are very few in number. Fortunately the 
British Museum has recently obtained a second specimen which 
supplies a number of blanks in our knowledge. 

The skull, on the whole, resembles that of Huparkeria in all 
essentials, so far as can be seen, though the cranial sutures are 
less easily made out in Ornithosuchus and the palate is unknown 
in Huparkeria. The skull of Ornithosuchus woodwardi is more 
slenderly built, and it differs in having apparently no inter- 
parietal, in the shape of the jugal, in “having a much larger 
antorbital vacuity, and in having the teeth differently arranged, 
In the restored side view of the skull which I give, and which 
differs only slightly from that given by Newton, the shape of 
the various openings and bones can readily be seen. The dental 
formula of Ornithosuchus appears to be 1.3, m.9 as against 
i.3, m.12 in Huparkeria; and there is the further difference 
that, while in the South African genus only one mandibular tooth 
overlaps the upper jaw-border, in Oriithosuchus there are two 


FOSSIL REPTILES. 625 


teeth which pass to the outside between the premaxillary and 
maxillary teeth. 

The palate of Ornithosuchus is well preserved but not com- 
pletely displayed in front. The restoration of it which I give is 
hypothetical only in the prevomerine region and in the basi- 
occipital. The pterygoids are large, as are also the palatines 
and ectopterygoids, and these three “bones of the two sides forma 
large vaulted bony roof to the mouth. On each side there is a 
pair of oval openings—one between the palatine, the ectopterygoid, 
and the pterygoid, and the other between the pterygoid and the 
palatine. This latter is regarded by Newton as the posterior 
nares. In this, I think, he is in error. The opening has the 
borders quite flat, and was probably covered by membrane in life 
and did not transmit any structure of importance. The ptery- 
goid sends forward a long slender process along the inner side of 
the palatine which doubtless meets the prevomer as suggested 
in the figure. 

The shoulder-girdle and anterior limb, though lost from the 
type, are fairly well preserved in the second specimen. The 
scapula differs from that of Huparkeria in being very narrow in 
the middle and much more expanded at the base. It is much 
more Dinosaurian in appearance. The coracoid has a rather 
small foramen, and the posterior border below the glenoid cavity 
is deeply notched. The interelavicle is narrow and: not expanded 
in front apparently. The clavicles are also slender. 

The humerus measures 57°5 mm. in length. There is a well- 
developed deltopectoral crest, and the general appearance of the 
bone suggests a comparison with that of the Theropoda. The 
radius and ulna are not very well preserved. The former 
measures 47 mm. and the latter 49 mm. Only afew bones of the 
manus are preserved, and these are apparently displaced. A 
cast of a specimen obtained by Mr. Watson shows rather more 
of the manus. There are pretty certainly three well-developed 
digits, and, I think, evidence of a fourth. The carpus is badly 
preserved: possibly it was imperfectly ossified. 

Ihave given a restoration of the pelvis for most of which, 
I think, there is clear evidence in the specimen. The pubis is 
essentially similar to that of Huparkeria, but much more elongated. 
It has a large single pubic foramen. There is evidently a com- 
plete symphysis. 

The bones of the hind foot of the type, though much displaced, 
have been separated out and identified by Newton with marvellous 
care, and the foot is seen to resemble that of Huparkeria pretty 
closely, the main difference being that the fifth metatarsal shows 
less of the peculiar Rhynchocephalian development, and the 
phalanges of this, too, are feebler. 


ORNITHOSUCHUS TAYLORI, sp.n. (PI. LX XVII. fig. 11.) 


In 1904, Mr. G. A. Boulenger described some rather badly 
preserved ‘reptilian remains from Hligin as a large example 
of Ornithosuchus woodwardi. Myr. Taylor, when sending the 


626 DR. R. BROOM ON 


specimen to the British Museum, recognised that it was a near ally 
of Ornithosuchus woodwardi, though about 24 times as large. 
Mr. Boulenger discusses at some length the question whether the 
specimen is merely an older example of O. woodwardi or a new 
species, and comes to the conclusion that there are no other 
differences than those that might be accounted for by difference 
of age. My comparison of the specimens has led me to con- 
clude that the animals, though allied, are at least distinct 
species, and I have therefore much pleasure in naming the form 
after the discoverer. 

The specimen shows most of the skull. The jugal is probably 
perfect but is partly hidden by matrix, and the quadrato-jugal 
is also nearly perfect. The squamosal and postorbital are much 
crushed, but the side view of the back half of the skull can be 
restored with moderate accuracy. Most of the upper side of the 
skull is preserved, but partly broken and not well displayed. A 
fairly complete snout with maxilla and premaxilla of what may 
perhaps be a second individual isalso preserved. And as we have 
the remains of the last maxillary tooth in the first specimen, and 
the corresponding tooth in the snout-fragment, we can make a 
fairly complete restoration of the skull. When this is done, the 
very marked differences between it and the skull of Ornithosuchus 
woodwardi are apparent, ‘The arches are seen to be massive 
‘instead of slender bars, and the snout relatively much more 
powerful. The fact of the two animals being distinct is further 
confirmed by the fact that the British Museum has recently 
obtained from Elgin two new specimens—the one exactly 
corresponding in size to the type of Ornithasuchus woodwardi, 
and the other, so far as can be made out, agreeing in size 
with the large form. In the new specimen of O. taylori part 
of the palate is preserved, and it agrees essentially with that of 
O. woodwardi. There are two similar openings in the back part, 
and the pterygoid sends forward a similar but relatively larger 
anterior process. 


HERPETOSUCHUS GRANTI Newton. 


This small Pseudosuchian from Elgin is less satisfactorily pre- 
served than Ornithosuchus woodwardi, but there is sufficient to 
show that it is a fairly near ally. The skull differs considerably 
in its proportions, but probably the essential structure is similar. 
The palate is narrower, and the anterior process of the pterygoid 
is relatively broader, and there is only a single opening in the 
pterygoid region. The shoulder-girdle and anterior limb are 
differently proportioned, being much more slender, and indicate 
that the habits of the two genera were different. An interesting 
point revealed by this specimen is the fact that the manus has 
five well-developed digits. In the carpus there are at least three 
elements. The five metacarpals are well preserved, but most of 
the phalanges are missing. 


FOSSIL REPTILES. 627 


BROWNIELLA AFRICANA, gen. et sp. n. ' (Pl. LX XIX. fig. 21.) 


In Mr. Brown’s collection there are portions of at least two 
skeletons of an animal rather larger than HLuparkeria capensis 
but closely allied to it. The only parts well preserved are the 
shoulder-girdle, pelvic girdle, and femur. These indicate an 
animal nearly a half larger than the better known African 
form. 

The following measurements will illustrate the differences in 
size of the two forms :— 


Huparkeria . Browniella 
capensis, africana. 
F mim. mim. 
Width of base of scapula ...... 15 20 
enguhoh coracoidy ese eee 24 30 
Pens theotsisehnumy eee. nee at ee 36 about 50 
Wemethvolstomuiy ees. a aes 56 74 


In general structure there is a close similarity between the 
bones, but those of Browniella are more massive, and there are 
numerous minor differences. 

The ischium differs in shape in being constricted near its 
middle, and the pubis, besides being much broader and stouter, 
differs in having only a single pubic foramen. This difference 
seems of sufficient importance to justify the placing of this 
species In a separate genus, and I have much pleasure in naming 
it after Mr. Alfred Brown, the veteran collector, to whom Science 
owes such a deep debt. 


MEsosucHUSs BROWNI Watson. (Pls. LXXVIII., LXXIX. 
figs. 12-15, 23.) 

Though Mesosuchus brown? differs considerably from Huparkeria 
and is probably not a Pseudosuchian, it has nevertheless some 
interesting affinities, and, both from its association with Ha- 
parkeria and from the fact of the two forms having been confused, 
it seems well to consider the one along with the other. 

As already mentioned, Watson, in his description of MWesosuchus 
browint, had regarded some of the imperfect skeletons associated 
with it as belonging to the same form. As we now know that 
most of these skeletons belong to the similar sized but distinctly 
different Huparkeria capensis, it is necessary to redefine J/eso- 
suchus to some extent. 

The type specimen consist§ of a badly crushed skull showing 
the premaxilla and maxilla of the left side in good condition, 
with most of the lower jaw and most of the palate badly crushed. 
Much of the rest of the skull is present, but so badly distorted 
that a restoration is difficult. Of the rest of the skeleton of the 
type individual almost the whole of the vertebral column from the 
head to about the middle of the tail is preserved. Of the shoulder- 
girdle there is practically nothing preserved, though much of the 

Proc. Zoou. Soc.— 1913, No. XLII. A2 


628 DR. R. BROOM ON 


right arm is seen. The pelvis is badly preserved, but both ischia 
ave well seen and most of the left hind leg. 

Tie pcremaxilla is particularly interesting in having no inter- 
nasal process, and in this resembling the bone in Rhynchosaurus 
and Hyperodapedon. It has two well-developed acrodont teeth, 
which ave round and blunt. The maxilla is long and narrow, 
and has 13 irregularly arranged rounded blunt acrodont teeth. 
A disarticulated quadrate, which is not that of Zuparkeria and 
pretty certainly that of Mesosuchus, is nearly as broad as long, 
and uch more massive than the quadrate of Huparkeria. here 
are two moderately distinct condyles. Watson's description of the 
palate cannot at present be added to. “ Pterygoid of remarkable 
shape with a deep posterior ramus applied to the inner side of 
the quadrate, small external ramus (not well exposed) and long 
anterior ramus which bears a closely-set series of small pointed 
teeth. Vomer apparently narrow, with a series of small poimted 
teeth articulated with anterior end of pterygoid. Other bones 
of palate not shown. Epipterygoid widened with a deep notch 
for the optic nerve, touching the top of the deep posterior ramus 
of the pterygoid. Parasphenoid very large and placed high up in 
the skull.” Most of the postcranial skeleton described by Watson 
are really bones of Huparkeria, while the supposed scapula is 
really the ischium of Browniella africana. 

The lower jaw is fairly well preserved in the type specimen. 
Tt differs from the jaw of Huparkeria in having a relatively 
small lateral opening and in having the part of the jaw behind 
the opening larger than the dentary portion. The surangular 
forms more than the upper half of the outside of the back of the 
jaw. and the rest is mainly formed by the angular. 

The vertebrz are not well preserved. They are of about the 
same size as those of Huparkeria capensis. ‘The cervicals have 
fairly long spines, and the whole neck 1s relatively longer than in 
Eupurkeria—probably 11 vertebie may be cervical. Altogether 
there a pest to be, as in Huparkeria, 26 pre-sacral vertebree, and 
apparently 2 sacral. 

The humerus, radius, and ulna are much more massive than in 
Kuparkeria, but not very well preserved. The humerus measures 
37 mm. in length. 

The ilium differs considerabl y from that of Kuparkeria, and 
resembles much more closely that of Howesia. hough imper fectly 
preserved, the upper part of the ium is manifestly about twice 
as deep as in Hupark ervd. ‘The ischium also differs markedly from . 
that of HMuparkerva in being relatively much shorter, and having 
only a short symphysis. The pubes are very badly pre esenveds 
but have manifestly been much broader than i Kuparkeria, 
though essentially similar in type. The illustrations given show 
the specimen as preserved and the pelvis restored. 

The femur, tibia, and fibula are not unlike those of Huparkeria. 
The femur probably measures 49 mm. in length, and the tibia 
47mm. The tarsus has the bones displaced, but 1s apparently 


FOSSIL REPTILES. 629 


better ossified than in Huparkeria. There are three large 
elements and at least two small ones. ‘The rest of the pes is 
probably of the Rhynchocephalian type. 

There seem to have been no dermal ossifications in Mesosuchus. 


SCLEROMOCHLUS TAYLORI Smith Woodward. 


One of the most remarkable of the double-arched reptiles from 
the Elgin sandstone is the little form named by Dr. Smith: 
Woodward Scleromochlus taylori. Though there are preserved in 
the British Museum the remains of five individuals, and though 
something is known of most of the skeleton, the animal is too 
small to be well preserved in the coarse sandstone, and hence our 
knowledge of the detailed structure is very imperfect, Smith 
Woodward’s description and restoration give practically every- 
thing one can be sure about. The skull seems to be essentially 
similar to that of Ornithosuchus. ; 


Affinities of the Pseudosuchia. 


Mr. EK. T. Newton, in describing Ornithosuchus and Herpeto- 
suchus, discusses the affinities of the forms. He calls attention to 
the marked resemblances of the forms to Aétosaurus, and even 
thinks 1t possible —as we now know to be the fact—that détosaurus 
has an infratemporal vacuity, and also discusses the relationships 
with Stagonolepis and others of the Parasuchia. A comparison is 
also made with the Dinosaurs Compsognathus, Anchisaurus, and 
others, and with the skull resemblances in the Pterosaurian 
Scyphognathus. In summing up the evidences he says:—‘ The 
many points of resemblance between the Parasuchia and certain 
of the forms usually included among the Dinosauria, have also 
been noticed by other writers; and the difficulty of separating 
the two groups is increased by a study of this new Elgin reptile, 
which holds, as I think, a more intermediate position between 
the two series than any form hitherto described, for although the 
characters of its skull and teeth find their nearest counterpart 
among the Dinosaurs, and the pelvis and limbs might belong to 
either a Theropodous Dinosaur, or a’ Parasuchian, the form of 
the free astragalus is more Crocodilian than Dinosaurian. While 
acknowledging the ditliculty of assigning this new reptile to either 
of these groups, it seems most in accordance with the facts to 

lace it provisionally with the Dinosaurs.” 

Mr. G. A. Boulenger, when describing the large specimen of 
Ornithosuchus, criticises Newton’s conclusions. He says:— 
“ Much as I admire Mr. Newton’s description of Ornithosuchus 
....I1 cannot... . endorse his conclusions as to the systematic 
position of the reptile.’ “ Dr. Smith Woodward was nearer the 
truth when he compared it with Aéfosawrus.” Boulenger argues 
that Ornithosuchus should be placed with Belodon, Stagonolepis, 
and dAétosaurus in the order named by Owen, Thecodontia—a 


495 


630 DR. R. BROOM ON 


group which agrees quite as much with the Rhynchocephalia and 
the Carnivorous Dinosaurs as with the Crocodilians. 

Though these two opinions seem at first sight to be at variance 
they are really pretty similar. Practically, it amounts to this, 
that in the Pseudosuchia we have a group of primitive reptiles 
which, while they do not fit into any of the later specialised 
orders, have affinities with quite a number of other groups. 

There cannot, I think, be the shghtest doubt that the Pseudo- 
suchia have close affinities with the Dinosaurs, or at least with 
the Theropoda. This has been recognised by Marsh, v. Huene, 
and others. In fact there seems to me little doubt that the 
ancestral Dinosaur was a Pseudosuchian. The skulls of such types 
as Huparkeria or Ornithosuchus are practically Dinosaurian even 
in detail, and the skulls of the early Dinosaurs, Such as Anchi- 
saurus, differ less from the skulls of Pseudosuchians than those 
of the early Dinosaurs do from many of the later types. And 
there is nothing in the post-cranial skeleton that is not just what 
we should expect to find in the Dinosaur ancestor. The shoulder- 
girdle is more primitive in retaining clavicles and interclavicle, 
but these are elements which we know from the history of other 
groups are very variable and readily lost. The pelvis is almost 
Dinosaurian, and differs only in having the acetabulum closed. 
This is an important character ; but when we consider the con- 
dition in the two nearly allied Monotremes—the one with the 
acetabulum closed, the other with it open—we see how easily even 
this character may change. ‘The hind limb is almost Dinosaurian 
in Huparkeria. The ankle is less specialised and the fifth toe is 
still well developed and retains the Rhynchocephalian characters. 
Huparkerva is ia my opinion potentially bipedal, and was probably 
partly bipedal in its habits. The fourth toe of the hind foot is 
more feebly developed than the third and the axis of the foot is 
down the third toe, which would seem to indicate that the feet 
were at least not so laterally placed as in lizards, and that the 
animal possibly ran on its hind feet. The relative shortness of the 
toes also seems to confirm this view, as well as the feebleness of 
the fore limbs. I believe Huparkertia fed on some large forms of 
insects like locusts, and captured them with its front feet. 

Ornithosuchus was probably very similar in habit to Huparkeria 
and was even a little better adapted for running on its hind feet ; 
the large species O. taylor could hardly have had the same habits 
as the small O. woodwardi. tis too massively built, and probably 
had become largely carnivorous, adding to thelarger insects various 
small vertebrates, and perhaps, like the vulture, the flesh of 
animals too large for it to kill. 

The aflinities of such small Pseudosuchians as Huparkeria, 
Ornithosuchus, and Aétosaurus with the Belodonts through such 
an intermediate form as Hrythrosuchus is very manifest. And as 
Boulenger stated, the Pseudosuchians are about as near to the 
Parasuchians as to the Dinosaurs. The series of dermal plates 
down the back, though suggesting affinity is, however, not a 


FOSSIL REPTILES. 631 


character on which too much weight must be placed. As we 
see in the Lacertilia, dermal ossifications are subject to great 
variation in even allied forms. 

Another group to which the Pseudosuchians seem to have 
affinities as suggested by Newton, isthe Ornithosauria. In general 
proportions the Pterodactyles differ very greatly, but the form 
from which they arose must have been very much like that seen 
in Pseudosuchians. ‘The Pterodactyl and Pseudosuchian skulls are 
almost exactly similar in essentials. As pointed out by Newton, 
theskullin Scyphognathus resembles pretty closely that of Ornitho- 
suchus. The Pterodactyl manus is simply an ordinary reptilian 
manus with the 5th digit lost and the 4th greatly specialised, 
and there can be no doubt that the 5th digit was lost before the 
wing-membrane was developed. The pelvis of the Pterodactyl 
is not thoroughly known beyond doubt, but seems to be a modi- 
‘fication of the ordinary plate-like type with the prepubis ossified. 

Seleromochlus is a very lightly built Pseudosuchian evidently 
adapted for taking leaps, and not for bipedal progression on the 
ground. The limbs are long and slender, and the length and 
slenderness of the fore limb suggest that possibly there was a 
membrane stretched between the fore and hind limbs and perhaps 
between the hind limbs and tail, which would enable the little 
animal to take sustained leaps like Petauwrus. 

Although Scleromochlus is already too specialised in the hind 
foot structure to have been in any way ancestral to the Ptero- 
dactyls, it may suggest how they have arisen, just as Galeopithecus 
suggests how the bats arose. 

There is still another group to which some Pseudosuchian has 
probably been ancestral, namely, the Birds. For a time one or 
other of the Dinosaurs was regarded as near the avian ancestor. 
The resemblance of the hind limb and pelvis seemed to make this 
extremely probable, and Huxley, Marsh, Cope, and others have 
all favoured this view. Others, however, were more impressed by 
the apparently avian characters in the skeleton of the Ptero- 
dactyls, and especially in the striking avian appearances in the 
brain, and have argued in favour of a close affinity between 
the Birds and the Pterodactyls. Osborn, while recognising 
the affinities to both groups, and especially to the Dinosaurs, 
believed that the Birds and the Dinosaurs had a common ancestor, 
probably in the Permian. Seven years ago, when describing the 
skeletogenesis of the Ostrich, I argued that the bird had come 
from a group immediately ancestral to the Theropodous Dinosaurs. 
The Pseudosuchia, now that it is better known, proves to be just 
such a group as is required. In those points where we find the 
Dinosaur too specialised we see the Pseudosuchian still primitive 
enough. The bird pelvis has probably developed from a type 
like that of Ornithosuchus by the pubis turning further back and 
the symphysis becoming lost. Whether the union of the meta- 
tarsals is a primary or a secondary character is a debatable point. 
The question is really whether the bird ancestor was a hopping 


632 DR. R. BROOM ON 


bipedal animal before it flew, or if it only hopped after the wing 
had become specialised. Jam strongly of the opinion that it was a 
hopping animal first, and that the metatarsus became strengthened 
to support the weight of the body entirely borne by the hind feet. 
It is easy to understand a hopping animal taking to an arboreal 
life and ultimately developing a wing out of a four-toed hand, 
while it seems unlikely that the hind foot could ever have 
developed by arboreal habits. It is interesting to note that 
while the ancestor of the Pterodactyls had four toes in the manus, 
there is very clear evidence from the skeletogenesis of the bird 
that the latter also had a four-toed ancestor. 

A Pseudosuchian which through a bipedal habit had developed 
a strengthened ankle-joint and a firm metatarsus, and had lost 
the 5th “digit from the manus would meet all the requirements of 
the avian ancestor. 

We know at present too little to discuss the relationship of the 
Pseudosuchians with Sphenodon and with Gnathodonts, nor can 
we say whether Howesia and Jesosuchus should be placed with 
the Pseudosuchians. ‘There is evidence of a generalised Permian 
Rbynechocephaloid order which gave rise to the more specialised 
‘Triassic groups, but at present we know too few forms and very 
few even of these are well known, and until our knowledge has 
much advanced it seems unwise to attempt any further classifi- 
cation. In South Africa we can trace through the Lower Triassic 
and Upper and Middle Permian beds forms that may be ancestral 
to the Pseudosuchians, and when these are better known a 
satisfactory classification will be possible. 

Tam much indebted to Mr. EH. 8. C. Dyke for the photographs 
of Kuparkeria. 


References to Literature. 


(1) E. T. Newron.—“ Reptiles from the Elgin Sandstone. 
Description of two New Genera.” Phil. Trans. vol. 185, 
1894, p. 573. 

(2) G. A. Boutencer.—“ On Reptilian Remains from the Trias 
of Elgin.” Phil. Trans. vol. 1968, 1903, p. 175 

(3) A. 8. Woopwarp.—“‘ On a new Dinosaurian Reptile (Sclero- 
mochlus taylori) from the Trias of Lossiemouth, Elgin.” 
Owe Gasavolsixan, W907.) o. VAs 

(4) F. v. Huzne.—“ Die Dinosaurier der Europiischen Tyrias- 
formation.” Geol. u. Pal. Abhand., Jena, 1907. 

(5) D. M. 8. Warson.—* Mesosuchus panne. gen. et spec. nov.” 
Rec. Albany Mus. vol. il. pt. iv. p. 296. 

(6) R. Broom.—‘ Note on Mesosuchus brownt Watson, and on 
a new South-African Triassic Pseudosuchian (Huparkeria 
capensis).” Ree. Albany Mus. vol. 11. pt. v. 1913, p. 394. 


A full list of other papers will be found in y. Huene’s work. 


FOSSIL REPTILES: 633 


EXPLANATION OF THE PLATES. 


Ang. Angular. B.o. Basioccipital. Cl. Clavicle. Co. Coracoid, D. Dentale, 
Fr. Frontal. JZ.cl. Interclavicle. J.P. Interparietal. Ju. Jugal, 2. Lacryinal, 
Mx. Maxilla. Wa. Nasal. Pa. Parietal. Pa.o. Paroccipital. Pma. Premaxilla. 
Po.F. Posttrontal. Po.O. Postorbital. Pr.F. Prefrontal. Q. Quadrate. @./, 
Quadrato-jugal. S.Ang. Surangular. Se. Scapula, 8,0, Supraoccipital. Sy, 
Squamosal. Sé. Sternum. 


Pyatr LXXV, 
Kuparkeria capensis, 


. Side view of type specimen. 4 nat. size. 

Top of skull of type specimen. Nat, size, 

. Left dentary of a second specimen. Nat. size. 
Right hind foot of this second specimen, Nat. size, 


Fic. 


02 bo ju 


PratE LXXVI. 
Euparkeria capensis, 


. Side view of skull, Nat. size. 

. Upper view of skuil. 

. The quadvate viewed obliquely from behind, showing the relations to the 
adjoming bones. Simall portions of the squamosal and opisthotic are 
broken off. 

8. Shoulder-girdle, slightly restored and with the probable cartilaginous sternum 

added in dots. Nat. size. 


“TD on 


Prave LXXVIT, 


Fig. 9. Side view of skull of Ornithosuchus woodwardi. Nearly nat, size. 
10. Under view of skull of Ornithosuchus woodwardi.. Nearly nat. size, 
11. Side view of skull of Ornithosuchus taylori. About 2 nat. size. Restored 


trom the type specimens. 
Prate LXXVIII. 


Fig. 12. Side view of imperfect skull of Mesosuchus browni. Nat, size. 
13. lnpression of the teeth of the right maxilla of Mesosuchus browni. Nat, 
size. 
14, Fragment of ilium, and left hind limb of Wesosuchus browni, Nat. size. 
15. Fragmentary pelvis of Wesosuchus browni. Nat. size. 
16. Shoulder-girdle and fore lnnb of Ornithosuchus woodwardi. About + nat, 
size. Slightly restored. 
17. Top of lett ihum of Huparkeria capensis. Nat. size. 
18. Acetabular portion of left illum of Huparkeria capensis. Nat, size, 
19. Portions of three lower dorsal ribs of Huparkeria capensis showing unci- 
nates. Nat. size, 
20, Right temur, tibia, and fibula of Huparkeria capensis. Nat, size, 


Pratt LXXTX, 


. Right pubis, left ischium, portions of both ilia and of two dorsal vertebra 
ot Browniella africana, Nat. size. 

. Right pubis of Huparkeria capensis. Nat, size, 

. Pelvis of Mesosuchus browni. Nat. size. 

|. Pelvis of Huparkeria capensis. Nat. size. 

. Pelvis of Ornithosuchus woodwardi. About > nat, size, 

. Pelvis of Gryponyx africanus, Much reduced, 


i>) 
ie 
ug 
to 

a 


to bw br 
Boe wr 


634 MR. G. E. BULLEN ON 


41. On some Cases of Blindness in Marine Fishes. 
By G. H. Butien, the Hertfordshire Museum, St. Albans *. 


[Received April 24, 1913: Read May 20, 1913. ] 


Of recent years considerable attention has been given by 
numerous ichthyopathologists and fish-culturists to the sporozoic 
and bacterial diseases affecting those species of fish which in a 
live state form an article of commerce. 

The important work of Hofer+, and the Monograph now 
appearing from the pen of de Drouin de Bouvillet, are perhaps 
two of the best known contributions of a general character 
upon the subject. There are, moreover, numerous more or less 
important works upon those organisms giving rise to disease: the 
Myxosporidia, for example, have received considerable notice 
from such investigators as Balbiani§, Thelohan |), Doflem 4, 
and Gurley **, to quote but a few of the authors upon the 
subject. 

Work which has been performed, even within the past decade, 
upon the bacterial aspect of fish diseases is of too extensive a 
character to be cited, however brietly, here. The same applies 
to Saprophytic and other fungoid growths attendant upon 
bacterial lesions. 

The investigations of J. Hulme Patterson and certain others 
upon Salmon disease are well known and important contributions 
to economic science in this respect. But with the exception 
of certain papers by G. H. Drew and Patterson’s findings, so 
far as they apply to Salmon installed in sea-water, most of 
the literature dealing with disease in marine fishes is confined 
almost entirely to the cecology of pathogenic organisms. For, 
whereas there is no inconsiderable amount of the literature 
directed solely to the pathology of freshwater fishes, there is 
at present a remarkable paucity dealing with those of marine 
habitat. 

The present paper, therefore, is put forward largely with a 
view to associating the pathological aspect of certain forms of 
eye diseases observed in marine fishes with those already 
described, in more or less general terms, in the case of freshwater 
species. 


* Communicated by Joun Hopkinson, F.LS., F.Z.S. 

+ ‘Handbuch der Fischkrankheiten.’ 

t+ ‘Les Maladies des Poissons d’eau douce d’ Europe.’ 

§ ‘ Lecons sur les Sporozoaires,’ Paris, 1884, etc. 

|| “ Recherches sur les Myxosporidies,” Bull. Soc. France et Belgique, ser. 4, vol. v. 
Paris 1894. ete. 

4 ‘Die Protozoen als Parasiten und Krankheit Serrigers,’ Jena, 1901. “Studien 
zur Naturgeschichte der Protozoen,” Zoologische Jahrb. xi. 1889, ete. 

** “ Myxosporidia, ete.” Rep. U.S. Commissioner for Fish and Fisheries, 1892. 
Washington, 1894. ; 


BLINDNESS IN MARINE FISHES. 635 


The several subjects described hereinafter were received by 
the writer from the Superintendent of the Brighton Marine 
Aquarium, Mr. EK. W. Cowley, to whom thanks are due. 
Acknowledgment of much assistance in the preparation of this 
paper is also accorded to Mr. John Hopkinson and to the late 
Mr. G. Harold Drew. 


Hofer * traces the progress of the more usual form of blindness 
in freshwater fishes in the following manner: (a) Corneal 
“cataract” due to injury or pathogenic bacteria attacking the 
integument; (3) gradual necrosis of the cornea-epithel and 
outer surface of the cornea; (vy) perforation of the cornea. 
Complications which may secitte ave “ Keratoglobus” in the 
earlier stages, and Saprophytic growth attendant upon the 
bacterial lesions. 

In the case of several freshwater fishes, the same author 
describes a not uncommon form of blindness due to parasites 
(Diplostomum larve), but of all the marine fishes which herein 
receive notice, none has been affected in this manner. More- 
over, it may be recorded that no evidence of Saprophytic growth 
occurred on even the worst examples. 


1. Traumatic Corneal Opacitis in a Conger Eel 
Conger vulgaris). 
fo) 5 


The subject of this note was a Conger Kel which measured 
3 feet 7 inches. It had lived in the Aquarium for nearly six 
years, retaining complete health until November 1912, when, 
according to the Superintendent of the Aquarium, Mr. E. W. 
Cowley, it went completely blind within the space of a week. 
It was destroyed immediately, when, from an observation of its 
behaviour, both eyes appeared to have become wholly useless. 

An examination of the head of this specimen showed that the 
right cornea-epithel was perforated in two places and the left in 
five, each perforation being just large enough to admit a seeker 
-5 mm. in diameter. Extensive penetration of sea-water followed 
probably by bacterial lesions had occurred throughout the under 
surface of the cornea-epithel, those on the left side extending to 
the adjacent epidermis of the gill-covers, destroying the colour- 
cells, and rendering the part affected opaque white. The eye 
itself, when superficially examined, was so completely dimmed 
as to render observation of the characteristic yellow iris difficult, 
except in strong light. 

The term ‘corneal opacitis” would be somewhat loosely 
applied in this instance, since upon removal of the integument 
the cornea proper was found to be completely free from injury, 
and but slightly affeeted with opacitis. 


* Op. cit. p. 291 etc. 
+ For a determination of this fact the writer’s thanks are due to Miss Lorraine 


Smith, of the British Museum. 


636 MR. G. E. BULLEN ON 


The condition termed by Hofer * ‘ Keratoglobus,” 7. e. out- 
ward bulging of the cornea away from the pupil, did not affect 
either eye, the lenses retaining their normal position. 

Blindness, therefore, in this instance was due apparently to 
mortification and bacterial lesions of the integument of the head 
passing over the eye attendant upon perforation of the cornea- 
epithel. 

Hofer? remarks: “In the case of a fish weakened by disease, 
or in a dying condition, the organs of sight are deeply sunk 
in their orbits, whereas in a healthy fish the eyes somewhat 
protrude.” In the present example this condition was wholly 
apparent, the eyes presenting the flabby appearance to be seen in 
a fish some hours after death. 

It is highly probable, therefore, that perforation of the integu- 
ment above the cornea, resulting in blindness, was largely due, 
in the present instance, to the susceptibility of the fish, alr eady 
weakened by graver organic disease, to injury ; and that incipient 
corneal opacitis may sometimes be taken as a diagnostic of other 
disorders. 


Corneal Opacitis in a Greater Weever 
(Trachinus draco). 


The chief difference between the condition of the eyes in the 
present example and that of the Conger Hel already described, 
lay in the fact that apparently no lesion of the cornea-epithel 
appeared to have taken place. The subject under consideration 
had lived for some years in the tanks at Brighton, and had 
attained the considerable measurement of 15? inches. In this 
case partial blindness only had come about at the time when the 
specimen was examined. The left eye was less affected than 
the right, this showing a more or less evenly distributed film of 
dimness, as if the specimen had been immersed for a short time 
in weak formalin. The right eye, however, not only showed 
distinct spots of necrosis unevenly distributed throughout the 
surface of the cornea, but there was also a pronounced “ kerato- 
globe.” With reference to this latter feature, it may here be 
remarked that a gathering of aqueous humour in front of the 
pupil, causing the cornea to bulge outward to a considerable 
extent, may occur, doubtless as the result of concussion, in fish 
the eyes of which are otherwise apparently unaffected. At the 
Brighton Aquarium occasionally numbers of a shoal of mackerel 
may be observed with both eyes protruding in this way, but 
their undoubted lack of effectual vision appears in no way to 
interfere with their capability for maintaining position in a 
moving shoal. But it is a noticeable fact that they appear 
much thinner than their fellows consequent upon a more limited 


* Op. cit. p. 293. 
+ Op. cit. p. 290. The translations adhere as nearly as possible to the original 
text. 


BLINDNESS IN MARINE FISHES. 637 


absorption of food. Not unfrequently, moreover, such examples 
present a certain broken-backed appearance, which, so it is stated 
by the attendant in charge of the tanks, is due to the fish, when 
first installed, persistently hammering themselves against the 
glass! 


3. Corneal Opacitis and Cataract in Pollack 
(Gadus pollachius), 


The Pollack forming the subject of the present note were four 
young specimens ranging from 74 to 83 inches in length. At 
the time when they came under the writer’s observation (in Sept. 
1912) they had lived in the Aquarium, in a tank apart from 
other fish, for a period of about nine months. When first 
installed their sight appeared to be normal; but it seems that 
these particular fish had been taken on a position close inshore 
off Shorehain, where there is an outflow of chemical waste. 

Mr. E. W. Cowley informed the writer that fish which were 
known to have come from this locality were as a rule refused for 
stock purposes, in view of the fact that, after a short period in 
the Aquarium, they invariably showed signs of disease. Hofer * 
states : “ Trout, etc., have been fr equently, attacked by perforation 
of the cornea and cataract through existing in waters impreg- 
nated with iron from drains”; and goes on to say, * It is not yet 
established whether these consequences are to be ascribed to 
iron salts in general or to any particular one.” A lack of 
similar observation, viz.: the effect of industrial wastes, 
sewerage, etc., upon those marine fishes often frequenting 
estuaries and tracts of water in which such impurities are known 
to oceur, prohibits any comparison. Yet it is a noticeable fact 
that the examples about to be described exhibit a form of 
blindness most frequently met with in various freshwater fishes 
(e. g. Pike, Perch, Carp, ete.). 

At the time when the present specimens came under the 
writer's observation, they were without doubt totally blind, both 
eyeballs of each example presenting the opaque white appearance 
similar to that seen in the case of fish which have been boiled. 
The cornea in every case was but slightly affected, no apparent 
lesion being discer nible on any part of the integument covering 
aiveameine general transparency, moreover, was consider: ably gr eater 
than it was in the case of either of the two preceding examples. 
On dissection it was found that the cornea protruded in front of 
the lens to a considerable extent (cir. 24 mm.). A transverse 
section, taken through the centres of both eyeballs from two of 
the specimens, showed that the extent of necrosis attacking the 
interlocking fibres was practically the same, viz., equivalent to 
two-thirds of the entire diameter of the section. Vertical 
sections, ¢. é. from anterior to posterior aspect of the lens, showed 


*Opcitaps 201. 


638 ON BLINDNESS IN MARINE FISHES. 


a slight increase on the anterior margin, the total extent, how- 
ever, being slightly less than that observed in the transverse 
sections, amounting roughly to slightly less than half the diameter 
in affected tissue. The term “cortical cataract,” as applied to 
the human subject, would perhaps aptly describe the condition 
observed when in its earlier stages *. 

The behaviour of these fish in the Aquarium is a matter 
deserving of some comment. Habitually they were observed to 
swim in mid-water, maintaining one position often for several 
minutes together, and seldom exhibiting much activity in move- 
ment. It is a somewhat remarkable fact, but one which the 
present writer, by an observation extending over several weeks, 
can personally vouch for, that on no occasion were any of the 
specimens seen to come in contact with each other, the sides 
of the tank, or the glass. Pollack with normal vision installed 
In an adjacent tank were easily attracted when a moistened 
finger-tip was rubbed rapidly up and down the glass causing 
vibration thereby, less freely by gentle tapping. The blind fish, 
even when situated within a few inches from the glass, made no 
effort to respond to this attraction. 

When food, in the form of finely-cut fish, molluscs, ete., 
was offered, it was allowed to sink to the bottom and remain 
there for several minutes before a search after it was made. 
This took the form of a wholly haphazard grubbing about on the 
bottom, the fish often making a vigorous “ bite” immediately 
beside a fragment, this operation being repeated until the morsel 
was secured. Although a sufficiency of food was placed in the 
tank daily, and on one occasion one individual fish was observed 
within the space of 15 minutes to secure three fragments of 
moderate size, all the specimens when examined post-mortem were 
found to be very ill-nourished and considerably below normal 
weight. 


Nore :—The writer is indebted to Mr. J. O. G. Ledingham, of 
the Lister Institute of Preventive Medicine, for the following 
report respecting the histological condition of the lens in one of 
the above specimens. This gentleman writes “I cut Celloidin 
sections of the fish’s eye you sent to me in formalin. They were 
very difficult to cut owing to the fact that the lens proved to be 
completely calcareous. This was the only obvious lesion I could 
detect. I saw no micro-organisms in the sections; but, as you 
will understand, the sections were not satisfactory for the 
demonstration of micro-organisms.” 


* Vide L. B. Harman, “ Aids to Ophthalmology,” p. 88. 


STR ARTHUR H. CHURCH ON TURACIN. 639 


42, Notes on Turacin and the Turacin-Bearers. 


By Sir Artnur H. Cuurcu, K.C.V.O., F.R.S.* 
[Received May 26, 1913: Read June 3, 1913.] 


In bringing, at the suggestion of the Secretary, these notes 
before the Zoological Society I have a threefold object.: First, 
to give a summary history of our knowledge of the occurrence in 
the Plantain-eaters of turacin, the remarkable (1 may say, the 
unique) decorative pigment which is distinctive of the three genera 
Musophaga, Turacus, and Gallirex ; secondly, to describe the 
properties of the pigment; and, thirdly, to correct some prevalent 
errors on the subject of turacin—errors due in part to initial 
mistakes made by observers, and in part to incorrectness in the 
transmission cf facts by copyists. 

Although the earliest observation as to the occurrence in 
certain Turacoes of a singular red pigment seems to be assignable 
to the year 1818, yet I have not found any publication of the fact 
by the original observer until more than half a century later. 
Then, on January 17th, 1871, a Corresponding Member of this 
Society, M. Jules Verreaux, described at a meeting his observa- 
tions and experiments on the subject, referring them back to 
the year 1818. Verreaux’s remarks are to be found in the 
‘Proceedings of the Zoological Society’ for 1871 (pp. 40 eé seq.); 
they refer to the Corythaix albicristata of Strickland. Many of 
these Turacoes were met with in the Knysna district of the Cape 
of Good Hope, and were carefully observed by Verreaux. He 
noticed that, in the torrential rains that prevailed at the time, 
these birds left the tree-tops and sought refuge in the dense lower 
branches. One bird, which he had seized by the wing, escaped, 
and he says that great was his surprise to see the inside of his 
hand coloured a blood-red. Some days later the experiment was 
repeated with three more specimens caught in a soaked condition ; 
it was easy to remove the pigment from the feathers by friction 
and to reduce them to a pale rose hue. What M. Verreaux here 
adds to his observation as to the solubility of the red pigment in 
rain-water is, | venture to think, hard to believe. He wrote in 
French, and I give his own words :—‘‘ Mais ce qui nous surprit 
le plus, fut de voir cette méme coloration rouge vif revenir des 
que loiseau était complétement séché. Nous avons renouvelé 
cette opération deux fois par jour, et chaque fois nous avons eu 
le méme résultat.’” M. Verreaux further extends his observa- 
tions to other species of Turacoes, finding in them the same 
colouring-matter, possessed of the same mobility and of the same 
power of renewal when the feathers became dry. He even 
affirms such recovery of colour in the case of birds that have been 
killed as well as of those that have been captured. One cannot 


* Communicated by the SrecRETARY. 


640 SIR ARTHUR H. CHURGH ON TURACIN. 


but ask, how is it possible for a mature feather to be again 
supplied with this localized red pigment after it has lost its 
original charge and after the vessels in the quill have dried up, 
their function having been fulfilled? The opinion of VY. Fatio, 
as to an oil making its way up the mature feather and dissolving 
and distributing pigmentary deposits already present, does not 
apply to turacin which is insoluble in oily media. As to the case 
now being considered, [ have not been able to obtain any con- 
firmation of this renewal of the pigment (apart from a moult) 
from any trustworthy naturalist. The late J. J. Monteiro, the 
late Dr. Benjamin Hinde (of Bathurst on the Gambia), and many 
other competent observers of these birds in their native haunts 
and in captivity have expressed themselves strongly on this 
point. It may be admitted that in the folded pion which has 
had its pigment moved by soaking, some of the red solution may 
travel from the feathers beneath to the surface as it dries; thus 
we should have a case of transference of pigment, not of its 
renewal. 

Amongst those who, after Verreaux, have observed this want 
of fixity in the red pigment of the Turaco I may name the late 
Rowland Ward, the late W. B. Tegetmeier, also J. J. Monteiro 
and Dr. B. Hinde.. The last-named, in a letter dated May 1865, 
said the moment soap touches the feathers the colour runs, 
although it is difficult of extraction by pure water. But he 
added ‘“‘ the birds which I sent home washed themselves neazly 
white in the water given them to drink.” In my memoir of 
1869, I mentioned Mr. Tegetmeier as having introduced the 
subject now being discussed to my notice. His own attention 
had been drawn to the matter by a correspondent of ‘ The Field, 
who sent him a washed-out feather. Mr. Tegetmeier forwarded 
this feather to me and asked me to ascertain if there were any 
possibility of its having been dyed. 

Such, in brief, was the position of the problem when in 1866 I 
began my researches. The chief results of these were published 
in the Phil. Trans. of 1869 and 1892, while im 1894 a Friday 
Evening Discourse in the Royal Institution presented a digest of 
the whole enquiry *. Although it would be unbecoming in me, 
a chemist, to enter into classificatory questions before a company 
of zoologists, I may venture to remind you that, according to 
many systematists, the Order Cuculiformes includes two suborders 
of equal rank, the Cuculide or Cuckoos and the Musophagide or 
Plantain-eaters. The former is an extensive suborder and 
cosmopolitan in range, while the latter contains less than 40 species 
and is confined to the Ethiopic region of Continental Africa. Of 


* <“Turacin, a new Animal Pigment containing Copper,’ ‘Student,’ i. (1868) 
pp- 161-168; with a coloured plate. 

“Researches on Turacin, an Animal Pigment containing Copper,” 
clix. (1869) pp. 627-636 (with 4 figs. of spectra). 

“Researches on Turacin, an Animal Pigment containing Copper,’ Part IL., 
Phil. Trans. clxxxii. A. (1892) pp. 511-530 (with 9 figs. of spectra). 

Proc. Roy. Inst. xiv. (1894) pp. 44-49. 


hil. Trans. 


SIR ARTHUR H. CHURCH ON TURACIN. 641 


the six genera of Musophagide three contain the crimson pigment 
which I named turacin. These are:—WMusophaga, 2; 7 Ur aCUS, 
21; Gallirex, 3; or 26 turacin-bearers in all. The ‘oe ee remain- 
ing genera from which turacin is absent are :—Corytheola, 1 ; 
Chizerhis, 4; and Gymnoschizorhis, 2; or 7 in all. There are 
two remarks that may be interposed in this connection. The first 
is that the crimson pigment is identical in all the species, and 
occurs not only in from 6 to 18 of the primary and secondary 
pinion-feathers, but also in all other similarly-coloured feathers 
or parts of feathers which are found on some of these birds—for 
example, in the head-feathers of Mwsophaga violacea, in the crest- 
feathers of Zuracus donaldsowi, 7. fischeri, and 7’. meriani, and 
in the head-feathers of Gallivex johnstoni behind the crest. The 
second remark refers to the pinion-feathers of three out of four 
species of Chizerhis, which have white patches bare of any pig- 
ment pretty much in the same position as the crimson patches in 
the turacin-bearers, although mostly confined to one side of each 
shaft—a curious coincidence, at all events. 

I may now turn to the properties of turacin. We havealready 
seen that it is soluble in water. Not, we may add, in hard water, 
but in clean rain water or, better, in distilled water. It is still 
more easily dissolved by weak alkaline liquids, extremely dilute 
ammonia being the best solvent. From this crimson solution, the 
colouring-matter, the turacin, is precipitated as gelatinous flocks 
on neuti: alization by hydrochloric or other acid. The flocks that 
separate when collected and dried form a deep red amorphous 
mass, crimson by transmitted light when in thin layers, but 
exhibiting a surface-lustre of a purplish hue, not unlike that 
reflected from erystals of potassium permanganate. Thus the 
appearance of solid turacin cannot be said to be accurately de- 
scribed when it is spoken of as ‘“‘a metallic red or blue powder,” ” as 
in the Enc. Brit. 11th ed. vol. x. p 226a@. Turacin is insoluble 
in alcohol, ether, chloroform, petroleum-spirit, benzol, and the 
usual solvents of resins and oils. In order to obtain it in a pure 
state special procedure is required to prevent its contamination 
with the natural oil of the feathers, while it is not possible to free 
it completely from all traces of non-essential mineral matter. 
The action of heat on dry turacin presents several points of 
interest. It suffers no change at a temperature rather above that 
of boiling-water, but at or near the boiling-point of mercury it is 
pr ofoundly modified without loss of w eight and becomes insoluble 
in alkaline liquids. If this altered ‘yu acin be now exposed to a 
high temperature in the presence of air its combustible con- 
stituents burn away, leaving a black ash, amounting in the purest 
samples to about 9°8 per cent. of the original weight. This ash 
consists almost entirely of oxide of copper, the amount corre- 
sponding to at least 7 per cent. of that metal in the turacin itself. 
Here again the statement in the Ene. Brit. (Joc. cit.) needs correc- 
tion, for this is a fixed percentage, not one that varies from “5 to 
8.” In further describing the action of heat upon dry turacin it 


642 SIR ARTHUR H. CHURCH ON TURACIN. 


must be mentioned that this colouring-matter, when suddenly and 
strongly heated, yields a volatile, red, copper-containing derivative, 
which, undissolved by weak ammonia-water, is not only soluble 
in, but may be crystallized from, ether. It is the vapour of this 
substance catching fire which gives rise to the green flame so con- 
spicuous when a particle of turacin is heated in theair. Hf I 
were discussing the nature and relationships of turacin from a 
chemical view-point, much would have to be brought forward as 
to its composition and probable formula. Here let it suffice to 
say that analysis gives these percentage-numbers :—— 


Warsbomireren.- Pre th 53°69 
JalyGleoyxeinl jcaanaeesAnaooncoc 4-60 
CIS} 0) SETS: 5 AAA RRA (Ou 
INTMUROPYEIM yo nqanossenecesG50 6:96 
Osxaime men iien a saceeeeeeeee 27°74 


These figures, though deduced from many careful determina- 
tions, do not lead unmistakably to one definite empirical formula. 
With a colloidal compound like turacin, which does not admit of 
purification by distillation or crystallization, there is always the 
chance of a disturbing factor being present in the shape of a trace 
of some impurity. Possibly such an expression as C,,H,,.N CuO,, 
is worth suggesting. In this the ratio of nitrogen to metal is as 
4:1, the same ratio which occurs in hematin from blood between 
nitrogen and iron, and between nitrogen and magnesium in sore 
chlorophyll constituents and derivatives. With all these bodies 
turacin is also brought into relation when its absorption-spectrum 
is considered, especially in regard to that broad band in the violet 
and ultra-violet (between the lines 4 and M) which is common to 
all these colouring-matters, and which was figured and described 
by the late Prof. Arthur Gamgee in a paper read before the Royal 
Society in 1896—a paper in which the author confirms my results, 
especially those relating to the discovery of turacoporphyrin. 

The spectrum of turacin may now engage our attention: in 
the feather itself the pigmented web shows two broad bands, one 
with its centre at wave-length at or near 585, the other with its 
centre near 538. <A solution in water containing a trace of 
ammonia shows the same two bands somewhat shifted towards 
the violet end of the spectrum; there is also seen a very faint 
band about the solar line F, but nearly as broad as band No. 2. 
When turacin which has been precipitated from its alkaline 
solution by strong hydrochloric acid is again dissolved in weak 
ammonia-water, and the solution spectroscopically examined, a 
narrow and faint fourth band situated on the less refrangible side 
of D makes its appearance. It is almost certain that this band is 
due to an alteration-product of the original turacin. The two 
well-marked bands, as they are shown when examining a feather 
with the spectroscope, closely resemble those of the oxy-hemo- 
globin of the blood; while the corresponding bands seen in the 


SIR ARTHUR H. CHURCH ON TURACIN. 643 


spectrum of an alkaline solution of turacin are not unlike those 
of CO-hemoglobin. However, the absorption-spectrum of turacin 
is sufficiently characteristic to enable one to use it as a criterion 
of the presence of this pigment in the red feathers of birds. So 
when Dr. C. F. W. Krukenberg announced the occurrence of 
turacin in a species not belonging to the Musophagide, and not 
even African, it became necessary to test the assertion. The bird 
in question is a Cuckoo from the Philippines, Dasylophus super- 
ciliosus. I obtained a skin, then an assurance of identity from 
the Bird Department of the British Museum, I removed the 
sparse red feathers from the head of this Cuckoo and submitted 
them to spectroscopic scrutiny. They did show an absorption- 
band, but it was situated midway between the two bands of 
turacin. Moreover, the colouring-matter, which is orange-red, not 
crimson, cannot be extracted by ammonia and contains no copper. 
Hitherto, therefore, turacin has not been found to occur outside 
the Musophagide. It is right to mention here that Dr. Kruken- 
berg’s statement as to its occurrence in a Dasylophus was made in 
consequence of a report addressed to him at his request by the 
authorities of the Senckenberg Museum in Frankfurt, and was 
not the result of an experiment made by himself*. It has 
unfortunately found its way into various articles and books, e. g. 
Ene. Brit. x. p. 226 a, where we read ‘‘ But turacin is not, as was 
supposed, confined to the feathers of the Plantain-eaters, since it 
has been obtained from a Cuckoo, Dasylophus superciliosus.” 1 
have been able to secure a threefold disproof of this statement, 
for although the occurrence of this pigment in any bird other than 
a Plantain-eater seemed unlikely, the closely-allied Cuckoos might 
have supplied an instance. 

The occurrence of so large a percentage of copper as 7 in 
isolated turacin needs a word of comment. It does not imply a 
large amount of this metal in the plumage of a single bird; a 
high estimate gives less than 14 of a grain of the metal. And it 
must be remembered that many recent analyses of vegetable 
and animal matters show the wide distribution of copper in both 
kingdoms. Two of the latest researches in this direction were 
made by Mr. J. W. Dougal7y in 1911 and by M. B. Guérithault 
in 1912. Im fact, there is no difficulty in accounting for the 
presence of copper in Turacoes and for its amount. 

I purposely exclude from the present paper any discussion of 
the curious green pigment named turacoverdin by Krukenberg ; 
of the relationships between hematoporphyrin and the turaco- 
porphyrin obtained from turacin by the action of acids; and of 
the supposed synthesis of turacin by treating hematoporphyrin 
with an ammoniacal copper solution. 


* ©. F. W. Krukenberg, Vergleichend-Physiologische Vortriige (1886), Bd. i. 


p. 152. f 
+ Pharm. J. 1911 (4) xxxii. pp. 405-7. 
+ Bull. Sci. Pharmacol. 1912, xviii. pp, 633-639. 


Qs 


Proc. Zoou. Soc.—1913, No. X LILI. 4 


644 DR. P. CHALMERS MITCHELL ON THE 


43. Observations on the Anatomy of the Shoe-bill (Baleni- 
ceps rex) and allied Birds. By P. CHatmers MircHett, 
M.A., D.Sc., LL.D., F.R.S., Secretary of the Society. 


[Received and Read June 3, 1913. ] 
(Plates LX XX.-LXXXIII.* and Text-figures 119-132.) 


INDEX. 

Anatomy : Page 
Pterylosis iene tdaks ccs nee teed. ce oes See noe een) 
Foo tiande@law gay. svat: ts :o. eieesekpeeeee: eee OLS 
Syrinx.. RAO CEE OA BeD se aNHOHUR oho bo ode aucicodobo eee ii GIL 
Carotid Arteries... Penne w saree Sele ee REPRE REE EE eR OD, 
pee Canal.. pea eeanan oAduaticeniadtienondatia yor 
Cloaca .. JL DUC ANDER ODES A Re ESSER At vonctde aug udonna cance ee (OO) 
Muscles" DE ee re jen nia SRE ROE ERE OOS 
Osteolooysiecs cas carci. coe ACSC GSS 

Syistematicucnaieskonn «suis Cie eae aes ele aS 


In the end of June, 1912, a young example of the Shoe-bill 
(Baleniceps rex) arrived at the Gardens as a gift from the 
Sirdar, Sir Reginald Wingate, G.C.V.O., Honorary Member of 
the Society. It was not quite adult, but stood between three 
and a half and four feet in height. It was placed in a warmed 
enclosure communicating with a grassy paddock containing 
a small pond, and it fed well, although its appetite was small, 
on fish, frogs and pieces of meat. It appeared to thrive through 
the winter, but in the end of March, 1913, it showed difficulty 
in breathing. It was removed to the Sanatorium and kept for 
a few days in a warm, moist atmosphere, but died. The post 
mortem examination showed the presence of mycosis in the 
lungs, although not in sufficient quantity to cause mechanical 
obstruction to respiration, and the interior of the larynx and 
bronchi was nearly free, but subsequent microscopic examination 
by Mr. Plimmer, F.R.8., the Society’s pathologist, revealed 
extensive infiltration of the tissues by the mould. As Dr. 
Beddard, the Society’s Prosector, was engaged on other work, I 
gladly took the opportunity of making some observations on the 
anatomy of this rare bird, and give the results here. My work 
was to a certain extent limited by the necessity of not injuring 
the skeleton, which was destined for the British Museum 
(Natural History). Certain portions of the viscera were sent to 
the Museum of the Royal College of Surgeons. 

T am indebted to my colleague Mr. D. Seth-Smith, the Society’s 
Curator of Birds, for leave to reproduce on Plate LX XX. the 
excellent photograph of this bird, which was an immature male. 


Pterylosis—A. D. Bartlett (1) discovered and described the 
powder-down patches, and Professor Giebel (21) has written a 


* For explanation of the Plates see p. 708. 


Bale & Danielsson, I*4 imp, D, Seth- Smith, photo: 


BALANICEPS REX. 


t ‘ 
1 
5 
2 
4 


By Zs Se US, Teil, ILOOR 


D. Seth- Smith, photo. Bale & Damelsson, Lt? amp, 


SKULLS OF: 1, BALA‘NICI 


[7] 
U 
W 
DO 
(ap) 
Q 
O 
ag) 
S 
Ww 
cs) 
FA 
be 

: 
: 
ep) 


ar 


seen tert 
buy 
vik Fetal 


at 


ia 


P, Zo So Is, Il, ILOoOX IU 


D. Seth- Smith, photo, Bale & Damelsson, Lh‘ imp. 


SKULLS! OF: 1, DISSURA. 2. TANTALUS. 3, CANCROMA: 


een 
i Hey 
, Sa aaie mn 

eign! 


er 


oe he 


PP 4. S&S, UNS, Wl ILecoui. 


D. Seth- Smith, photo. 


Bale & Danielsson, Lt? imp. 


SIMUILLG Ors 1, NRDIBA, 2, IPISWECANIOS., Si, IALOUUS- 


OO. 


ch 
ny 


ANATOMY OF THE SHOE-BILL. 645 


careful account of the external characters and pterylosis, but 
fortunately I finished my examination and notes of the pterylosis 
before having the opportunity of consulting Professor Giebel’s 
paper, and find that my observations and inferences follow his 
very closely. 

The down feathers are distributed practically uniformly over 
the pteryle and the apteria as in Steganopods and Storks, where- 
as in Scopus and Herons they are found only on the apteria. 
The oil-gland bas a tuft of feathers as in nearly all the members 
of the group. The contour feathers have a small aftershaft ; 
this structure is present in Herons and Scopus, variable and 
sometimes absent in Storks. 

The feathered tracts generally are in marked contrast with those 
of Herons and resemble those of Storks in being relatively wide 
and fading off into the apteria, those in Herons being remarkably 
narrow and very sharply defined at their lateral and posterior 
edges. The neck is continuously feathered for almost the whole 
of its length, and shows no trace of the long lateral apteria 
characteristic of Herons and present in Scopus. Far back, near 
the base of the neck, there is a lateral space at each side. ‘The 
spinal tract forks between the shoulders, leaving two parallel bands 
of feathers continuous with tne neck area, and separated by a 
wide median apterion. ‘These dorsal bands are wider than those 
of Herons and cease about the level of the axilla. Parallel with 
them at each side is a very broad and strong humeral tract, wider 
even than in Storks and much wider than in Herons. ‘The dorsal 
tracts reappear again as a narrow diamond-shaped band, weaker 
in the middle line, and stretching between the powder-down 
patches to the oil-gland. The powder-down patch at each side is 
an enormous, elongated oval area extending from a little way 
behind the axilla to the rectrices, and reaching some way down 
the side of the ramp. ‘The patches are separated in the middle 
line. The feathers are long, very thickly set, and break down 
into a yellowish powder. When the bird had been in spirit, they 
formed unpleasant, greasy masses. ‘Their discovery led Bartlett 
to associate Baleniceps with Herons as opposed to Storks. As, 
however, there are only two patches in Baleniceps as compared 
with four to six in Herons generally, eight in Cancroma, 
as they are absent in Scopus and exist in many other birds, 
e.g. Hurypyga, Rhinochetus, Mesites, some Tinamus, Birds of 
Prey, Parrots, and Goatsuckers, it is difficult to attach any 
systematic weight to their presence. Nitzsch’s observation that 
they may disappear with age (e. g. Gypaétus) still further destroys 
their systematic importance. 

The femoral tracts bordering the powder-down patches are 
rather badly defined. The arrangement on the ventral surface is 
almost identical with that given by Nitzsch for Ciconia. The 
median apterion is rather indefinite except at the base of the 
neck and near the vent. On each side of it is a very broad 


feathered tract. 
43* 


646 DR, P. CHALMERS MITCHELL ON THE 


The rectrices are 12 in number as compared with from 8 to 12 
in Herons, and 12 in Scopus and Storks. 

The primary quills of the wing are 11, not 10 as Giebel states. 
Of these six are on the metacarpal, three on phalanx 1 of digit 
JI and two on phalanx 2 of digit I, the last bemg much smaller 
than the others. There are 11 in Herons, 10 in Scopus and 11 or 
12 in Storks. There are 19 secondaries as compared with 11 to 
18 in Herons and 14 to 25 in Storks. 


Text-fig. 119. 


Wing-stracture of Baleniceps. 


Diagram of the distal secondary quills and coverts, showing the diastataxic condition. 
Right wing; external view. The quills are in outline, the major coverts are 
shaded ; the transverse rows are represented by dots, showing the insertion. 


8. First Secondary. C.R. Carpal remex. 
aw. Diastataxic gap. P. First Primary. 
C.C. Carpal covert. 


In the figure (text-fig. 119) I give a diagram of the distal 
secondaries, showing the arrangement known as “ aquintocubital ” 
until W. P. Pycraft and I showed simultaneously (28, 36) that it 
was not due to the loss of a secondary feather, after which my term 
“ diastataxic ” has been used. The major coverts are inserted 
proximally to the quills and cross over them, and this arrange- 
ment is repeated in the case of the degenerate carpal covert and 
carpal remex, whereas the major coverts of the primary quills 
are distally placed. ‘The diastataxic gap is very evident and the 
major covert, which occupies the gap, is tied down to the 
membrane supporting the quills by a special slip. In the same 
fashion the carpal remex has a membranous slip supporting it. 


ANATOMY OF THE SHOE-BILL. 647 


There is nothing peculiar in this arrangement; the diagram 
resembles closely the similar diagrams that I have given for 
other diastataxie birds (¢.g., 32, fig. 23). There is some difference 
of opinion as to whether the secondary quills, major and minor 
coverts, and the feathers nearer the outer border of the wing 
represent horizontal or transverse series. Most writers, following 
the obvious lead given by the quills and major coverts, have 
preferred to regard the rows as horizontal. I have taken the 
other view, seeing in the quills merely the enlarged members of 
the transverse rows which happen to lie along the margin of the 
wing and to have become the flight feathers, and the points of 
insertion of the feathers in the plucked wing have always 
appeared to me to lie in transverse or rather diagonal rows 
stretching upwards from the quills and reappearing on the under 
side of the wing in the feathers with reversed surfaces. These 
diagonal rows were plain in the wing of Aaleniceps, but I was 
surprised to find what I have not noticed in any other bird, 
although I am by no means prepared to say that it does not 
occur, that there was a transverse row too many. The row 
corresponding te the carpal covert and remex curved upwards and 
backwards ; the next most proximal row had a similar curve and 
belonged to the distal secondary quill; then there appeared to be 
an extra row in front of, and not behind the second secondary 
quill. More proximally the rows were in regular correspondence 
with the quills and gradually changed their inclination. I tried 
to correlate the arrangement with what W. P. Pycraft has called 
the “‘intercalary row” (a transverse row which in his opinion is 
associated with the mode of origin of the diastataxic gap by 
“faulting” of the horizontal rows in development), but was 
unable to make anything of it. No one appears to have pub- 
lished any observations on the theory of diastataxy since Pycraft 
and myself. In our communications to the Linnean Society (28, 
36) we showed that the condition was not due to the loss of a 
feather, Pycraft arguing from ontogeny and I from comparative 
anatomy. I showed that whereas most pigeons were diastataxic, 
a few were eutaxic and had arrived at this condition by a secon- 
dary closing of the diastataxic gap. I also showed that the 
eutaxic pigeons were in other respects more specialized than 
their diastataxic allies. In later papers (29, 31) I showed that 
similar conditions existed amongst Kingfishers and amongst 
Gruiform birds. The general inference seems to be clear: that 
the eutaxic groups are more specialized birds and that in 
association with their general specialization they have lost the 
primitive diastataxic arrangement. My argument, however, 
may be anatomically sound with regard to the groups presenting 
both conditions, and yet not applicable to birds generally. All 
birds may have been eutaxic originally ; certain families may 
have become diastataxic, and amongst these certain members 
may have secondarily reverted to the eutaxic condition. Pycraft 
assumes that the eutaxic condition was primitive, and if he be 


648 DR. P. CHALMERS MITCHELL ON THE 


correct, my facts would have to be interpreted according to the 
alternative I have just given. But I am not convinced by 
Pycraft’s deduction from his observations on the ontogenetic 
changes in the wing. The development of the individual is 
partly a process of latent differences becoming visible, and 
because a diastataxic bird appears to be eutaxic at a very early 
stage, it does not follow that eutaxy was primitive. J am much 
more impressed by the general view that passerine birds are 
plainly the most specialized of all birds, that they are eutaxic, 
and that the members of diastataxic groups which have become 
eutaxic are in other respects most passerine-like. In the absence 
of any convincing theory of the phylogenetic origin of diastataxy, 
all peculiarities in the arrangement of feathers are interesting 
and may come to have significance, and so I have digressed 
with regard to the wing of Baleniceps. 

It is plain that the pterylosis of Baleniceps is of the same 
general character as that in Storks, Herons and Scopus. To my 
eye, the general appearance and coloration suggest affinity with 
Herons rather than with Storks. The sedate habit of standing 
slently on any little eminence, the absence of the habit of 
clattering with the beak, which we noticed in the Gardens, and 
the reported heron-like bending of the neck in flight, confirm 
this view. But the actual details of the feathering do not 
confirm it; without any doubt, so far as pterylosis can be relied 
upon as indicating affinity, baleniceps is more Stork-like than 
Heron-like. 

Petherick, however, (34) has recorded that the young ran about 
with extended wings making a “yrattle-like noise produced by 
the snapping of their bills.” 

Foot and Claw.—The hind toe (hallux) is usually carried 
pointing backwards, but is freely movable in every direction. It 
is on the same level as the other toes, as in Herons and Scopus, 
not slightly elevated as in Storks. There is no trace of a web 
uniting any of the toes, whereas in Herons and Scopus there is 
usually a distinet web uniting the third and fourth toes, and in 
Storks all three front toes are united by web. 

A good deal of confusion, which I am able to dispel, has crept 
into the literature with regard to the condition of the claw of the 
third digit of the foot. Itis well known that the inner edge of 
this claw is pectinated in Herons and unbroken in Storks. 
Professor Reinhardt (37, p. 378) stated that Baleniceps had not 
a comb on the middle claw, adding that this absence afforded ‘a 
strong warning not to class it with the Boatbill, as this peculiar 
serrature never fails in any member of the Heron tribe.” Gadow 
(16, p. 137) divided the Arde into Family 1. Ardeide, dividing 
the latter into the sub-families Ardeine and Balenicipitine, 
mentioning as a character of the former ‘“‘ Mittelkralle gezihnt,” 
and of the latter “ Mittelkralle nicht gezihnt,” and Family 2, 
Scopide, including in the definition of the latter, ‘* Mittelkralle 
gezihnt.” Beddard (3, p. 289) cites Professor Reinhardt’s opinion 


ANATOMY OF THE SHOE-BILL. 649 


that ‘as the middle claw is not pectinated, Baleniceps cannot 
be referred to the Herons,” adding ‘‘ Professor Reinhardt would 
associate Baleniceps particularly with Scopus,” but omitting to 
mention that in the same paper Reinhardt stated that the claw 
was pectinated also in Scopus. Mr. A. H. Evans (8, p. 87) 
divides the Sub-order Ardez into the Families Ardeide (in 
which he places Baleniceps) and Scopide, and states that in the 
Sub-order the “claw of the middle digit is toothed on the inner 
side, save in Baleniceps.” It is surprising that as Gadow, 
Beddard, and Evans all seem to have thought the matter worth 
comment, they should have omitted to notice Giebel’s (21, p. 351) 
very definite description. Guiebel stated that in the two examples 
ot Baleniceps he had examined the pectination was clear and 
sharp, the actual teeth being not so small, numerous and deeply 
incised as in Cancroma and Nycticorax, but larger, separated by 
wider intervals, and, beginning at the point, reaching nearly the 
middle of the nail. He described them as closely corresponding 
with those of Scopus, adding that the latter had not received full 
attention from systematists. 

I hope that the drawings reproduced in the figure (text-fig. 
120), which were made by Mr. Berridge from the actual specimens, 
will explain the matter. Ina large number of birds belonging 
to different groups, the claw of the third toe of each foot is not 
symmetrical about the middle line; the ulnar or abaxial edge is 
relatively straight, and the radial edge, that nearest the body, is 
curved out into a sharp-edged scoop. The foot of the White Ibis 
(fig. 120, 1) shows this condition well. In Baleniceps (fig. 120, 2), 
owing to the lateral compression of the claws, the scooped edge is 
not quite so conspicuous in a dorsal or lateral view, but it exists. 
Moreover, as Giebel described, this sharp edge is marked by a few 
serrations between each of which there is rather a wide space, 
but which are so conspicuous on the claw that they can be felt 
not only along the edge but as slightly ribbing the surface. In 
Scopus (fig. 120, 3) the condition of the claw is almost exactly 
similar to that in Baleniceps. In the example I examined and 
from which the drawing was made the serrations were cut a 
little deeper than in Baleniceps, but according to Giebel, in the 
specimens of Baleniceps he examined, the serrations were deeper. 
The exact amount of serration is no doubt subject to individual 
variation. In the corresponding claw of the left foot of the 
Umbre I examined, each tooth was much narrower and more 
pointed, resembling the Laleniceps condition more closely. In 
the Little Bittern (fig. 120, 4) and in the Goatsucker (fig. 120, 5), 
the teeth are much more numerous, regular, smaller and more 
closely set together, and when the comb is highly developed, 
it may stand out conspicuously from the edge of the claw, 
sometimes by not being developed along the whole edge, and 
sometimes because the thin edges of the teeth give the comb a 
yellow semi-transparent coloration, readily visible against the 
duller and more opaque unbroken part. Almost every gradation 


650 DR. P. CHALMERS MITCHELL ON THE 


between the sharp unbroken edge and a fully formed comb can 
be found in nearly allied birds, and it must be a matter of 
individual taste at what particular stage of elaboration it is 
possible to describe the serration as a comb. It is at least certain 
that Baleniceps cannot be separated from Scopus and the Herons 
and associated with the Storks because of absence of pectination. 


Text-fig. 120. 


AWariasweeuneneee 


ARAN 


Modified Claw in some birds. 


Claw of the third toe of the right foot, in each case the left-hand figure showing the 
dorsal aspect, the right-hand figure the axial side. 


Red-billed White Ibis (Hudocimus longirostris). 

. Shoe-bill (Baleniceps rex). 

. Tufted Umbre (Scopus wnbretta). 

. South American Little Bittern (Butorides cyanurus). 
. Nacunda Goatsucker (Podargus nacunda). 


oo NO 


or 


It seems obvious, however, that the formation of a comb falls 
in the category of what I have termed “ multiradial apocentri- 
cities,’ modifications from the normal or primitive type, which 
as they have occurred repeatedly and independently, afford no 
information as to the systematic position of the animals in which, 
they occur. By the kindness of Mr. Seth-Smith I have looked 
through a number of the skins in the collection of the Society, 
and with the assistance of Mr. Ogilvie-Grant a still larger number 


ANATOMY OF THE SHOE-BILL. 651 


in the Bird Department of the British Museum. Most of the 
Steganopods have a scoop-like edge forming a primitive stage 
like that shewn in the figure (fig. 120, 1), but some of them, for 
instance the common Cormorant, havea well-formed comb. The 
small claws of Storks seem to have no trace even of the initial 
asymmetry, but Ibises are certainly more nearly allied to Storks 
than to Spoon-bills, and although the White Ibis (fig. 120, 1) has 
only the scoop-like edge, the “Glossy Ibis (Plegadis falcinellus) 
has a well-formed comb. Hur, ypyga and Rhinochetus have the 
curved edge but no serration; Cursorius, Dromas and Glareola 
have weil-marked combs. Among the Strigide, Scops and some 
of the smaller owls have the cur ved cutting-edge without serra- 
tion ; Aetwpa is in the same condition. Bubo hasa well developed 
cutting-edge with a few slight serrations ; Striw flammea has a 
well-marked comb occupying the upper part of the edge of the 
claw. By a curious accident of nomenclature I was led to 
examine some of the Birds-of-Paradise. Fiirbringer mentions 
Falcinellus as a genus in which the claw is pectinated, and as I 
had forgotten that that name had been used for a genus of Ibis, 
and knew that it was used for a Bird-of-Paradise, I examined the 
latter, and found that the claw in the Rifle-birds usually showed 
a sharp cutting-edge and that occasionally (e. g. Pédlor his and 
Epimachus) there were slight nicks in it. 

H. R. Davies (7, p. 368) in discussing the function of this organ 
remarks that the ‘ pectinated claw should not be regarded as a 
structure peculiar to nightjars, owls, herons, cormorants and 
gannets, and different from anything found in any other bird, 
but merely as a highly modified form of a structure found in a 
less modified form in many birds.” There seems to be no doubt 
but that the chief use of the modification of the claw is for 
scratching, possibly for removing parasites, and its presence 
may be compared in a general way with the condition in the 
mammalian Dassies, in which all the digits are protected by 
flat nails, except the inner digit on each hind foot which is 
provided with a sharp claw used in scratching. 

Rhamphotheca.—The horny covering of the beak is compound in 
Baleniceps, the premaxillary portion being separate, as in Scopus. 
In Storks and Herons it is simple. The edge of the horny lower 
jaw is delicately serrated. 

The Syrinw.—This has been studied and figured by Beddard (3), 
and I have to add to his description only that the first two 
incomplete bronchial rings are partly calcified, that the bronchi 
are relatively rather long, and that the distal bronchial rings 
are practically complete. On the most careful examination, I 
could find no trace of anything corresponding to what Beddard 
took to be fibrous vestiges of the intrinsic muscles present in 
Herons ; they were as completely absent as in Storks. I cannot 
follow Beddard, moreover, in his view that the structure of the 
syrinx is “‘conclusively in favour of regarding Balaniceps as a 
Heron and not as a Stork.” So far as the syrinx of Balceniceps 


652 DR. P. CHALMERS MITCHELL ON THE 


agrees in structure with that organ in Herons and in Scopus, it 
conforms not with a structure that is specially Ardeine, but with 
one that is found in so many different groups that Beddard 
himself has spoken of it as the typical avian syrinx. Storks, as 
he himself has shown, display a series of stages most conveniently 
regarded as degenerations in different degrees from the typical 
avian syrinx. Daleniceps shows degeneration, if not complete 
absence of the intrinsic muscles ; it has not degenerated so far as 
most of the Storks, but the fact is that as there is no typical 
ciconine and no typical ardeine syrinx, the condition in 
Baleniceps affords no clue to its relationship with either of these 
groups. 

Carotid Arteries —The normal condition, present in a large 
number of birds of different groups, and what development and 
comparative anatomy would seem to indicate as the primitive 
avian condition, is the presence of both right and left carotids, 
separate and well developed. This is the condition usual in 
Steganopods, Herons, Scopus and Storks. I was surprised there- 
fore to find that only the right carotid was present in Baleniceps, 
and although I sear ched carefully, IT could find no trace of even a 
degenerate ‘left artery. I do not attach systematic importance to 
the condition of the carotids, as this often varies within a Family. 
In Botauwrus, for instance, the two carotids fuse very close to 
their origin, and in another member of the Ardeide (Ardetta) 
the right carotid only is present. 


ALIMENTARY CANAL. 


The tongue is only a vestige as in Storks, Scopus and Cancroma, 
whereas it is long in Herons. 

Stomach.—The stomach is a capacious, elongated, rather thick- 
walled sac, extending posteriorly in the Ime of the cesophagus, 
from which it is to be distinguished externally only by a gradual 
increase of calibre. Distally it ends in a blunted angular point, 
and just proximal of this it gives off, on the right side, a globular 
chamber from which the duodenum arises (text-fig. 122, P, p. 657). 
Externally there is no trace of any specially tendinous area, and 
no constriction to mark off a proventriculus from a gizzard. The 
interior of the stomach, including the chamber at the pyloric 
end, is lined with a thin but well-marked layer consisting of the 
hardened secretion of the gastric glands. The cavity of the 
stomach is distinctly marked off from that of the csophagus by 
the corrugated edge of the membrane-like layer of secretion. The 
general cavity is sharply marked off from that of the pyloric 
chamber by a well-marked constriction (text-fig. 121). The 
greater part of the interior of the larger chamber is marked by 
longitudinal folds, but towards the posterior end these pass into 
irregular corrugations which are continued into the pyloric 
cavity. When the internal surface is scraped and the wall 
squeezed, the large glandular apertures may be seen to be 
distributed over the whole area of the stomach and pyloric cavity, 


ANATOMY OF THE SHOE-BILL. 653 


with a slight tendency to be arranged in longitudinal bands, but 
there is no indication of separation into a glandular proventri- 
culus and a muscular gizzard, and no trace of aggregation into 
specialized patches or areas. 


Text-fig. 121. 


Stomach of Baleniceps. 


The distal end of the stomach and the pyloric chamber have been laid open to show 
the constriction separating the general cavity from the cavity of the pyloric 
chamber and the minute aperture, A, into D, the duodenum. 


We have to recognize in the first place that the absence of 
distinction between proventriculus and gizzard gives no clue to 
the position of Baleniceps in the assemblage of Pelargo-Colym- 
biform birds. F. 8. Leuckart (23) discussed this formation in 
1841, citing the earlier authors, such as Blasius and Cuvier, who 
had called attention to it, described it in a number of birds and 
associated it with diet. It is tempting to associate such an 
undifferentiated condition with a primitive structure, but I do 
not think that such a view is tenable. At one time I myself 
thought that it might be possible to derive information useful for 
systematic purposes from the condition of the stomach, and I 
examined and made drawings of the organ in a large number of 
birds. But throughout the group, from Divers to Eagles, the 
extent to which gizzard may be separated from proventriculus by 
external or internal configuration, by specialization of muscle and 
tendon, or by aggregation of glandular areas, varies so irregularly 
as to suggest adaptation to habit rather than genetic tendency. 
The typical fish-eaters on the whole have a bag-like sac, weakly 
muscular and diffusely glandular ; those that live more on flesh 


654 DR. P. CHALMERS MITCHELL ON THE 


or on mixed diet have the muscular portion more specialized and 
a tendency to the concentration of the glands. Hven a formation 
so remarkable as the aggregation of the proventricular glands into 
two large circular masses has apparently little or no systematic 
significance. Within the group generally, the arrangement of 
the proventricular glands is diffuse over the whole area, but 
there is a tendency for them to lie in longitudinal bands, which 
miy be numerous, as for instance in the Sea-eagles and in 
Baleniceps, ov in two bands one anterior and one posterior. I 
found these bands rather short and rounded off in the American 
Grebe (“chmophorus major) and even more definitely rounded 
off in Garden’s Night-heron (Vycticorax gardeni). The condition 
I described and figured for the African Vantalus (Pseudotantalus 
ibis), where the glands are in a couple of rounded bosses, and 
which I noted as occurring also in Leptoptilus crumeniferus and 
L. argala, and in Carphibis spinicollis (25), is obviously a simple 
derivative from the pair of rounded bands. The state of affairs 
noted by Garrod in Levaillant’s Darter (20) and by Forbes in the 
Indian Darter (11) differs from that in the Storks only by 
the circular form of the two patches being a little more advanced 
and by a slight tendency for the circular masses to retreat into 
eversions of the stomach wall, a condition which is completed in 
Plotus anhinga (19) by the two patches having retreated into 
a rounded diverticulum. It is clear that the absence of such 
circular patches in Laleniceps tells us nothing as to the place of 
that bird in the system. 

When I found that there was a well-marked pyloric chamber, 
a fact which appears not to have been recorded before, I at once 
remembered the existence of such a chamber in the Pelican from 
my own notes, and in Plots from the observations of Garrod and 
Forbes. But as Leuckart (loc. cit.), Gadow (14), Cazin (6) and 
many others have shown, a pyloric chamber in varying degrees 
of completeness of separation occurs in many birds, notably in 
Herons, Storks and Darters; and even if we try to follow Cazin 
in limiting the term to cases where the constriction from the 
larger chamber is very well-marked, its presence gives no sure 
eround for associating Baleniceps more closely with any one of 
the groups of allied birds. 

Liver.—As Beddard (3) has noted, the right lobe of the liver is 
very much larger than the left,a condition which he was inclined 
to think showed affinity with the Herons rather than with the 
Storks. In a later work (4), however, he states that the ‘“ relative 
sizes of the liver lobes appear to be of no importance syste- 
matically ”—an opinion with which I concur. There is a large 
gall-bladder, and the cystic and hepatic ducts open nearly 
together, but the cystic duct distad of the hepatic duct, just 
beyond the end of the straight distal limb of the duodenal loop 
of the intestines, the disposition being very like that figured by 
Beddard in the case of the Indian Darter (4, p. 32) except that 
the hepatic duct passes through the substance of the lobulated 


ANATOMY OF THE SHOE-BILL. 655 


pancreas. On referring to my own notes, I find that a closely 
similar arrangement occurs in a very large number of birds 
belonging to widely separated groups. 

Intestinal Tract.—The most notable general feature of the 
alimentary canal is the extremely small bulk it occupied in the 
body compared with the size of the bird. The actual specimen 
stood nearly four feet high; when the alimentary tract and 
mesentery had been removed by cutting the duodenum close to 
the stomach and the rectum close to the cloaca, the little handful 
of viscera placed on the dissecting board was not so large as the 
similar mass from a duck. The calibre of the whole tract was 
narrow and fairly uniform, except that the cecum and large 
intestine were rather wider. The aperture leading from the 
stomach (7. e., from the pyloric chamber of the stomach) to the 
duodenum was excessively small (text-fig. 121, A, p. 653); 
grain of millet would have had difficulty in passing through. ‘The 
minuteness of this aperture is no doubt an adaptation similar in 
purpose to the hair-like brush found by Garrod in Plotus anhinga 
and the similarly placed plug found by the same anatomist in 
Levaillant’s Darter (Garrod, 19 and 20), which he surmised to be 
devices for preventing the passage of fish-bones into the gut. 

The minuteness of the exit from the stomach and still more 
the further guarding of the aperture by a plug of hair-like 
structures may have another advantage than prevention of the 
passage of fish-bones. The hair-plug occurs also in the Turkey- 
buzzard (Cathartes aura), which is certainly not a habitual fish- 
eater. In his “Last Journal” (12) under the date Aug. 20, 
Forbes mentions dissecting an example of Plotus levaillanti and 
finding the stomach full of nematodes, none of which, however, 
had penetrated beyond the plug, although several had been caught 
init. I found a number of nematodes in the stomach of my 
specimen of Baleniceps. We know now that intestinal parasites 
may do much damage to the animals they infest, and it is 
possible that the plug of hairs in the Darters and the very small 
exit from the stomach in Baleniceps, serve the useful purpose of 
preventing nematodes, which have been eaten with the food, 
from entering the intestines, keeping them in the stomach where 
they may eventually be killed. 

The aperture by which the distal end of the small intestine 
communicates with the cecum and large intestine (text-fig. 123, B, 
p. 658) is only just a little larger than the opening into the duo- 
denum. The structure of the alimentary tract shows that the 
Shoe-bill is adapted to make the most of a limited diet, consist- 
ing probably entirely of animal food, fish, frogs, or even small 
mammals; that the food must be retained for a considerable time in 
the stomach until it is very well macerated, for large lumps could 
neither enter the intestines from the stomach, nor even if they 
reached the intestines leave them by passing into the cecum and 
large intestines. From much unpleasant experience in dissecting 
she alimentary tract of birds and mammals, I have learned to 


656 DR. P. CHALMERS MITCHELL ON THE 


note differences in the odour on the dissecting table. In the case 
especially of vegetarian and omnivorous creatures where the 
digestive tract is bulky, and large quantities of food are taken, 
there is usually a very offensive odour, showing that putrefaction 
attends the processes of intestinal digestion. In other cases, and 
amongst birds, notably in birds-of-prey and many fish-eaters like 
the Divers, the odour of the alimentary tract behind the stomach 
is rarely offensive, and, sometimes, even attractive and aromatic. 
Although I was unable to examine this specimen of the Shoe- 
bill until it had been dead for several days, the contents of the 
intestinal tract were not offensive. 

In the figure (text-fig. 122) I represent the course of the ali- 
mentary canal, from the stomach to the cloaca, dissected out in a 
fashion which, as I have described in former memoirs (26, 30), 
seems to me to give much information as to its morphology and to 
afford a useful basis of comparison with the conditions existing in. 
the different groups of birds. The secondary foldings and modes 
in which the gut is packed in the body-cavity are naturally not 
shown by this method, but the relation of the gut to the primitive 
mesentery, the portions of it which have been expanded into 
loops, and the configuration of these loops, appear with diagram- 
matic clearness. The first specialized loop is the duodenum ; it 
is relatively not quite so long asin Herons generally, but it shows 
on its distal limb a minor expansion, represented in the drawing 
as two short folds, and comparable with the condition which I 
have figured in Wycticorax and Ardea. Then follows a second 
definite loop with a minor loop on its proximal limb, then a well 
marked loop, and then a few irregular twists, after which comes 
the portion bearing the remnant of Meckel’s diverticulum (text- 
fig. 122, m.), the vestige of the yolk-sac, lying in the line of axis of 
the main branch of the portal vein. In Baleniceps this was very 
slender, and bound closely to the inferior edge of the gut by a 
ventral mesentery. It might quite easily have been overlooked, if 
it had been sought for in the usual fashion, merely by running 
the gut through the fingers, but when the tract was laid out in 
the way I recommend, so that the blood-vessels were visible and 
the mesentery undisturbed, it was at once obvious. The part of 
Meckel’s tract between this diverticulum and the usual position 
of the ceca is thrown first into a series of short ill-defined loops 
and then into a long and definite supra-duodenal loop (text- 
fig. 122, S.D.F.) closely attached to the duodenum in the un- 
disturbed condition and supplied from the duodenal vein by what 
I have termed a “bridging” or short-circuiting vessel, which 
traverses the mesentery and must be cut through in process of 
laying out the gut (text-fig. 122 x, x). Then follows a rather 
irregular piece of gut forming two of the folds which I have 
described as ‘‘ supra-ceecal” kinks, the presence of at least one 
of these being characteristic of the great assemblage of birds 
containing the eagles and vultures, herons and storks, penguins 
and petrels. 


ANATOMY OF THE SHOE-BILL. 657 
Just behind the distal extremity of these kinks, the mesentery 


narrows so as to bring the distal end of Meckel’s tract very 
close to the proximal end of the duodenum. This almost circular 


Text-fig. 122. 


Intestinal Tract of Baleniceps. 


S. Stomach. 
P. Pyloric chamber. 
D. Duodenal loop. 
1, 2, 3. The three loops of Meckel’s tract anterior to the diverticulum, 


typical of the Pelargo-Colymbomorphine Brigade. 
m. Meckel’s Diverticulum. 
S.D.F. Supra-duodenal loop. 
x,«. Cut ends of bridging vessels from duodenal loop to supra- 
duodenal loop. 
K1,K2. Supra-cexcal kinks, 
Ca. Caecum. 


expanse of mesentery grows from the simple primitive loop of 
which the main branch of the portal vein is the axis and which 


658 DR. P. CHALMERS MITCHELL ON THE 


earries the yolk-sac at its extreme point. If, as happens in many 
birds belonging to widely separated groups, Meckel’s tract is 
twisted in the course of growth, the twist in the mesentery is 
seen here with the result that in the dissection of the gut as 
shown in my figures, the mesentery carrying the posterior region 
of the gut may be tucked or folded under the mesentery carrying 
Meckel’s tract, so that although the two are morphologically 
continuous, the continuity may ‘not be apparent except where the 
whole gut is short or very simple. In my diagrams, I have 
simplified this region, showing the morphological continuity, as 
the secondary twisting was not a part of my argument. 

The ceca (or caecum) are to be looked for at this point, which 

marks the transition from Meckel’s tract to the hind gut at the 


Text-fig. 123. 


SI. 


Cxcum of Baleniceps. 


S. I. Cut small intestine. 
L. 1. Cut large intestine. 
B. The lateral wall of the cecum has been cut away to show at B, the 
aperture into the small intestine, lying distally of the point 
where the intestine appears to pass into the cecum. 


beginning of the area drained by the posterior branch of the 
portal vein. In Baleniceps only one is present, as shown in the 
general diagram (text-fig. 122, Ca.). The end of Meckel’s tract 
meets the large intestine almost at a right angle, and the cecum 
is in continuation of the line of the large intestine. The calibre 

of the large intestine is rather greater than that of Meckel's tract, 
the difference being greater than is represented in the general 
diagram and rather less than in the enlarged figure (text- 
fig. 123), which was drawn from the specimen after it had been 
washed out and slightly stretched in the process of opening. ‘The 


ANATOMY OF THE SHOE-BILL. 659 


aperture from the small intestine to the large intestine (text- 
fig. 123, B) is relatively extremely small and is considerably more 
posterior (nearer the cloaca) than the region where the one portion 
of the gut joins the other, the actual passage running in the 
conjoined walls for a certain distance. There is not more than 
the very slightest fold or bulge on the wall of the large intestine, 
which may be imagined rather than definitely stated to be a relic 
of an originally paired condition of the ceca. 

The presence of a single cecum in Baleiiceps has already been 
noted by Forbes (10) from a prepared specimen of that region 
of the gut mounted in the Museum of the Royal College of 
Surgeons, and this specimen corresponds in every particular 
with the example which I have dissected and figure in this 
communication to the Society. Beddard, however, (38) wrote as 
follows :—‘‘In the intestines I could not discover any trace of 
exca at all; I believe that the single cecum which characterizes 
the Ardeide (there are two in the Ciconiz) may be extremely 
minute, and might therefore easily escape recognition in the 
spirit-preserved alimentary tract.” I have probably examined 
carefully at least as many ceca of birds as any other anatomist, 
and Dr. Beddard’s supposition seemed to me extremely improbable 
on general grounds. When I found that the specimen in the 
College of Surgeons’ Museum was extremely like my own dis- 
section, it seemed still more improbable that a structure so 
definite and peculiar could be present or absent in different 
individuals. Dr. Beddard examined viscera which had been 
preserved in spirit and which had been previously handled by 
some other investigator. On consulting with him, he was able 
to add to the information given in his memoir, that the late 
Professor Stewart was rather unwilling that so rare a specimen 
should be cut about too much, and he agreed with me that it was 
quite possible that the portion of the gut to which the cecum is 
attached had: been removed before he examined it. Thanks to 
the kindness of Mr. R. H. Burne I have now had the opportunity 
of comparing the gut from the example I dissected with the 
actual material examined by Dr. Beddard in 1888. Dr. Beddard’s 
material was in three pieces and the greater part of the mesentery 
had been cut away, but enough of the latter had been left to 
enable me to identify with complete certainty the general dis- 
position of the gut, to recognize the duodenal loop followed by 
the subsidiary loops into which the hepatie ducts open, the large 
loop with its proximal minor loop, exactly as in the diagram from 
my specimen, the short loop with the remnant of Meckel’s 
diverticulum (which Dr. Beddard, apparently, had not noticed) in 
precisely the same relative position on the loop and pointing 
forwards, the short twists preceding the supra-duodenal loop, and 
the latter loop. It was evident, moreover, that the remainder of 
the intestinal tract had been cut away, and that Dr. Beddard had 
failed to find the cecum because he had not quite the whole of 
the small intestine before him, and no part of the large intestine. 


Proc. Zoon, Soc.—i913, No. XLIV. 44 


660 DR. P. CHALMERS MITCHELL ON THE 

Examination of the preparation of the cecum in the College of 
Surgeons’ Museum, which Forbes had seen, as cited by Beddard 
in a footnote, made it most probable that that was the portion of 
the intestinal tract removed from the specimen Beddard examined. 
There is therefore no evidence in favour of Dr. Beddard’s 
suggestion that the cecum in Saleniceps may be absent, or so 
small as to be unnoticed, 

The large intestine from the cecum to the cloaca is relatively 
long in Baieniceps and is rather wider in calibre than the small 
intestine; it is thrown into a series of short irregular loops, 
threaded, so to say, on a mesentery which is much shorter than 
the course of the gut itself, and which in the usual way is drained 
by a large branch of mesenteric vein. 

I have shown on a former occasion (30) that the characters 
of the intestinal tract are capable of affording a large amount 
of information as to the inter-relationships of the groups of 
birds. The primitive gut may be regarded as a tube not much 
longer than the length of the body-cavity it traverses and 
suspended from the dorsal body-wall by an antero-posterior 
mesentery. Jt is fixed at its anterior end, where it joins the 
stomach, at its posterior end, where it enters the cloaca, and near 
the middle of its length, on its ventral surface, where it is con- 
tinuous with the yolk-sac. The great embryonic vein forms the 
chief radius of this crescentic loop, running up to the dorsal wall 
from the yolk-sac, and receiving a large tributary, which runs 
parallel with the dorsal wall, from the hind end of the body. In 
the course of growth the gut becomes much longer than the 
distance separating its anterior and posterior fixed parts, and the 
lengthening takes place by the outgrowth of subsidiary loops from 
the primitive gut. The position of these loops with regard to the 
fixed points of the gut, and their number and character, differ in 
different cases, with the result that when the alimentary tracts 
are laid out on the dissecting board in the fashion in which I have 
described, they form definite patterns. In their main features, 
these are constant in individuals of the same species, closely 
similar in the species of a genus, and show definite relationships 
in the families and greater groups. A type of pattern persists 
through the large divisions and shows a gradual increase in 
definiteness and specialization in the different members of these 
groups, with the result that the patterns can be arranged in 
family trees. These correspond so closely with infor mation that 
can be derived from other anatomical characters, that I regard 
them as being an extremely useful guide to the relationships of 
birds. Obviously patterns are more easy to place in the system 
when they are highly specialized and complex, but even the 
shortened guts of fruit-eaters may retain marked indications of 
pattern. 

In the great assemblage of birds which Gadow (16) has 
called the Pelargomorphine Legion, in which Baleniceps must 
certainly be placed, the pattern ‘of the gut is relatively complex. 


ANATOMY OF THE SHOE-BILL 66. 


The duodenum is a definite loop, tending to be rather wide 
and further sub-divided in the Falconiformes, to be excessively 
long, twisted and rolled up with the first loop of Meckel’s tract 
in the Ciconiiformes, to be very long, narrow and with a minor 
loop on its proximal limb in the Ar ‘deiformes. In Baleniceps, i 
is not highly specialized, but the differentiation it displays is 
more like that in Herons than in Storks, especially in the forma- 
tion of the secondary expansions at the base of its distal limb. 

In the same assemblage Meckel’s tract 1s long, and is suspended 
round the circumference of an almost circular expanse of mesen- 
tery which grows out from a very short portion of the primitive 
dorsal mesentery It is roughly symmetrical about the main axis 
which runs out to the remnant of Meckel’s diverticulum, the 
latter being invariably present. On the proximal half of the 
tract there is a tendency to the formation of three minor loops 
between the duodenum and Meckel’s diverticulum. These tend 
to remain comparatively simple and similar in the Steganopods 
and in the Faleoniformes. In the Ciconiiformes the tendency is for 
the first to become very long and secondarily twisted up with the 
duodenal loop, the second and third becoming nearly obliterated. 
In the Ardeiformes the first and second are generally rather long 
and definite and may form minor loops, the third is often reduced 
to a little bunch of small loops closely set together. In Baleeniceps 
the three loops are rather more distinct than in either the Storks 
or the Herons, remaining in the more generalized Steganopod con- 
dition, but such differentiation as exists approaches the Ardeine 
pattern and shows no trace of the Ciconine peculiarities. 

In the assemblage Meckel’s diverticulum is actually or very 
nearly at the extremity of the axial loop of the tract. When it 
is not quite terminal, it 1s always on the proximal side of the 
loop and is then bent parallel with it, the free tip pointing 
proximally. The axial loop may grow out to a great relative 
length. ‘These dispositions are distributed very irregularly 
throughout the whole assemblage which I am discussing. The 
axial loop, for instance, is long and has the diverticuium at the tip 
in some Storks and in many of the smaller Hagles and Falcons. 
It is short, but usually has the diverticulum at the tip in some 
of the Herons and in Scopus, Cathartes and Polyborus. It is 
short and bears the diverticulum proximal to its apex in Phaethon, 
Pelecanus, Fregata, Pseudotantalus, Serpentarius and Neophron. 
The latter condition exists in Baleniceps, and it is plain that its 
occurrence has no systematic significance. 


(=) 

In the Pelargomorphine Legion the distal part of Meckel’s tract 
is less highly differentiated, and therefore affords fewer dis- 
criminating characters. The ‘portion of it that follows Meckel’s 
diverticulum is usually thrown into a bunch of short, irregular 
loops, and then follows a well-marked supra-duodenal loop, 
supplied from the mesenteric vein, and one or more definite 
supra-cecal kinks. In Baleniceps, there is first the irregular 
region, then a definite short loop, then a long typical supra- 


44% 


662 DR. P. CHALMERS MITCHELL ON THE 


duodenal loop, and then two supra-cecal kinks. This region 
supphes no definite information which might help to place 
Baleniceps inside the Legion. 

In the Legion the colie ceca are much reduced and apparently 
practically functionless except in the Anseriformes, in most of 
which they are very large and functional. Although reduced in 
the Steganopods, they are rather less so than in the Herons 
and Storks and occasionally contain fecal matter. I think the 
presence in Baleniceps of a single cecum, by no means so large 
as either of those in the Anseriformes, but definitely functional, 
communicating with the hind-gut and containing fecal matter, 
may be taken to be established. The presence of one cecum, 
instead of the normal pair, associates Baleniceps with the Herons. 
I should be disposed to guess that the loss of one cecum of the 
pair had taken place whilst both were functional, as there seems 
no particular reason why one of two vestigial organs should be 
suppressed, except as an occasional abnormality, and that the 
condition in the Herons, where there is a single functionless 
cecum, 1s secondary to that in Baleniceps. 

The characters of the large intestine in Birds generally are not 
sufficiently differentiated to afford much information of systematic 
value. There seems to have been a general tendency to the 
reduction of this area to an extremely short and straight course 
from the ceca to the cloaca, a tendency which has been inde- 
pendently followed by the higher members of a large number of 
groups. Baleniceps has a relatively long and capacious large 
intestine, and in so far has remained in a rather more primitive 
condition than most of the members of the Pelargomorphine 
Legion. 

To sum up, the characters of the intestinal tract of Baleniceps 
are those of the Pelargomorphine Legion, and such specialization 
as it displays associates it with Ardeine birds rather than with 
Ciconine birds. 

In a communication to this Society, Dr. Beddard (5) has 
made some additions to or corrections of my observations, 
particularly with regard to the presence of a specialized supra- 
duodenal loop in birds in which I did not record it, which are the 
more valuable as my work stretched over a number of years, as 
material was available, and it was only in its course that I began 
to recognize the significance of the various points and what had 
specially to be looked for. Dr. Beddard also on several grounds 
throws doubt on the value of my mode of displaying and com- 
paring the intestinal tract patterns. These grounds are due to 
misapprehension. He thinks that my method of figuring the 
tract gives ‘an appearance of simplicity that is misleading, with 
the result that birds which are separated by marked characters 
are represented as being almost identical.” Certainly the patterns 
(even if correct) do not in every case afford enough information 
to place clearly, or to separate clearly cases where the patterns 
are very simple. I was rather careful to insist on this point in 


ANATOMY OF THE SHOE-BILL. 663 


my memoir. Healso thinks that I do not distinguish sufficiently 
between what he terms “ fixed loops definitely formed by a narrow 
mesentery, and the irregular folds into which any mobile coil of 
the intestine may fall when disposed on the dissecting board.” 
The whole tract 1s of course supported by mesentery, and 
Dr. Beddard’s “fixed loops” and irregular folds are merely the 
beginning and final result of differentiation. The loops to which 
I have called attention are fixed by their morphological position, 
and they may be wide or narrow, long or short. More serious, 
however, is Dr. Beddard’s s misapprehension of the morphology of 
the gut which leads him into very curious comparisons. He 
states “that there are no essential differences between the 
intestinal tract in Birds and Crocodiles.” He accepts as “‘ per- 
fectly correct” a diagram I gave (26, p. 137) of the alimentary 
tract in the Alligator, in which the canal is displayed as showing 
a series of almost exactly similar loops from the stomach to 
the cloaca, suspended on a crescentic fold of mesentery. As 
the pancreas les in the first of these loops, the latter may by 
analogy be called the duodenum, but it is simply the first of a set 
of regular loops. Meckel’s tract and the delimitation between 
that and the large intestine are not shown ; it is quite clear that 
Meckel’s tract is not differentiated. Comparison with the next 
figure, that of the tract in an embryonic pheasant, shows the 
essential difference. Immediately posterior to the duodenum a 
mesenteric area, corresponding to a very short length of the whole 
distance from the stomach to the cloaca, grows out into an 
enormous nearly circular tract, of which the great vein from the 
yolk-sac forms nearly a diameter. This region is Meckel’s tract, 
and from the point where it returns to the dorsal line again and 
where the. ceca, if present, are given off, the large intestine 
begins, and corresponds to a much larger part of the primitive 
distance from the pyloric extremity of the stomach to the cloaca, 
than the combined length of origin of the duodenum and Meckel’s 
tract. This mode of development of the gut dominates its adult 
morphology. So also Dr. Beddard does not appreciate the mor- 
phological importance of the position of Meckel’s diverticulum, 
the remnant of the yolk-sac. Fortunately it persists throughout 
life in most of the different groups, and its presence rules out 
such comparisons as Dr. Beddard makes between particular 
loops in Rhea and a Tinamu (with a Passerine intervening in 
the argument !). He is trying to identify different morphological 
material, belonging to different somites of the embryo, and this 
error makes his conclusions invalid. It would be of great interest 
to examine young chicks of those birds in which the rudiment of 
the yolk-sac does not usually persist, and this would clear up some 
of my dubious cases. But so far as they go, the gut-patterus afford 
an amazingly conclusive body of evidence as to “the Avian system. 

Croaca.—The rectal portion of the large intestine expands 
suddenly to enter the large cloaca. The first chamber of the 
cloaca, called the coprodeeum by Gadow, is separated by a thin 


664 DR. P. CHALMERS MITCHELL ON THE 


transverse fold (text-fig. 124, A) from the second, much narrower 
urodeum. The paired ureters open into the latter (text-fig. 124, U) 
on the dorsal surface but rather laterally placed. The actual 
apertures are situated on a small ridge and were very dificult to 
see, considering the size of the bird. I found them eventually by 
passing a horse-hair backwards through the ureter, and I think 


Text-fig. 124. 


Cloaca of Baleniceps. Ventral view. 


The sphincter has been cut through im the middle ventral line and folded outwards ; 
portions of the dorsal wall of the gut have been removed. 


LI. Cut end of large intestine. 
A. Fold separating Coprodzeum from Urodeum. 
B. Fold separating Urodeum from Proctodeum. 
U. Aperture of Ureter. 
G. Genital aperture. 
C. Glandular crypts. 
D. Cut sphincter muscle. 
F. Aperture of Bursa Fabricii; «, Dorsal wall of Bursa. 


it possible, although I am not certain, that there were several 
very small apertures at each side instead of one large one. 
Rather lateral and slightly posterior to the ureters were the 
minute openings of the vasa deferentia (text-fig. 124, C). There 
was no genital papilla, but the specimen was a very immature 
male, and the testes were minute. The vas deferens accompanied 


ANATOMY OF THE SHOE-BILL. 665 


the ureter until the latter nearly had reached the wall of the 
cloaca, and then twisted outwards. A transverse fold also sepa- 
rated the urodeum from the proctodeeum (text-fig. 124,B). In 
the proctodeum, at each side and just at the edge of the sphincter, 
were four or five little glandular apertures leading into small 
cavities lined with ir regular ridges. I find in my notes of dissec- 
tions of Ostriches, both male and female, that similar glandular 
crypts are present in that bird. In the middle line of the procto- 
deum, just behind the fold separating that chamber from the 
urodeum, lies the large, elongately oval aperture of the Bursa 
Fabricii (text- -fig. 124, “B. The bursa is a very large chamber, 
lying above the cloaca, running forwards almost to the rectum. 
The inner wall is lined by irregular, heavy ridges, making it 
resemble the reticulum of a ruminant stomach. In the figure, 
part of the dorsal wall of the coprodeeum and urodeum is repre- 
sented as cut away to show the cavity. The bursa was empty. 
There was no trace of a penis. 

Our knowledge of the Bursa Fabricii is due chiefly to Forbes, 
later writers having added very little to his observations and 
conclusions (9). In Struthious birds, especially when they are 
young, there is practically no constriction separating the procto- 
deum and the bursa, the latter being simply a forwardly 
directed and dorsally placed continuation of the cavity of the 
posterior division of the cloaca. In the different groups of birds 
there appears to be a general tendency for a convergent modifica- 
tion of this simple arrangement ; the constriction between procto- 
deeum and cloaca becomes more and more pronounced, until the 
bursa becomes a tubular or pyriform sac opening by a very small 
pore into the dorsal wall of the cloaca. This progressive change 
is most marked in Passerines and in those birds in other groups 
which most nearly mimic the passerine type,and may lead to the 
complete disappearance of the aperture and of the bursa. There 
is of course no reasonable doubt but that the Passerines present 
the most specialized results of avian evolution. ‘To a certain 
extent, ontogenetic changes in the bursa show a similar course of 
change, the aperture of the bursa narrowing, and the bursa ttself 
tending to contract and even to disappear with age. There is 
probably, therefore, no special significance in the condition of the 
bursa in the example of Baleniceps I dissected, its large size and 
wide aperture being perhaps due to youth. Forbes, however, 
states that in the Storks and Herons he examined, the bursa was 
large and its aperture small. He also mentions the absence in 
these birds of the reticulum of ridges in the lining wall of the 
bursa, although he found them in Steganopods much as I describe 
them in Baleniceps. I cannot draw any systematic conclusions 
from these facts. 

A small penis is stated to be present in Storks, absent in 
Herons, so that in the absence of that organ balewniceps resembles 
the latter group, but I attach no systematic value to this. 


666 DR. P. CHALMERS MITCHELL ON THE 


Muscutar ANATOMY. 
Muscurs oF Heap. 


Dermo-temporalis.—This thin but extensive sheet of muscle 
arose at each side by a fleshy, narrow head, behind and above the 
temporal fossa and close to the origin of the biventer maxille, 
and spread out on the skin of the ventral surface and sides of 
the neck, precisely as in the Pelican. 

Biventer maxille was very large and strong, arising from a well- 
marked area on the back of the head behind the quadrate and 
inserted to the posterior end of the lower jaw. As in the 
Pelican, I could not separate this from the underlying digastric 
or depressor mandibule. 

Temporal.—External portion. A very strong mass, arising 
from the dorsal part of the temporal fossa back to the biventer, 
runs downwards and forwards, dipping under the ramus of the 
maxilla to be inserted to the outer and upper surface of the lower 
jaw opposite the orbit. 

Pyramidal portion. Strong fan-shaped muscle arising trans- 
versely under the post-orbital process and converging toa rounded 
tendon which runs forwards and outwards to the inner side of the 
lower jaw. Parallel with this, and possibly a separate portion of 
it, is a flat band of muscle running from close to the articulation 
of the quadrate, alongside the tendon of the pyramidal portion to 
be inserted just anterior to it. 

Quadrato-mandibular portion. Very strong fleshy muscle 
running transversely from the whole of the anterior surface and 
forward process of the quadrate to the ramus. 

Quadrato-orbital portion. Long muscle from just behind the 
optic foramen across to the whole inner edge of the orbital 
process of the quadrate. 

Pterygoid——An enormous mass of muscle, partly separable 
into layers, on the lower surface of the jaw, from the posterior 
angle and ventral posterior portion of the mandible running 
forwards to the pterygoids. 

The temporal and pterygoid muscles are practically identical 
with what I have found in the Pelican. 

Biventer cervicis.—I examined this muscle to see if there were 
any trace of the peculiar formation described by Garrod in the 
case of the Darters, but found that the muscle with its anterior: 
and posterior bellies was quite normal. 


Hyor Muvsctes. 


Mylohyoid anterior —As in the Pelican a very slender and thin 
sheet of muscular fibres superficial to the other muscles of the 
under surface of the jaws, and running transversely across from 
the ramus of the jaw to spread out on the interspace between the 
two rami, but without meeting its fellow in a median raphe. 

Mylohyoid posterior.-—Arises as a broad strap from the outer 
surface of the angle of the jaw just below the ear and divides into 


ANATOMY OF THE SHOE-BILL. 667 


a thin wide sheet of fibres which runs over the ventral surface, 
meeting its fellow of the other side and forming a platysma 
myoides, and a better defined band which runs across to be in- 
serted to the ceratohyal, superficial to both divisions of the 
geniohyoid. 

Geniohyoid.—Two well-marked divisions. A very strong 
division arises from the last joint and cartilaginous end of the 
ceratohyal, round which it is wrapped, and runs forwards to the 
outer surface of the lower jaw just under the orbit. The second 
division arises from the first joint of the ceratohyal and runs 
straight forwards as a sheet of fibres which meets the corres- 
ponding fibres of the same division of the other side. These 
fibres can be traced up to the junction of the rami of the 
mandible. 

Genioglossus.—Probably in relation with the degeneracy of the 
tongue, this muscle was absent. 

Ceratoglossus.—Fleshy from the outer side of the first joint of 
the ceratohyal to the tip of the tongue. There was no separate 
tendon. 

The hyoid muscles, like those of the head, of Baleniceps were 
excessively like those of the Pelican, but I attach no systematic 
importance to the similarity, as I have very little material with 
which to compare these muscles in a number of different groups, 
and the material J have shows that, apart from obviously adaptive 
features, these muscles are much alike in widely separated groups. 


CAUDAL MUSCLES. 


Pubo-coccygeus externus.—A flat band of muscle from the 
posterior dorsal margin of the end of the pubis, narrowing to its 
insertion on the under surface of the sheath of the external 
rectrix. 

Pubo-coccygeus internus.—This is a much wider and thinner 
muscle, deep of the externus, and arising from a greater area of 
the pubis with a reach on to the ischium. It is inserted to the 
heemapophyses of the posterior caudal vertebree. 

Levator coccygis.—The two levators form a strong diagonal 
mass of musculature on the dorsal surface of the tail, anterior to 
the oil-gland. They arise from the ilium and the lateral pro- 
cesses of the caudal vertebre and are inserted by a series of 
tendinous slips to the spinous processes of the caudals and to the 
membrane covering the rectrices. 

Depressor coccygis.—Avises from the transverse process of the 
last sacral vertebra by a strong tendon just at the articulation 
with the ilium, and from the transverse processes of the first 
three free caudals; insertion to the transverse processes and 
hemapophyses of the posterior caudals. 

Ilio-coccygews.—Only the outer of the two slips which usually 
represent this muscle is present. It arises from the illum just 
dorsal to the origin of the depressor coceygis, and is inserted to 
the outer surface of the capsule of the external rectrix. 


668 DR. P. CHALMERS MITCHELL ON THE 


I have no standard for comparison in the case of the caudal 
muscles. ‘The chief difference from the condition in Leptoptilus 
is the absence of the inner slip of the ilio-coccygeus. 


MUSCLES OF THE SHOULDER AND WING. 


Cucullaris.—Vhe cervical portion is well developed, forming a 
definite sheet of circular fibres which stop abruptly in line with 
the proximal edge of the rhomboideus externus where they are 
inserted along the clavicle. 

Lthomboideus externus.—Orvigin tendinous from the neural 
crests of five and a half vertebre beginning at just opposite the 
junction of the scapula and clavicle. The fibres run outwards 
nearly transversely to all the scapula except the down-turned 
posterior end and forwards to part of the clavicle (text-fig. 125, 
Rh. 2). 

LRthomboideus profundus or internus.—Origin tendinous, a little 
short of the externus proximally and reaching just beyond it 
distally. The fibres run outwards and backwards to no part of 
the clavicle but to the whole length of the scapula including the 
down-turned end (text-fig. 125, Rh. 1). 

The two rhomboid muscles are nearly equal in thickness. 
These two muscles, according to Fiirbringer, and my own obser- 
vations confirm his view, are in process of creeping forwards. 
Their condition in Baleniceps shows a considerable degree of 
specialization, but I have not material to compare the condition 
in allied birds. 

Latissimus dorsi anterior.—A broad strap of muscle arising 
from the anterior dorsal vertebre only and running downwards 
and forwards, dipping under the anconeus, to a fleshy insertion to 
the shaft of the humerus distal of the insertion of the posterior 
division of the muscle and unconnected with it (text-fig. 125, L.A.). 

Latissimus dorsi posterior,—Fleshy origin, the anterior edge of 
which touches but is not fused with the posterior edge of the 
anterior division. Origin wider than that of the anterior 
division, but not reaching quite as far back as the proximal edge 
of the ilium. Its fibres converge to form a band about the same 
width as the lat. dorsi anterior (text-fig. 125, L.P. 1), pass under 
that muscle with a more proximal slope, to be inserted along a 
strong tendon (text-fig. 125, L.P. 2) which is inserted to the 
scapula under the scapular anchor of the anconzus, proximal to 
the insertion of the lat. dorsi anterior, and which joins the 
anconeus belly distally. 

Latissimus dorsi metapatagialis.—Absent. 

The anterior division is like that in Leptoptilus. The connec- 
tion of the tendon of insertion of the posterior division with the 
anconeus occurs also in Leptoptilus, but I have noted a somewhat 
similar arrangement in Bubo maximus. The loss of the meta- 
patagial division has been noted by Fiirbringer in Plotus, but it 
is usually present in the Herons, Storks, and Steganopods. 


ANATOMY OF THE SHOE-BILL. 669 


Serratus superficialis anterior.—¥rom the ventral end of the 
last cervical rib and the first dorsal rib converging to a flat 


Text-fig. 125. 


Rh... 
Ve —— Dela) 
TH Ui inane Del. 2 
WO SH YK WN \\\ oly hing DY Rrra € 

fis iat mi i yk Ri So Seah 
yf ft! = TARR 
fi) ANT 

AN 


Shoulder-muscles of Baleniceps. 


Right wing, external aspect. Muscle striped: tendon dotted. 


Rh. 1. Rhomboideus profundus, cut across. 

Rh. 2. Rhomboideus externus, cut across. 

Del. 1. Tendinous anchor of Deltoides major. 

Del. 2. Cut surface of Deltoides major reflected. 

Del. 3. Distal portion and insertion to humerus of Deltoides major. 

S.P. Scapulo-humeralis posterior. 

Pee.m. Tendon of insertion of Supra-coracoideus (Pectoralis minor or secundus). 
$.C. | External scapular head of Sub-coraco-scapularis. 

A.S,.  Anconzeus scapularis, showing scapular origin, and anchor to humerus. It 


LO 


has been divided to show the Latissimus dorsi. 

Anconeus scapularis, part of the belly. 

Anconeus humeralis. 

Latissimus dorsi anterior, insertion. 

Cut edge of Latissimus dorsi posterior. : 

Tendon of insertion of Lat. dorsi posterior from humerus to junction with 
Anconeeus scapularis. 

Vestige of Expansor secundariorum. The distinctness of this is exaggerated 
in the drawing. 


670 DR. P. CHALMERS MITCHELL ON THE 


tendon which is inserted to the postglenoid scapula between the 
two parts of the sub-coraco-scapularis, its insertion being quite 
covered by the outer part of that. \ 

Serratus superficialis posterior.—From the first two uncinate 
processes and area of their ribs in line with them to about three- 
quarters of an inch of the posterior inferior border of the scapula ; 
covered .by the third portion of the serratus superficialis. 

Serratus superficialis metapatagialis —Origin from the same 
two ribs as the ser. super. posterior, but entirely from below the 
uncinate processes, and reaching down almost to the sternum. 
Insertion to the metapatagium, Cwith a strong tendinous slip to 
the tip of the scapula. 

The first of the three serrati, according to Fiirbringer, is very 
variable even within families. ‘The second is constant in many 
families, variable in others. The limitation of the origin to 
dorsal of the uncinates is somewhat rare but has been noted in 
Phenicopterus. The insertion of the serratus metapat. partly to 
the scapula appears to be extremely rare. Fiibringer has noted 
it in Crex, where the condition 1s much as I find it to be in 
Baleniceps, and in Fulmarus and Bucorvus, where the scapular 
insertion alone occurs. The relation to the scapula is probably a 
vestige of the origin of this muscle as a separated portion of the 
serratus superfic. posterior. 

Serratus profundus.—From the last two cervical and first two 
dorsal ribs to the scapula in four digitations. According to 
Fiirbringer, this arrangement is normal in Herodii. 

Biceps brachii.—This arises by a flat narrow tendon from the 
acrocoracoid (text-fig. 128, B. 1, p. 675), alongside but not covered 
by the origin of the coracobrachialis externus and separated by 
that muscle from the tendon of insertion of the supracoracoideus 
(pectoralis secundus). It passes under the insertions of the 
pectoralis major without being connected with them, and passing 
into a rounded belly (text-fig. “128, B. 2) runs down parallel with 
the humerus to be inserted toa knob on the ulnar face of the 

radius (text-fig. 127, Bi., 1, p. 672). It is then continued across 

to the opposite face of the ulna (text-fig. 127, 4, 5) by a deep 
broad tendon and a narrow more superficial tendon, first sending 
a strong slip (text-fig. 127, 2) to the radial end of a radio-ulnar 
ligament. 

The biceps obviously presents a highly specialized condition in 
Baleniceps, the specialization consisting of the complete loss of 
the usual humeral head. The two divisions are well separated 
at their origin, and the radial and ulnar tendons of insertion 
separate rather high up in most Steganopods, Storks and Herons. 
As Furbringer has pointed out, the humeral head in such cases 
ean be traced to the radial insertion. As both radial and ulnar 
insertions are well marked, indeed rather unusually complex in 
Baleniceps, 1 infer that the loss of the humeral head is com- 
paratively recent. 

Biceps patagialis —This slip to the patagial tendons is absent 


ANATOMY OF THE SHOE-BILL. 671 


in Baleniceps, as in Storks, Herons, Scopus, and mest Steganopods. 
The absence, however, is not of much value; the slip is present 
in some Steganopods, in Spoonbills, in Phenicopterus, and is 
present or absent within the same family in a number of cases. 
Deltoides propatagialis (text-fig. 126, Del. pat.). 
I have already mentioned that there is no biceps propatagialis. 


Text-fig. 126. 


cry A 
cs 


Patagial muscles and tendons of Baleniceps. 


Del.m. Deltoides major. Ane. Anconzus scapularis. Hum. Humerus. Bie. 
Biceps. Del.pat. Deltoides propatagialis. Pec. Pectoralis major cut across. 
P.l. Pectoralis slip to longus tendon. P.b. Pectoralis slip to brevis tendon. 
Lon. Longus tendon. Brev. Brevis tendon with the slips named «, 3, and 
y by Fiirbringer. Ex. Extensor metacarpi radialis. 


672 DR. P. CHALMERS MITCHELL ON THE 


There is no cucullaris propatagialis. The patagial tendons arise 
solely from the deltoides propatagialis and from the pectoralis 
propatagialis, and there are no anchors to the humerus. 

The origin of the deltoides patagialis is from the clavicle, 
acrocoracoid ligament and part of the scapula, the latter origin 
not being found in Herons. Distally it sphts into two peaks, a 
smaller from which the longus tendon arises and a larger for the 
brevis tendon. This is a more specialized condition than in 
Steganopods, Storks and Herons generally, where even the longus 
and brevis tendons have a short common course, but in Scopus 
and Leptoptilus there ave separate peaks for the tendons. 


Text-fig. 127. 


Bo. 


Insertion of Biceps hgament. 


R. Radius. U. Ulna. Bi. Biceps tendon: 1, msertion to radius; 2, insertion to 
radio-ulnar ligament; 4, insertion to ulna; 5, second superficial insertion to 
ulna ; 3, radio-wnar ligament. L.h. Humero-ulnar hgament. 


The longus tendon (text-fig. 126, Lon.), after being reinforced 
by a slip from the pectoral, enlarges in width and becomes elastic, 
this portion being doubled, and being anchored by a very faint 
(much fainter and more diffuse than would appear from the 
drawing in fig. 126) set of fibres from the distal portion of the 
brevis. 

The brevis tendon (text-fig. 126, Brev.) is highly specialized. 
The main mass of the muscle passes into a strong rounded tendon 
which is reinforced by the pectoralis slip and represents the con- 
joined a and /3 slips of Furbringer, the beta slip being the direct 
continuation of the muscle, but giving off half-way down the 
patagium a broader and weaker alpha slip. The latter itself 
becomes doubled distally, gives off a weak anchor to the elastic 
portion of the longus tendon and is inserted to the extensor 
metacarpi radialis tendon. ‘The beta slip broadens out as it 
reaches the fascia over the extensor, sends forward a stout anchor 
which covers and is fused with the tendinous head of the extensor 
metacarpi, and sends downwards a branch which forms a forked 
fan reaching the distal edge of the forearm. The gamma slip of 


ANATOMY OF THE SHOE-BILL. 673 


Fiirbringer is the most proximally placed, and is distinct although 
very flat and weak throughout its whole length from its origin at 
the proximal side of the brevis muscular peak to its insertion to 
the recurrent beta slip. 

The course of evolution of the brevis tendon appears to have 
been from a wide rather diffused band to first a specialization of 
portions of that band into the slips distinguished by Firbringer, 
then to a separation of these slips, and finally to the loss of one or 
more of them. In the Storks, Herons and Scopus the slips are 
at least separate distally ; in Storks and Herons the separation of 
alpha and beta is only distal and does not begin so high up as in 
Baleniceps, in which, although the actual separation occurs only 
about half-way down the patagium, the identity of beta can be 
traced right up to its origin. Beddard figures an almost 
similar condition for Scopus (2, fig. 2). So also the very com- 
plete separation of gamma and beta occurs in Scopus and 
Baleniceps, and is much less distinct except distally in Storks and 
Herons. The resemblance between Baleniceps and Scopus appears 
to be rather close; the most important differences being the 
‘greater distinctness of the anchor to the longus in Scopus, and 
the presence of an anchor to the humerus in the same bird. 

Delioides major (text-fig. 125, Del. 1, Del. 2, Del. 3, p. 669; 
text-fig. 126, Del.m.; text-fig. 128, De.).—This large muscle arises 
fleshy ‘from the scapula but with a distal tendinous anchor just 
external to that of the anconzeus, and is inserted fleshy to nearly 
half-way down the humerus. It is very nearly divided into 
the two portions visible in Leptoptilus and other storks. The 
tendinous anchor occurs in the Herons and Storks that I have 
dissected, and Beddard has recorded it in Scopus. 

Deltoides minor.—This muscle, possibly owing to the large size 
of the deltoides major, is not to be distinguished as a separate 
muscle: probably it is absent. In Storks it is small and quite 
separate. 

Scapulo-humeralis anterior.—This small muscle is absent. In 
Steganopods, Storks and Herons it les very close to the teres 
major, so that it is possible that it may have fused with this in 
Baleniceps. Beddard does not mention it in his description of 
the shoulder muscles of Scopus, so that possibly it may also be 
absent in that bird. 

Scapulo-humeralis posterior (Teres major) (text-fig. 125, SP).— 
A strong but relatively rather small muscle arising from about 
the distal half of the scapula and inserted to the humerus between 
the two heads of the anconeus. A relatively narrow insertion, 
according to Furbringer, also occurs in Steganopods, Storks 
and Herons. In Saleniceps it has no accessory anchors or 
attachments. 

Swb-coraco-scapularis.—The coracoid head (Coracobrachialis 
brevis of Garrod) is single and much smaller than the scapular 
heads. It arises only from the proximal half to third of the 
inner face of the coracoid, as in Storks and Herons, and converges 


674 DR. P, CHALMERS MITCHELL ON THE 


to join the scapular heads near their tendon of insertion. The 
external (text-fig. 125, 8.C., p. 669) and internal scapular heads 
arise from about the second fifth of the under surface of the 
scapula, where they are separated by the insertion of the serratus 
superficialis anterior. The three heads unite to form a strong 
rounded tendon inserted to the median tubercle of the humerus. 


‘ 


AnCOnCUS. 


Anconeus scapularis.— Origin by a strong forked tendon 
from scapula (text-fig, 125, A.S.1, A.S. 2), passes into a rounded 
muscular belly which sends an anchor to the humerus near the 
insertion of the latissimus dorsi and receives a strong tendon 
from the latissimus dorsi posterior. Passes into a strong flat 
tendon just before reaching the elbow. 

Anconeus humeralis (text-fig. 125, A.H.).—Origin from the 
whole length of the humerus, the origin being cleft proximally. 
Passes into a tendon at the distal end of the humerus, and this 
runs parallel with but united only by membrane to the tendon of 
insertion of the anconeus scapularis. Insertion to the olecranon 
of the ulna. 

The forked head of the scapular portion, the anchor to the 
humerus, and the general relations of the two divisions of the 
muscle are very much like what I have observed or find recorded 
in Storks and Herons. The absence of any extension of the 
scapular head to the clavicle or coracoid is rather a primitive 
feature. 

Anconeus caput coracoideum (Hupansor secundariorunr) (text- 
fig. 125, 8.)—At the elbow there was a slip of muscular fibres 
connected with the feathers and giving rise to a very delicate 
tendon which I traced up the under surface of the skin close to 
the anconzeus, but which then appeared to become diffuse and be 
lost in the subdermal fascie. There was no trace of it passing 
through the edge of the teres major, as usually happens when it 
is well developed, or in the axilla. 

This was one of the muscles to which Garrod paid great atten- 
tion, hoping to find it useful in classification, but further obser- 
vations have not justified his anticipations, as it 1s present or 
absent in very closely allied birds. It is usually absent in 
Steganopods, but present in a few cases. It is present in Storks, 
and in Herons except Ardetta and Cancroma. According to 
Beddard it is absent in Scopus. Its vestigial presence in Balent- 
ceps is therefore interesting but of no systematic value. 

Pectoralis thoracicus.—The great pectoral (text-fig. 126, Pec., 
p. 671; text-fig. 128, Pec.) in Baleniceps is an enormous mass of 
muscle arising from the clavicle, the membrane between the clavicle 
and coracoid, from the whole of the keel and from all the posterior 
part of the sternum with a considerable overlap to the ribs. I 
could not trace any definite horizontal division of the muscular 
mass. The insertion is by two very distinct tendons, which 
cross each other in a remarkable fashion. The greater and more 


ANATOMY OF THE SHOE-BILL. 675 


proximal mass of the muscle converges to a strong flattened 
tendon (text-fig. 128, 1), which is inserted to the humerus rather 
distally and not far from the posterior end of the deltoid inser 
tion. The more distal portion of the muscle converges to a mucl. 
broader tendon, which forms the posterior border of the whole 
muscle and then dipping under the first tendon of insertion runs 
in to the humerus proximally of it (text-fig. 128, 2). There is 
also a strong anchor to the humerus, shown as cut and reflected 
in the figure (text-fig. 128, 3). 


Text-fig. 128. 


Pectoral muscle of Baleniceps. Tendon dotted ; muscle striped. 


Acro. Acrocoracoid process. Hu. Humerus. 

Pec. Pectoralis majer, cut across. 

1, 2. Insertion tendons of pectoralis to humerus. 8. Anchor to humerus, 
divided and reflected. 

P.pt. Pectorales propatagiales. 

Su. Tendon of insertion of supracoracoideus (Pect. minor). 

De. Deltoides major. 

Cor. ex. Coracobrachialis externus. 

B.1. Tendon of origin of biceps. B.2. Cut belly of biceps. 


The large area of origin of the great’ pectoral is of course 
associated with a powerful wing, and is probably purely adaptive 
Proc. Zoou. Soc.—1913, No. XLV. 45 


676 DR. P. CHALMERS MITCHELL ON THE 


The strong proximal anchor (text-fig. 128, 3) ito the humerus also 
occurs In many Steganopods, mm Storks, Herons, and is probably 
vepresented in Scopus, 12 which bird Beddard mentions a strong 
insertion to a fibrous aponeurosis attached to the crista of the 
humerus and covering the biceps. It is characteristic of Storks. 
as opposed to Herons, that in the former birds the great pectoral 
is completely divided into two msucles, as it 1s in the Pelican 
and some other Steganopods. ‘The insertions of these portions as 
described by Weldon correspond almost exactly with the double 
insertion in Baleniceps, and I was able to separate the mass of 
muscle quite easily into portions corresponding with these inser- 
tions, although, in the absence of the separate insertions, I should 
not have described the muscle as doubled. But, whatever the 
distinction be worth, the great pectoral muscle of Daleniceps is 
more Ciconine than Ardeme. Beddard’s description of the condi- 
tion in Scopus is not sufficiently detailed to follow in this matter, 
but he speaks of it as ‘‘ partly doubled,” and the humerus shows 
marks of a double insertion. 

Pectoralis propatagialis (text-figs. 126, 128, pp. 671, 675).—-As IT 
have already stated, there are separate slips from the pectoral for 
the longus and brevis tendons. Both slips are entirely tendinous, 
and that for the longus is smaller and more superficial (text-fig. 
126, P.1., P.b.; text-fig. 128, P.pt.). In my dissections of Herons, 
I find similarly cdistinet slips for the brevis and longus from the 
pectoral ; Beddard mentions them for Scopus, but-in the case of 
Storks the usual arrangement appears to be the more primitive 
condition of a single slip, which joins the patagial tendon before 
that has divided into longus and brevis. 

Pectoralis abdominalis.—This is absent in Baleniceps as in 
Storks. It is present in Herons, but Beddard does not refer to: 
its presence or absence in the case of Scopus. 

Supracoracoideus (Pectoralis minor).—This is a small and 
rather narrow muscle elongately oval, with a centrally placed 
tendon like the mid-rib of a leaf. Its fleshy origin is limited to: 
a very small part of the sternum, including no part of the keel, 
and part of the coracoid and the membrane between the coracoid 
and clavicle. It is widely separated on the coracoid from the 
origin of the coracobrachialis posterior (pectoralis tertius) and its 
tendon of insertion (text-fig. 128, Su.) is free from any fibres. 
that could represent a deltoides minor. Its general relations and 
small size are closely paralled in Herons and Storks. 

Coracobrachialis externus or anterior.—A strong muscle arising 
fleshy from the acrocoracoid only (text-fig. 128, Cor. ex.) and 
covered only at the extreme edge by the tendon of origin of the 
biceps. Insertion on a bro adly oval area to the planum: bicipitale- 
of the humerus. 

Coracobrachialis internus (Pestoralis tertius).—A very stout 
almost doubled mass of muscle from the distal dorsal two-thirds 
of the edge of the coracoid opposite the origin of the supracora- 


ANATOMY OF THE SHOE-BILL. 677 


coideus, but quite separate from that. Its fibres converge to a 
strong tendon inserted to a peak of the median tubercle of the 
humerus. 


Muscurs oF ForREARM AND NECK. 


Brachialis inferior.—A very strong flat sheet of muscle with « 
fleshy origin and insertion, occupying the angle between the 
humerus and ulna, the insertion to the ulna being twice the 
width of the origin from the flexor aspect of the humerus, 

Pronator sublimis or brevis.—Short muscle from the inner 
condyle of the humerus to the first quarter of the radius. This 
insertion 1s rather shorter than in Leptoptilus. 

Pronator profundus or longus.—As in Leptoptilus a larger 
muscle, from the inner condyle of the humerus to a little beyond 
the surface of the radius covered by the brevis and more on the 
ulnar aspect than the brevis. 

Entepicondylo-ulnaris.—Absent, as in Leptoptilus, but accord- 
ing to Gadow present only in Rasores and Tinamus. 

Hetepicondylo-ulnaris.—From outer condyle of the humerus to 
first third of ulna on its radial face. A very thick and strong 
muscle, closely united with the flexor digitorum profundus, As 
in Leptoptilus. 

Eetepicondylo-radialis.—A thin muscle arising by a flat tendon 
from the outer condyle of the humerus along with the extensor 
digitorum communis, inserted to a quarter of the radius; as in 
Leptoptilus, except that the insertion is shorter in the latter 
bird. 

Flexor carpi ulnaris.—From the inner condyle of the humerus 
with a sesamoid ; runs down the inner surface of the ulna to the 
great tuberosity of the ulnar carpal. Arising as a fleshy belly a 
thinner tendon connected with the quills runs down to end on 
the carpal alongside the great tendon. As in Leptoptilus. 

Ulni-metacarpalis ventralis.—Fleshy from the last third of the 
ulna on the radial face; tendon crosses over a slide on the radial 
carpal and is inserted on a hump of the second metacarpal near 
the attachment of the pollex. 

Ulni-metacarpalis dorsalis.—Short muscle arising by a tendon 
from the distal end of the ulna on its lateral face ; it divides into 
a shorter portion running straight across to the upper part of 
metacarpal ITT and a broader portion inserted to about two- 
thirds of the upper surface of metacarpal III, where that is free. 
Similar in Leptoptilus, except that the first portion is tendinous, 
the second fleshy, while both are fleshy in Baleniceps. 

Hetensor metacarpi radialis—Two heads, outer tendinous, 
inner fleshy, from the outer condyle of the humerus. The outer 
belly is quite separate from the inner belly and is connected with 
the brevis tendons of the patagium (text-fig. 126, Ex., p. 671). 
Insertion to the base of metacarpal I, the tendons from the two 

45* 


678 DR. P. CHALMERS MITCHELL ON THE 


bellies remaining separate until their insertion, so that the muscle 
is completely double. In Leptoptilus the tendons fuse distally. 

Extensor metacarpi ulnaris.—Avises from the external condyle 
of the humerus by a tendon superficial to that of the ectepi- 
condylo-ulnaris ; then a long fleshy belly, then a thin tendon 
passing over a groove in the distal end of the ulna from which it 
receives a strong anchoring slip, absent in Leptoptilws, to its in- 
sertion on metacarpal IT just where metacarpal IIT is given off. 

Flexor digitorum sublimis.—A strong band of tendon runs 
from the inner condyle of the humerus to the ulnar carpal, from 
the upper side of which the flexor digitorum sublimis arises as a 
delicate fleshy muscle giving rise to a slender tendon which passes 
over a groove in the ulnar carpal and is inserted to the base of 
phalanx 2 of digit II, but with first an insertion to the base of 
the first phalanx of that digit, which I do not find recorded in 
my notes on Leptoptilus. 

Flevor digitorum profundus.—Arises fleshy from the second 
and third fifths of the ulna, very closely connected with the in- 
sertion of the ectepicondylo-ulnaris. The tendon begins where 
the origin from the ulna ceases, and runs down the radial face of 
the ulnar carpal under the ligament from the radius to meta- 
carpal IT, and then follows the tendon of the superficial flexor to 
be inserted just beyond it to phalanx 2 of digit IT. It receives 
a strong slip from the short extensor of the thumb, which I did 
not record in the case of Leptoptilus. Gadow mentions somewhat 
similar relations with the thumb in the case of Owls and 
Feliornis. ae 

Extensor digitorum commuinis.—Arises tendinous from the 
external condyle of the humerus and passes into a slender belly a 
quarter way down the forearm, but receives no fibres from the 
ulna. Its tendon of insertion passes througha groove in the end 
of the ulna and then sends a branch to the base of phalanx 1 of 
digit Iand a stronger tendon to phalanx 2 of digit IT. As in 
Leptoptilus. 

Extensor pollicis longus.—Two slender fleshy heads from the 
adjacent surfaces of the radius and ulna at their proximal ends, 
with accessory fibres froma large part of the length of the radius 
on its ulnar face. Tendon unites with that of the extensor 
metacarpi radialis at its insertion. As in Leptoptilus. 

Extensor indicis longus.—One head fleshy from the distal half 
of the radius and a second much smaller, tendinous from the 
distal end of the radius and from radial carpal. Insertion to the 
second phalanx of digit IT, but attached by fascia to the first 
phalanx. As in Leptoptilus. 

Tnterosseus dorsalis.—Arises fleshy from the opposite faces of 
metacarpals II and Il]; fibres run to a centrally placed tendon, 
like the midrib of a leaf, and this is inserted to the base of the 
second phalanx of digit II. As in Leptoptilus. 

Interosseus palmaris.—More ventral and stronger than the 
foregoing muscle but with similar origin and arrangement. 


ANATOMY OF THE SHOE-BILL. 679 


Tendon inserted to phalanx 2 of digit IT. In Leptoptilus I have 
noted it as reaching only the first phalanx. 

Abductor indicis.—Strong muscle arising fleshy from the whole 
of the radial side of metacarpal II. Inserted to the base of the 
phalanx | of digit IT. As in Leptoptilus. 

Flexor digitt [1T.—Arises fleshy from the ulnar side of meta- 
carpal ITI and is inserted to the base of the first phalanx of the 
corresponding digit. As in Leptoptilus. 

Adductor pollicis—A strong muscle from the metacarpal to 
the tip of the pollex. 

Hetensor pollicis—This muscle, which is usually described as 
single, is represented by two distinct and well-developed muscles, 
a condition which has been described in the ease of Struthio, but 
not in other birds. Most probably, if it were carefully looked 
for, it would be found elsewhere. The first of the two isa strong 
slip from metacarpal I and the tendon of the extensor metacarpi 
radialis to the radial side of the thumb. It is this muscle that 
gives off a slip to the flexor digitorum profundus. The second 
muscle is from the head of metacarpal II to the base of the 
thumb. 


MuSscLEs oF THE THIGH AND Lec. 


Iio-tibialis internus (Sartorius).—A large and strong strap 
arising from the anterior and lower border of the ilium and from 
the fasciee over the gluteus medius, and closely united along its 
distal border with the gluteus maximus. Normal insertion to 
the tibia. I find no notable difference as compared with Storks 
and Herons. 

Ilio-tibialis (Gluteus maximus).— The origin is entirely ten- 
dinous and a median tendinous area separates the fleshy anterior 
and posterior borders. Posteriorly the origin extends backwards 
half-way over the origin of the biceps, that is to say what Garrod 
called the post-acetabular part of the muscle, the ilio-tibialis pos- 
terior, is present. Garrod tried to use the presence or absence of 
this in his systematic arrangements, but without much success. 
It is absent in. the Steganopods generally, usually absent in Storks 
but present in Ciconia, absent or very slightly developed in 
Herons. So far as I can judge, the presence of the post-acetabular 
portion of this muscle is a primitive condition, and it has been 
lost or reduced independently in many groups of birds. 

Ilio-trochanterici.— The externus (text-fig. 129, Gl.a) and the 
posterior (text-fig. 129, Gl.2) are both present in the normal 
condition. The minimus and quartus are represented by a single 
tendon of origin and muscular belly (text-fig. 129, Gl.3). In 
Leptoptilus I found these quite distinct in their origin and inser- 
tion; in Vycticorax they had a common tendon to the femur but 
separate insertions to the ilium. These muscles, however, vary so 
much from bird to bird that I cannot attach any significance to 
their distinctness or fusion. 


680 


DR. P. CHALMERS MITCHELL ON THE 


Text-fig. 129. 


Va: 
"EL. Pro. 
Diagram of Muscles of the leg in Baleniceps. 
Left leg, external aspect. Tendon is dotted. 


FEMUR. Femur. FIB. Fibula. 
Gl.a. llio-trochantericus externus (Gluteus anterior). Gl. 2. Il.troch. 
posterior (Gluteus minor). GI. 3. Il.troch. anterior et medius 
(Gluteus minimus and quartus). 
Ob.ex. Obturator externus. 


Ob.in. Obturator internus, surrounded by Gem., Gemellus. 

FE.C. Origin of Femoro-caudal. 

A.D. Adductor longus (the upper muscle) and Adductor magnus. 

BIC, Insertion of Ilio-fibularis or biceps, passing through a sling. 

Gas. External head of Gastrocnemius, cut and reflected to show relation 


to short arm of the Biceps sling. 

F111, Fl1.10, Fl.1.1V. Tendons to respective toes of the Perforated 
Flexor muscles. 

Amb. Ambiens head of Perforated Flexors. 

Fl.1.E. External head of Perforated Flexors. : 

FI.2.11, F1.2.11I. Perforated and Perforating Flexors of digits II & ITT. 

Fl. Hal. Flexor longus hallucis. 

Fl. Pro. Flexor profundus. 

Ex.Com. Extensor communis. 


ANATOMY OF THE SHOE-BILL. 681 


Ilio-femoralis internus (Pectineus).—Strong, little fleshy slip 
from the ventral edge of the preacetabular portion of the ilium to 
the back of the femur below the neck. As in most birds. 

Femori-tibiales (Crureus and Vastus).—The external muscles 
are fused to form a common mass; the vastus internus is large, 
arising from nearly the whole length of the femur. 

Caud-ilio-femoralis (lKemoro-caudal).—This is a broad thin 
strap of muscle arising fleshy from the femur (text-fig. 129, F-H.C.) 
and running upwards and backwards under the biceps and be- 
tween the semitendinosus and the semimembranosus and passing 
to the underside of the tail, where it becomes a thin tendon 
spreading out into a sheet which meets its fellow of the other 
side, the combined insertion being to the tendon of the depressor 
ecoccygis where that is inserted to the hzemapophyses of the 
posterior caudal vertebree. There is no accessory femoro-caudal, 
The accessory fem.-caud. is, I believe, invariably absent in Stega- 
nopods, Herons and Storks, although it is present in Spoonbills 
and the Flamingo. The fem.-caud. itself tends to be degenerate. 
It is usually present in Storks, but is very slender in /%ssuia and 
absent in Leptoptilus; it is weak in the Herons and absent in 
several genera. 

Caud-ilio-flecorius (Semitendinosus and Accessory semitendi- 
nosus).—Origin fleshy from the ischium behind the biceps and 
extending on to the fascia posterior to the ischium; meets the 
rather small but distinct accessory or femoral head in a tendinous 
raphe, and the combined muscles are inserted to the middle belly 
of the gastrocnemius (text-fig. 130, C.J]. 1 & 2). The muscle is 
much weaker than the semimembranosus. 

Ischio-flecorius (Semimembranosus).—This ig the usual broad 
strap underlying the semitendinosus, and in this case much 
thicker and wider than the latter. It has a wide origin from the 
lower edge of the ischium and the fascia over the obdurator 
externus, is unconnected with the semitendinosus, but receives a 
strong tendinous slip (text-fig. 130, Sl.) from the inner adductor 
and then is inserted to the tibia by a flat tendon. 

Gastrocnemius.—There are the usual three heads of which the 
tibial head is the strongest. The outer head arises from the 
external condyle of the femur in common with the short arm of 
the biceps sling (text-fig. 129, Gas.). The middle head is the 
smallest and arises from between the condyles of the femur by a. 
flat tendon. The tibial head is enormous and arises from the 
tendon of the ilio-tibialis and from the cnemial crest of the tibia. 
The three heads unite in the usual way rather less than half-way 
down the leg to form the tendo achillis. 

Relations of the Caud-ilio-flexorius, [schio-flexorius, and Gastro- 
cnemius.— Weldon (38) called attention to the varying relations 
of these muscles and the differences they presented in Storks, 
Ducks, and Phenicopterus. I have noted them in a number 
of Storks and Herons and paid a good deal of attention to 
them in Gruiform and Limicoline Birds (31 & 32). With 


minor variations as to the precise interconnections of the 


682 DR. P. CHALMERS MITCHELL ON THE 


semitendinosus and semimembranosus, the condition which I 
found in Laleniceps seems to be typical in Herons and Storks 
(text-fig. 130). ‘The middle head of the gastrocnemius is joined 
by the semitendinosus just after the latter has been met by 
its accessory. ‘The accessory origin from the femur is parallel 
to but separate from that of the middle head. The accessory is 
present in Herons, Storks, Seopus, and in Daleniceps ; it is 
frequently absent among the Steganopods. In my opinion the 
middle head of the gastrocnemius is a separated portion of the 
accessory semitendinosus, and separation of the two, with sub- 
sequent disappearance of one or of both, is a secondary or 
specialized condition. 

Tlio-fibularis (Biceps)— Very strong fleshy origin from the 
whole of the post-acetabular ridge of the ilium to the beginning 
of the origin of the semitendinosus. The strong belly converges 
to a rounded tendon which is inserted to the fibula after passing 
through a sling in the usual way (text-fig. 129, BIC.). The short 
arm of the sling has a strong anchor to the fibula, which I happen 
to have noted in Herons, but which is present also in many birds. 
belonging to widely separated groups. 

Ischio-femoralis (Obdurator eaternus).—Arises by strong tendon 
from external condyle of the femur (text-fig. 129, Ob.ex.) and 
inserted fleshy to surface of the ischium. 

Obdurator (Obdurator internus).—Origin by a strong tendon 
surrounded by a gemellus muscle (text- fig. 129, OB.in.Gem.) from 
the external condyle of the femur proximal to the obdurator 
externus. Garrod (18) believed that in most cases the insertion 
of this muscle to the inner aspect of the pubis and ischium 
could be distinguished as oval or triangular, and attached some 
systematic value to the condition. He described it as oval in 
Steganopods and Storks and triangular in Herons; in Baleniceps 
it is plainly oval. 

Pub-ischio-femorales (Adductor longus and Add. nagnus).—The 
external or longus is only, about half the width of the inner 
or magnus, but. their origins and insertions are practically co- 
extensive. In my notes i find that they were nearly equal in 
Herons and Storks, but I have not paid special attention i the 
point. The slips ta ‘om the magnus to the tibia (text-fig. 130, Sl.) 
and the slp to the semimembranosus (text-fig. 130) I nave noe 
noted in Storks or Herons. 

Peroneus superficialis (longus).—Strong muscle from crest of 
tibia and fascia over the tibialis anticus ; usual insertion by broad 
tendon to the fascia of the ankle and a long tendon running down 
to join with the tendon of the perforated flexor of the third toe. 
Precisely the same relations exist in Storks and Herons, but also 
in so many other birds that no systematic importance can be 
attached to them. 

Peroneus profundus.—A. short but stout muscle from the tibia 
below the fibula; its tendon passes over the ankle-joint to be 
inserted to a knob on the outer side of the tarsus-metatarsus. 


- ANATOMY OF THE SHOE-BILL. 683 


According to Weldon and my own notes, this muscle is absent in 
Leptoptilus, but: I found it present and with an extension of its 
origin to the fibula in Herons. 

Tibialis anticus.—The outer head arises by a strong tendon 
from the external condyle of the femur and runs in a deep groove 
to join the fleshy head from the tibial erest. Insertion by a 
forked tendon to the tarsus-metatarsus, in a pit about an Inch 
below the joint. The conditions are practically the same in 
Herons and Storks. 


Text-fig. 130. 


[sf K—= ary rT 
il 


Gaatrocnemius and its relations in Baleéniceps. 


Ad. Adduetor magnus SI. Tendinous slips from adductor to tibia. 
Isf. Ischio-flexorius (Semimembranosus). 
C.11.1. Femoral head of Caud-ilio-flexorius (Accessory Semitendinosus). 
C. 11.2. Belly of Caud-ilio-flexorius (Semitendimosus). 
Ge. 1, external, Ge.2, middle, Ge. 3, tibial portion of Gastrocnemius. 


Soleus.—This little muscle has the usual relations, but is 
relatively rather stronger than in Storks and Herons; it arises 
fleshy from the inner side of the tibia and is inserted to the 
annular cartilage of the ancle-joint. 

Extensor communis digitorum: arises fleshy from the crest and 
external surface of the shaft of the tibia (text-fig. 129, Ex.com.). 
‘The strong tendon passes through a bony and a fibrous bridge 
and runs down to the digits where it divides symmetrically into 


684 DR. P. CHALMERS MITCHELL ON THE 


two, each branch again dividing into two, the four tendons 
running respectively to the second digit, to each side of the third 
digit and to the fourth digit. The arrangement is practically 
identical in Storks and Herons. 

Flexor perforans et perforatus (Flexor secundus) of digit I1.— 
Origin is from the external condyle of the femur distal to the 
short arm of the biceps sling (text-fig. 129, Fl. 211, p. 680) and 
from the fascia over the knee-joint, immediately superficial to the 
corresponding flexor of the third digit with which it is closely 
connected. The tendon passes in the normal fashion to the 
second digit, perforating the tendon of the flexor primus and 
being perforated by the branch of the tendon of the flexor 
communis. 

Flexor perforans et perforatus (Flexor secundus) of digit I1I.— 
This has two heads, one just deep of the corresponding flexor of 
the second digit and practically common with it, and a second 
from the edge of the fibula. Its tendon receives a strong slip 
from the tendon of the perforated flexor (flexor primus) of its own 
digit and then is inserted to digit IIT in the same fashion as the 
corresponding flexor of digit II. I did not record the existence 
of the second head of this flexor in my notes on Storks and 
Herons, but otherwise the flexores secundi have identical relations 
in Baleniceps, Storks and Herons. It must be noticed, however, 
that these relations are found in a very large number of birds 
belonging to different groups. 

Flexores perforati (Flexores primi) of digits II, III, IV; 
Rudiment of Aimbiens-—The perforated flexor muscles (text- 
fig. 129, Fl. 1. 11, Fl. 1. ILI, Fl. 1. LV) are very closely united. 
Distally the tendons for the respective digits separate out ; 
proximally the common muscular belly arises from three distinct 
heads and the arrangement is such that fibres to each tendon can 
be traced to each head. The largest head is fleshy from the 
intercondylar notch of the femur; there is an outer rather broad 
tendinous head, superficial to the biceps tendon and arising from 
the head of the fibula (text-fig. 129, Fl. 1. E). The third head is 
a round and very distinct tendon, passing under the biceps tendon 
and running partly te the head of the fibula and partly to the 
fascia of origin of the flexores secundi (text-fig. 129, Amb.). The 
inner fleshy head is normal and oceurs in practically identical 
form in all birds that I have dissected. The outer tendinous 
head also is usually present, and exists in Storks and Herons, the 
chief differences it presents being in the extent to which it is 
muscular. In Baleniceps, the tendinous portion is longer and 
the muscular portion relatively shorter than in Storks and Herons, 
thus showing a degenerate condition. The rounded tendon under- 
lying the biceps is more interesting. In birds where the curious 
muscle known as the ambiens is present, the tendon of that 
muscle passes through the knee-joint, passes under the biceps 
tendon, sometimes with an anchor to the edge of the fibula, and 
then forms a third head of origin of the perforated flexors, 


ANATOMY OF THE SHOE-BILL. 685 


precisely similar in position and relations to the rounded tendon 
under the biceps in Balwniceps. But for the fact that the 
tendon stops short at the head of the fibula and does not pass 
through the knee capsule to a normal ambiens muscle, it can- 
not be distinguished from the ambiens head of the perforated 
flexors. Garrod (17) first called attention to the interest of the 
ambiens muscle and regarded it as a major key to the classi- 
fication of birds. He divided the Class into two Subclasses, the 
Anomalogonate, containing the Piciformes, Passeriformes, and 
Cypseliformes in which the ambiens is never present, and the 
Homalogonate, containing all the other groups of birds and 
showing that in them the ambiens was normally present. Among 
the Anomalogonatz there is no species in which the ambiens has 
been found; among the Homalogonate there are families and 
genera in which it is absent, and Garrod believed that in 
such cases it had been secondarily lost. In a much later con- 
tribution to the subject (24), I showed that in the Night Heron 
and in Helectus, birds without an ambiens but belonging to 
Garrod’s Homalogonatee, there existed an ambiens head to the 
perforated flexors, absent in the Anomalogonate, and plainly 
suggesting that it was a remnant of the ambiens muscle. In 
a memoir on the anatomy of the Hoatzin (27) I was able to 
describe from dissections of different examples of that bird, a case 
of this possible degeneration in actual progress. Garrod had 
dissected both legs in three examples of the bird and in all cases 
found the ambiens small but normal above the knees, but in five 
out of the six legs it was lost at the knee-joint. He does not 
appear to have had his attention called to the importance of the 
ambiens head of the perforated flexors. I examined each leg in 
two examples, and found in every case an ambiens head to the 
perforated flexors, but the ambiens muscle in some instances 
absent above the knee, in others small and lost at the knee-joint. 
It may therefore be taken as established that the ambiens head 
of the perforated flexors represents a vestige of a complete 
ambiens muscle, and its existence in Baleniceps is of morpho- 
logical rather than systematic importance. The ambiens is 
usually present in the Steganopods, present in the Spoonbills, 
present in some genera of Storks absent in others, absent in 
Scopus, absent in Herons and Saleniceps, but in the last two 
cases its recent loss is shewn by the existence of the vestige to 
which I have now called attention. 

Flexor profundus or perforans and Flexor longus hallucis.—The 
deep flexor as in Storks and Herons arises by fleshy digitations 
(text-fig. 129, Fl. Pro.) from the side of the fibula and from down 
the shaft of the tibia to form a strong round tendon, The flexor 
longus hallucis comes from the inner surface of the outer condyle 
of the femur (text-fig. 129, Fl. Hal.) and similarly forms a round 
tendon. The two tendons pass down to the flexor surface of the 
foot in the usual way. The deep flexor (text-fig. 131, B) breaks 
up into a branch for digits 2, 3, 4, and the hallucis tendon, 


686 DR. P. CHALMERS MITCHELL ON THE 


crossing over the profundus, runs to the hallux, but sends a long 
slender branch which joins the profundus tendon just before 
that divides for the digits. The condition corresponds with what 
Gadow (16, p. 195) calls type I. The hallucis tendon crosses 
over the profundus to reach the hallux, but sends a vineulum to 
it. In Storks and Herons the condition is essentially simular, but 
in the former group the vinculum is stronger and may be in 
separate slips; in Scopws and the Herons the vinculum is much 
more slender and may be absent. For comparison | figure the 
condition in a Stork (text-fig. 131, 8), a Heron (text-fig. 131, A), 
and in Baleeniceps (text-fig. 131, B). 


Text-fig. 131. 


a 


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£5 
f¢ 


— 
Roe 


pa 


53 


Diagram of Deep Flexor Tendons in A, Nyecticorax; B, Baleniceps : 
8, Leptoptilus. 
The longus hallucis tendon is in outline, the flexor profundus is shaded. 
1. Hallux. 2,3 &4, 2nd, 3rd & 4th digits. 


Popliteus.—There was only one of these little muscles stretch- 
ing across between the head of the fibula and the tibia. In 
Leptoptilus I noted two. 


SUMMARY OF MuscutAar ANATOMY. 


Garrod’s hope, excited by his extraordinarily interesting pioneer 
work, that muscular anatomy would furnish a sure clue to the 
classification of birds has not been fulfilled. Garrod relied chiefly 
on the presence or absence of certain muscles which he found to 
vary from group to group. Gadow, who has attempted on a 
complete scale to apply to the system Garrod’s group of muscles, 
using the additional facts made known by Beddard and other 
writers, appreciated that as these muscles were a common heritage 
of all birds, the presence of any of them in any group of birds 
could not be taken as a guide to the systematic position of that 


ANATOMY OF THE SHOE-BILL. 687 


group. He was disposed, however, to attach value to the loss of 
any of these muscles, and accordingly regarded the loss of this or 
that muscle as one of the characters to be employed in judging of 
the relationships of groups. Even this cautious use seems to me 
to be going too far. At present I do not know of any reason 
why we é should suppose that a particular muscle may not have 
been lost independently many times; that is to say of any reason 
why a bird that has lost its femoro-caudal muscle should be more 
nearly related to another bird with a similar loss than to a bird 
which has retained the possession once common to all three. The 
loss is what I have described as a multiradial apocentricity. 
Possibly when we know as much of the development and mor- 
phology of the muscles used by Garrod, as Fiirbringer has taught 
us in the case of the shoulder and wing muscles, we shall be able 
to make more definite use of muscular anatomy in systematic 
ornithology. As, however, muscular anatomy has been used 
freely, I may give a summary, of the chief facts from which more 
confident anatomists would draw inferences 


Comparison of Herons, Storks, Scopus and Baleniceps. 


Peculiar to Baleniceps. 
Absence of latissimus dorsi metapatagialis. (? Scopus.) 
Absence of humeral head of biceps brachii. (Unique.) 
Absence of deltoides minor. (¢ Scopus.) 
Absence of teres minor. (¢ Scopus.) 
Origin of serratus superficialis posterior confined to dorsad of 
uncinate processes. (Same in Phenicopterus.) 
Accessory origin from tip of scapula of serratus metapatagialis. 
Common to Laleniceps and Scopus. 
Condition of deltoides patagialis and patagial tendons. 
Expansor secundariorum vestigial or absent (so also in most 
Steganopods). 
Common to Baleniceps and Herons. 
Presence of peroneus profundus. 
Ambiens reduced to a distal vestige (said to be absent in 
Scopus, present in Storks). 
Deep flexor tendons. 
Common to Laleniceps and Storks. 
Pecuhar arrangement of tendon of insertion of latissimus 
dors. post. 
Practical doubling of pectoralis major (also in some Steganopods). 
Presence of post- acetabular portion of gluteus maximus (at 
least in some Storks; ? Scopus). 
Oval origin of obdurator internus (also in most Steganopods). 


Common to Baleniceps, Scopus, Herons, Storks and most Stega- 
nopods (but also in many other groups). 
Absence of biceps slip to patagium. 
Absence of accessory femoro-caudal. 


688 DR. P. CHALMERS MITCHELL ON THE 


Clearly, so far as the evidence from muscular anatomy goes, 
Baleniceps is an ally of Scopus, Herons and Storks, and shares 
many characters with these birds and Steganopods. The two 
facts that impress me most are the close similarity of the rather 
highly specialized patagial muscle and tendons in the case of 
Baleniceps and Scopus, and the very remarkable condition of the 
pectoralis thoracicus found in Baleniceps and Storks, but which 
probably also exists in Scopus, and certainly in Pelecanus and 
some other Steganopods. 


OstTEOLOGICAL NOTES. 


As the osteology of Baleniceps has been described at length 
in W. K. Parker’s well-known monograph (83), and as I have 
not the time at my disposal to make the elaborate study of 
the different types of Storks and of Herons which is necessary 
before final conclusions can be drawn from the skeleton of 
Baleniceps, | must content myself with a few notes on some of 
the salient points which struck me as requiring special study. 

Occipital condyle.—In Baleniceps this, seen from in front and 
below, has much the appearance of a moderately dolicocephalic 
human cranium. It is sessile, elongated antero-posteriorly, and 
its posterior margin, where it projects slightly into the foramen 
magnum, is convex. In the Herons the condyle is transversely 
elongated, and the posterior margin, where it projects into the 
foramen magnum, 1s the broadest part, is concave and slightly 
erooved, as if to form the beginning of two condyles. In Scopus 
the condyle is also transversely elongated but not so much as in 
Herons, and its foraminal margin is abruptly truncated. In 
Anastomus, Dissura, Xenorhynchus, and Ciconia the long axis is 
transverse, and the foraminal margin is concave or notched. In 
Tantalus, on the other hand, the condyle is nearly spherical, and 
although a notch may just be indicated, the foraminal margin is 
convex. The configuration of the condyle in Lalwniceps is, there- 
fore, unlike Scopus, Storks or Herons, but it is most nearly 
approached by the Z'antalws Storks. 

Paroceipital processes.—The broad, thin and shell-like pro- 
cesses which bend down over the articulation of the quadrate 
are repeated on a smaller scale in Z’antalus, where however the 
lamine are relatively thicker and less extensive. In other Storks 
they are replaced by similarly situated, thick and curving ridges. 
In Scopus and Herons all appearance of the shell-like arrange- 
ment is absent. 

Basitemporal plate.—In Baleniceps the anterior margin of this 
has a erescentic free edge which nearly meets at each side a curved 
lamina projecting from the basisphenoid, so that the Eustachian 
tubes are nearly floored in below. Parker wrote that ‘in the 
Heron these parts are essentially a miniature” of those in Bale- 
niceps, adding that this ‘is certainly not a faint and superficial 
mark of affinity.” But the similarity extends to Scopus and 


ANATOMY OF THE SHOE-BILL 689 


Storks. In Herons the anterior margin is slightly pointed, and 
in Scopus and the Storks, including Zantalus, it is rather more 
sharply pointed, but the general relations and particularly the 
relations to the laminze from the basisphenoid, which I shall now 
describe, are more like those of Laleniceps in the case of Storks 
than of Herons. 

Basisphenoid.—Seen from below this has the usual “T-shaped” 
appearance, the cross bar of the “'T” being contiguous with the 
anterior edge of the basitemporal, the main limb forming the 
rostrum. In Baleniceps delicate crescentic lamelle project back- 
wards nearly meeting the front edge of the basitemporal plate and 
with it forming a floor for the Eustachian tubes. The condition 
of these lamine in Zantalus most closely resembles that of 
Baleniceps; in the other Storks the laminz are less complete, 
and they are least complete in the Herons*and Scopus. The 
rostrum from the “T” cross bar to the attachment of the 
pterygoids is a stout, broad beam of nearly equal width through- 
out its length in Laleniceps. The other birds in the set 1 am 
considering present a series ranging from aleniceps through 
Tantalus, the typical Storks, and Scopus, to the Herons which 
present the end of the series most remote from Laleniceps. The 
rostrum gradually in the series changes from an even beam to 
a sharply contracting, almost triangular outline, and its smooth 
ventral curved surface becomes first slightly ridged, and then 
strongly carmate as in Herons. 

Orbital septum.—This is completely ossified in Baleniceps, 
Scopus, all the Storks, including Zantalus; very incomplete in 
the Herons, including Cancroma. 

Lacrymal.—As Parker has described, the lacrymal of Baleniceps 
is highly peculiar, although no doubt'the peculiarity is partly 
adaptive in relation to the enormous beak. It is a stout vertical 
strut forming the anterior wall of the orbit, firmly anchylosed 
below with the jugal and maxilla, and above with the nasal. On 
the roof of the skull it forms the external portion of the fronto- 
maxillary hinge which runs as a transverse suture across the 
forehead, being thus entirely anterior to the hinge. A thin 
vertical lamina projects from it into the cavity of the orbit, which 
is pierced by a large lacrymal foramen, external to the nasal 
cavity. In Scopus the lacrymal is a vertical beam scooped out 
on its orbital face for the lacrymal canal, but hanging down 
freely along the front of the orbital cavity, until it almost meets 
but does not actually touch the jugal. It has no contact with 
the maxilla and depends from the orbital edge of the frontal, 
behind the fronto-maxillary hinge, and with no more than the 
minutest overlap to the nasal on the distal aspect of the hinge. 
Jn Storks of the genus Vantalus the lacrymal is suspended from 
the orbital edge of the frontal behind the hinge, with just a 
trace of overlap across it to the nasal. From this point of 
suspension the flat external face hangs vertically downwards, 
gradually narrowing, and free from the maxilla and not reaching 


690 DR. P. CHALMERS MITCHELL ON THE 


the jugal below; it has a stout lamina projecting into the orbital 
cavity transversely to the long axis of the skull and pierced for 
the lacrymal canal. In the typical Storks and in Anastonus, 
the external face of the lacrymal is roughly triangular, the convex 
basal line being attached to the orbital edge of the frontal, but 
definitely extending forwards across the hinge to form a very 
loose connection with the nasal, not more than a fifth of the 
whole dorsal edge of the bone. From this, the rapidly narrowing 
triangle hangs down in front of the orbit and is far from reaching 
the jugal and has no connection with the maxilla, The inwardly 
projecting flange is a very thick beam pierced for the lacrymal 
canal. In the Herons, the relations are a still further exaggera- 
tion of the difference between Storks and baleniceps. The outer 
surface of the lacrymal is triangular with a very broad base of 
attachment to the frontal behind the hinge, a small loose overlap 
to the nasal in front of the hinge, and with no connection with 
jugal or maxilla. The inwardly directed flange is absent and 
there is no lacrymal canal. 

In the case of the lacrymal bone, therefore, Baleniceps and 
Ardea stand at opposite ends of a series, Zantalus being nearest 
to Baleeniceps. 

Nasal region.—The nostrils in aleniceps are impervious, a 
stout nasal septum being developed. Gadow states that they are 
pervious in Scopus, but this is a mistake; a very thin lamina 
of bone very slightly fenestrated separates them. In all the 
Herons and Storks they are pervious, the cavity from one nostril 
to the other being large and quite open. The skull is holorhinal 
as in Scopus, Ardea, Cancroma and all the Storks including 
Tantalus, but in many, especially the larger Storks, the proximal 
end of the nasal bone shows a line of weakness running up 
towards the naso-frontal hinge and leading to the schizorhiny 
seen in /bis. From the anterior border of the nostril a groove 
runs along the surface of the beak to the extreme anterior 
end, only the hook of the beak projecting beyond it. Beddard 
appears to lay some stress (4, p. 484) on this point because he 
says that the groove is “precisely like that of Scopus and Can- 
croma.” It is like that of Scopus, but in Cancroma the groove 
is much wider and more shallow and does not reach the extreme 
anterior end. It is much more exactly repeated in the Pelican, 
the Ibis and the Flamingo. In Ardea it is represented by a groove 
which runs about half-way from the nostril to the anterior end 
of the beak; and in most of the Storks it is represented by a 
line of weakness in the bone reaching about half-way to the tip 
of the beak. This is specially well marked in Z'antalus. 

The nasal processes of the premaxille are so firmly fused with 
the adjacent nasals that their exact outline cannot be seen. It 
is clear, however, that they do not invade the frontal region but 
terminate distad of the fronto-maxillary hinge. This also is the 
ease in Cancroma and Storks, including Z'antalus, but in Ardea 


ANATOMY OF THE SHOE-BILL. 691 


and Scopus the nasals break the transverse line of the hinge, 
fitting into the frontals. 

The anterior tip of the premaxilla is produced in Daleniceps 
to form the strong down-turned hook of the bill. Scopus repeats 
this on a smaller scale ; in Cancroma the sharp point is not bent 
into a hook. In the other Herons and in Storks it is straight. 

Palate—The palate is desmognathous in Salwniceps, the 
maxillo-palatines being fused in the middle line, and the vomer 
represented by a triangular, very thin, ossification, the knife-like 
base of which divides the internal nares when seen from below. 
The condition in Scopus is almost identical, but at its proximal 
end, where it touches the central lamine of the palatine, the 
rather larger vomer shows a broader edge with the faintest 
suspicion of doubling. In the Storks, the vomer is relatively 
smaller even than in Laleniceps and there is no trace of forking. 
In Cancroma and Ardea the vomer is relatively very much larger 
and its edge is quite distinctly cleft between the palatines, each 
blade being attached to the palatine lamina contiguous with it. 
Tn this respect Baleniceps and the Herons are at the opposite 
ends of the series. With regard to the palatines, the most 
striking feature in Laleniceps, fully described by Parker, is the 
coalescence of the internal laminz to form a strong keel stretch- 
ing back from the posterior nares to the pterygoid articulations. 
Allowing for differences in shape and proportion, the similarity 
with Scopus is close. The median keel is still more strongly 
marked in the Pelican and in Plotus; it is represented in Storks 
by a delicate median ridge, but in Cancroma and Ardea the 
internal lamine of the palatines remain completely separate. 

Pterygoids.—I notice no significant. differences between the 
pterygoids of Daleniceps and those of Scopus, Storks and Herons, 
Basipterygoid articular processes are absent in all, and I have 
not found even any rudimentary trace such as is common in the 
Pelican. The ventral distal end of each pterygoid is smoothly 
rounded in Laleniceps and Ardea; in Cancroma, Scopus and 
most of the Storks it shows a sharp keel running out as if to 
meet the outer lamina of the palatal. 

Quadrate.—This is substantially alike in Balewniceps. Scopus, 
Herons and Storks, but the orbital process in Balwniceps is almost 
triangular, the blunted apex projecting into the orbital cavity. 
In Scopus the orbital process is rather blunter ; in Zantalus move 
acute, but in Storks generally it tends to expand to a spatulate 
end, and in the Herons, including Cancroma, the apex is much 
expanded. 

Quadratojugal bar.—This is enormously stout in Baleniceps, 
and the separate portions of which it is composed cannot be dis- 
tinguished. In all the other birds I am considering, it forms a 
slender, much elongated rod. 

Temporal cavity.—The boundaries of the temporal cavity 
present interesting modifications in baleniceps and its aliles. 


Proc. Zoot. Soc.—1913, No. XLVI. 46 


692 DR. P. CHALMERS MITCHELL ON THE 


Unfortunately, owing to the very complete union of the bones 
concerned, these cannot be interpreted completely without the 
examination of very young skulls, but comparison of adult skulls 
shows a good deal worth noting. In Baleniceps the size of the 
brain is small in proportion to the size of the bird, with the 
result that the skull is very short between the orbit and the 
posterior end, Attachment for the powerful temporal muscle is 
increased by the strength and size of the postfrontal process 
(Pl. TOOL, fig. 1), which depends as a triangle of bone behind 
the orbit, with the external face strongly ridged. The blunted 
apex of the triangle reaches nearly half-way down to the quadrato- 
jugal bar and is continued to a strong tubercle on the latter by a 
fibrous band. The central portion ‘of the band bas a separate 
ossification, and it would not be surprising to find, in an old bird, 
that calcification of the fibrous band had joined this central ossi- 
fication with the postfrontal above and the jugal below, to form 
a complete beam of bone separating the orbital and temporal 
cavities externally. Owing to the complete fusion of the bones, 
it is impossible to be certain as to the exact composition of the 
postfrontal. A more primitive skull like that of Dromews (in 
which the proportion of the temporal region to the rest of the 
skull closely resembles that in Laleniceps) shows that the ali- 
sphenoid contributes the main portion of the postfrontal, and 
that the frontal grows down over it only about half-way. In 
Baleniceps it appears as if the frontal covered the alisphenoid 
right down to the lower end of the postfrontal, and on the 
poster ior face the alisphenoid may itself be covered by a process 
of the squamosal. On the other side of the temporal cavity in 
many birds the squamosal sends forwards and downwards from 
just over the quadrate articulation a stout beam of bone p1o- 
jecting tow ards the point of the postfrontal. In LBaleniceps this 
is represented only by a narrow edge pr ‘ojecting over the quadrate 
articulation. Scopus (Pl. TOOL, fe, 2) has like Baleniceps also 
a small brain and narrow temporal space. The postfrontal has 
almost exactly the relations of that of Laleniceps, but it does not 
reach nearly so far towards the jugal. The spur of the squamosal 
is minute, and allowing for differences in the strength of the 
muscular attachments, this region is almost the same in Scopus 
and Baleniceps. 

In a large Stork like Yenorhynchus (Pl. LXXXI. fig. 3) there 
is an arrangement strikingly different in appearance but which, 
none the lass, can be interpreted easily. The brain is still small 
and the temporal cavity narrow. The postfrontal triangle narrows 
very rapidly and is continued downwards as a slender bar which 
stops short long before the jugal is reached. Clcse serutiny seems 
to show that “the frontal contributes a superficial splint-lke 
factor, running down almost to the tip on the anterior face, and 
that the squamosal forms the greater portion of the lower and 
posterior part, but how much Saye alisphenoid contributes it is 
impossible to say. The spur of the squamcsal from over the 


ANATOMY OF THE SHOE-BILL, 693 


quadrate articulation is enormous and forms a powerful process 
which runs downwards and forwards to meet and fuse with the 
tip of the postfrontal. Ina smaller skull, like that of the Dissura 
(Pl. LXXXIT. fig. 1), the arrangement is practically identical, 
but the postfrontal, although it meets the squamosal spur, is 
much more slender, and the frontal factor does not appear to 
reach more than half-way down, 

In Tantalus (Pl. LXXXII. fig. 2) the brain is relatively 
shghtly longer and larger. ‘The squamosal spur is exactly as in 
NXenorhynchus and Dissura, but two things have happened to 
the postfrontal. In the first place it is much shorter, and does 
not reach the squamosal spur, In the second place the triangular 
base is very much wider and is deeply notched in front, with the 
result that it has an anterior and smaller portion corresponding 
exactly to the anterior margin in all the other birds I have been 
describing here, but certainly with no squamosal factor, and a 
longer portion running down towards the point of the squamosal 
spur, corresponding with the posterior part of the postfrontal 
in other birds and certainly consisting chiefly of alisphenoid and 
squamosal factors. 

The condition in Cancroma (Pl. LX XXII. fig. 3) can now be 
followed easily. The brain is still larger relatively ; the squa- 
mosal spur is reduced, and the separation between the two parts 
of the postfrontal, only just apparent in Zantalus, is well marked. 
The purely frontal, anterior portion is the stouter of the two. 
In Ardea (Pl. LX XXIII. fig. 1), where again the brain is still 
larger, the squamosal spur is relatively rather small, but the 
separation between the two parts of the postfrontal is very 
wide indeed. 

As this matter appears to be of some interest, and as I 
have not found it discussed, I shall continue the description 
outside the immediate relatives of Baleniceps. The Pelican 
(Pl. LX XXIII. fig. 2) shows a further extension of the series. 
The squamosal spur is as in Ardea, but the two portions of the 
post-frontal are even further separated, and the posterior of the 
two is reduced to a mere tubercle, intermediate in position be- 
tween the squamosal spur and what would normally be taken to 
be the postfrontal. Im Plotus (Pl. LX XXIII. fig. 3), which has 
a very large brain indeed, the squamosal spur is small, there is a 
mere stump to represent the posterior portion of the postorbital 
process and this is actually nearer the squamosal spur than 
it is to the anterior representative of the postfrontal. 

I do not suggest that the series, as I have arranged it, is 
phylogenetic, but it is a striking example of the differences that 
identical morphological material may exhibit in allied birds, and 
a warning against the hasty drawing of conclusions as to sys- 
tematic position from the comparison of one or two presumably 
allied birds. So far as this point goes, Baleniceps and Scopus 
stand together as birds with small brains, with the squamosal 
spur slight and the postfrontal process simple. Storks form a 

46* 


694 DR. P. CHALMERS MITCHELL ON THE 


second group also with small brains, with the squamosal spur 
very highly developed, frequently reaching the postfrontal, and 
with the latter simple, but in Zantalus showing the beginning 
of cleavage. Cancroma and the other Herons form a third 
group, characterized by larger brains, with the squamosal spur 
reduced as in the first group but with the postfrontal split into 
two separate processes of which the anterior tends to become the 
more important. 

Mandible.—Vhe fusion of the component parts is so complete 
that I could not see any trace of sutures. The most notable 
feature is the absence of the projecting spur of the angle, an 
absence which Baleniceps shares with Scopus, all the Storks and 
Cancroma, in all of which the end of the mandible behind the 
articular cavities for the quadrate is as if abruptly sawn off, 
while in Ardea and typical Herons it is produced backwards as 
a long rounded spur. 

Vertebral column.—As Parker (33) and Gadow (16, p. 76) have 
pointed out, there are 17 cervical vertebre in Baleniceps, 16 in 
Scopus, 17 or 18 in Storks, and 18 to 20 in Herons. The carotid 
canal is complete in all this group of birds. The individual 
vertebree are very much shorter antero-posteriorly in Laleniceps 
than in the Herons and Scopus; the Storks are intermediate 
between Laleniceps and Herons in this respect, which no doubt 
is purely adaptive, but it suggests at least that Baleniceps is 
not very closely related with Herons. Parker states that there 
are no foramina for the vertebral arteries on the sides of 
the atlas in Baleniceps and in its allies. They are certainly 
absent in Baleniceps, but present, although small and limited to 
the anterior lateral part of the atlas, in Ardea, Cancroma and 
Scopus, and present and large in Storks. 

The thoracic vertebre have no hemapophyses in Baleniceps, 
Cancroma, Ardea and Xenorhynchus, but there is a very small 
unpaired process in Tantalus. 

Sternum.—The posterior lateral processes are very long in 
Baleniceps, projecting well behind the metasternum ; in Scopus, 
Herons and Storks, they are short, not projecting behind the 
metasternum. ‘The notch separating the posterior lateral process 
from the metasternum is rounded in Saleniceps, Scopus and 
Storks, angular in Ardea and Cancroma. Parker figured a small 
posterior intermediate process, and compared it with that of the 
Ibis, but it was completely absent in the skeleton I examined, as 
well as in Herons, Storks and Scopus. 

The internal spine of the sternum 1s absent in Baleniceps as in 
Scopus, Storks and Herons, but the external spine is also absent 
in Baleniceps, small in Scopus, small or absent in Storks (best 
developed in Vantalus), large and prominent in Herons including 
Cancroma. 

Furcula.—In Baleniceps the clavicles are joined to form a 
very short-stemmed ‘“‘ Y,” the stem of the Y as well as the distal 
ends of the diverging arms being strongly anchylosed to the 


ANATOMY OF THE SHOE-BILL. 695 


projecting anterior end of the keel of the sternum. There is no 
trace of a median process opposite the hypocleideum between the 
arms of the furcula. Anchylosis with the keel may be regarded 
as an adaptive character. It is as complete in the Pelican as in 
Baleniceps: it exists 1 some of the large Storks, but in Storks 
and Herons generally the connection is by ligament. In Scopus 
the furcula is far short of reaching the keel. In Scopus, Storks 
and Herons including Cancroma, the furcula is more ‘‘U ”-shaped; 
the median forward process between the arms of the “U” is 
absent in Scopus and Storks as in Baleniceps, but is well-marked 
in Herons. 

The proximal end of each clavicle where it reaches the coracoid 
has a very strong flange (text-fig. 132, A, Cl.a.) which articulates 


Text-fig. 132. 


B 


Shoulder-girdle Articulation in Baleniceps and Scopus. 


A. Baleniceps. B. Scopus. 


Ac. Acrocoracoid process of Coracoid. 
Cor. Coracoid. 

Pe. Procoracoid process of Coracoid. 
Cl. Clavicle: 

Cl.a. Acrocoracoid process of Clavicle. 
Sc. Scapula. 


' 


directly with the anterior border of the coracoid. A similar 
flange is present in Scopus (text-fig. 132, B, Cl.a.), and in its place 
there is a minute flat articular facet in Zantalus, but it is absent: 
in Storks generally and in Herons including Cancroma, the 
clavicle being attached to the acrocoracoid only by strong liga- 
ments. The development of this flange in Baleniceps is a very 
strongly marked character, but too much weight cannot be laid 
on this similarity with Scopus, for the acrocoracoid flange of the 
clavicle is equally well marked in the Pelican, in Plotus, in 
Cormorants and Gannets, and in Birds-of-Prey. 


696 DR. P, CHALMERS MITCHELL ON THE 


The epicleideal plane of the clavicle passes across the median 
face of the acrocoracoid and nearly reaches the scapula, with 
which, however, it is united only by ligament (text-fig. 132, A, CZ.). 
Its tip, however, does not extend so far as to cross the acro- 
coracoid,fand it is therefore wholly concealed when it is viewed 
from the outer side, the opposite aspect from that shown in text- 
figure 132. In Scopus the tip of the clavicle projects slightly 
further. In all the Storks and Herons that I have examined the 
tip of the clavicle projects completely across the acrocoracoid so 
that its hgamentous connection with the end of the scapula is 
freely visible when the shoulder-girdle is viewed from the outside. 

Coracoid.—The basal ends of the coracoid, where they articu- 
late with the sternum, do not quite meet in Baleniceps ; they 
either do not meet, or just meet in Storks ; in Scopus and Herons 
they overlap. 

The procoracoid process (text-fig. 132, A, Pe.) is very large in 
Baleniceps, with a curved border nearly reaching an emar- 
gination in the clavicle, with which, however, it is united only 
by ligament. This process varies in Storks, being almost as 
large but not expanded in some, small in others (e.g. Tantalus). 
It is quite small in Scopus (text-fig. 132, B, Pc.) and Herons 
including Cancroma. 

Pelvis.—Parker lays some stress on the narrowness of the 
pelvis in Baleniceps, comparing it in this respect with Herons 
and contrasting it with Storks. Certainly such a difference does 
exist, but as it was not so striking to my eye as Parker found it,: 
JT measured a number of pelves of Storks and Herons, reduced 
the measurements of the narrowest and widest portions of each 
to a common standard and averaged the results, 


Width of Pelvis in percentage of length. 


Narrowest part. Widest part. 


Baleniceps...... wis 20 50 
PAROLE) Viera: = eyes Ne 25 50 
Cancroma ......... 28 57 
NXenorhynchus ... 35 60 
Tantalus ......... 3D 56 
ISCOPUS relents sees oF 62 


It will be seen that the width at the widest part differs much 
less than the width at the narrowest part. In other respects I 
find that the pelvis of Baleniceps is rather more like that of 
Scopus and Storks than of Herons. There is a well-marked 
notch separating the distal ends of the ilium and ischium in 
Baleniceps, Scopus, and Storks which is absent in Herons, in- 
cluding Cancroma. In Herons, including Cancroma, the dorso- 
lateral edge of the post-acetabular ilium projects outwards as a 
sharp horizontal ridge, which is very slightly marked in Scopus 
and is absent in Baleniceps and Storks. 


ANATOMY OF THE SHOE-BILL. 697 


Humerus.—A good deal of attention has been paid to the 
anterior surface of the proximal end of the humerus in birds, and 
I have compared the conditions presented by Baleniceps, Scoprus, 
Storks and Herons. Saleniceps is markedly different in this 
respect from the others and resembles much more closely the 
structure shown by the same bone of the Pelican, which has 
been adequately figured by Pyeraft (35, fig. 6, p. 90). The 
expanded end of the humerus is very flat and rather symmetrical. 
The coraco-humeral groove is extremely faint, rather better 
marked than in the Pelican but entirely different from the deep 
transverse groove shown by Scopus, Storks and Herons. The 
evista inferior is hardly raised above the surface; only the 
faintest distal groove separates it from the shaft of the bone. 
In the Pelican it is much more salient; Pycraft’s figure rather 
under-represents its prominence. In Herons the crista is rather 
like that of Laleniceps, but more prominent; in Scopus it is still 
more prominent, and in Storks, including Z'amwtalus, it is even 
more prominent. ‘The pectoral crest is still longer and better 
marked in Laleniceps than in the Pelican, but is generally 
similar, and in both the oval impression for the insertion of the 
second division of the pectoral is very clear. There is no trace 
of thisin the Herons. In the Storks, including Zantalus, it is 
well-marked. This region of the humerus in Scopus is charac- 
terized by the enormous development of the pectoral crest, which 
is very much larger than in any other of the birds with which I 
am dealing here, and there is a faint oval impression which 
certainly seems to imply the existence of a divided pectoral in 
that bird, as in Storks and baleniceps (see supra, p. 676). 

Hand.—The hand of birds is usually described as containing 
the representatives of three digits, of which that corresponding 
with the index finger (on the usually accepted supposition that 
the pollex is present) is the largest. The proximal phalanx of 
this digit has usually a broad flange on the ulnar side on which 
one or more of the primary quills rest. In dissecting this region 
in Baleniceps | was struck by the way in which the arrangement 
of muscles suggested that this phalanx represented the phalanges 
of two adjacent digits united by a narrow sheet of bone. On 
examining the skeleton in Laleniceps itself, Scopus, Ardea and 
a number of Storks, the suggestion is still more striking. The 
broadened phalanx 1s obviously thickened along the ulnar and 
radial borders, and if these borders were the phalanges of two 
digits united by an ossified sheet of fibre the structure would be 
intelligible. The point requires investigation, both by com- 
parative anatomy and embryology, and I mention it here, only 
to direct attention to it, but I shall be surprised if it does not 
turn out that the hands of these and many other birds show four 
not three digits. 

Tibial bridge.—The bony canal for the tendon of the extensor 
muscle of the digits is complete in Laleniceps as in Scopus, Storks 
and Herons. 


698 DR. P. CHALMERS MITCHELL ON THE 


Tarsal bridges.—The hypotarsus is complex in Baleniceps, 
Scopus and Herons; that is to say the proximal end of the shaft 
is provided with two bony bridges through which the flexor 
tendons pass; in Storks these bridges are absent, and the tendons 
lie in a groove. 


SUMMARY or OsTEoLoGICAL Nores. 


There is a strong general resemblance between Baleniceps, on 
the one hand, and Scopus, Herons and Storks on the other. 
Baleniceps stands alone in its general proportions, in the struc- 
ture and relationship of the lacrymal, quadratojugal bar, atlas, 
posterior lateral process of the sternum, and the humerus. 

Baleniceps and Scopus agree, opposed to the others, in the 
impervious nostrils, the union of the inner plates of the palatines, 
she processes bounding the temporal cavity, the acrocoracoid 
articulation of the clavicle, and the shortness of the tip of the 
clavicle. 

Baleniceps, Scopus and Storks agree, opposed to the others, in 
the basi-temporal plate, the complete interorbital septum, the 
very small vomer, the shape of the posterior notch of the ster- 
num, the absence of a spina interna, and of a median process 
between the diverging arms of the furcula, and the presence of 
a notch separating the posterior extremities of the ischium and 
lium. 

Baleniceps and Storks agree, opposed to the others, in the 
large procoracoid, and the smooth edge of the post-acetabular 
ium. Of the Storks, Z’antalus shows the closest agreement with 
Baleniceps in the occipital condyle, the paroccipital processes 
and the basisphenoid. 

Laleniceps and Herons and Scopus agree, to the exclusion of 
the others, in the proportions of the pelvis, and the complex 
hypotarsus. 

T have been unable to find any osteological points in which 
Baleniceps agrees with Herons to the exclusion of the others. 


Sysvematic Posrrion or BAL-ENICEPS. 


John Gould (22) named and diagnosed this bird in 1851. He 
relied on external characters and compared it carefully with 
Pelecanus, Grus, Ardea and Cancroma. He came to the con- 
clusion that it was the “Grallatorial type of the Pelecanide.” 
He rejected alliance with Ardea and Cancroma apparently because 
in the latter the nail of the central toe is pectinated, a character 
which he thought to be wholly absent in Baleniceps. I have 
shown (supra, p. 648) that much reliance cannot be placed on 
this character. Undoubtedly there are many marked anatomical 
characters common to baleniceps and the Pelican. I may 
mention the presence of a pyloric chamber of the stomach ; 
the division of the great pectoral muscle; the condition of the 


ANATOMY OF THE SHOE-BILL, 699 
tendons of the wing; the absence of intrinsic muscles in the 
syrinx ; the long lacrymals; the fusion of the internal lamine 
of the palatines; the shell-like paroccipital processes ; the anchy- 
losis of the clavicle to the keel of the sternum; the shape 
of the head of the humerus. But these can probably be best 
explained either as convergent modifications in birds which, 
after all, are not very far apart im the system, or the common 
inheritance of Steganopods and their immediate alhes. Sub- 
sequent writers on Baleniceps appear to have been dominated 
by the wish to prove Gould incorrect. J. Reinhardt (37) in 
1860 came to the conclusion that the nearest ally of balew- 
niceps was Scopus, and that Scopus and Baleniceps together 
were nearer the Storks than the Herons. He appreciated that 
the pterylosis, especially of the neck, was similar in Laleniceps, 
Scopus and Storks and markedly differed from that of Cancroma 
and the other Herons. He attached importance to the pectina- 
tion on the claw of Cancroma and the Herons: noted that 
there was an approach to it in Scopus, but did not recognize 
that it also occurred in Laleniceps. He noted that the inter- 
orbital septum was complete in Scopus and Storks, incomplete in 
Cancroma and Herons, but had not information on that point 
in the case of Baleniceps. He compared the bills in considerable 
detail, and pointed out essential points in which baleniceps and 
Scopus agreed, and differed from Cancroma. Parker (33) read his 
great paper before Bartlett’s contribution, but as the latter was 
actually published first, it is more correct chronologically to take 
Bartlett (1) first, although, curiously enough, the authors quote 
one another, and each relies on the other’s opinion. Bartlett in 
1861 made the interesting discovery that Laleniceps had a pair 
of large powder-down patches on the back, and this for him 
settled the Ardeine aftinities of the bird. He included, however, 
Hurypyga amongst the Herons, and I have shewn (supra p. 645) 
that in the present state of our knowledge, the presence of 
powder-down patches is no conclusive evidence as to affinity. 

T cannot understand why it has been assumed and stated by 
writers who had an opportunity of reading the memoir, that 
W. K. Parker’s osteological examination proved LDaleniceps to be 
a Heron and net a Stork. In the Introduction to his memoir, he 
made the following general statement :—‘‘ It is to the stilted, 
wading group of scavengers that Baleniceps belongs, being one 
of the Ardew affines, and therefore intimately related to the 
White Stork, the Marabout, and the Adjutant. Its nearest 
relations, however, are the South American Boat-bill (Cancroma 
cochlearia) and the Little South African Umbre (Scopus wmbretta).” 
In a footnote to his paper he refers to Bartlett’s discovery as 
“having proved beyond al! dispute, that the Balaniceps, like the 
Boat-bill, is essentially a Heron.” But these are ex cathedra 
statements. He also gives a list of what he regards as ‘“‘ Ardeine 
genera ”;—including Ciconia, Leptoptilus, Mycteria, Anastomus, 
Aramus, Ardea, Botaurus, Herodias, Nycticorax, Scopus, 


700 DR. P. CHALMERS MITCHELL ON THE 


Cancroma and Baleniceps. He selected the Heron as the type- 
form as ‘it is best known, and has the characters of the family 
moderately, but markedly developed.” In his detailed examin- 
ation, he is quite as concerned to prove that Laleniceps is like 
Scopus or like a Stork as like a Heron. He came to the very 
definite conclusion that Baleniceps was not a Pelican, but was 
Axdeine, in his broad sense of the word. Quite certainly he did 
not prove it to be a Heron as opposed to a Stork; and in this 
I entirely agree, after having gone through his own observations 
with the skeletons before me, and having made some further 
comparisons myself. 

C. G. Giebel (21) in 1873 re-opened the matter, chiefly after a 
full examination of the pterylosis and external characters. He 
corrected Gould and Reinhardt (not quoting the latter, however) 
as to the pectination, describing it carefully and showing that it 
was present in aleniceps 11 much the same form as in Scopus. 
He showed also that these birds agreed in pterylosis, where 
they differed from Cancroma, and came to the conclusion that 
Baleniceps was much more closely allied with Scopus than with 
Cancrona. 

¥. KE. Beddard in a special memoir and in his treatise on 
the anatomy of birds (3 and 4) assumed that Parker’s osteo- 
logical researches had proved Baleniceps to be a Heron, not 
a Stork, overlooked the evidence supplied by Reinhardt and 
Giebel as to the Stork-like pterylosis of Baleniceps and Scopus, 
and although he referred to Giebel’s paper, did not note that 
Giebel showed Baleniceps and Scopus to be alike in the peetin- 
ation of the claw. He placed Baleniceps definitely among the 
Herons as opposed to the Storks, relying apparently chiefly 
on the syrinx. I have already stated (supra p. 651) that his 
argument cannot support his conclusion. Possibly it may show 
that Baleniceps is not a Stork; it does not show that it is 
a Heron. 

H. Gadow (16) placed both Baleniceps and Scopus under the 
Ardez as opposed to the Ciconie, but placed Baleniceps merely 
as a sub-family of the Herons, distinguished from the true 
Herons by the absence of pectination and various minor characters. 
He was misled, I think, by the literature, and does not claim to 
have made independent observations. 

Those who have followed the old and new facts regarding 
Baleniceps that I have been able to bring together must be 
impressed by the number of characters in which Scopus and 
Baleniceps agree, and by the much greater number of points in 
which these two birds agree with Storks than with Herons. If 
we adopt the method made familiar by many of those who have 
written most copiously on the anatomy of birds, take characters 
on their face value, and regard those birds as most nearly allied 
which have the greatest number of characters in common, then 
the sub-order Arde will contain the single family Ardeide, and 
the adjacent sub-erder Ciconie will comprise the families Scopide, 


ANATOMY OF THE SHOE-BILL. 701 


Balenicepidide, Ciconiide and Ibidide. But if we desire that 
classification should represent phylogeny, we must await further 
knowledge as to the value of the various characters which 
anatomists have tried to use. It is no use bringing pectination 
or powder-down patches, or a completely feathered neck into an 
argument between Storks and Herons, if these characters have 
been independently acquired or independently lost in the case of 
many different groups. It is no use for systematic purposes to call 
a particular form of syrinx Ardeine if that be a common type in 
many different kinds of birds, or to call a doubled pectoral muscle 
Ciconine if that be shared by many Steganopods. The fact is 
that Steganopods, Scopus, Baleniceps, Storks and Herons have a 
large common heritage, consisting partly of actual common 
structures (some of which they share with a very much larger 
assemblage of birds) and partly of the capacity to be modified 
in certain definite directions. ‘The characters latent and patent 
composing this common heritage are distributed irregularly 
amongst them, partly for reasons that we do not know and partly 
in response to similar habits. Until the meaning and history 
of each set of characters have been worked out very fully we have 
less than no idea as to their real value in indicating affinity. 
The only set of characters on which I have sufficient knowledge to 
have any confidence is shown in the disposition of the alimentary 
tract. In that respect Baleniceps seems to me to have passed 
through the condition common to Storks and Herons, and to 
display specialization in the same direction as the Herons. But 
until we have further knowledge, not so much of Baleniceps in 
particular, but of the value of anatomical characters, the safe 
course is to regard Laleniceps as the representative of a group of 
equal value with Storks and Herons. 

In my opinion, however, the relation of Storks and Herons 
to the Steganopods requires revision. Pending this, I must add 
that John Gould’s description of Laleniceps as the “ Grallatorial 
type of the Pelecanide ” is at least as happy as the more confident 
statements of later writers. 


List of References. 


1. Barruert, A. D., On the Affinities of Baleniceps rex. P.Z.S. 
ISOM JO. Ika sk 


. BepparD, F. E., Anatomy of Scopus umbretta. P.Z.S. 1884, 
p- 943. 
: a Visceral Anatomy of Baleniceps rex. P.Z.S. 


1888, p. 284. 
The Structure and Classification of Birds. 
London, 1898. 
3 99 On the Alimentary Tract of Certain Birds. 
JP fais HOTT Fos 24 
‘ CAZIN, Maurice, L’Appareil eastrique des Oiseaux. Ann. 
Sci. Nat. iv. p. 177 (1887). 


Oa oa - wo WD 


SI 
Lo 


CO COS 


10. 
nts 
12. 
13. 
14. 


15. 


16. 
17. 
18. 
atts), 
20. 
21. 
22. 
23. 


24. 
25. 
26. 
27. 
28. 


29. 


DR. P. CHALMERS MITCHELL ON THE 


. Davies, H. R., Pectination. Nature, 1891, p. 367. 
. Evans, A. H., Birds, in the Cambridge Natural History. 


London, 1889. 


. Forses, W. A., On the Bursa Fabricii in Birds. P.Z.S. 


1877, p. 304. 


5 On the Pterylosis of Mesites. P. Z.S. 1882, 
footnote p. 269. 

53 On some Points in the Anatomy of the 
Indian Darter. P.Z.S. 1882, p. 208. 

a The Last Journal of W. A. Forbes. Ibis, 
1883, p. 494. 


Fursprincer, M., Water ‘suchungen zur Morphologie und Sys- 
tematik der Vogel. Amsterdam, 1888. 

Gapvow, H., Versuch einer vergleich. Anatomie des Ver- 
dauungssystemes der Vogel. Jen. Zeitsch. 
Xl. po. 91S 79): 

55 Remarks on the Cloaca and on the Copulatory 
Organs of the Amniota. Phil. Trans. 
elxxvul. p. 5 (1888). 
Vogel in Bronn’s Thier-reich (1891). 

Garrop, A, H., On certain Muscles of the Thigh of Birds, 
and on their Value in Classification. P. Z. S. 
1873, p. 626. 


4 On the Anatomy of Chaunaderbiana. P.Z.S. 
1876, p.189: 

a On the Anatomy of Plotus anhinga. P.Z.S. 
IRSA, JO 31330). 

4, On the Anatomy of Levaillant’s Darter. 


123 Agiss UiSiish, Os O08); 

GieBEL, C.G., Baleniceps rew. Zeits. fur Gesam. Naturw. 
vol. xli. p. 350 (1873). 

GouLp, Joun, On a new and most remarkable Form in 
Ornithology. P.Z.S. 1851, p. 1. 

Levcxart, F. S., Ueber eine zusammengesetztere Magenbild- 
ung bei verschiedenen Vogeln. Zoolog. 
Bruchstiicke, ii. p. 64 (1841). 

Mircuet, P. Cuatmers, On the perforated Flexor Muscles 
in some Birds. P.Z.S. 1894, p. 495. 


a On the Proventricular Crypts of Pseudo- 
tantalus ibis. P.Z.S. 1895, p. 271. 

5 On the Intestinal Tract of Birds. P.Z.S. 
1896, p. 136. 

a Anatomy of the MHoatzin (Opisthocomus 
cristatus). P.Z.S. 1896, p. 618. 

up Quintocubitalism in the Wing of Birds. 
Journ. Linn. Soc., Zool. vol. xxvii. p. 210 
(1899). 

os Anatomy of Kingfishers. The Ibis, 1901, 


Da oie 


30. 
31. 
32. 
33. 
34. 


35. 
36. 


37. 
38. 


ANATOMY OF THE SHOE-BILL. 703 


MircuHett, P. CHatmers, On the Intestinal Tract of Birds. 
Trans. Linn. Soe., Zool. viii. p. 173 (1901). 


5 Anatomy of Gruiform Birds. P.Z.S. 1901, 
p. 629. 

f Anatomy of Limicoline Birds. P.Z.8. 1905, 
p. 155. 


Parker, W. K., The Osteology of Baleniceps rex. Trans. 
Z.8. iv. p. 269 (1860). 

PETHERICK, JOHN, Memoranda on the Hippopotamus and 
Baleniceps recently imported to England. 
P.Z.8. 1860, p. 195. 

Pycrart, W. P., Contributions to the Osteology of Birds. 
Part I. Steganopodes. P.Z.S. 1898, p. 82. 


a Aquintocubitalism in the Bird’s Wing. 
Journ. Linn. Soc., Zool. vol. xxvu. p. 236 
(1899). 
REINHARDT, J., On the Affinities of Baleniceps. P.Z.8. 1860, 
p. 377. 


Wetpon, W. F. R., The Anatomy of Phwiicopterus. P.Z.S. 
1883, p. 638. 


EXPLANATION OF THE PLATES, 
Prats LXXX. 


Baleniceps rex. From a living young male. 
Photo by D. Seth-Smith, F.Z.S. 


Prare LXXXI. 


Fig. 1. Side of Skull in Baleniceps. 
2. = 5 Scopus. 
3. ss if NXenorhynchus. 


Pratt LXXXII. 


Fig. 1. Side of Skull in Dissura. 
a 35 Tantalus. 
3. 5 3 Cancroma. 


Pirate LXXXIII. 
Fig. 1. Side of Skull in Ardea. 


5 pH Pelecanus. 


a 5 i Plotus. 


704 PROF. A. DENDY AND MR. R. W. H. ROW ON 


44, The Classification and Phylogeny of the Calcareous 
Sponges, with a Reference List of all the described 
Species, systematically arranged. By ArrHur Denpy, 
D.Sc., F.R.S., F.Z.8., Professor of Zoology in the 
University of London, and R. W. Haroip Row, B.8c., 
F.L.S., Assistant Lecturer and Demonstrator in Zoology 
at King’s College. 


[Received May 30, 1913: Read June 8, 1913. | 
(Text-figure 133.) 


“ Le groupe ne se définira plus par la possession de certains charactéres, 
mais par sa tendance a les accentuer.”— Bergson. 


INDEX. 

Page 
JES faeh OND} ORGHTNICOhc He oHeeReatemnerctsonciosucdacodana sea seboanouseapas echO! 
PRINCIPLES OF CLASSIFICATION ........2..-c00-020.008--. 09 
SysTEMATIC ARRANGEMENT OF THE CALCAREA ...... 715 
Family 1. Homoceelide, fam. nov. ..........6....00-.. 716 
Baas ICUCASCLO MCR ee eee re ee Ce RIL. 
oy Ob LECTORINGIEIES NENING WOW noo sonsanccassea0a60s 736 
» 4. Minchinellide, fam. nov. .................. 739 
» 9. Murrayonide, fam. NOV. ......-0...ccees 741 
st Ie ORRASITRECLAAOLIZ AN ob Gane Semede dablouneataudebeaoosad socauain (Le 
Bn. Us Sbeterapidd enact Neben cocoa Reeser 750 
pe USI CU AiO HOU eRe speacn ade obsbae ana dab ban soo reebe dor: 1 ZOE. 
Ge ZU UNOPISCIUE soonscaeasn ba xcesno spaa0acoo0en “hail 
eel OMMelapeidcen Tam Mover seec reer ao 
Genera and Species “‘ Incertze Sedis” ............00.605 786 
List of rejected Generic Names ..................0..... 786 
PEVOGENY OF DHE CATCAR HAS ay.t cen seecceseecheaceennn: Lod 
Bir mnssry (OYE UND) 2 O'd ones Alaa aaah arp Ronan eu bod aeacadcaa oso ndodstiae, i 1s01) 

INTRODUCTION. 


The first Caleareous sponges, ‘‘ Spongia ciliata” and “ Spongia 
compressa,’ were described in 1780 by Fabricius, but it was 
not until much later that the essential differences between the 
Calcareous and Non-calcareous Sponges were recognised. Fleming, 
in 1828, however, proposed the genus Grantia for the former 
group, including in it all the forms whose skeleton consisted of 
ealeium carbonate. (Risso’s earlier genus, Sycon, and Gray’s 
Seypha were diagnosed differently.) 


CALCAREOUS SPONGES. 705 


The next important advance in the history of the group was 
the erection of the genus Leucosolenia by Bowerbank in 1866, 
for certain sponges which we now include in the Homoccelidee, in 
addition to other genera no longer employed. From that time 
forward numerous investigators studied the group, and many 
new species and several new genera were described, but no really 
serious attempt to deal with the question of the classification of 
these sponges was made until the time of Haeckel, who in 1870 
published his ‘ Prodromus,’ and in 1872 his famous Monograph 
of the group, with extremely detailed, though somewhat un- 
successful descriptions of all the then known species, including 
many which he described for the first time. Haeckel’s so-called 
“natural ? system, with its three families of Ascones, Leucones, 
and Sycones, based upon the type of canal system, and its twenty- 
one genera based upon the types of spicules present, is so well 
known, and has been so often criticised, that it needs no further 
description by us, especially as it proved extremely artificial, and 
expressed only to a very limited extent the true phylogeny of 
the group. 

The scheme proposed by Poléjaeff in 1883 was considerably 
more successful, and his primary division of the group into 
Homoca@na and Hererocata has been made the basis of almost 
every Classification since proposed. Weare now beginning to 
realise, however, that this division also is of a very arbitrary 
character. 

The next scheme of classification we need notice is that of 
Vosmaer, in Bronn’s ‘ Klassen und Ordnungen des Thierreichs ” 
[1887], which is almost identical with that of Poléjaeff, with 
the addition of the Pharetronide as a fourth family of the 
HETEROCGLA. 

In 1891 von Lendenfeld proposed a modification of Haeckel’s 
system, erecting a fourth family, the Sylleibide, intermediate in 
canal-system between the Leucones and Sycones, and reducing 
the number of genera in each family to two, according to the 
presence or absence of oxea. This was undoubtedly a con- 
siderable improvement upon Haeckel’s system, but again it failed 
to interpret the interrelationships of the members of the group 
correctly, and it has since been almost entirely abandoned, though 
certain spongologists, notably Breitfuss, retained it with but 
little modification for a considerable time. 

During the years 1891-1893 there was published by Dendy 
[1891 A, 1892 B, 1893 A| a scheme of classification based on 
almost wholly different lines. Retaining Poléjaefi’s Orders 
Homocana and Hererocana, and, like that author, including in 
a single genus, Leucosolenia, all the species of Homocaa, he 
divided the Hrrrrocaia into five families, whose differentiating 
characters were based far more on the structure and arrangement. 
of the skeleton than on the canal system; and although this 
system has not been accepted by all writers, yet we ourselves feel 
that it embodies a more natural arrangement of the group than 


706 PROF. A. DENDY AND MR. R. W. H. ROW ON 


any of its predecessors, and we have made it the basis of the classi- 
fication here proposed. 

In 1896 Minchin published a paper entitled “Suggestions for 
a Natural Classification of the Asconide,” which may fairly be 
said to mark a new departure in the taxonomic study of the 
Calearea, in that it introduces for the first time the idea of the 
position of the nucleus in the collared cell as a character of 
taxonomic importance, a character which has since proved, 
our opinion, to be of great value. We have not been able, how- 
ever, to follow Minchin completely in those modifications of 
Dendy’s classification of the group which he proposed, partly in 
the paper referred to, and partly in his well-known article in 
Lankester’s ‘ Text-book of Zoology ’ [1900]. 

In 1898 Bidder, in a paper on “The Skeleton and Classification 
of Caleareous Sponges,” proposed to carry out Minchin’s ideas 
with regard to the nucleus of the collared cell to their logical 
conelusion, and to divide the Calearea into two great groups 
acecordingly—CaLcaRoNEA and Caucingea. Although not actually 
adopting this division, which we consider to be somewhat pre- 
mature in the present state of our knowledge, we have ourselves 
followed much the same line of cleavage. Although he accepts to 
a large extent, with regard to his families, the system proposed 
by Dendy [1892 B], Bidder makes certain rearrangements which 
do not appear to be altogether satisfactory. He does good 
service, however, in indicating for the first time the relation- 
ship of Carter’s Clathrina tripodifera, for which he proposes 
the genus Dendya, to Haeckel’s Leucaltis clathria (= Hetero- 
pegma nodus-gordii Poléjaeft). In the same paper he discusses 
the position of the crystalline optic axis of the radiate spicule 
systems, and endeavours to assign taxonomic value to this 
character also, but whatever may be the theoretical value of his 
conclusions, which have since been accepted by Minchin [1909], 
we cannot consider that such a character is of any practical use 
to the systematist. 

In 1908 Jenkin erected two new families, the Chiphoride and 
Staurorrhaphide, supposed to be differentiated by the presence of 
what he considered to be a new type of spicule, the “ chiactine,” 
from all previously recognised families. The peculiarity of these 
spicules was believed to consist in the orientation of the various 
rays both in relation to one another and to the other parts of the 

skeleton, and a special method of development was suggested for 
them. Finally, in 1909, one of us (Row) still further elaborated 
the ‘* Chiact Theor y, as it was called, and proposed yet another 
family, the Grantillide, in which more primitive, but similar, 
‘‘nrochiacts ” were supposed to be present, and which was made 
by him the starting-point from which the Heteropiide were 
supposed to have been derived. As we shall show later, however, 
we do not now think that the spicules in question are more than 
very slight modifications of ordinary ty pes, ¢ and we have abandoned 
all three families. 


CALCAREOUS SPONGES. 707 


It is now more than forty years since any attempt was made 
at a complete revision of the Calcarea, and in the interval 
the number of known species has increased from 111, described 
in Haeckel’s ‘Kalkschwimme,’ to 436 recognised by us at 
the present time. Having recently been engaged in examining 
collections of Calearea of considerable importance and extent, 
we have had impressed upon us the necessity for a complete 
systematic catalogue of all the known species, the descriptions 
of which are scattered through an immense number of separate 
memoirs. In order to supply this want we have had to go 
through practically the whole of the literature of the group, 
and there is not a paper, to the best of our belief, which contains 
a reference to a new species which we have not seen. It is with 
considerable satisfaction that we find, as a result of this work, 
that the scheme of classification proposed by one of us twenty 
years ago [Dendy, 1892 B] is still applicable in its essential 
respects, and requires comparatively little revision in order to 
bring it up to date. 

In the present memoir we propose to give diagnoses of all the 
families and genera employed, and to enumerate all the known 
species under the genera to which we assign them. We have 
decided to confine ourselves to the consideration of living forms, 
since our knowledge of fossil sponges is at present so unsatis- 
factory, and the number of described species so great. 

We have had access, during the course of our work, to a large 
amount of material, comprising a large proportion of the known 
species. This has consisted chiefly of the collections in the 
Natural History Department of the British Museum, a large 
collection brought by one of us from Australia (see Dendy 
[1891 A and 1892 B]); the collections made by Mr. Cyril Cross- 
land in the Red Sea (see Row [1909]), Zanzibar (see Jenkin 
[1908 A]), and Cape Verde (see Thacker [1908]); a collection 
made by the ‘Sealark’ Expedition in the Indian Ocean (see 
Dendy [1913]); and the magnificent collection made by the 
Hamburg South-Western Australian Expedition in 1905 (see 
Row P1913 MS.]). In addition, one of us (Row) visited Berlin 
and Jena in 1912, and Sotned valuable information from the 
study of type specimens at those places. 


With regard to the difficult question of synonymy, and the 
limits which should be assigned to each species, we have, never- 
theless, been obliged to rely in most cases upon the published 
descriptions and determinations of the authors responsible, and 
we have unfortunately found that these descriptions are fre- 
quently very inadequate; while even where they are more com- 
plete, the fact that they have been prepared from the point of 
view of a different scheme of classification has sometimes made 
them difficult to use. We have, however, assumed that all 
descriptions are correct, except in cases where they have been 
shown by subsequent authors, from an investigation of type 


Proc. Zoou. Soc.—1913, No. XLVII. AT 


708 PROF. A. DENDY AND MR. R. W. H. ROW ON 


specimens, to be erroneous. Further, we have felt that we could 
not, merely on the basis of these published descriptions, under- 
take the responsibility of determining whether or not a species 
described as new by its author should more properly have been 
allocated to a previously known species. Thus we have proceeded 
on the principle that all species described as new. must be con- 
sidered to be distinct, unless they have been proved to belong to 
a previously known species by the subsequent reinvestigation of 
type specimens or otherwise. We have only given such synonyms 
and references as seem absolutely necessary, but where one species 
has been merged in another by any authority, and this result 
accepted by us, we give the synonym and the appropriate 
authority under the species in which it is now placed. 

It is thus highly probable that, of the 486 species which we 
enumerate, a considerable proportion will ultimately prove to be 
identical with one another. This question, however, can only be 
decided by a very thorough study of the specific characters and 
range of variation in each case, probably necessitating in many 
cases a reinvestigation and comparison of the original types. 
Professor Minchin, in his paper on ‘“'The Characters and 
Synonymy of the British Species of Sponges of the genus Lewco- 
solenia” [1905], has set a good example of the manner in which 
this critical revision of the group ought to be carried out. 

We have, as a generai rule, taken no notice of varieties as 
distinct from species, but certain of Haeckel’s so-called “ Specific 
Varieties,’ to which he has already given distinctive names, 
appear to us, after careful consideration of his descriptions, to 
deserve to rank as separate species. 

In the preparation of this paper we have throughout borne in 
mind the requirements of the systematist, and it is hoped that 
its publication will greatly assist the determination and arrange- 
ment of species in the future. We have had to set aside a great 
number of published generic names as synonyms, but it is hoped 
that the appended list will enable the student to trace them in 
the present system. 

We have indicated in the list of species under each genus that 
species which we recognise as the type, our method of procedure 
being to take, in the case of old genera, that species, of those 
which we assign to the genus, to which the name of the genus 
was first applied, while in the case of new genera we have 
chosen as our type the species which seems to exemplify best 
the special characters on which we have founded the genus. 

As the publication committee of the Zoological Society has 
decided against the use of brackets around the names of authors 
of species in all cases, it must be understood that the authors 
cited are responsible for the specific but not necessarily for the 
generic names employed by us. 


_ CALCAREOUS SPONGES. 709 


PRINCIPLES OF CLASSIFICATION. 


It appears to us that the chief point to be borne in mind im 
attempting to arrive at any natural system of classification is the 
importance of utilising as many characters as possible. A classifi- 
cation based upon a small number of characters must necessarily 
be arbitrary and artificial, and characters which are of great 
importance in some cases may be of comparatively little use in 
others. here can be no doubt that there are certain significant 
characters which do indicate genetic relationships, but these 
characters are by no means the same in all cases, and they have 
to be carefully sought for and distinguished amidst a host of less 
important features. Sometimes it is the canal system that affords 
the best clue, sometimes the arrangement of the skeleton, some- 
times the form of the spicules, and sometimes even the position of 
the nucleus of the collared cells. There must also be remembered 
the undoubted fact that the phenomenon of convergence has 
played a large part in the evolution of the Calcarea, and has led 
in many cases to totally deceptive resemblances, as, for example, 
between the genera Leucetta and Leucandra. 

It seems likely, however, that the collared cell, or choanocyte, 
which itself is by far the most characteristic histological con- 
stituent of the sponge organisation, may ultimately prove, as 
suggested by Bidder [1898], to afford a means of dividing the 
whole of the Calcarea into two main branches, one having the 
nuclei of these cells placed basally, and the other having them 
apical. Indeed, the acceptance of this principle, if only in a 
tentative manner, constitutes the chief difference between our 
present views on the subject and those which we previously held ; 
but in the present state of our knowledge it is a principle which 
must not be pushed too far, and we have only been able to make 
use of it as subsidiary to more easily determined characters. The 
acceptance of this principle, however, necessitates the wide sepa- 
ration of the Lelapiide from the other Pharetronid sponges with 
which they have hitherto been associated. 

We shall discuss this question in some detail later on, but it 
may serve a useful purpose if we give at once a list of all the 
species of Calcarea in which we have been able to determine the 
position of the nucleus, and the results of our determination. 
This list includes no less than 75 species, as follows :— 


Nuclei apical. Nuclei basal. 


Family Homoca@Lipa. 


Leucosolenia bella Row. Leucosolenia falcata Haeckel. 

Leucosolenia complicata Montagu, Leucosolenia stolonifer Dendy. 
de Minchin. Leucosolenia ventricosa Carter. 

Leucosolenia lucasi Dendy. Leucosolenia gardinert Dendy. 


47° 


710 PROF, A. DENDY AND MR. R. W. H. ROW ON 


Nuclei apical. 


Nuclei basal. 


Family Homoca@1i1p&% (continued). 


Leucosolenia variabilis Haeckel, Leucosolenia coriacea Montagu, 


fide Minchin. 


fide Minchin. 
Leucosolenia depressa Dendy. 
Leucosolenia cavata Carter. 
Leucosolenia pelliculata Dendy. 
Leucosolenia proxima Dendy. 
Leucosolenia pulcherrimaDendy. 
Leucosolenia vitrea Row. 
Dendya tripodifera Carter. 
Ascute uteoides Dendy. 


Family Levucascip#&. 


Leucascus simplex Dendy. 
Leucascus insignis Row. 
Leucascus clavatus Dendy. 
Leucetta chagosensis Dendy. 
Leucetta expansa Row. 
Leucetta microraphis Haeckel. 
Leucetta prolifera Carter. 
Leucetta pyriformis Dendy. 
Pericharax heteroraphis Polé- 
jaeff. 
Pericharax pexiza Dendy. 


Family Levcautip&. 


Leucaltis clathria Haeckel. 
Leucettusa dictyogaster Row. 


Family MINcHINELLID2. 


Minchinella lamellosa Kirk- 
patrick. 


Family MurRAYoNIDs. 


Sycon boomerang Dendy. 


Sycon cartert Dendy. 


Sycon gelatinosum de Blainville. 
Sycon gigantewm Dendy. 


Sycon lendenfeldi Row. 


Sycon ramsayt von Lendenfeld. 
Sycon raphanus O. Schmidt. 
Sycon setosum O. Schmidt. 


Sycon verum Row. 


Murrayona phanolepis Kirk- 
patrick. 


Family Sycerrips. 


CALCAREOUS SPONGES. 711 


Nuclei apical. Nuclei basal. 
Family HErerRopiips, 


Grantessa erinaceus Carter. 
Grantessa hastifera Row. 
Grantessa hispida Carter. 
Grantessa poculum Poléjaeff. 
Grantessa polyperistomia Carter. 
Grantessa sacca von Lendenfeld. 
Grantessa intusarticulata Carter. 
Heteropia glomerosa Bowerbank. 
Heteropia simplex Row. 
Vosmaeropsis dendyt Row. 
Vosmaeropsis depressa Dendy. 
Vosmaeropsis macera Dendy. 
Vosmaeropsis primitiva Row. 
Vosmaeropsis wilsont Dendy. 


Family GRANTUD. 


Grantia compressa Fabricius. 
Grantia genuina Row. 

Grantia vosmaeri Dendy. 
Grantia indica Dendy. 
Teichonopsis labyrinthica Carter. 
Grantiopsis infrequens Carter. 
Ute syconoides Dendy. 

Ute spiculosa Dendy. 

Synute pulchella Dendy. 
Leucandra hispida Carter. 
Leucandra australiensis Carter. 
Leucandra echinata Schuffner. 
Leucandra meandrina von Lendenfeld, 
Leucandra minima Row. 
Leucandra phillipensis Dendy. 
Leucandra thulakomorpha Row. 
Aphroceras cataphracta Haeckel. 


Family AMPHORISCIDA. 


Amphoriscus oblatus Row. 
Leucilla australiensis Carter. 
Leucilla princeps Row. 


Family LELAPIID#. 
Lelapia australis Gray. 


With the exception of this important feature, which in the 
main harmonises very well with our previous conclusions, the 
principles that we have followed in arriving at the classification 
set forth in the present paper are almost exactly those which 


712 PROF. A. DENDY AND MR. R. W. H. ROW ON 


were expounded by one of us some twenty years ago | Dendy 
1891 A, 1893 A], as the following review of our present position 
will show. 

The canal system, including the form of the flagellate chambers, 
is, we are convinced, of comparatively little taxonomic value in the 
higher Calcarea. In the lower forms it necessarily determines the 
arrangement of the skeleton, which must he in the walls of the 
ascon tubes, however these may be arranged, and in the Sycet- 
tide the arrangement of the radial tubes has undoubtedly been 
the determining factor in the development of the articulate tubar 
skeleton. With the appearance of a definite dermal cortex, how- 
ever, the arrangement of the skeleton begins to vary more or less 
independently of the canal system, so that with an identical 
eanal system we find such different types of skeleton as that 
of the Grantiidee, the Heteropiude and the Amphoriscide. In 
each of these families, while the type of skeleton remains fairly 
constant, the canal system ranges from syconoid to leuconoid, or 
at least sylleibid. The syconoid type is again met with on a 
totally different line of descent in the homocel genus Dendya, 
and also in Lewcaltis, and again gives rise to a leuconoid type 
both in the Leucascide and the Leucaltide. 

Our view that it is the canal system rather than the skeleton 
that has repeatedly undergone convergent evolution is strongly 
supported by the distribution of the different types of spicules 
and of the two types of collared cells. The form of the spicule, 
however, must be used with great caution as a guide to genetic 
relationships, for it is largely a question of adaptation. The tri- 
radiate is undoubtedly the fundamental spicule form in the group, 
but one might almost say that it tends to become quadriradiate 
on the slightest provocation. Thus we almost invariably find 
quadriradiates in the gastral cortex, whose inwardly directed 
apical rays are undoubtedly of great value as a protection against 
enemies, such as small crustaceans, approaching through the 
osculum. Then, again, the ordinary triradiates of the dermal 
cortex not infrequently develop a more or less conspicuous, 
centripetally directed, apical ray ; and this latter tendency appears 
to have led, in the case of the Amphoriscide, to a constant 
skeletal character which forms the most characteristic feature of 
the group. Similarly with regard to the distinction between 
equiangular and sagittal triradiates, we find that the latter can 
always be developed, when the situation in the sponge demands 
this form, by the bending back of the oral rays during growth. 
This nearly always takes place, for example, in the oscular collar, 
where there is no room for the oral rays to extend forward at the 
asual angle. 

Whether or not there is a fundamental difference between an 
equiangular triradiate, however its rays may be bent, and an 
alate one in which the primitive oral angle is really different 
from the paired angles, and in which there is a correspond- 
ing difference in the position of the crystalline optic axis, as 


CALCAREOUS SPONGES. 713 


maintained by Bidder and Minchin, is another and much more 
difficult question to decide. The presence of regular equiangular 
triradiates seems most certainly to be very characteristic of the 
Leucascid-Leuealtid line of descent, and we have made use of it 
as one of the distinguishing features of the members of those 
groups. It is, however, extremely difficult in practice to distin- 
guish between a sagittal spicule which owes its sagittal character 
merely to the backward bending of the oral rays, and one which 
is sagittal owing to a real inequality between the primitive 
angles. 

There can be no question that a superficially sagittal condition 
may be attained in different ways, and one of the most interesting 
results at which we have arrived in the preparation of the present 
paper is that the so-called subdermal sagittal (pseudosagittal) 
spicules of the Heteropiide have a quite different origin from 
the ordinary sagittal form, the basal ray not being homologous 
in the two cases. 

With regard to Jenkin’s [1908 B] supposed families Chiphorid 
and Staurorrhaphide, we have come to the conclusion that these 
are based upon purely imaginary distinctions. It will be re- 
membered that Jenkin maintained that in these families a special 
type of spicule, the ‘“chiactine,” constitutes the first (or only) 
joint of the tubar skeleton. It seems highly improbable, from 
purely @ priori reasons, that this joint should be differently con- 
stituted in different syconoid sponges. As a matter of fact, no 
one, so far as we are aware, has demonstrated how it arises in 
ordinary cases, such as Sycon or Grantia, but everybody has been 
content to speak of it as being composed of subgastral sagittal 
triradiates. It is, moreover, well known that these triradiates 
may develop an apical ray, as they do in many species (e. g. 
Sycon ensiferum, Sycon verum and Grantia indica), and thus 
become quadriradiates. Jenkin thought that he had demon- 
strated that in certain cases the subgastral quadriradiates arise 
by rotation and re-orientation of the basal rays of gastral quadri- 
radiates, owing to the pressure of the developing chambers, and 
regarded this mode of origin as distinguishing his so-called 
chiactines from other subgastral tri- and quadriradiates. As, 
however, he made no attempt to show how the subgastral tri- 
radiates amd quadriradiates arise in other cases, this distinction 
cannot be regarded as having any value; and we ourselves are of 
opinion, from our own observations, that the spicules which con- 
stitute the first joint of the tubar skeleton probably arise in the 
same way in all cases, although we are not disposed to accept 
without further evidence the exact mode of origin described by 
Jenkin. 

It is certain that, in some cases at any rate, the spicules at the 
growing margin of the osculum have not yet assumed their defi- 
nitive or jentacian but exhibit a confused arrangement. Some of 
them gradually become oriented as dermal cortical spicules, with 
their three rays lying parallel to the surface, others as subgastral 


714 PROF. A. DENDY AND MR. R. W. H. ROW ON 


spicules, with their basal rays centrifugally directed and their oral 
rays lying in the deeper part of the gastral cortex. The spicules 
of the other joints of the tubar skeleton, on the other hand, 
appear to arise in the walls of the chambers themselves, and this 
difference in mode of origin probably accounts for the difference 
which undoubtedly exists between them and the subgastral 
spicules. 

We consider that the abolition of the families Chiphoride and 
Staurorrhaphide will effect a much needed simplification in the 
classification of the group, and also that it is highly desirable to 
do away with them from the point of view of practical con- 
venience, for to draw a real distinction in practice between a 
so-called chiactine and an ordinary subgastral sagittal quadyri- 
radiate is quite impossible. The bending of the apical ray, 
whereby it is brought to he nearly or quite in the same straight 
line as the basal ray, is merely a question of degree, as may be 
seen from the examination of the apical rays of ordinary gastral 
quadriradiates in various species. 

The family Grantillide, proposed by one of us [ Row, 1909] 
bas, of course, also been abandoned by us. The rejection of 
Jenkin’s chiact theory and the fact that we attach little import- 
ance to the mere presence of subdermal quadriradiates, have 
removed both the characters on which the family was founded, 
and it has consequently been merged in the Heteropiide. 

The presence, however, of subgastral sagittal spicules (tri- 
radiates or quadriradiates) appears to be very characteristic of 
the Sycettid as contrasted with the Leueascid-Leucaltid line of 
descent. 


The distribution of oxea in the Calcareous sponges presents an 
extremely difficult problem, as species possessing them occur side 
by side with species that lack them in almost all the large genera 
throughout the group. Asa result we have found it impossible 
to assign to this character any such important place in our scheme 
of classification as previous authors have suggested, though as a 
matter of practical convenience we have used it asa basis for 
arranging the species of a genus in sections. Certainly the ability 
of some sponges to produce oxea may be looked upon as differ- 
entiating them, at any rate to some extent, from others which 
either have lost this power, although descended from oxea-bearing 
ancestors, or else have never possessed it. 

Further, we have found that two types of oxeote spicules 
can be distinguished—the comparatively large, usually radially 
arranged form, and a much smaller for which we employ the term 
‘microxea.’ In typical cases the latter are less than 0-1 mm. in 
length, and they are usually of a very definite hastate shape, 
with an enlargement at a short distance from the distal extremity. 
They thus form very characteristic and well-defined skeletal 
elements, and it is remarkable to find them recurring in so many 
perfectly distinct genera, belonging to most of the families within 


CALCAREOUS SPONGES. 715 


the group. It would obviously be impossible, in any natural 
systematic arrangement, to associate together all the species 
which possess microxea, for these species differ amongst them- 
selves in nearly every other respect. It is noteworthy that in 
some cases these spicules occur in very small numbers, so that 
they might easily be overlooked, while in others they are ex- 
tremely numerous; and it seems by no means impossible that 
some individuals of a species may possess them while others do 
not. Nevertheless, as a matter of convenience, we have decided 
to make use of the presence or absence of these spicules for the 
purpose of distinguishing sections of genera. 

In this connection, however, it must be observed that we do not 
include, in our conception of the term microxea, those long, hair- 
like spicules frequently found surrounding the osculum, or some- 
times echinating the surface. These we believe to be merely 
slightly modified or imperfectly developed large oxea, and we 
include them under that head, under the term ‘ trichoxea.’ 

One is tempted to explain the sporadic distribution of oxea by 
speculations which, in the present state of our knowledge, are 
perhaps unjustifiable; but we may perhaps venture to suggest 
that the presence of oxea constituted a characteristic feature of 
some remote ancestor, and that the faculty of producing them has 
never been entirely lost, but requires special genetic conditions 
of which we know nothing before it can become active in any 
particular species. It is quite possible that our sections are 
somewhat artificial, but a grouping of the species by easily 
recognisable characters, especially in the larger genera, can hardly 
fail to be of use to the systematist. 


Further discussion of the principles of classification may con- 
veniently be left until we come to deal with the various sub- 
divisions of the group. 


SYSTEMATIC ARRANGEMENT OF THE CALCAREA. 


Class and Order CALCAREA. 


Diagnosis. Sponges in which the spicules are composed of car- 
bonate of lime (calcite), and consist of either triradiate or 
quadriradiate systems, or are oxea (monaxons). 


For many years past it has been the almost universal practice 
amongst spongologists to divide the class CALCAREA into two 
sharply contrasted orders, viz. Homoca@ia, in which the whole 
of the gastral cavity is lined by collared cells, and Hererocana, 
in which the collared cells are confined to special flagellate 
chambers, a practice which was first initiated by Poléjaeff in 


716 PROF. A. DENDY AND MR. R. W. H. ROW ON 


1883. It appears to us that the time has come, owing to our 
greatly increased knowledge of the group, to abandon this primary 
division, and we now propose to consider the class as consisting 
of a single order only. 

One reason for taking this step lies in the discovery of various 
intermediate forms. Such are the species of Dendya, which in 
the radial arrangement of the ascon tubes approach closely to the 
Leueascid type of Hrrrrocana, and the species of Lewcascus itself, 
which are but slightly modified from homoccel ancestors and 
form the starting-point of a distinct evolutionary series within 
the group Herrrroca:ta; while von Lendenfeld’s Homoderma 
sycandra is obviously merely a Sycon in which the collared cells 
persist in the central gastral cavity throughout the life of the 
sponge. 

Another even more important reason lies in the fact that, as 
noted above, the group Hrrmroca:ta is at least diphyletic in 
origin, the genus Dendya and the family Sycettide forming two 
distinct starting points from which the evolution of the higher 
Leuconoid forms has proceeded. 

We therefore propose to divide the class CALCAREA straightway 
into families, of which the first will be the Homoceelide, practi- 
cally co-extensive with the Homocaa of Poléjaeff. His group 
Hererocana, on the other hand, is here definitely abandoned, 
being represented by the families Leucascidee, Leucaltide, Min- 
chinellide, Murrayonide, Sycettide, Heteropiide, Grantiide, 
Amphoriscidee and Lelapiide. 


Family 1. HOMOCCELID AE nov. 


Diagnosis. The whole of the gastral cavity and its various out- 
growths lined by collared cells throughout the life of the 
sponge. Sponge colony rarely radiate, and, if so, the central 
individual retains the primitive ascon structure, with a 
lining of collared cells and without a special gastral cortex. 
No true dermal membrane or true dermal cortex is ever 
developed. 


In 1872 Haeckel proposed seven genera of ‘‘ Ascon” Calcarea, 
based upon the permutations and combinations of triradiate, 
quadriradiate, and oxeote spicules. ‘This constituted his so-called 
“natural” system, but he also had an “ artificial” system based 
upon the type of colony formation. Both systems have shown 
themselves far from satisfactory in practice and have long since 
been abandoned. 

In 1883 Poléjaeff, recognising the extreme difficulty of sub- 
dividing the group, placed the whole of the species in the genus 
Leucosolenia of Bowerbank, which takes priority over all Haeckel’s 
genera and is the only genus recognised by Poléjaeff in his order 
Homocaa. 


CALCAREOUS SPONGES. lua 


In 1891 von Lendenfeld proposed a modification of Haeckel’s 
“natural” system, retaining only two genera, Ascetta and 
Ascandra, the former genus Jacking oxea, ane lnuien possessing 
them. 

In the same year Dendy, in his “‘ Monograph of the Victorian 
Calcarea Homoceela,” while accepting Poléjaeff’s conclusion that 
only a single genus could be recognised, proposed to divide that 
genus into sections and subsections, according to the type of 
colony formation and canal system. Three sections were re- 
cognised, Simplicia, Reticulata and Radiata, and the Leticulata 
were further subdivided into /ndivisa and Subdivisa, according to 
the absence or presence of an endogastric network. Of these 
sections the Radiata now constitute the genus Dendya of Bidder, 
while the other two are of little value to the systematist. 

In 1896 Minchin proposed to distinguish three genera of 
Homocasa, Clathrina, Leucosolenia and Ascandra, and in 1900, 
in Lankester’s ‘Text- Book of Zoology,’ he recognised two distinct 
families, Clathrinidze and Leucosoleniide, and gave (p. 110) the 
following classification and diagnosis :— 


“Grave A. HOMOC(@LA, Pol., s. Ascones, H. 


‘“‘Gastral layer continuous. 


“ Waminy |. Cruararinip&®, Minchin. Form reticulate. Tyri- 
radiate systems always present, equiangular; monaxons present 
or absent. Collar-cells with nucleus at base. Larva a parenchy- 
mula, Genera—Clathrina, Gray (= Ascetta, H., pars, Ascaltis, H., 
pars, etc., and Leucascus, D.); Figs. 2, 6, 7, 8; Ascandra, H., 
emend. (= Homandra, Liaf., for Ascandra falcata, H.); Dendya, 
Bidder, for Clathrina tripodifera, Crtr. Faminry 2. Lruco- 
SOLENUDM®, Minchin. Form erect; monaxons always present ; 
triradiates, 1f present, alate; collar-cells with nucleus apical ; 
larva an amphiblastula. Genera—Ascyssa, H.; Leucosolenia, 
Bwk. (= Ascandra, H., pars, etc.) ; Figs. 3, 4, 5.” 

In 1909, Zool. Anzeiger, xxxv. p. 230, in response to criticisms 
by Hammer [1908] and Dendy as to ane position of the nucleus 
in the collared cells, he emended his diagnoses as follows :— 


“Class CALCAREA. Sponges with the skeleton composed of 
calcite, in the form of spicules either monaxon, triradiate or 
quadriradiate in form. 

“Grade 1. Homocana. Calcarea with the gastral layer of collar- 
cells continuous, not forming separate flagellated chambers. 

“Family 1. Crarurinip&. Oscular tubes generally short, 
arising as shallow vents from the network of tubes, form of the 
body typically reticulate.  Triradiate spicules always present, 
equiangular, and with the crystalline optic axis vertical to the 
facial plane of the rays; monaxon spicules present or absent. 
Collar-cells with the flagellum arising quite independently of the 


718 PROF. A. DENDY AND MR. R. W. H. ROW ON 


nucleus, which is spherical in form, and situated at the base of 
the cell. Larva a parenchymula. 

“ Family 2. LeucosoLentip®. Oscular tubes long, arising as 
distinct individuals from the stolon-like system of basal tubes ; 
form of the body erect. Monaxon spicules always present ; tri- 
radiates, if present, typically bilateral in form, with two paired, 
and one unpaired angles, and with the crystalline optic axis never 
vertical, but always inclined, to the facial plane of the rays. 
Collar-cells with the flagellum arising directly from the pear- 
shaped nucleus, which is situated at, or near, the apex of the cell. 
Larva an amphiblastula.” 


Without entering into a long discussion as to the theoretical 
value of these diagnoses, we may point out that in some respects 
they are in actual practice very difficult of application. Only 
very few of the numerous described species of homoccel sponges 
have been examined with reference to the mode of origin of the 
flagellum in the collared cells, the nature of the larva, or the 
direction of the crystalline optic axis in relation to the facial 
plane of the spicule. If it were necessary to investigate these 
very obscure characters in every case, the classification of the 
group would indeed make slow progress. 

The more obvious characters which Professor Minchin first 
made use of for the subdivision of the group, viz., the equiangular 
or alate character of the triradiates and the position of the nucleus 
of the collared cell, together with the erect or reticulate form of 
the colony, lose their value when we extend our investigations 
beyond the familiar British species. The Australian species, 
Leucosolenia lucasi, L. stolonifer and Ascute uteoides all have 
the characteristic non-reticulate, ‘‘ Leucosolenia” form, and all 
possess oxea (monaxons) ; L. stolonifer and A. uteoides, however, 
have collared cells with basally placed nuclei, while in LZ. lucast 
the nuclei are apical, though unfortunately the position of the 
basal granule is—as is always the case in specimens preserved 
without very special precautions—-indeterminable. In L. stolonifer 
and ZL. lucast, again, some at any rate of the triradiates are 
apparently equiangular and indistinguishable from clathrinid 
spicules. The test concerning the direction of the optic axis is 
far too difficult to apply accurately to be of any general value. 

As to the larvee, again, not only are these rarely met with in 
the Homoccelids, but Professor Minchin himself has shown that 
there is a transition from the one type of larva (parenchymula) 
to the other type (amphiblastula). He says’ (Lankester’s ‘ Text- 
Book of Zoology,’ part i. p. 75), “The type of parenchymula 
larva exemplified by Clathrina reticulum (Fig. 59, 7), affords an 
easy transition to the so-called amphiblastula found in Leuco- 
solentide, and in the great majority of the Hrrrrocana.” 

The genus Clathrina of Gray [1867] was originally based on 
the reticulate form of the sponge colony, and this is still almost 
the only character which could be made use of in practice as a 


CALCAREOUS SPONGES. 719 


distinction from Leucosolenia, but here again we know that no 
sharp separation can be drawn between the two types of external 
form, for one and the same colony may be reticulate in its lower, 
and non-reticulate in its upper portion. 

In short, we do not think that any spongologist who has 
examined a large and representative collection of Homoccelidze 
would be prepared to maintain Professor Minchin’s subdivision 
of the group into Clathrinidee and Leucosoleniidee. It appears to 
us that the most that can be done at present is to pick out and 
diagnose in a more or less satisfactory way certain more or less 
isolated and well-characterised generic forms, and to leave the 
vast bulk of the species in the genus Leucosolenia. 

At the same time we are of opinion, as already pointed out in 
the introduction, that Professor Minchin has indicated some 
characters at any rate which will in the future prove to be of 
very great value for taxonomic purposes, and we ourselves have 
made extensive use of the position of the nucleus in the collared 
cells, as roughly determined from the spirit material, in support 
of our views as to the evolution of the heteroccel Calcarea. The 
reason why this character appears to be of less value amongst the 
Homoceelidee will be discussed in the section dealing with the 
phylogeny of the group. 


There can be no doubt that the Homoccelide have all been 
derived from a common Olynthus-like ancestor, from which a 
number of lines of descent have branched out in various directions. 
Colony formation seems to have played the chief part in the 
process of evolution and many different types of colony have thus 
arisen. 

At present we are only able to distinguish four genera in this 
family, Leucosolenia, Ascute, Ascyssa and Dendya. ‘Three of 
these, Ascute, Ascyssa and Dendya, are easily and clearly definable, 
possessing well-marked characters, but the remaining genus, 
Leucosolenia, is distinguished almost entirely by negative cha- 
racters, and contains a very large number of species, presenting 
a very great diversity amongst themselves, both in colony-form 
and spiculation. 


Genus 1. LeucosotentA Bowerbank [1864-1882]. 


Diagnosis. Diverticula of the gastral cavity, if any, never radially 
arranged around a central tube. Skeleton composed of tri- 
radiate or quadriradiate spicules, to which oxea may be 
added. No uteoid dermal skeleton. Nucleus of collared 
cells basal or apical. 


For illustrations of this genus see Dendy [1891 A]. 


The external form in species of this genus ranges from simple 


Olynthus-like individuals, which may be connected together by a 


720 PROF. A. DENDY AND MR. R. W. H. ROW ON 


basal stolon as in ZL. ducasi and L, stolonifer, to complex reticulate 
colonies which have acquired by integration a new individuality 
of a higher type, sometimes with pseudogaster, pseudosculum, 
pseudoderm and pseudopores, as in L. ventricosa, and sometimes 
with reversal of the canal system as in LZ. cavata. It may happen 
that one and the same colony exhibits a reticulate structure in 
one portion, and a non-reticulate in another, as in several forms 
figured by Haeckel [1872]. For further information as to the 
variations in the mode of colony formation the reader is referred 
to Dendy [1891 A]. 

Haeckel’s Ascandra falcata, which we include in the genus 
Leucosolenia, has been made by both Minchin [1896] and von 
Lendenfeld [1891] the type of a special genus (with the generic 
names of Ascandra and Homandra respectively) on account of 
the peculiar appearance of the gastral layer, which is thrown into 
folds in such a way that in transverse section there appear to be 
present a series of shallow radial tubes. These endodermal ridges 
are always supported by the large apical rays of the gastral 
quadriradiates, which have either pushed the layer of collared 
cells out before them as they grow, or formed a foundation upon 
which the collaved cells have spread. It may be pointed out that 
other Homoccelid: also show this type of structure, e.g. LZ. canar- 
vensis and L. gegenbaurt. We ourselves do not consider that this 
character is of sufficient importance, per se, to take generic rank ; 
nor do we consider that there is any relationship between the 
“pseudoradial” character thus given to the gastral layer, and a 
true syconoid canal system, since, in the radial tubes of Sycon, for 
example, both dermal and gastral layers are folded, while in 
Leucosolenia falcata the gastral layer only is affected. Further, 
embryology shows that the radial tubes of Sycon are outgrowths 
from the central gastral cavity, and not formed by ingrowths 
into it. Had the syconoid type of canal system originated from 
some such condition as that of Lewcosolenia (Ascandra) falcata, 
the sycon person would have been provided with a dermal cortex 
ab initio, whereas the more primitive Sycettide have the ends 
of the chambers freely projecting on the surface of the sponge. 

The position of the nucleus of the collared cells in this genus 
is not, so far as we are aware, correlated with any particular 
type of spiculation or canal system, and we cannot, if only for 
practical reasons, make use of this character by itself for sub- 
dividing the genus. Asa means of grouping the various species 
of the genus into sections, however, we may, as in the higher 
forms, adopt the criterion afforded by the presence or absence of 
oxea. 

We recognise the following species as belonging to this 
genus :— 

Section A. Oxea present. 


1. L. ama@porweEs Haeckel. 
Ascandra complicata, var. ameboides Haeckel [1872]. 


CALCAREOUS SPONGES. 721 


2. L. aneuLata von Lendenfeld. 


Ascandra angulata von Lendenfeld [1891]. 


3. L. arAcHNotpES Haeckel. 


10. 


ie 


12. 


13. 


14. 


15, 


16. 


Gel be 


Ascandra variabilis var. arachnoides Haeckel [1872]. 


ARMATA Haeckel. 
Olynthus pocillum Haeckel [1870], fide Haeckel [1872]. 
Asculmis armata Haeckel [1872]. 


The earlier name is a nomen nudum. 


5 by 


5 IU 


a 


ATLANTICA Thacker. 
Leucosolenia atlantica Thacker [1908]. 


BELLA Row, 
Leucosolenia bella Row [1913 MS8.]. 


BOTRYOIDES (Hllis and Solander). Type species of the genus. 
Spongia botryoides Hllis and Solander [1786 }. 
Leucosolenia botryoides Minchin [1905]. 


. Borrys Haeckel. 


Ascandra botrys Haeckel [1872]. 


. CERVICORNIS Haeckel. 


Ascandra variabilis var. cervicornis Haeckel [1872]. 


. CLARKIL Verrill. 


Ascortis clarkit Vervill [187 iI(6 


. COMPLICATA Montagu. 


Spongia complicata Montagu [1812]. 

Grantia botryoides Lieber kithn [1859], fide Haeckel [1872]. 
Olynthus hispidus Haeckel [1859], fide Haeckel [1872]. 
Leucosolenia complicata Minchin [1905]. 


. CONFERVICOLA [aeckel. 


Ascandra variabilis var. confervicola Haeckel | 1872]. 


. contorTA Bowerbank. 


Leucosolenia contorta Bowerbank [1864-1882]. 
Ascandra contorta Haeckel [1872]. 


. CORALLORHIZA Haeckel. 


Sycorhiza corallorhiza Haeckel [1870]. 
Auloplegma haeckeli O. Schmidt MS. a Haeckel [1872]. 
Ascortis corallorhiza Haeckel [1872]. 


. CoRDATA Haeckel. 


Ascandra cordata Haeckel [1872]. 


. DENSA Haeckel. 


Tarrus densus Haeckel [1870]. 
Nardopsis gracilis Haeckel [1870], fide Haeckel 11879}. 
Ascandra densa Haeckel [1872]. 


722 PROF. A. DENDY AND MR. R. W. H. ROW ON 


17. L. piscoveryt Jenkin. 
Leucosolenia discoveryi Jenkin | 1908]. 


18. L. pusra Dendy. 
Leucosolenia dubia Dendy |1891 A}. 


19. L. ecuiata Ark. 
Leucosolenia echinata Kirk | 1893). 


20. L. scurnorEs Haeckel. 
Leucosolenia echinoides Haeckel [1870]. 
Olynthus cyathus Haeckel [1870], fide Haeckel [1872]. 
Ascandra echinoides Haeckel [1872]. 


21. L. eLeanor Urban. 
Leucosolenia eleanor Urban [1905]. 


22. L. FaBRict O. Schmidt. 
Lewcosolenia fabricti O. Schmidt [1870]. 


23. L. ratcata Haeckel. 
Ascandra falcata Haeckel [1872]. 


24, L, rraciiis Haeckel. 
Ascortis fragilis Haeckel [1872]. 
Leucosolenia botryoides James-Clark [1869], fide Haeckel 


[1872]. 
Leucosolenia thamnoides Haeckel [1870], fide Haeckel 
[1872]. 


Haeckel’s earlier name is a nomen nudum. 


25. L. nermest breitfuss. 
Ascandra hexmesi Breitfuss [1896 B]. 


26. L. nispipisstmA Haeckel. 
Ascandra variabilis var. hispidissima Haeckel [1872]. 


27. L. norrwa Haeckel. 
Nardopsis horrida O. Schinidt MS., fide Haeckel [1872]. 
Ascortis horrida Haeckel {1872}. 


28. L. incerta Urban. 
Leucosolenia incerta Urban [ 1908). 


29. L. IRREGULARIS Jenkin. 
Leucosolenia irregularis Jenkin [1908 A }. 


30. L. nacunosa Johnston. 
Grantia lacunosa Bean MS., fide Johnston | 1842). 
Grantia lacunosa Johnston | 1842). 
Ascortis lacunosa Haeckel [1872]. 


31. L. naxa Kirk. 
Leucosolenia lawa Kirk [18995]. 


32. L. 


33. L. 


34. L. 


30. L. 


36. L. 


37. L. 


38. L. 


43. L. 


44. L. 


Proc. 


CALCAREOUS SPONGES. (2433 


LIEBERKUHNI O. Schmidt. 
Grantia botryoides Lieberktihn [1859], fide O. Schmidt 
[1862], and Haeckel [1872]. 
Grantia leberkiihniit O. Schmidt [1862]. 
Leucosolenia robusta Haeckel [1870], fide Haeckel [1872]. 
Ascandra lieberkiihnit Haeckel [1872]. 


Lucas! Dendy. 
Leucosolenia lucasi Dendy [1891 A}. 


MINCHINI Jenkin. 
Leucosolenia minchini Jenkin [1908 B]. 


nitipa Haeckel. 
Olynthium nitidum Haeckel [1870]. 
Olynthium splendidum Haeckel [1870], jide Haeckel 
[1872]. 
Ascandra nitida Haeckel [1872]. 


Panis Haeckel. 
Ascandra panis Haeckel [1872]. 
pinus Haeckel. 
Leucosolenia botryoides Lacaze-Duthiers MS., fide Haeckel 
[1872]. 
Ascandra pinus Haeckel [1872]. 


RETICULATA Haeckel. 
Tarrus reticulatus Haeckel [1870]. 
Ascandra reticulum, var. reticulata Haeckel [1872]. 


. RETICULUM O. Schmidt. 


Nardoa reticulum O. Schmidt [1862]. 
Ascandra reticulum Haeckel [1872]. 


4. SERTULARIA [Haeckel. 


Ascandra sertularia Haeckel | 1872). 


. STOLONIFER Dendy. 


Leucosolenia stolonifer Dendy [1891 A}. 


. TENUIPILOSA Dendy. 


Leucosolenia tenuipilosa Dendy [1905]. 
Leucosolenia canariensis ‘Thacker [1908], pars, fide 
Row [1909]. 


TENUIS Schuffner. 
Ascandra tenuis Schuffner [1877 ]. 


VARIABILIS Haeckel. 
Leucosolenia variabilis Haeckel [1870]. 
Ascandra variabilis Haeckel [1872]. 
Leuconia somesi Bowerbank [1864-1882], fide Minchin 
[1896]. 
Leucosolenia variabilis Minchin [1905]. 
Zoou. Soc.—1913, No. XLVITI. 48 


724 
45. LL. 


46. L. 


48. L. 


49. L. 


50. L. 


+ ie OP 


52. L. 


53. L. 


54, L. 


od. L. 


PROF. A. DENDY AND MR. R. W. H. ROW ON 


VENTRICOSA Carter. 
Clathrina ventricosa Carter [1885-1886 ]. 
Leucosolenia ventricosa Dendy [1891 A]. 


Srcrrion B. Without oxea. 


Acassizit Haeckel. 
Ascaltis lamarchii var. agassizii Haeckel [1872]. 


. BLANCA Michlucho- Maclay. 


Gaancha blanca Michlucho-Maclay [1868]. — 

Olynthus (dc.) guancha Haeckel [1870], fide Haeckel 
[1872]. 

Ascetia blanca Haeckel [1872]. 


CANARIENSIS Michlucho-Maclay. 

Nardouw canariensis Michlucho- Maclay | 1868]. 

Nardoa sulphurea Michlucho-Maclay [1868], fide Haeckel 
[1872]. 

Nardoa rubra Michlucho-Maclay [1868], jide Haeckel 
[1872]. 

Ascaltis canariensis Haeckel [ 1872). 

Leucosolenia nanseni Breitfuss [1896], fide Thacker 
[1908]. 

Ascaltis compacta Schuftner [1877], fide Thacker [1908]. 


Leucosolenia canariensis ‘Thacker [1908]. 


CANCELLATA Verrild. 
Leucosolenia cancellata Verrill [1873]. 


cAROLI Haeckel. 
Ascaltis darwinii, var. caroli Haeckel [1872]. 


CAVATA Carter. 
Clathrina cavata Carter | 1885-1886]. 
Leucosolenia cavata Dendy [1891 A]. 


CEREBRUM Laeckel. 
Ascaltis cerebrum Haeckel | 1872]. 


CHALLENGERI Poléjaeff. 
Leucosolenia challengeri Poléjaeff [1883 ]. 


CHARYBDMA Haeckel. 
Ascaltis gegenbauri var. charybdea Haeckel [1872]. 


CLATHRATA Carter. 
Leucetta clathrata Carter [1883]. 
Clathrina tripodifera var. gravida Carter [1885-1886], 
jide Row [1913 MS. ]. 
Grantia cliftont Bowerbank MS., fide Row [1913 MS.]. 
Leucosolenia intermedia Kirk [1895], fide Row [1913 MS.]. 
Leucosolenia clathrata Row (1913 MS. ]. 


56. 


57. 


58. 


59. 


67. 


68. 


69. 


. GARDINERI Dendy. 


i) | 


CALCAREOUS SPONGES. 72 


. CLATHRUS O. Schmidt. 


Grantia clathrus O. Schmidt [1862]. 

2 Clathrina sulphurea J. Ki. Gray [1867], fide Haeckel 
[1872]. 

Tarrus labyrinthus Haeckel [1870], fide Haeckel [1872]. 

Nardoa labyrinthus O. Schmidt MS., fide Haeckel [1872]. 

Ascetta clathrus Haeckel [1872]. 


. CONVALLARIA Haeckel. 


Ascilla gracilis var. convallaria Haeckel [1872]. 


. CORTACEA Montagu. 


Spongia coriacea Montagu [1812]. 

Grantia multicavata Bean MS., fide Johnston [1842]. 
Olathrina sulphurea Carter [1871 A], fide Haeckel [1872]. 
Ascetta coriacea Haeckel [1872]. 


. DARWINIL Haeckel. 


Leucosolenia darwintt Haeckel [1870]. 
Ascaltis darwintt Haeckel [1872]. 


. DECIPIENS Haeckel. 


Ascaltis cerebrum var. decipiens Haeckel [1872]. 


. DEPRESSA Dendy. 


Leucosolenia depressa Dendy {1891 A}. 


. picrvowEs Haeckel. 


Leucosolenia dictyoides Haeckel [1870]. 
Ascetta primordialis var. dictyoides Haeckel [1872 /. 


. FALKLANDICA breitfuss. 


Leucosolenia falklandica Breitfuss [1898 EK}. 


he 
. FLEXILIS Haeckel. 


Ascettu flewilis Haeckel [1872]. 


Leucosolenia gardinert Dendy [1913]. 


. GEGENBAURI Haeckel. 


Leucosolenia gegenbauri Haeckel [1870]. 
? Nardoa spongiosa Kolliker [1864], fide Haeckel [1872]. 
Ascaltis gegenbaurt Haeckel [1872]. 


. GoETHEL Haeckel. 


Leucosolenia goethet Haeckel [1870]. 
Ascaltis goether Haeckel [1872]. 


. cractuis Haeckel. 


Ascilla gracilis Haeckel [1872]. 


. GRANTIL Haeckel. 


Leucosolenia grantii Haeckel [1870]. 
Ascaltis solanderti (=Ascaltis botryoides var. solanderit) 
Haeckel [1872], fide Haeckel [1872]. 
: AR 


726 


71. 


(DR, 


73. 


74. 


75. 


76. 


17. 


78. 


79. 


80. 


Sill 


82. 


83. 


PROF. A. DENDY AND MR. R. W. H. ROW ON 


. HIMANTIA Johnston. 


Grantia botryoides var. himantia Johnston [1842]. 
Ascetta coriacea var. himantia Haeckel [1872]. 


. JAPONICA Haeckel. 


Ascilla japonica Haeckel [1872]. 


. LAMARCKIL Haeckel. 


Leucosolenia lamarckii Haeckel [1870]. 
Aulorhiza intestinalis Haeckel [1870], fide Haeckel oe 
Ascaltis lamarckii Haeckel [1872]. 


. LocULOSA Haeckel. 


Ascetta primordialis var. loculosa Haeckel [1872]. 


. MACLEAYI von Lendenfeld. 


Ascetta macleayt von Lendenfeld [1885 A]. 


. MINORICENSIS Lackschewitsch. 


Leucosolenia minoricensis Lackschewitsch [1886 ]. 


. MULTIFORMIS Greitfuss. 


Leucosolenia multiformis Breitfuss [1898 B]. 


. osScULUM Carter. 


Clathrina osculum Carter [1885-1886]. 


. PEDUNCULATA von Lendenfeld. 


Leucopsis pedunculata von Lendenfeld [1885 B]. 


. PELLICULATA Dendy. 


Leucosolenia pelliculata Dendy [1891 A]. 


_ PHILLIPINA Haeckel. 


Ascetta blanca var. phillipina Haeckel [1872]. 


. POTERIUM Haeckel. 


Ascetia primordialis var. poterium Haeckel [1872]. 
Ascandra conulata von Lendenfeld MS., jide Breitfuss 
[1897]. 


. PRIMORDIALIS Haeckel. 


Prosycum primordiale Haeckel [1870]. 

Olynthus simplex Haeckel [1870], fide Haeckel [1872]. 

? Grantia pulchra O. Schmidt [1862], fide Haeckel [1872]. 

Nardoa arabica Michlucho-Maclay MS., fide Haeckel 
[1872]. 

Ascetta primordialis Haeckel [1872]. 


. PROTOGENES Haeckel. 


Ascetta primordialis var. protogenes Haeckel [1872]. 
Ascetta procumbens von Lendenfeld [1885 B], fide Dendy 
[1891 Aj, 


' CALCAREOUS SPONGES. TG 


84. L. proxima Dendy. 
Leucosolenia proxima Dendy [1891 A}. 


85. L. PSAMMOPHILA Row. 
Leucosolenia psammophila Row [1913 MS.]. 


86. L. putcHERRIMA Dendy. 
Leucosolenia pulcherrima Dendy [1891 A}. 


87. L. RosEA Kirk. 
Leucosolenia rosea Kirk [1895]. 


88. L. saarrrarta Haechel. 
Ascetia sagittaria Haeckel [1872]. 


89. L. sceprrum Haeckel. 
Ascetta sceptrum Haeckel [1872]. 


90. L. spinosa von Lendenfeld. 
Ascetta spinosa von Lendenfeld [1891]. 


91. L. stiprrata Dendy. 
Leucosolenia stipitata Dendy [1891 A}. 


92. L. vestcuta Haeckel. 
Ascetta vesicula Haeckel [1872]. 


93. L. virrea Row. 
Leucosolenia vitrea Row [1913 MS.]. 


94. L. witsont Dendy. 
Leucosolenia wilsont Dendy [1891 A]. 


The following species are of doubtful value :— 


95. L. LAMINOCLATHRATA Carter. 
Clathrina laminoclathrata Carter [1885-1886]. 


Too imperfectly described to be recognisable. 


96. L. putcHRA O. Schmidt. 
Grantia pulchra O. Schmidt {1865}. 
Possibly identical with LZ. primordialis Haeckel, fide Haeckel 
[1872]. 
97. L. sponeiosa Kolliker. 
Nardoa spongiosa Kolliker [1864]. 
Possibly identical with LZ. gegenbauri Haeckel, fide Haeckel 
[1872]. 
98. L. SULPHUREA Gray. 
Clathrina sulphurea J. Ki. Gray (non Carter) [1867 ]. 
Possibly identical with LZ. clathrus O. Schmidt, fide Haeckel 
[1872]. 


728 PROF. A. DENDY AND MR. R. W. H. ROW ON 


Genus 2. Denpya Bidder [1898]. 


Diagnosis. Sponge colony consisting of a large central individual 
lined by collared cells, from which radially arranged diver- 
ticula are given off. Skeleton composed of equiangular 
triradiates to which quadriradiates may be added. Subgastral. 
sagittal radiates never present. Nuclei of collared cells 
probably always basal. 


For illustrations of this genus see Dendy [1891 A]. 


Carter’s Clathrina tripodifera was included by Dendy [1891 A] 
in the genus Lewcosolenia, as the sole representative of the 
“ Radiate” section of that genus. Bidder [1898] proposed for 
its reception a new genus, Dendya, and the recent discovery by 
the ‘Sealark’ Expedition of a closely allied, but quite distinct, 
species in the Indian Ocean seems to justify the retention of 
Bidder’s genus. Unfortunately Bidder associated his genus with 
Poléjaefi’s Heteropegma (= Leucaltis) in a new family Hetero- 
pegmidx, of which Dendya was made the type genus. While 
admitting a certain degree of relationship between Dendya and 
Leucaltis, we cannot agree that this is so close as to justify 
placing them in the same family, for not only is Lewcaltis 
corticate, while Dendya is non-corticate, but Leucaltis is also 
heteroceel, while Dendya is homoceel. 

The chief interest attaching to the genus Dendya lies in its 
radiate structure, which, at first sight, seems to suggest a possible 
starting point for the Sycettid as well as for the Leucascid- 
Leucaltid line of descent. We no longer consider, however, that 
Dendya stands very near the origin of the Sycettide, from the 
simplest of which it differs widely in the structure of the skeleton, 
especially in the absence of subgastral sagittal radiates, in the 
fact that the radial tubes tend to cinastbonn ase. and in the basal 
position of the nuclei of the collared cells. The tendency of the 
radial tubes to form reticulations is indeed a difficulty in the way 
of separating the genus sharply from ZLeucosolenia, a fact well 
illustrated by Carter’s Leucetta clathrata (= —Leucosolenia clath- 
rata), which is intermediate between the two as regayds the 
canal system while resembling Dendya in the presence of the 
characteristic dermal tripod spicules. [ Row, 1913 MS. ] 


We recognise the following species as belonging to this 
genus :— 


1. D. prouirerA Dendy. 
Dendya prolifera Dendy [1913]. 


2. D. TRIPODIFERA Carter. Type species of the genus. 
Olathrina tripodifera Carter |1885— 1886]. 
Leucosolenia tripodifera Dendy [1891 A}. 
Dendya tripodifera Bidder [1898]. 


CALCAREOUS SPONGES. 729 


Genus 3. ASGUTE nov. 


Diagnosis. The diverticula of the gastral cavity, if any, never 
radially arranged around a central tube. With a uteoid 
dermal skeleton of colossal longitudinal oxea. Nuclei of the 
collared cells (? always) basal. 


As the presence of a uteoid dermal skeleton is considered to 
form a good generic character in other families, we see no reason 
why it should not be used in the same way amongst the Homo- 
ceelidee, and therefore propose this genus for Dendy’s Lewcosolenia 
uteoides, with which Carter’s Aphroceras asconoides is doubtless to 
be associated. The nucleus of the collared cells is basal in the 
former species, but its position is not known in the latter. 

In both the known species of this genus the sponge has the 
form of a group of simple ascon persons, attached to one another 
by their bases, and without any anastomoses in the colony. 


We recognise the following species as belonging to this 
genus :— 
1. A. AsconorbEs Carter. 
Aphroceras asconoides Carter | 1885-1886 }. 


2. A. urEoIDES Dendy. ‘Type species of the genus. 
Leucosolenia uteoides Dendy [1892 C}. 


Genus 4. Ascyssa Haeckel [1872]. 


Diagnosis. Diverticula of the gastral cavity, if any, never radially 
arranged around a central tube. Skeleton consisting entirely 
of oxea. 

For illustrations of this genus see Haeckel [1872]. 

We consider the entire absence of radiate spicules as affording 
sufficient reason for the generic separation of Haeckel’s two 
species of Ascyssa from all the other Homoccelide. 


We assign the following species to this genus :— 


1. A. acurera Haeckel. 
Ascyssa acufera Haeckel [1872]. 


2. A. TRoGLODYTES Haeckel. Type species of the genus. 
Ascyssa troglodytes Haeckel [1872]. 


Family 2. LEUCASCIDAL Dendy [1892 B] (emend.). 


Diagnosis. Sponge typically forming a massive colony, usually 
with several or many oscula, but sometimes integrated into 
a single individual with definite external form. Without 
any large central gastral cavity lined by collared cells, but 
with an exhalant canal system devoid of collared cells. 


730 PROF. A. DENDY AND MR. R. W. H. ROW ON 


Flagellate chambers ranging from long and possibly branched, 
with a tendency to radial arrangement round the exhalant 
canals, to small, approximately spherical, and scattered. 
With a distinct and independent dermal membrane (or 
cortex) pierced by true dermal pores. Skeleton consisting 
mainly of equiangular and equiradiate spicules, which may 
become sagittal at the oscular margins. Radiates of the 
chamber! ayer without definite arrangement, but irregularly 
scattered in the walls of the elongated chambers, or between 
the small, scattered chambers. No subgastral sagittal 
radiates. Nuclei of collared cells probably always basal. 


This family was provided by Dendy [1892 B] for the reception 
of the genus Leucascus with its two species, L. simplex and 
L. clavatus. Minchin [1900] refused to recognise either the 
genus or the family, and included the two species in his Clathrina, 
apparently ignoring the fact that none of the known species of 
Clathrina, or indeed any other homoccel sponge, possess an 
independent dermal membrane or cortex. It is true that many 
Clathrinas develop a pseudoderm, but this is invariably formed 
from the outermost tubes of the reticulation, and therefore 
includes a layer of endoderm (gastral layer). In Lewceascus, on 
the other hand, the dermal membrane is formed exclusively of 
ectoderm and mesoglea (dermal layer), and does not consist 
merely of the outer tubes of the reticulation. Moreover, the 
radiate and non-reticulate arrangement of the elongated chambers 
in Leucascus indicates a relationship with Dendya rather than 
with the Clathrinoid Leucosolenias. 

We here extend our conception of the family Leucascide to 
include, not only the genus Leucascus and allied genera with a 
similar type of canal system, but also a number of species with 
a leuconoid type of canal system, which we have hitherto assigned 
to the genera Leucandra and Leucilla. The skeleton of these 
species, in the absence of all traces of syconoid ancestry such as 
subgastral sagittal triradiates, clearly imdicates a wide phylo- 
genetic separation from the typical Leucandras and Leucillas, 
such as Leucandra aspera and Leucilla amphora, and closely 
resembles that of Lewcascus. 

It will be remembered that Haeckel, in “ Die Kalkschwimme ” 
(vol. 11. p. 122, [1872]), placed his Leucetta primigenia (one of the 
species which was formerly assigned to Leucandra, but now placed 
in the Leucascide) at the beginning of his Leucones, which he 
derived directly from an Ascon ancestry. He says: “ Leucetta 
primigenia, als die wahrscheinliche Stammform der Leuconen, 
steht in ihre Skeletbildung der gemeinsamen Stammform aller 
Kalkschwimme, der Ascetta primordialis, so nahe, dass man sie 
unmittelbar von der letzteren ableiten kann.” We agree with 
these views so far as the relationship to the ‘‘ Ascones”’ is con- 
cerned, but we can no longer agree that the more advanced types 
of ‘‘ Leucones” (such as Leweandra) have had a leucettid ancestry. 


CALCAREOUS SPONGES. 731 


There is certainly no indication whatever of the genus Leucetta 
having passed through a syconoid stage in its evolution, as we 
believe to be the case with the true Leucandras and Leucillas, 
and we therefore propose to re-establish this genus for certain 
leuconoid Leucascidee. 

We have also placed in this family the genus Pericharax 
Poléjaeff [1883], on account of the similarity of the skeleton and 
the basal position of the nucleus in the collared cells, while the 
genus Leucomalthe can only be placed here provisionally. 


It should perhaps be mentioned here that von Lendenfeld’s 
genus Leucopsis [1885 B] was also supposed by its author to 
represent a ‘Transition form between Asconide and Leuconide.” 
The genus, with its single species, Leucopsis pedunculata, was, 
however, so imperfeetly ‘described that it is impossible to form 
any definite opinion as to its systematic position; it is very 
possibly merely a reticulate Lewcosolenia with well developed 
mesoglea and pseudogaster, an opinion which is supported by an 
examination of some of von Lendenfeld’s type specimens, which 
one of us was able to make in Berlin. At all events, it appears 
to differ widely from any of our Leueascide, and it has been 
placed by us among the Leucosolenias. 

We have been able to determine the position of the nucleus in 
no less than ten species of this small family, including five species 
of Leucetta, and in all cases have found it basal, a fact that affords 
strong support to our view that the genus Leucetta should be very 
widely separated from Lewcandra, in which the nucleus is, so far 
as we are aware, always apical. 


Genus 5. Leucascus Dendy [1892 B]. 


Diagnosis. Flagellate chambers greatly elongated, tubular, and 
sometimes copiously branched. 


For illustrations of this genus see Dendy [1893 A]. 
We recognise the following species in this genus :— 


Section A. Without oxea. 


1. L. rnstenis Row. 
Leucascus insignis Row [1913 MS. ]. 


2. L. stmpLeEx Dendy. Type species of the genus. 
Leucascus simplex Dendy | 1892 B}. 
Secrion B. With large radially arranged oxea. 


3. L. cravatus Dendy. 
Leucascus clavatus Dendy [1892 B}. 


CBX PROF. A. DENDY AND MR. R. W. H. ROW ON 


Genus 6. Leucomaurie, Haeckel [1872] (emend.). 


Diagnosis. Colony individualised, with definite external form 
and large central gastral cavity opening by a large single 
osculum. Flagellate chambers greatly elongated, tubular, 
copiously branched. Skeleton consisting of regular radiates, 
large longitudinally placed oxea scattered throughout the 
sponge body and not confined to the cortex, and minute, 
irregularly hastate microxea. 


For illustrations of this genus see Haeckel [1872]. 


The name Lewcomalthe, originally applied to one of Haeckel’s 
subgenera, 1s retained for his Lewcandra bomba, which presents 
many peculiarities distinguishing it from the rest of the species 
of that genus. Of these, the most important from the point of 
view of our present classification is the very unusual type of canal 
system figured by Haeckel, which shows a number of large and 
very much branched flagellate chambers radiating from a central 
gastral cavity. This is very different from the normal leuconoid 
canal system, and is fairly similar to that of Zewcascus. On this 
account, and on account of the regular triradiates of the skeleton, 
we have placed this species among the Leucascide, though, it 
must be acknowledged, on somewhat doubtful grounds, and the 
peculiarities of its spiculation have necessitated the provision of a 
special genus to receive it. 


The only known species is :— 


1. L. sompa Haeckel. 
Leucandra bomba Haeckel [1872]. 


Genus 7. Leucerra Haeckel [1872] (emend.). 


non Leucetta Poléjaetf [1883]. 
non Leucetéa von Lendenfeld [1891]. 


Magnosis. Canal system leuconoid, with small, spherical or sub- 
spherical flagellate chambers irregularly scattered through 
the chamber layer. 


For illustrations of this genus see Haeckel [1872] and 
Dendy [1913]. 


Haeckel [1872] proposed the genus Leucetta for calcareous 
sponges with a leuconoid canal system and a skeleton composed 
of triradiate spicules only, taking Z. primigenia for his type 
species. 

Poléjaeff [1883] abandoned Haeckel’s classification, but re- 
tained the name Lewcetta in an entirely different sense, equivalent 
to our Leucettusa, taking one of Haeckel’s species, LZ. corticata, 
for the type species of his genus. 


« 


CALCAREOUS SPONGES. TBO 


Von Lendenfeld [1891] used the genus Leucetta to include 
“‘Teuconide with triacts or tetracts or both.” 

Dendy [1893 A] included Haeckel’s species of Leucetta in the 
genus Leucandra, on the ground that the mere absence of quadri- 
radiate or oxeote spicules, or both, could not be regarded as of 
generic significance. 

Asa result of further study of the Calearea, we are convinced 
that the genus ZLewcandra, as used hitherto by Dendy, is not a 
natural one, but must be subdivided, although on lines different 
from any that have previously been suggested, It appears to us 
that certain species, namely those which we now assign to 
Leucetta, have originated, quite independently of the remainder, 
from the homoccl sponges through a leucascid ancestry, and 
have consequently never passed through a Sycon stage in their 
phylogeny. The species in question are characterised by thei 
equiangular triradiates, and by the absence of any trace of the 
typical Sycon skeleton. 

As regards the canal system, it appears that the same course of 
evolution, from a condition with elongated and more or less 
radially arranged flagellate chambers, to a condition with small 
scattered spherical chambers, has taken place in both cases, so 
that there has been a very complete convergence between the 
genera Leucettw and Leucandra as now understood by us. The 
true Leucandras, however, are distinguished by more or less 
distinct traces of the skeletal structure exhibited by their 
syconoid ancestors. 

This view, so far as our present information goes, is supported 
by histological evidence ; for, as already indicated, the position 
of the nucleus of the collared cells in Leucetta is basal, while in 
typical Leucandras it appears to be apical. 


We also include in the genus Lewcetta, as now conceived, 
certain species which we have hitherto regarded as belonging 
to the genus Leucilla. These species are characterised by the 
development to a varying extent of an inwardly pointing apical 
ray on some of the trivadiates of the dermal cortex. They are to 
be distinguished from the true Leucillas on the same grounds as 
those on which the other Leucettas are distinguished from the 
true Leucandras, and we do not consider the mere presence of 
such apical rays by itself as a sufficient justification for generic 
separation. 

As we are taking Haeckel’s type species of Leucetta as the 
type of the genus as understood by us, we consider that we ave 
justified in retaining his name for the genus, although that name 
has been used in different senses by subsequent writers. 


We allocate the following species to this genus :— 


Secrion A. Without oxea. 


1. L. cuacosEnsis Dendy. 
Leucetta chagosensis Dendy [1913]. 


734 PROF. A. DENDY AND MR. R. W. H. ROW ON 


2. L. rroripana Haeckel. 
Leucaltis floridana Haeckel [1872]. 


3. L. INFREQUENS Row. 
Leucetta infrequens Row [1913 MS. ]. 


4, L. microrapuis Haeckel. 
Leucetta primigenia var. microraphis Haeckel [1872]. 
Leuconia dura Poléjaeff [1883], fide Dendy [1892 Bj. 


5. L. primicenta Haeckel. Type species of the genus. 
Sycothamnus fruticosus Haeckel [1870], fide Haeckel 
1872}. 
ae clausu Haeckel [1879], fide Haeckel [1872]. 
Leucetta primigenia Haeckel [1872]. 
Leuconia fruticosa Poléjaeff [1883 }. 
Haeckel’s two earlier names are nomina nuda, as they were 
not accompanied by diagnoses, and we have rejected them, 
although Poléjaeff has revived one of them. 


6. L. pyrtrormis Dendy. 
Leucetta pyriformis Dendy [1913]. 


7. L. PROLIFERA Carter. 
Teichonella prolifera Carter | 1878]. 
Leucilla prolifera Dendy [1892 B]. 
Leucetta prolifera Row [1913 MS. |}. 


8. L. souipa O. Schmidt. 
Grantia solida O. Schmidt [1862]. 
Leuconia nivea J. HK. Gray [1867], fide Haeckel [1872]. 
Leucaltis solida Haeckel [1872]. 


9. L. tricgona Haeckel. 
Leucetta trigona Haeckel [1872]. 


Section B. With large, usually radially arranged oxea, 
but without microxea. 
10. L. cArtERI Dendy. 
Leucaltis floridana var. australiensis Carter [1885-1886]. 
Leucandra cartert Dendy [1892 B]. 


Section C. With both large oxea and microxea. 
11. L. expansa Low. 
Leucetta expansa Row [1913 MS.}]. 
The following species is doubtfully assigned to Leaucetéa :— 


12. L. nomorapuis Poléjaeff. 
Pericharax carteri var. homoraphis Poléjaeff [1883]. 
The name cartert cannot be used for this species, as it is 
already employed in this genus, 


CALCAREOUS SPONGES. 735 


Genus 8. PEricHarax Poléjaeff [1883]. 


Diagnosis. Sponge colony individualised, with large central 
cavity (probably a pseudogaster) opening by a wide vent 
and surrounded by a very thick wall. Canal system leuco- 
noid, with sub-spherical, scattered, flagellate chambers, and 
with subdermal cavities whose walls are supported by a 
special skeleton derived partly from the inturned rays of 
tangential dermal triradiates. Skeleton of the chamber 
layer confused, composed of equiangular triradiates of two 
very different sizes. 


For illustrations of this genus see Poléjaeff [1883] and Dendy 
[1913]. 


This genus was proposed by Poléjaeff [1883] for a species, 
Pericharax carter, from Tristan da Cunha, but recent authors, 
for the most part, have not accepted it. The discovery by the 
‘Sealark’ Expedition of the original species, and of a new one 
closely allied to it, has enabled us to make a careful study of 
the question, and we are convinced that the genus is valid. 
The position which we assign to it is justified, not only by the 
character of the skeleton, but also, as in the case of Leucetta, by 
the basal position of the nucleus of the collared cells in both 
species. 

The dermal triradiates in both species are very curious spicules, 
with a strong tendency to irregular curvature of the rays, often 
resulting in one or more of them dipping down deeply between 
the subdermal cavities. 

Pericharax heteroraphis (Poléjaett’s P. carteri var. heteroraphis) 
must be taken as the type of the genus. The same author's 
P. carteri var. homoraphis, as we have noted above, appears to be 
quite distinct, and is not a Pericharax at all, but probably a 
Leucetta. Pericharax poléjaevi Breitfuss [1896 A], placed by its 
author in this genus on account of the presence of subdermal 
cavities, is really a typical Leucandra, for the presence of sub- 
dermal cavities without a special supporting skeleton cannot be 
regarded as of generic importance. Poléjaeff also included in the 
genus Haeckel’s Lewcandra cucumis, the position of which is 
discussed by us under the genus Paraleucilla. 


We recognise the following as species of this genus :— 


1. P. HETERORAPHIS Poléjaef. ‘Type species of the genus. 
Pericharax carteri var. heteroraphis Poléjaeft [1883]. 


2. P. pezizA Dendy. 
Pericharax pexiza Dendy [1913]. 


736 PROF, A. DENDY AND MR. R. W. H. ROW ON 


Family 3. LEUCALTIDA‘ nov. 


Diagnosis. Sponge colony tubular and ramified, or even anas- 
tomosing, with many oscula, or individualised with large 
central cavity and single osculum. Wall of colony composed 
of at least two distinct layers, namely, a dermal cortex with 
strongly developed skeleton of tangential radiates, and a 
chamber layer with a skeleton greatly reduced or even 
absent. A thin gastral cortex or membrane may or may 
not be present. Skeleton composed, mainly at any rate, of 
equiangular radiates. No sub-gastral sagittal radiates. 
Nuclei of collared cells probably always basal. 


The members of this family appear to have been derived from 
a Dendya-like ancestor by the development of a thick dermal 
cortex with a strongly developed cortical skeleton, and the con- 
sequent more or less complete reduction of the no longer necessary 
skeleton of the chamber layer. As in other families of Calcarea, 
the flagellate chambers range from greatly elongated and even 
branched, and more or less radially arranged, to small, sub- 
spherical and scattered. 

Bidder [1898] has already pointed out that the nuclei of the 
collared cells in Lewcaltis clathria Haeckel (Heteropegma nodus- 
gordit Poléjaeff) are basal in position, and we are able to confirm 
this observation and to add that they are basal also in Leucetiusa 
dictyogaster Row [1913 MS. ]. 

The reduction of the skeleton of the chamber layer, correlated 
with the development of a thick dermal cortex with a special 
cortical skeleton, finds its parallel in the genus Grantiopsis 
amongst the Grantiide, but in that case the syconoid ancestry 
is very clearly indicated in the remains of an articulate tubar 
skeleton, while in Lewcaltis the vestigial skeleton of the chamber 
layer shows no indications whatever of an articulate origin. 

The characters above mentioned, In our opinion justify the 
close association of Leucaltis with Leucettusa, and the wide 
separation of these genera from both Leucandra and Leucilla 
in our scheme of classification. 


We prefer to derive the Leucaltide directly from a Dendya- 
like ancestor, rather than indirectly through Leucascus, because 
Leucaltis still preserves the more primitive type of radial colony 
formation with what we presume to be a true central gastral 
cavity, while Leweascus has adopted a massive type of colony 
formation in which the exhalant canals are possibly to be regarded 
as pseudogastral in nature. 


CALCAREOUS SPONGES. 737 


Genus 9. Leucanris Haeckel [1872] (emend.). 
Heteropegma Polejaeff [1883]. 


Diagnosis. Sponge colony tubular, ramified and anastomosing, 
with many oscula. Flagellate chambers elongated and 
branched, more or less radially arranged round the central 
gastral cavities of the tubes. 


For illustrations of this genus see Poléjaeff [1883] and Dendy 
[1893 A]. 


Dendy [1892 B, 1893 .A] placed this genus in the family Am- 
phoriscide, on account of the large subdermal quadriradiates 
possessed by the only known species. He also regarded the 
vestigial triradiates of the skeleton of the chamber layer as the 
remnants of an articulate tubar skeleton, being misled by the 
radial arrangement of the flagellate chambers. We are now 
convinced that in both these respects he was wrong, and that 
the genus is probably, as Bidder [1898] maintained, related 
to Dendya, from which it has been directly derived without 
passing through an intermediate syconoid stage. 

In arriving at this conclusion we lay great stress upon the 
regular and equiangular form of the triradiates of both the 
chamber layer and the dermal cortex, and also upon the basal 
position of the nuclei of the collared ‘cells. The genus may, in 
fact, almost be regarded as a Dendya with a thick dermal cortex. 


Dendy [1913] has shown that only one species can be re- 
cognised in the genus. A re-investigation of the type specimen 
of Haeckel’s Leucaltis clathria has convinced us that it is not 
only generically, but also specifically identical with Poléjaeft’s 
Heteropegma nodus-gordu, and the latter name thus becomes a 
synonym of Leucaltis clathria Haeckel. As the other species of 
Haeckel’s genus Leucaltas must be removed to older genera, 
L. clathria must be taken as the typical species, and we are 
therefore unable to retain Poléjaefi’s name Heteropegma. Again, 
Carter’s Clathrina latitubulata is only a variety of Leucaltis 
clathria, differing in some slight details of spiculation. Mr. 
Carter seems to have been led into provisionally placing his 
species in the genus Clathrina by the external form of the whole 
colony, inh resembles a reticulate Clathrina on a gigantic 
scale. The reticulation, however, is not composed of simple 
ascon tubes as in Clathrina, but of a colony of a higher order, 
with numerous true ascon tubes lying in the thickness of the 
wall. Finally, Ridley’s Leucaltis bathybia var. mascarenica is 
evidently, from his description, and from the slides which we 
have been able to examine, nothing but the same species. 


The extent to which apical rays are developed on the tangential 
radiates of the dermal cortex varies greatly in different in- 
dividuals, 


738 PROF. A. DENDY AND MR. R. W. H. ROW ON 


We recognise only the following species in the genus :— 


1. L. craruria Haeckel. 

Leucaltis clathria Haeckel | 1872). 

Heteropegma nodus-gordii Poléjaeff [1883], fide Dendy 
[1913]. 

Clathrina latitubulata Carter [1885--1886], jide Dendy 
(L913s]: 

Leucaltis bathybia var. mascarenica Ridley [1884], fide 
Dendy [1913]. 

Leucaltis clathria Dendy [1913]. 


Genus 10. Leucerrusa Haeckel [1872] (emend.). 
Leucetta Poleéjaeff [1883]. 


Diagnosis. Sponge colony individualised, with definite external 
form and large central cavity opening by a large single 
osculum. Canal system leuconoid. 


For illustrations of this genus see Poléjaeff [1883] under the 
name Leuceita, and Row [1913 MS. ]. 


The remarkable reticulate type of colony formation found in 
Leucaltis prevents us from regarding that genus as directly 
ancestral to Leucettusa, but as regards canal system the two are 
related in the same way as Grantia and Leucandra. 

Within the limits of the genus Leucettusa, however, we find 
considerable variation with regard to the form of the flagellate 
chambers. Poléjaeff’s figure of the canal system of Leucettusa 
(Leucetia) vera shows the flagellate chambers in the outer part of 
the chamber layer elongated and radially arranged, while those 
in the inner part are subspherical and scattered, so that this 
species appears to be intermediate between Lewcaliis and the 
more typical Leucettusas in this respect. 

Leucettusa (Leucetta) haeckeliana, on the other hand, has the 
flagellate chambers all small and subspherical, and lying in the 
irregular trabecule of the chamber layer, which are separated by 
very wide, irregular exhalant lacune. In Leucettwsa dictyogaster 
Row [1913 MS8.] the trabecule bearing the flagellate chambers 
form a network which almost completely blocks up the central 
gastral cavity as an altogether askeletal layer. A section of this 
askeletal chamber layer, isolated from the cortex, would be almost 
indistinguishable from a similar section of Oscarella, which also 
has large collared cells with basally placed nuclei, so that the 
possibility presents itself that Oscarella may be nothing but a 
calcareous sponge which has lost the whole of its skeleton. 

We also find in the genus Lewcetiusa various stages in the 
development of the subdermal quadriradiate spicules, which are 
entirely absent in L. corticata and L. dictyogaster, very sparse 
in L. haeckeliana and large and very numerous in LJ. vera. As 


- CALCAREOUS SPONGES. 739 


already indicated, we no longer consider the presence of sub- 
dermal quadriradiates, taken by itself, to be a character of generic 
importance, being convinced that the addition of an apical ray 
to a triradiate spicule may take place whenever and wherever it 
may be required. 

Tt will have been noticed that our genus Leucettusa is identical 
in scope with Poléjaeft’s Leucetta [1883], but as we have found it 
necessary to retain Haeckel’s name Leucetta for another genus, 
as previously explained, we have been obliged to adopt his 
subgeneric name for the group of species which Poléjaeff quite 
rightly separated out. 


We recognise the following species as belonging to this genus:— 


Suction A. Without oxea. 


1. L. corticara Haeckel. Type species of the genus. 
Leucetta corticata Haeckel [ 1872). 


2. L. HAECKELIANA Poléjaeff. 
Leucetta haeckeliana Poléjaeft [1883]. 


3. L. mprrFecta Poléjaeff. 
Leucetta imperfecta Poléjaeft [1883]. 


4, L. samBucus Preiwisch, 
Leucetta sambucus Preiwisch [1904]. 


5. L. vera Poléjaeff. 
Leucetta vera Poléjaeff [1883]. 


Srcrron B. With microxea but without large oxea. 


6. L. DICTYOGASTER Low. 
Leucettusa dictyogaster Row [1913 MS8.}. 


Family 4. MINCHINELLIDA/ nov. 


Lithonina Doderlein [1892]. 
Lithonine Kirkpatrick [1911 A}. 


Diagnosis. Canal system leuconoid (in all known forms and 
presumably always so). Main skeleton composed of quadri- 
radiates cemented together in various ways by calcareous 
cement. Apparently without subgastral sagittal radiates. 
Nuclei of collared cells (probably always) basal. 


We have been able, owing to the kindness of Mr. Kirkpatrick, 
to examine preparations of Minchinella and Murrayona, in which 
the collared cells are sufficiently well preserved to enable us to 
determine the position of the nucleus. We find that this is basal 
in both cases, whereas in Lelapia we find 1% to be apical. As 

Proc. Zoou. Soc.—1913 No XULIX 49 


740 PROF, A, DENDY AND MR. R. W. H. ROW ON 


Lelapia differs widely in other characters also from both Min- 
chinella and Murrayona, we are forced to the conclusion that the 
so-called Pharetronid sponges are at least diphyletic in origin, 
and we have removed the Lelapiide to a position in the Sycettid 
line of descent. We shall, however, discuss the question further 
under the head of phylogeny. 


The genera Minchinella, Petrostroma and Plectroninia all 
agree in the possession of a stony skeleton composed of fused 
quadriradiates, and thus differ widely from Murrayona. We 
therefore unite them in one family under the name Minchinellide, 
while relegating Murrayona to a special family of its own. 


Genus 11. Mincurnetyia Kirkpatrick [1908]. 


MNagnosis. Sponge lamellar, with pore-bearing chimneys on one 
side and oscular chimneys on the other. The quadriradiates 
of the main skeleton cemented together.into a compact 
network and completely embedded in the enveloping cement. 
Dermal skeleton of radiates, including tuning-fork spicules, 
and oxea. 


For illustrations of this genus see Kirkpatrick [1908]. 
The only known species of the genus is :— 


1, M. LAMELLOSA Kirkpatrick. 
Minchinella lamellosa Kirkpatrick [1908]. 


Genus 12. Perrostroma Doderlein [1892]. 


Diagnosis. 'The quadriradiates of the skeleton of the chamber- 
layer fused together laterally by calcareous cement into a 
network. Dermal skeleton of separate quadriradiates and 
triradiates and bunches of tuning-fork spicules. 


For illustrations of this genus see Déderlein [1897]. 
The only known recent species of this genus is :— 


1. P. scHutzer Doderlein. 
Petrostroma schulzei Doderlein [1892]. 


Genus 13. PLecrroninta Hinde [1900]. 


Diagnosis. Quadriradiates of the main skeleton with their facial 
rays truncated or expanded terminally and fused end to end 
with the facial rays of adjacent spicules, while the apical 
rays remain free and pointed. Dermal skeleton of separate 
radiates, including tuning-fork spicules, and oxea. 


For illustrations of this genus see Kirkpatrick [1900 B]. 


CALCAREOUS SPONGES, 7A) 


‘The only known recent species of this genus are :— 


1. P. peansit Kirkpatrick. 
Plectroninia deanstt Kirkpatrick [1911 A}. 


2. P. winpE1 Kirkpatrick. 
Plectroninia hinder Kirkpatrick [1900 B]. 


The type of the genus is P. halli Hinde [1900], a fossil species 
from the Eocene of Victoria. 


‘Family 5. MURRAYONIDN nov. 
Murrayonine Kirkpatrick [1911 A]. 


Diagnosis. Canal system presumably always leuconid. Skeleton 
of the chamber layer a rigid calcareous network, not com- 
posed of spicules. No subgastral sagittal radiates. Dermal 
skeleton composed chiefly of overlapping calcareous scales. 
Nuclei of collared cells basal. 


This family seems to mark the culminating point of the 
Leucascid-Leucettid line of evolution, and there is no other 
known calcareous sponge with a skeleton so highly specialised as 
WMurrayona. 


It is at any rate possible that the aspicular main skeleton of 
Murrayona is derived from the cement-covered fibres of spicules 
which are known to occur in many of the fossil forms, by a 
gradual disappearance of the spicular core, analogous to what we 
find among the Chalinine, during the evolution of the group. 
It is, however, a far cry from the one to the other, and we do not 
doubt that there are many other ways in which the Murrayonid 
skeleton may have arisen. 


Genus 14. Murrayona Kirkpatrick [1910]. 


Diagnosis. With a definite pore-zone in which the dermal skeleton 
consists of small trivadiates. Tuning-fork spicules present 
beneath the dermal scales. 


For illustrations of this genus see Kirkpatrick [1910]. 
The only known species of this genus 1s :— 


1. M. pHanoeris Kirkpatrick. 
Murrayona phanolepis Kirkpatrick [1910]. 


4g* 


742, PROF, A. DENDY AND MR. R. W. H. ROW ON 


Family 6. SYCETTIDA Dendy [1892 B}. 


Diagnosis. Flagellate chambers elongated, arranged radially 
around a central gastral cavity, their ends projecting more 
or less on the dermal surface and not covered over by a 
continuous dermal cortex strengthened by tangential dermal 
spicules. “Tubar skeleton articulate, with subgastral sagittal 
radiates. Collared cells usually confined to the radial 
chambers in the adult, and probably always with apical 
nuclei. 


The Sycettide, in our opinion, form the starting point of a 
distinct line of evolution, embracing the great majority of the 
recent heteroccel Calcarea. This view is supported by the fact 
that in all the 44 species of Sycettide, Heteropiide, Grantiide, 
Amphoriscidee and Lelapiidee in which it has been determined, 
the position of the nuclei of the collared cells is apical. 

It would appear therefore that this line of evolution must 
have originated from homoccel ancestors with apical nuclei. Such 
forms are known to occur amongst the simple (7. e. non-reticulate) 
species of Leucosolenia, as for example L. lucasi and-L. bella. 

Why the transition from the simple homoccel to the radial 
heteroccel condition originally took place we can only guess. 
That it was by the outgrowth of radial buds is clearly indicated, 
however, by the ontogeny of the genus Sycon (compare Schulze 
[1875]). No real intermediate forms are known, however, for we 
can no longer consider that Dendya is on the same line of 
ancestry, while von Lendenfeld’s Homoderma sycandra is already 
a highly specialised Sycon with the typical Sycon skeleton, and 
differing from other Sycons only in the persistence of the collared 
cells in the central gastral cavity. According to Minchin the 
Sycettidz have arisen from his homoccel family Leucosoleniide, 
but we have already endeavoured to shew that in the present state 
of our knowledge it is impracticable to distinguish this family 
from his Clathrinide. 


The presence of sagittal radiates in which the oral angle is 
wider than the lateral angles appears to be a very constant 
character of the Sycettide and their derivatives. It would seem, 
further, that the occurrence of such spicules in the articulate 
tubar skeleton is correlated with the development of very definite 
radial tubes, but it is extremely difficult, at any rate in practice, 
to distinguish such spicules individually from equiangular 
radiates which have become sagittal by bending of the oral rays. 
The presence of subgastral sagittal radiates is, however, especially 
characteristic of this line of evolution. The oxeote type of 
spicule also tends to assume more importance in the skeleton of 
this family than in the Leucascidee and Leucaltide, in which 
families it but rarely occurs. 


CALCAREOUS SPONGES. | 143 


Genus 15. Sycerra Haeckel [1872] (emend.). 


Diagnosis. The radially arranged flagellate chambers always 
completely separate from one another, and never possessing 
tufts of oxea at their distal ends. With no properly defined 
inhalant canals leading to the prosopyles. 


For illustrations of this genus see Haeckel [1872] under 
Sycaltis conifera and Sycetta primitiva. 


This is the simplest of all the genera with syconoid canal 
system, its primitive nature being shown by the absence of fusion 
between the flagellate chambers and by the absence of tufts 
of oxea at their distal ends, unless, indeed, this absence is due to 
the disappearance of ancestral oxea. 

Sycetta asconoides Breitfuss [1896 B], to judge by the published 
description, apparently occupies an intermediate position between 
Sycetta and Sycon, as these genera are understood by us. The 
description states that the flagellate chambers are fused together 
laterally, but that they have no tufts of oxea at their distal ends. 
During a recent visit to Berlin, however, one of us (Row) had 
the opportunity of examining one of the type slides of this 
species, and found that Breitfuss had overlooked the presence of 
a few tangentially placed dermal triradiates. The occurrence of 
these spicules, of course, places the species in the genus Grantia in 
the family Grantiide, of which it is one of the simplest forms. 


We include the following species in this genus :— 


1. S. contFERA Haeckel. 
Sycaltis conifera Haeckel [ 1872). 


2. S. primitiva Haeckel. ‘Type species of the genus 
Sycetta promitiva Haeckel [1872]. 


3. 8. sacirriFERA Haeckel. 
Sycetta sagittufera Haeckel [1872]. 


Genus 16. Sycon Risso [1826] (emend.). 


Diagnosis. Radial chambers usually more or less united at places 
where they come into contact with one another, and always 
crowned distally with tufts of oxeote spicules. Properly 
defined inhalant canals usually present, the outer ends of 
which may be covered by a thin pore-bearing dermal 
membrane without special skeleton. 


For illustrations of the structure of this genus see Schulze 
[1875] and Dendy [1893 A]. 


As pointed out by Dendy [1893 A] the most characteristic 
feature of this genus is afforded by the tufts of oxeote spicules 
which crown the distal ends of the radial chambers, taken in 


744 PROF. A. DENDY AND MR. R. W. H. ROW ON 


conjunction with the absence of a dermal cortical, as distinct: 
from a tubar, skeleton. In the more specialised species a pore- 
bearing dermal membrane stretches between the distal ends of the 
radial chambers, covering over the ends of the inhalant canals, but 
this contains no special skeleton of its own. The flagellate 
chambers may also exhibit a considerable amount of branching 
towards their distal extremities, but they never lose thei 
elongated character and radial arrangement. 

The genus is sharply distinguished from Sycetta by the presence 
of the tufts of oxea at the distal ends of the chambers, less. 
sharply from Grantia by the absence of a special dermal cortical 
skeleton. 

We include in our conception of the genus Sycon von 
Lendenfeld’s genera Homoderma and Sycantha, and Jenkin’s. 
Tenthrenodes antarcticus, Streptoconus australis and Hypodictyon 
longstafi. We have already mentioned that Homoderma is. 
merely a Sycon with persistent collared cells in the central 
gastral cavity. Jenkin [1908 B] has shown conclusively that 
Sycantha tenella is a typical Sycon, but has erected a new genus. 
Tenthrenodes for ‘‘Sycettidee with linked chambers,” an almost 
identical character with that on which the genus Sycantha was 
founded; and although Dendy [1893 A] retained Sycantha on 
these grounds, we no longer consider that such “linking” can be 
regarded as of generic importance. We may point out here that. 
Tenthrenodes scotti, the other species included by Jenkin in his 
genus, is placed by us in the genus (’rantia, on account of the 
presence of tangential triradiates in the dermal cortex. Strepto- 
conus australis and Hypodictyon longstaffi are “ chiact”’-bearing 
forms which were placed by Jenkin in his family Chiphoride. 


We assign the following species to the genus :— 


1. 8S. aLopecurus Haeckel. 
Sycum alopecurus Haeckel [1870]. 
Sycandra ampulla var. alopecurus Haeckel [1872]. 


2. S. ampuLLA Haeckel. 
Sycariun ampulla Haeckel [1870]. 
Sycon petiolatus O. Schmidt MS., fide Haeckel [1872]. 
Sycum petiolatum Haeckel [1870], fide Haeckel | 1872). 
Sycandra ampulla Haeckel [1872]. 


3. 8. ANTARCTIOUM Jenkin. 
Tenthrenodes antarcticus Jenkin [1908 B]. 


4, 8. arcticum Haeckel. 
Sycum arcticum Haeckel [1870]. 
Sycon raphanus O. Schmidt [1870], fide Haeckel [1872]. 
Sycandra arctica Haeckel [1872]. 


5. S. ASPERUM Gibson. 
Sycandra aspera Gibson | 1886}. 


10. 


Mae 


16. 


ie 


18. 


CALCAREOUS SPONGES. 745 


. AUSTRALE Jenkin. 


Streptoconus australis Jenkin [1908 B]. 


. BARBADENSE Schuffner. 


Sycandra barbadensis Schuffner [1877 ]. 


. BOOMERANG Dendy. 


Sycon boomerang Dendy [1892 B]. 


. BOREALE Schuffner. 


Sycandra borealis Schuffner [1877]. 


. CAMINATUM Thacker. 


Sycon caminatum Thacker [1908]. 


. CARTERI Dendy. 


Sycon carteri Dendy [1892]. 

Sycantha tenella von Lendenfeld MS., fide Breitfuss 
[1897]. 

Sycon carter’ Row [1913 MS. ]. 


. CILIATUM Labricius. 


Spongia ciliata Fabricius [1780]. 

Sycum gigantewm Haeckel [1870], fide Haeckel [18721. 

Sycocystis oviformis Haeckel [1870], jide Haeckel 
[1872]. 

Sycodendrum ramosum Haeckel [1870], jide Haeckel 
[1872]. 

Sycandra ciliata Haeckel [1872]. 


. coactum Urban. 


ycandra coacta Urban [1905]. 


. comMuTATUM Haeckel. 


Sycandra coronata var. commutata Haeckel [1872]. 


. compactum Lambe. 


Sycon compactum Lambe [1893]. 


. cononaAtTUM “Hillis and Solander. 


Spongia coronata Ellis and Solander [1786]. 

Grantia ciliata Bowerbank [1864-1882], fide Haeckel 
[1872}. 

Sycandra coronata Haeckel [1872]. 

Sycon coronatum Dendy [1892 B]. 


- EGLINTONENSIS Lambe. 


Sycon eglintonensis Lambe [1900 B}. 


. ELEGANS Bowerbank. 


Dunstervillia elegans Bowerbank [1845]. 

Dunstervillia lanzerote Haeckel [1870], jide Haeckel 
[1872]. 

Sycandra elegans Haeckel [1872". 


746 


7S 


5 Se 


PROF. A. DENDY AND MR. R. W. H. ROW ON 


. ENSIFERUM Dendy. 


Sycon ensiferum Dendy [1892 B]. 
Sycon ensiferum Row [1913 MS. ]. 


rormosuM Haeckel. 
Dunstervillia formosa Haeckel [1870]. 
Sycandra elegans vay. formosa Haeckel [1872]. 


. GELATINOSUM de Blainville. 


Alcyoncellum gelatinosum de Blainville [1834-1847 ]. 

Grantia virgultosa Bowerbank MS8., fide Haeckel [1872]. 

Sycandra alcyoncellum Haeckel [1872], fide Dendy 
[1892 B]. 

Sycandra arborea Haeckel [1872], fide Dendy [1892 B]. 

Sycon gelatinosum Dendy [1892 B}. 


. GIGANTEUM Dendy. 


Sycon giganteum Dendy [1892 B]. 


. HELLERI von Lendenfeld. 


Sycandra helleri von Lendenteld (1891). 


. HUMBOLDTIL Lisso. Type species of the genus. 


Sycon humboldtii Risso [1826]. 

Dunstervillia corcyrensis O. Schmidt [1862], fide 
Haeckel [1872]. 

Dunstervillia schmidti Haeckel [1870], fide Haeckel 
[1872]. 

Sycandra humboldtii Haeckel [1872]. 


. IMPLETUM /aeckel. 


Artynas villoswm Haeckel [1870], fide Haeckel [1872]. 
Sycandra villosa var. impletum Haeckel [1872]. 


. INCONSPICUUM von Lendenfeld. 


Sycandra inconspicua von Lendenfeld [1885 B]. 


. INCRUSTANS brettfuss. 


Sycon incrustans Breitfuss [1898 E}. 


. KARAJAKENSE Breitfuss. 


Sycon karajakense Breitfuss [1897 }. 


. KERGUELENSIS Urban. 


Sycon kerguelensis Urban [1908]. 


. LAMBEL, sp. 2. 


Sycon asperum Lambe [1896]. 
The new specific name has been given to the above 


species on account of the fact that the name asperwm is 
already occupied in this genus. (See above.) 


31. S. LANCEOLATUM Haeckel. 


Sycum lanceolatwm Haeckel [1870]. 
Sycandra ciliata var. lanceolata Haeckel [1872]. 


AO. 


4]. 


44, 


45, 


A6. 


47. 


48. 


op) 


CALCAREOUS SPONGES. 


. LENDENFELDI Row. 


Sycon lendenfeldi Row [1913 MS.}. 


. LInGuA Haeckel. 


Sycortis lingua Haeckel [1872]. 


. LONGSTAFFI Jenkin. 


Hypodictyon longstagi Jenkin [1908 B]. 


. MAxImMuUM Haeckel. 


Sycandra arctica var. maxima Haeckel [1872]. 


. MINUTUM Dendy. 


Sycon minutum Dendy [1892 B]. 


. MUNDULUM Lambe. 


Sycon mundulum Lambe [1900 B}. 


. MUNITUM Jenkin. 


Sycon munitum Jenkin [1908 A]. 


. ORNATUM Kirk. 


Sycon ornatum Kirk [1897]. 


. ovatum Haeckel. 


Sycum ovatum Haeckel [1870}. 
Sycandra ciliata var. ovata Haeckel | 1872). 


. PARVULUM Preiwisch. 


Sycandra parvula Preiwisch [1904}. 


. PEDICELLATUM AUrk. 


‘ycon pedicellatum Kirk {1897}. 


. PeTIoLATUM Haeckel [1870]. 


Sycum petiolatum Haeckel [1870]. 
Sycandra ampulla vay. petiolata Haeckel [1872]. 


. POLARE Haeckel. 


Sycandra arctica var. polaris Haeckel {1872}. 


. PROBOSCIDEUM Haeckel. 


Syconella proboscidea Haeckel [1870]. 
Sycandra raphanus vay. proboscidea Haeckel [1872] 


. PROCUMBENS Haeckel. 


Sycum procumbens Haeckel [1870]. 


Sycandra raphanus var. procumbens Haeckel [1872]. 


. PRoTECTUM Lambe. 


Sycon protectum Lambe [1896]. 


. QUADRANGULATUM O. Schmidt. 


Syconella quadrangulata O. Schmidt {1868}. 
Sycandra quadrangulata Haeckel [1872]. 


747 


748 


50. 


51. 


52. 


53. 


54. 


55. 


_ 56. 


57. 


58. 


59. 


60. 


PROF. A. DENDY AND MR. R&R. W. H. ROW ON 


. S. RAMosUM Haeckel. 


Sycandra ramosa Haeckel [1872]. 
Leuckartea natalensis Michlucho-Maclay MS., fide Haeckel 
[1872]. 


S. RAMSAYI von Lendenfeld. 
Sycandra ramsayt von Lendenfeld [1885 A }. 


S. RAPHANUS O. Schmidt. 
Sycon raphanus O. Schmidt [1862]. 
Spongia inflata Delle Chiaje [1828], fide Haeckel [1872]. 
Sycarium vesica Haeckel [1870], fide Haeckel [1872]. 
Sycandra raphanus Haeckel [1872]. 


S. scumiptit Haeckel. 
Sycandra schmidtii Haeckel [1872]. 
This species must be distinguished from Dunstervillia 
schmidtii Haeckel, a synonym of Sycon humboldti Risso. 


S. SCHUFFNERI, sp. 1. 
Sycandra quadrata Schuffner [1877]. 
We propose this new name in order to avoid confusion 
with Haeckel’s variety quadrata of Sycon quadrangulatum 


(O. Schmidt). 


S. serosum O. Schmidt. 
Sycon setosum O. Schmidt [1862]. 
Sycandra setosa Haeckel [1872]. 


S. STAURIFERUM Preiwisch. 
Sycandra staurifera Preiwisch [1904]. 


S. SUBHISPIDUM Carter. 
Grantia subhispida Carter [1885-1886]. 


S. sycaAnpDRA von Lendenfeld. 
Homoderma sycandra von Lendenfeld [1885 A]. 
Leucosolenia (2) sycandra Dendy [1891 A}. 
See also Row [1913 MS.], under Sycon lendenfeldi. 


S. TABULATUM Schuffner. 
Sycandra tabulata Schuftner [1877]. 


Very probably identical with Haeckel’s variety tabulata of 
Sycon elegans Bowerbank. 


S. TENELLUM von Lendenfeld. 
Sycantha tenella von Lendenfeld [1891]. 
Sycon tenellum Jenkin [1908 B]. 


S. TeRGESTINUM Haeckel. 
Sycum tergestinum Haeckel [1870]. 
Sycandra raphanus var. tergestina Haeckel [1872]. 


CALCAREOUS SPONGES. . 749 


61. S. rEssELLATUM Bowerbank. 
Grantia tessellata Bowerbank [1864-1882 ]. 
Sycandra elegans var. tessellata Haeckel [1872]. 


62. S. resserARIUM Haeckel. 
Sycandra quadrangulata var. tesseraria Haeckel [1872}- 


63. S. ruBaA von Lendenfeld. 
Sycandra tuba von Lendenfeld (1891). 


64. S. ruBuLosum Haeckel. 
Sycandra coronata var. tubulosa Haeckel [1872]. 


65. S. veruM Row. 
Sycon verum Row [1913 MS.]. 


66. S. vittosum Haeckel. 
Sycarium villoswm Haeckel [1870]. 
Sycum clavatwm Haeckel [1870], fide Haeckel [1872]. 
Sycandra villosa Haeckel {1872}. 


67. S. vircuLtosum Haeckel. i 
Sycandra aleyoncellum var. virgultosa Haeckel [1872]. 


Genus 17. Sycanpra Haeckel [1872] (emend.). 


Diagnosis. The radially arranged flagellate chambers more or less 
united where they come into contact with one another. 
Gastral cavity traversed by strands of tissue containing 
bundles of parallel oxea and forming a more or less strongly 
developed endogastric network. Radially arranged dermal 
oxea present. 


For illustrations of this genus see Haeckel [1872]. 


We propose to use this generic name for O. Schmidt’s Ute 
utriculus (=Sycandra utriculus Haeckel), which is sufficiently 
sharply distinguished by its skeletogenous endogastric network. 
There is only one other species in which this character is known 
to occur, namely Leucettaga loculosa, a member of the family 
Grantiide. 

The speciés of Sycandra which precede S. utriculus in Haeckel’s 
monograph having been relegated to earlier genera such as Sycon, 
Ute and Grantia, this species becomes the type of the genus. 


The only known species is :— 


1. S. urricutus O. Schmidt. 
Ute utriculus O. Schmidt [1870]. 
Sycandra utriculus Haeckel [1872]. 


750 PROF. A. DENDY AND MR. R. W. H. ROW ON 


Family 7. HETEROPIIDA) Dendy [1892 B]. 


Diagnosis. With a distinct and continuous dermal cortex covering 
over the chamber-layer and pierced by inhalant pores. 
Subgastral sagittal and subdermal pseudosagittal radiates 
are present. Flagellate chambers varying from elongated 
and radially arranged to spherical and irregularly scattered. 
With or without an articulate tubar skeleton. Nuclei of 
collared cells probably always apical. 


This family is identical in scope with the family as originally 
proposed by Dendy [1892 B], and the difference now made in the 
diagnosis is due to the fact that our conception of the subdermal 
triradiates has changed. Up to the present we have considered 
the characteristic subdermal spicules in this family as being truly 
sagittal, with the basal ray centripetally directed. We have now 
convinced ourselves, however, by a careful examination of a 
number of species, that this is not the case, but that the 
inwardly pointing ray is really one of the oral rays, and that 
the original basal ray has taken on the appearance and position 
of an oral ray. In other words, we find the clearest evidence 
that these spicules are derived from ordinary distally situated 
triradiates of the articulate tubar skeleton, which have undergone 
rotation followed by the acquisition of a secondary pseudo- 
symmetry. We therefore propose for them the name of 
pseudosagittal. It will be remembered that Poléjaeff [1883] 
recognised, in the case of Grantessa (Anphoriscus) poculum and 
G. flamma, that the subdermal triradiates are not ordinary 
sagittal spicules and that the centripetal ray is really one of the 
lateral (=oral) rays and not the basal ray. He, however, con- 
sidered that they are triradiates of the dermal cortex which 
have undergone re-orientation, and not, as we maintain, tubar 
triradiates. 

Various species of the genus Grantessa show quite clearly how 
the change has taken place. In Grantessa hirsuta we have a 
primitive type with long chambers and an articulate skeleton of 
many joints. At the distal ends of the chambers are tufts of 
oxea, towards which the basal rays of the triradiates of the distal 
joint of the tubar skeleton are inclined, as indeed occurs also in 
the genera Sycon and Grantia. Moreover, the whole spicule has 
become tilted until in some cases one of the original oral rays has 
assumed a position at right angles to the surface, while the other 
has come to lie nearly parallel to the surface, where it probably 
serves to guard the entrance to the inhalant canal. In more 
advanced cases, such as Grantessa sacca and G. hispida, the great 
elongation of the now inwardly directed oral ray increases the 
resemblance to an ordinary sagittal spicule, but a characteristic 
asymmetry of the outwardly directed (apparent oral) rays, 
accompanied by a definite kink or angulation in one of them, 


. CALCAREOUS SPONGES. OD 


(which appears to be due to change of position during individual 
growth), affords a clear indication of what has really taken place. 
Finally, in the most advanced types, such as Grantessa intusarti- 
culata, we find the pseudosagittal subdermal spicules assuming 
great dominance, almost to the exclusion of the typical articulate 
tubar skeleton, so that we arrive at the so-called inarticulate 
type. | 

The development of these characteristic spicules appears to 
antedate the appearance of a definite dermal cortex, for we 
find in Sycon ensiferwm Dendy a similar canting of certain of 
the distal tubar triradiates, which renders this species almost 
indistinguishable from Grantessa. Indeed, it is this out-turning 
of one of the rays of the distal tubar trivadiates that has, in our 
opinion, led to the formation of a dermal cortex, probably by the 
drawing out of the soft tissues of the sponge with the rays in 
question. Thus the origin of the dermal cortex in this family 
would be intimately connected with the development of these 
subdermal pseudosagittal triradiates. In the Grantiide, on the 
other hand, the dermal cortex appears to have originated in the 
development of tangentially placed trivadiates in a previously 
aspicular pore-bearing dermal membrane. 

These views undoubtedly tend to bridge over the gap between 
the Sycettide and the Heteropiide, and ‘radioed the more primitive 
species of Grantessa are differentiated from Sycon and Grantia by 
very slight characters, and difficult to separate from them, but. 
the rotation of the trivadiates in question appears to have formed 
the starting point of a new line of skeletal evolution which seems 
to us to deserve recognition as marking a distinct family. 

We consider the views here put ferward as to the origin of the 
subdermal pseudosagittal spicples of the Heteropiide to be more 
in accordance with observed facts than those previously suggested 
by one of us (Row 1909) in regard to the ‘“‘subdermal secondary 
sagittal triradiates ” of Grantilla, which seem to be pseudosagittal 
spicules really similar to those of Grantessa. 


We have changed the spelling of the name of the family from 
Heteropide to Heteropiide, the latter being more in accordance 
with the usual practice. 


Genus 18. GranteEssA von Lendenfeld [1885 B] (emend.). 


Diagnosis. Canal system syconoid. No colossal longitudinally 
placed oxea. 


For illustrations of this genus see von Lendenfeld [1885 B] 
and Dendy [1893 A]. 


The tubar skeleton in this genus ranges from articulate, with 
very numerous joints, as in Grantessa sacca, G. erinaceus, G. hirsuta 
and G, hispida, to inarticulate or nearly so, as in G. glabra and 


752 PROF. A. DENDY AND MR. R. W. H. ROW ON 


G. polyperistomia. This character might imdeed be used as a 
basis for the subdivision of the genus, were it not for the 
impossibility of drawing a satisfactory line between the two types 
of tubar skeleton, 


We recognise the following species as belonging to this 
genus :— 


Secrion A. With large, usually radially arranged oxea 
but without microxea. 


1. G. compressa Carter. 
Heteropia compressa Carter [1885-1886 }. 


2. G. ERECTA Carter. 
Heteropia erecta Carter [1885-1886]. 


3. G. ERINACEUS Carter. 
Leuconia erinaceus Carter [1885-1886 | 


4, G. rtamMa Poléjaeff. 
Amphoriscus amma Poleéjaeft [1883]. 


5. G. GLABRA Low. 
Grantessa glabra Row [1909]. 


6. G HASTIFERA Low. 
Grantilla hastifera Row [1909]. 
Grantessa hastifera Dendy [1913]. 


7. G. nirsuta Carter. 
Hypograntia hirsuta Carter [1885-1886]. 
Grantessa hirsuta Row [1913 MS.]. 


8. G. HIsPIDA Dendy. 
Grantessa hispida Dendy {1892 B). 


9, G. LANCEOLATA Breitfuss. 
Lbnerella lanceolata Breitfuss [1898 B]. 


10. G. niTIDA Arnesen. 
Ebnerella nitida Arnesen [1901). 


11. G. paiaaica Ridley. 
Nardoa pelagica Ridley [1881 }. 


12. G. PLURIOSCULIFERA Carter. 
Heteropia pluriosculifera Carter [1885-1886]. 


13. G. pocutum Polejaeff. 
Amphoriscus poculum Poléjaeff [1883 ]. 
Heteropia patulosculifera Carter [1885-1886], fide Dendy 
[1892 B]. 
Grantessa poculum Dendy [1892 B]. 


14 


15. 


16. 


17 


.G 


G 


CALCAREOUS SPONGES. 753 


. POLYPERISTOMIA Carter. 
Heteropia polyperistonia Carter [1885-1886 ]. 
Grantessa polyperistonua Row [1913 MS. ]. 


. sacca von Lendenfeld. Type species of the genus. 
Grantessa sacca von Lendenfeld [1885 B]. 


G. SYCILLOIDES Schuffner. 


Sycortis sycilloides Schuftner [1877]. 


Section B. Without large oxea, but with microxea, 


. G. INTUSARTICULATA Carter. 


Hypograntia intusarticulata Carter [1885-1886 ]. 

Hypograntia medioarticulata Carter [1885-1886], fide 
Dendy [1892 B]. 

Grantessa intusarticulata Dendy [1892 B]. 


Section C. With large, usually radially arranged oxea 


and with microxea. 


18. G. KiKentTHALI Breitfuss. 


Hbnerella kikenthali Breitfuss [1896 A]. 


19. G, PREIWISCHI, sp. n. 


Ebnerella compressa Preiwisch [1904]. 


This new specific name has been given to the species, as 
compressa is already occupied. (See above.) 


20, G. spissa Carter. 
Heteropia spissa Carter [1885-1886]. 
21. G. rHompsont Lambe. 
Amphoriscus thompsoni Lambe [1900 B]. 
Section D. Without any oxea. 
22. G. ciactatis Haeckel. 
Sycaltis glacialis Haeckel [1872]. 
23. G. MURMANENSIS Greitfuss. | 
Amphoriscus murmanensis Breitfuss [1898 B]. 
24, G. stmpLex Jenkin. 
Grantessa simplex Jenkin [1908 A}. 
25. G, srauRiDEA Haeckel. 
Sycetta stauridea Haeckel [1872]. 
Djeddea violacea Michlucho-Maclay MS., fide Haeckel 
[1872]. 
26. G. ZANZIBARENSIS Jenkin. 


Grantessa zanzibarensis Jenkin [1908 A]. 


754 PROF. A, DENDY AND MR. R. W. H. ROW ON 


Genus 19. Hermropia Carter [1885-1886] (emend.). 


Diagnosis. Canal system syconoid. Dermal cortex with colossal 
longitudinal oxea. 


For illustrations of this genus see Row [1913 MS.]. 


This genus stands in precisely the same relation to Grantessa 
that Ute does to Grantia. It is noteworthy that, in all known 
species of this genus, as in the more highly developed species of 
Grantessa, the tubar skeleton has been reduced to the subgastral 
sagittal triradiates, supplemented by the subdermal pseudosagittal 
triradiates, and has thus become ‘‘ inarticulate.” 


The genus Heteropia was diagnosed by Carter in July 1886 as 
follows :—‘‘ Caleareous sponges in which the wall is simply 
composed of sarcode supported on large sagittiform triradiates, 
whose heads are fixed in opposite sides of it respectively, and 
whose long shafts, extending perpendicularly across it, more or 
less overlap each other.” 

Most of the species placed by Carter in this genus belong 
to the earlier genus Grantessa of von Lendenfeld. There is, 
however, one of his species, Heteropia ramosa, which is dis- 
tinguished by the presence of colossal longitudinal dermal oxea, 
and which may be regarded as the type of Carter’s genus. It is 
curious that Mr. Carter himself [1886] described it under the 
name of Aphroceras ramosa, whilst saying at the same time that 
it belonged to his genus /eteropia. 


We recognise the following species as belonging to this 
genus :— : 
Section A. Without microxea. 


1. H. GLomErosa Bowerbank. 
Leuconia glomerosa Bowerbank [1872-1876]. 


2. H. rAmosa Carter. Type species of the genus. 
Aphroceras ramosa Carter [1886]. 


3. H. stmpLex Low. 
Heteropia simplex Row [1913 MS. ]. 


Section B. With microxea. 


4. H. roperert Lambe. 
Heteropia rodgerit Lambe [1900]. 


Genus 20. Ampuiutr Hanitsch [1894]. 


Diagnosis. Canal system syconoid. Both gastral and dermal 
cortices with colossal longitudinal oxea. 


For illustrations of this genus see Hanitsch [1895]. 


CALCAREOUS SPONGES. 755 


This genus may be regarded as derived from some more 
primitive type of Grantessa by the addition of colossal longitudinal 
oxea to both dermal and gastral cortices. Those in the gastral 
cortex are probably to be regarded as having been derived from 
the oxea of the oscular fringe, by downward extension. In the 
only known species microxea are present, and the articulate tubar 
skeleton still persists. 


The only known species is :— 


1. A. pautint Hanitsch. 
Amphiute paulint Hanitsch [1894]. 


Genus 21. VosmArropsis Dendy [1892 B}. 


Diagnosis. Canal system sylleibid (or leuconoid ?). Skeleton of 
the chamber layer composed of the centrifugally directed 
rays of subgastral sagittal triradiates and the centripetally 
directed rays of subdermal pseudosagittal triradiates, which 
may be supplemented or partially replaced by confused 
triradiates. No colossal longitudinal oxea. 


For illustrations of this genus see Dendy [1892 B]. 


In all the known species of Vosnaeropsis we find that the 
canal system has not developed beyond the sylleibid condition, 
and there can still be distinguished in the chamber layer very 
clear indications of an inarticulate tubar skeleton ; in fact, the 
genus seems to have reached almost exactly the same level of 
evolution as JJegapogon in the Grantiide, when allowance is 
made for the different type of skeleton in the two families. 
This would at any rate seem to suggest that the family Heteropiide 
is of comparatively recent origin, and that more complex forms, 
comparable to the higher types of the Grantiide, have not yet 
made their appearance, unless, indeed, they have merely escaped 
observation. 


We recognise the following species in this genus :— 


Section A. With large oxea and microxea. 


1. V. pepressa Dendy. 
Vosmaeropsis depressa Dendy [1892 B}. 


2. V. MAcERA Carter. Type species of the genus. 
Heteropia macera Carter [1885-1886 }. 
Vosmaeropsis macera Dendy [1892 B}. 


3. V. witsont Dendy. 
Vosmaeropsis wilsont Dendy [1892 B]. 


Proc. Zoou. Soc.—1913, No. L. 50 


756 PROF. A. DENDY AND MR. R. W. H. ROW ON 


Srecrion B. With large, usually radially arranged oxea, 
but without microxea. 


Although the author’s description does not conform to the 
above diagnosis, we include Poléjaefi’s Leucilla connexiva in this 
section of the genus, for the figures given by him show oxea 
present, though no reference is made to them in the text, and 
our own examination of the type specimen revealed the presence 
of occasional trichoxea. 


4. V. connEXIvA Poléjaeff. 
Leucilla connexiva Poléjaetf [1883]. 


5. V. cyaruus Verrill. 
Leucandra cyathus Verrill [1873]. 


6. V. DENDYI How. 
Vosmaeropsis dendyi Row [1913 MS8.]. 


7. V. primitiva Row. 
Vosmaeropsis primitiva Row [1913 MS.]. 


8. V. spericatum Lidley. 
Aphroceras sericatum Ridley [1884]. 

This species has been placed in Vosmaeropsis as a result of an 
examination of the type specimen made by us at the Natural 
History Department of the British Museum, which revealed the 
existence of typical subdermal pseudosagittal triradiates, though 
the author’s original description does not mention them. 


Genus 22. GranrittA Row [1909] (emend.). 


Diagnosis. Canal system syconoid, 'Tubar skeleton ( ? always) in- 
articulate, composed of subdermal pseudosagittal triradiates 
and subgastral sagittal triradiates, supplemented by 
subdermal quadriradiates. No colossal longitudinal oxea. 


For illustrations of this genus see Row [1909]. 


This gens was originally proposed by Row for two species, 
G@. quadriradiata and G. hastifera, which were supposed to possess 
certain features that necessitated the provision of a new family, 
Grantillide. We now consider, however, that the characters in 
question do not represent any fundamental peculiarities of 
structure, and we have therefore abandoned the family, as already 
stated in the Introduction. 

One of the two species originally assigned to Grantilla, G. 
qguadriradiata, however, presents an association of subdermal 
quadriradiates with subdermal pseudosagittal triradiates, which is 
not known in any other species of calcareous sponge, and we 
therefore retain the name Grantilla for this species with an 
emended diagnosis. The development of subdermal quadri- 
radiates has evidently taken place repeatedly in the phylogeny of 


CALCAREOUS SPONGES. 757 


the Calearea. We have seen it already, for example, in Lewcetta, 
Leucaltis and Leucettusa, and have not in those cases considered 
the presence of such spicules as of generic value. In Grantilla, 
however, they seem to assume more importance, and to take a 
larger share in the formation of the skeleton of the chamber layer. 
Nevertheless, had the genus not been already in existence, we 
should have hesitated to propose it on this character alone. 


The only known species of the genus is :-— 


1. G. QUADRIRADIATA Row. 
Grantilla quadriradiata Row [1909]. 


Family 8. GRANTIID Ai Dendy [1892] (emend.). 


Diagnosis. With a distinct dermal cortex and a proper cortical 
skeleton of tangential radiates, sometimes supplemented by, 
and occasionally replaced by, oxea. Flagellate chambers 
ranging from elongated and radially arranged to small, 
spherical and irregularly scattered. Skeleton of the chamber 
layer ranging from regularly articulate to irregularly 
scattered. Typically with subgastral sagittal radiates. No 
subdermal pseudosagittal triradiates. Subdermal quadri- 
radiates, if present, always associated with a chamber-layer 
skeleton containing confused triradiates. Nuclei of collared 
cells probably always apical. 


It must frankly be admitted that the boundary line between 
the Sycettide and the Grantiide is by no means sharply defined. 
The great distinguishing feature is the presence in the latter of a 
distinct dermal cortex with its own proper skeleton. The develop- 
ment of such a cortex appears to have formed the determining 
condition for the further evolution of both the canal system and 
the skeleton, and it must therefore be regarded as of great system- 
atic importance. 

The first commencement of such a cortex is, however, so slight 
as to be almost indistinguishable from the mere pore-bearing 
dermal membrane of the most highly specialised Sycons. In 
Grantia compressa the cortex is so feebly developed that Dendy, in 
his early work [1892 B], included this species in the genus Sycon, 
laying more stress upon the presence of dermal tufts of oxea than 
we are now inclined to do in this connection. It appears to us 
that the line between Sycon and Grantia, and therefore between 
the Sycettidee and Grantiide, must be drawn at the appearance 
of a dermal cortical skeleton of tangential radiates distinct from 
the skeleton of the radial chambers, and in accordance with these 
views Grantia compressa is excluded from the genus Sycon. 
Moreover, it must be pointed out that G. compressa is not the 
only member of this family in which dermal tufts of oxea occur, 

‘as they are present also in Sycute dendyi Kirk. 
; 50* 


758 PROF, A. DENDY AND MR. R. W. H. ROW ON 


With the transition from the syconoid to the leuconoid type o 
canal system in this family, and the correlated replacement of th 
articulate tubar skeleton by irregularly scattered radiates, we ge, 
a close approach to the more advanced Leucascide, such at 
Leucetta and Pericharax, and we have here one of those cases o 
convergence which are so frequently met with amongst sponges» 
but we have already laid sufticient emphasis upon this point. 
Even in the genus Leucandra, however, subgastral sagittal tri- 
radiates are usually present, and when they are absent their 
absence must be regarded as secondary. 


So far as our experience goes the nucleus of the collared cells 
is always apical in position in this family. We have been able to 
determine it in 17 species, as enumerated in an earlier section of 
this paper. 


The family is a very large one, comprising no less than 25 out 
of the 51 genera of recent calcareous sponges which we recognise, 
and containing a great diversity of structural types within it. 
There are, however, very great difficulties in the way of dividing 
it into subfamilies, the chief of these being the fact that the 
possible methods of deriving the various genera from one 
another within the family are manifold, and it is impossible 
to determine satisfactorily which are the true lines upon which 
evolution has proceeded. We might, for example, place all 
those genera which have a syconoid canal system and colossal 
longitudinal oxea in the dermal cortex together in a subfamily 
Uteine; or we might separate the genus Uteopsis from the 
others, and unite it with Achramorpha and Anamiwilla in a 
subfamily characterised by the reduction of the tubar skeleton to 
a single joint. But neither of these two possible subfamilies 
would seem to be very sharply defined, and moreover, the cha- 
racters in question are not confined to members of the Grantiidee. 
In short, we feel that in the present state of our knowledge it is 
impossible to decide which method of grouping would express 
most correctly the real affinities of the genera concerned. ‘This 
is the case with almost all the possible methods of grouping the 
genera, and we have therefore decided not to attempt to split up 
the family, but merely to indicate the approximate relationships 
of the genera, so far as this is possible in a linear series, by the 
order in each we have arranged them, 

Although it seems probable that the majority of the genera in 
this family are descended from the genus Sycon, yet it is quite 
possible that some of them may be descended independently from 
Sycetta, and therefore that the family may be of diphyletic 
origin. 


We have changed the name of the family from Grantide to 
Grantiide in accordance with the usual practice of systematic 
zoologists. 


CALCAREOUS SPONGES. 759 


Genus 23. Grant1A Fleming [1828] (emend.). 


Diagnosis. Canal system syconoid. Colossal longitudinal oxea, 
if present, projecting from the surface. Tubar skeleton 
articulate, composed of radiate spicules, which may or may 
not be supplemented by oxea. 


For illustrations of this genus see Dendy [1893 A ]. 


It has beeri conclusively shown by Minchin [1896] that the 
type species of the genus Grantia is G'. compressa, and that the 
name Grantia must always be given to the group of species 
associated with G. compressa. We have already pointed out that 
this species has a definite, though slight, dermal cortex, and 
that its true position is in the present family, and in fact, in the 
genus Grantia as defined by Dendy in 1892. 

We may point out that G. intermedia Thacker stands alone in 
the genus, as far as is at present known, in the presence of apical 
rays on the tangential cortical radiates, and although we do not 
attach much importance to such spicules, we feel that their 
presence in this species indicates at any rate a possible starting 
point for the family Amphoriscidee. 


We recognise the following species as belonging to this 
3 § Sp ging 
genus :— 


Section A. With large, usually radially arranged oxea, 
but without microxea. 


1. G. artantica Ridley. 
Grantia atlantica Ridley [1881]. 


2. G. BrevipiLis Haeckel. 
Sycandra cupillosa var. brevipilis Haeckel [1872]. 


3. G. CANADENSIS Lambe. 
Grantia canadensis Lambe | 1896]. 


4, G. caprntosa O. Schmidt. 
Ute capillosa O. Schmidt [1862]. 
Sycandra capillosa Haeckel [1872]. 


5, G, CHARLACEA Jenkin. 
Dermatreton chartaceum Jenkin [1908 B]. 


6. G. CLAVIGERA O. Schmidt. 
Sycinula clavigera O. Schmidt [1870]. 
Sycandra clavigera (Sycandra compressa var. clavigera) 


Haeckel [1872]. 


7. G. comoxensis Lambe. 
Grantia comoxensis Lambe [1893]. 


760 
8. 


10. 


11. 


13. 


14. 


18. 


19, 


20. 


2l. 


G. 


Ge 


G. 


G. 


PROF. A. DENDY AND MR. R. W. H. ROW ON 


compressa Fabricius. Type species of the genus. 
Spongia compressa Fabricius [1780]. 
Sycum lingua Haeckel [1870], fide Haeckel [1872]. 
Sycarium rhopalodes Haeckel [1870], fide Haeckel [1872]. 
Sycandra compressa Haeckel {1872}. 
Sycon compressum Dendy [1892 B]. 


. FOLIACEA Montagu. 


Spongia foliacea Montagu [1812]. 
Sycandra foliacea (Sycandra compressa var. foliacea) 


Haeckel [1872]. 


. GENUINA Row. 


Grantia genuina Row [1913 MS.]}. 


. GRACILIS von Lendenfeld. 


Vosmaeria gracilis von Lendenfeld [1885 B]. 


. HODGSONT Jenkin. 


Dermatreton hodgsoni Jenkin [1908 B}. 


.INtTERMEDIA Zhacker. 


Grantia intermedia Thacker [1908]. 


. LOBATA Haeckel. 


Sycandra lobata (Sycandra compressa var. lobata) Haeckel 


[1872]. 


. toners Haeckel. 


Sycandra capillosa vax. longipilis Haeckel (1872). 


. MonsTRUOSA Dreitfuss. 


Grantia monstruosa Breitfuss [1898 B]. 


PENNIGERA Haeckel. 
Sycandra pennigera (Sycandra compressa var. pennigera) 


Haeckel [1872]. 


scorrt Jenkin. 
Tenthrenodes scotti Jenkin [1908 B]. 


. tenuis Urban. 


Grantia tenuis Urban [1908]. 


. VOSMAERL Dendy. 


Grantia vosmaeri Dendy [1892 B]. 


Section B. Without any oxea. 


AscoNoIbDES bretifuss. 
Sycetta asconoides Breitfuss [1896 B]. 


Our reasons for placing this species under Grantia rather 


than under Sycetta have been stated when discussing the 
latter genus. 


CALCAREOUS SPONGES. 761 


22. G. cuputa Haeckel. 
Sycetta cupula Haeckel [1872]. 


23. G. iInvENuSTA Lambe. 
Grantia invenusta Lambe [1900 B]. 


24, G. strositus Haeckel. 
Sycetta strobilus Haeckel [1872]. 


Section C. With large, usually radially arranged oxea, 
and with microxea. 


25. G. acuLEATA Urban. 
Grantia aculeata Urban [1908]. 


26. G. EXTUSARTICULATA Carter. 
Hypograntia extusarticulata Carter [1885-1886]. 
Grantia extusarticulata Dendy [1892 B]. 


27. G. inpica Dendy. 
Grantia indica Dendy [1912]. 


28. G. MIRABILIS [ristedt. 
Ascandra mirabilis Fristedt [1887]. 
Grantia mirabilis Lundbeck [1909]. 


29. G. tuBEROSA Poléjaeff. 
Grantia tuberosa Poléjaeff [1883]. 


Section D. With microxea, but without large oxea. 


30. G. pavieata Haeckel. 
Sycortis levigata Haeckel [1872]. 
Sycortusa levigata von Lendenfeld [1885 B]. 


31. G. painiipsir Lambe. 
Grantia phillipsii Lambe [1900 B]. 


The following are doubtfully assigned to this genus :— 
32. G. sINGULARIS Breitfuss. 
Sphenophorina singularis Breitfuss [1898 B]. 
The genus Sphenophorina is discussed at some length in the 
list of rejected genera. 


33. G. uRcEOLUS Miller. 
Spongia urceolus Miller [1788-1796]. 
Stated by Johnston [1842] to be very probably a variety of 
Grantia compressa. 


Genus 24. T'kICHONOPSIS nov. 


Diagnosis. Sponge consisting of a single stipitate person with 
enormously expanded gastral cavity and thin, much folded 


162 PROF. A. DENDY AND MR. R. W. H. ROW ON 


wall, whose convoluted edge represents the oscular margin. 
Canal system syconoid. Tubar skeleton articulate. Without 
colossal longitudinal oxea. 


For illustrations of this genus see Dendy [1891 B]. 


We propose this genus for the reception of the remarkable 
species 7’. labyrinthica, usually known as Grantia labyrinthica, 
which forms the subject of a special memoir by one of us, Dendy 
[1891 B]. We now consider that the very peculiar external form 
is of sufficient importance to justify generic separation. The 
species was originally placed by Carter in his genus Zeichonella, 
on account of some superficial resemblance to his 7. prolifera; 
but although the name TZeichonella has now been universally 
abandoned even for Leucetta (Teichonella) prolifera, we do not 
consider ourselves justified in reviving it for Grantia labyrinthica, 
for Mr. Carter himself subsequently dissociated this species from 
Teichonella and placed it in the genus Grantia [1885-1886]. 


The only known species is :— 


1, T. LABYRINTHICA Carter. 
Teichonella labyrinthica Carter | 1878]. 
Grantia labyrinthica Dendy [1891 B]. 


Genus 25. Grantropsis Dendy [1892 B]. 


Diagnosis. Canal system syconoid. Dermal cortex as thick as 
the chamber layer, with many layers of tangential triradiates. 
Tubar skeleton articulate, the proximal joint being composed 
of subgastral sagittal quadriradiates (? or triradiates), the 
other joints of sagittal triradiates practically reduced to 
the basal ray by suppression of the paired rays. Without 
colossal longitudinal oxea. 


For illustrations of this genus see Dendy [1893 A] and Row 
[1913 MS.]}. 


This genus was first proposed by Dendy [1892 B] for his 
Grantiopsis cylindrica, and was considered by him to be a sub- 
genus of Grantia. Jenkin [1908 Bj placed it as a distinct genus 
in his family Staurorrhaphide, on the ground that the subgastral 
spicules were “‘chiactines.” As we cannot accept the chiact 
theory, we again transfer the genus to the Grantiide, but 
consider it sufficiently distinct from Grantia to deserve generic 
recognition. 

We have recently discovered, as the result of our study of 
Mr. Carter’s MS. illustrations, in the possession of one of us, 
that that author’s “ Hypograntia infrequens (incertze sedis)” is 
undoubtedly a species of Girantiopsis, and the same species has 
recently turned up again in the collection made by the Hamburg 


CALCAREOUS SPONGES. 763 


South-Western Australian Expedition of 1905. As neither 
Mr. Carter’s Hypograntia nor the species H. infrequens were 
ever recognisably diagnosed, we do not consider it necessary to 
abandon the generic name Grantiopsis. The question will be 
more fully dealt with in the forthcoming report on the above- 
mentioned collection [Row 1913 MS.]. 


We recognise the following species in this genus :-— 


1. G. cyninprica Dendy. Type species of the genus. 
Grantiopsis cylindrica Dendy [1892 B]. 


2. G. INFREQUENS Carter. 
Hypograntia infrequens Carter [1885-1886 }. 
Grantiopsis infrequens Row [1913 MS.]. 


Genus 26. SycurTE nov. 


Diagnosis. Canal system syconoid. Dermal cortex provided with 
colossal longitudinally arranged oxea. ‘Tubar skeleton 
articulate. Distal ends of the flagellate chambers crowned 
with tufts of oxea lying between the colossal longitudinal 
oxea. 


For illustrations of this genus see Kirk [1894]. 


This genus has been provided for Kirk’s Sycon dendyi, a species 
which is curiously intermediate in character between Sycon and 
Ute, retaining the well-defined tufts of oxea which are charac- 
teristic of Sycon and at the same time possessing the colossal 
longitudinal oxea characteristic of Ute. 


The only known species is :— 


1 S. penpy1 Kirk. 
Sycon dendyi Kirk [1894]. 


Genus 27. Urs O. Schmidt [1862] (emend.). 


Diagnosis. Canal system syconoid. Tubar skeleton articulate. 
Dermal cortex well developed, containing colossal longi- 
tudinal oxea. No tufts of oxea at the distal ends of the 
flagellate chambers. 


For illustrations of this genus see Dendy [1895 A]. 
We allocate the following species to this genus :— 


Srcrion A. Without microxea. 


1. U. ensata Bowerbank. 
Grantia ensata Bowerbank [1864-1882]. 
Sycandra glabra var. ensata Haeckel [1872]. 


764 PROF. A. DENDY AND MR. R. W. H. ROW ON 


2. U. atapra O. Schmidt. Type species of the genus. 
Ute glabra O. Schmidt [1864]. 
Ute capillosa J. K. Gray [1867], fide Haeckel [1872]. 
Sycandra glabra Haeckel [1872]. 


3. U. rniaipa Haeckel. 
Sycandra glabra var. rigida Haeckel [1872]. 


4, U. SYCONOIDES Carter. 
Aphroceras syconoides Carter [1885-1886]. 


Section B. With microxea. 


dD. U. spENCERI Dendy. 
Ute spencert Dendy [1892 B]. 


6. U. sprcunosa Dendy. 
Ute spiculosa Dendy [1892 B]. 


Genus 28. Synure Dendy [1892 A]. 


Diagnosis. Sponge consisting of many Ute-like individuals com- 
pletely fused together, and invested with a common cortex 
containing colossal longitudinal oxea. 


For illustrations of this genus see Dendy [1893 A]. 


This genus represents the highest known type of integration 
met with amongst syconoid sponges. 


The only known species is :— 


1. 8. puncHELLA Dendy. 
Synute pulchella Dendy [1892 A]. 
Synute pulchella Row [1913 MS.]. 


Genus 29. Sycoporus Haeckel [1872] (emend.). 
Utella Dendy [1892 B}. 


Diagnosis. Canal system syconoid. ‘Tubar skeleton articulate. 
Gastral cortex with a layer of large longitudinally arranged 
oxea, but no oxea in the dermal cortex. 


For illustrations of this genus see Haeckel [1872]. 


Dendy proposed the genus Utella in 1892 for the reception of 
Haeckel’s Sycandra hystrix, and suggested that O. Schmidt’s 
Ute utriculus might also be included in it. As we feel that the 
laws of priority necessitate our using Haeckel’s subgeneric names, 
where possible, in preference to later ones, we propose to substitute 
Sycodorus for Utella, the species which precede S. hystria in the 
subgenus in Haeckel’s monograph having been assigned to earlier 
genera. For Ute utriculus we have retained the generic name 
Sycandra. 


CALCAREOUS SPONGES, 765 


The only known species is :— 


1. 8. nystrix Haeckel. 
Sycandra hystrix Haeckel [1872]. 


Genus 30. AcHrAMorPHA Jenkin [1908 B]| (emend.). 


Diagnosis. Canal system syconoid. Skeleton of the chamber 
layer reduced to the basal rays of the subgastral sagittal 
triradiates (which may become quadriradiates by the addi- 
tion of an apical ray), with radial oxea lying between the 
chambers and projecting from the surface. No colossal 
longitudinal oxea. 


For illustrations of this genus see Jenkin [1908 B]; and Breitfuss 
[1898 D] under Hbnerella schulzer. 


This genus was proposed by Jenkin for the three species 
glacialis, grandinis and nivalis, which resemble one another 
closely, and which all possess the so-called chiactines of his 
supposed family Staurorrhaphide. The fact that another species, 
Breitfuss’s Hbnerella schulzei, differs in no essential point except 
the absence of chiactines, affords strong evidence for our view 
that the latter are nothing but subgastral sagittal triradiates 
that have developed apical rays, and therefore not even of generic 
importance, since such spicules are known to occur in other 
Grantiide. These species, however, form a well-defined group, 
and we accordingly retain the generic name Achramorpha, with 
an emended diagnosis based upon what we believe to be more 
important characters. 


We assign the following species to the genus :— 


Section A. With microxea. 


1. A. enacianis Jenkin. 
Achramorpha glacialis Jenkin [1908 B}. 


2. A. GRANDINIS Jenkin. 
Achramorpha grandinis Jenkin [1908 B}. 


3. A. NIVALIS Jenkin. Type species of the genus. 
Achramorpha nivalis Jenkin [1908 B]. 


4. A. SCHULZEL Greitfuss. 
Hbnerella schulzei Breitfuss [1896 A]. 
Secrion B. Without microxea. 


5. A. rRuNcATA Topsent. 
Grantia truncata Vopsent [1907]. 


766 PROF. A. DENDY AND MR. R. W. H. ROW ON 


Genus 31. UTropsis nov. 


Diagnosis. Canal system syconoid. Tubar skeleton reduced to 
the basal rays of subgastral sagittal radiates, supplemented 
distally by radially arranged oxea. Dermal cortex well 
developed, and containing colossal longitudinal oxea, 


For illustrations of this genus see Poléjaeff [1883]. 


We propose this genus for Poléjaeff’s Ute argentea, which 
obviously differs widely from the other species of the genus Ute. 
The replacement of the distal portion of the tubar skeleton 
by oxea is a very unusual feature, and, from the analogy of 
Grantiopsis, we think it possible, but not probable, that these 
oxea are really radiates whose paired rays have been completely 
lost. It seems more probable that they are to be compared to 
the radial oxea of Achramorpha. 

The ‘‘tubar” quadriradiates referred to by Poléjaeff presumably 
belong to the exhalant canals of the chambers, and not to the 
chambers themselves. 


The only known species of the genus is :— 


1. U. arcentea Poléjaeff. 
Ute argentea Poléjaeff [1883]. 


Genus 32. ANAMIXILLA Poléjaeff [1883]. 


Diagnosis. Canal system syconoid. Tubar skeleton reduced to 
the outwardly directed basal rays of the subgastral sagittal 
radiates. Skeleton of the chamber layer otherwise con- 
sisting of large triradiate spicules, arranged without regard 
to the direction of the chambers. Dermal cortex well 
developed, but without colossal longitudinal oxea. 


For illustrations of this genus see Poléjaeff [1883]. 


As Dendy has previously pointed out [1893 A], this genus may 
be looked upon as a Grantia in which the ordinary tubar skeleton 
has been almost entirely replaced by the invasion of large tri- 
radiates from the dermal cortex. Thus the genus is of interest 
as indicating one method by which the confused chamber-layer 
skeleton of Lewcandra may have arisen. 


The only known species is :— 


1. A. rorrest Poléjaeff 
Anaminilla torresi Poléjaeff [1883]. 


CALCAREOUS SPONGES. 767 


Genus 33. Sycyssa Haeckel [1872]. 


Diagnosis. Canal system syconoid. Skeleton entirely composed 
of oxea. Dermal cortex well developed, but without colossal 
longitudinal oxea. Gastral cortex with a subgastral layer of 
oxea, arranged longitudinally. 


For illustrations of this genus see Haeckel [1872]. 


This genus is highly remarkable for the complete suppression 
of the radiate spicules. An analogous condition is met with in 
Ascyssa, Leucyssa, Trichogypsia and Kuarrhaphis. 


The only known species is :— 


1. S. suxtEey1 Haeckel. 
Sycyssa huxleyi Haeckel [1872]. 


Genus 34. Mrcaprocon Jenkin [1908 B] (emend.). 


Magnosis. Canal system sylleibid or leuconoid. Skeleton of the 
chamber layer retaining clear traces of the original articulate 
character and not confused ; composed chiefly of subgastral 
sagittal quadriradiates, with their apical rays projecting into 
the gastral cavity; with a few sagittal trivadiates arranged 
as usual. No gastral skeleton of tangentially placed radiates, 
except round the osculum. No colossal longitudinal oxea. 


For illustrations of this genus see Jenkin [1908 B]. 


Jenkin included in this genus five species, W/. cruciferus, 
M. villosus, M. raripilus, M. pollicaris and M. crispatus, and 
placed it in the family Staurorrhaphide on account of the 
presence of so-called chiactines. His figure of J/. villosus, 
however, 1s alone sufficient to indicate that the “ chiactines” 
are merely subgastral sagittal radiates which have developed 
apical rays, as In so many other cases, and we find it necessary 
to base the genus, which we believe to be a natural one, on other 
characters. 

The absence of tangentially arranged gastral radiates, combined 
with the presence of the so-called ‘“chiactines,” might be used as 
an argument for the validity of the chiact theory, on the sup- 
position that all the gastral tangential radiates had been converted 
into chiactines, but we must remember that in one species at any 
rate, J. raripilus, the so-called chiactines are associated with 
subgastral sagittal triradiates, which differ from them only in 
the absence of an apical ray, and there is no ground for supposing 
that the chiactines have any special significance. It is quite 
possible that all these subgastral sagittal spicules have been 
rotated into their present positions, as already pointed out in the 
Introduction, but this fact does not justify us in distinguishing 


768 PROF. A, DENDY AND MR. R. W. H. ROW ON 


the chiactines as fundamentally different from other subgastral 
sagittal radiates. 


Megapogon villosus, with its sylleibid canal system and almost 
syeonoid skeleton, exhibits a very interesting stage in the 
evolution of the leuconoid type, and the same is perhaps true 
of MW. pollicaris. 


We place the following species in this genus :— 


1. M. crispatus Jenkin. 
Megapogon crispatus Jenkin [1908 B]. 


2. M. crucirerus Poldjaeff. Type species of the genus. 
Leuconia crucifera Poléjaeff [1883]. 


3. M. PoniicaRris Jenkin. 
Megapogon pollicaris Jenkin [1908 B]. 


4, M. rAripiLus Jenkin. 
Megapogon raripilus Jenkin [1908 B}. 


5. M. vinLosus Jenkin. 
Megapogon villosus Jenkin [1908 B}. 


Genus 35. Leucanpra Haeckel [1872] (emend.). 


Diagnosis. Sponge usually a single person, or a colony of such 
persons in which the component individuals are readily 
recognisable. Canal system leuconoid. Skeleton of the 
chamber layer more or less confused, but frequently with 
vestiges of an articulate tubar skeleton in the form of sub- 
gastral or other sagittal trivadiates. Dermal skeleton of 
tangentially placed triradiates, which may sometimes develop 
an apical ray. Colossal longitudinally placed oxea, when 
occurring in the dermal cortex, never forming a smooth 
layer, but always projecting conspicuously from the surface. 


For illustrations of this genus see Vosmaer [1880] and Dendy 
[1893 A]. 


The genus Leucandra as here defined is much more narrowly 
circumscribed than it was by Dendy previously [1892 B]. In 
fact Dendy’s genus is here represented by no less than 10 genera, 
namely, Lewcandra, Baeria, Leucopsila, Aphroceras, Leucettaga, 
Lamontia and Hilhardia in the family Grantiide, and Leuco- 
malthe, Pericharax and Leucettusa in other families, while 
certain species have been transferred to Leucetta. On the other 
hand, we include in the present genus certain species which 
possess subdermal quadriradiates, and which on that account 
were placed by Dendy in the genus Leweilla; for, as we had 
occasion to point out with regard to both Leucetta and Grantia, 


CALCAREOUS SPONGES. 769 


we do not consider that the mere presence or absence of a fourth 
ray on a radiate spicule in the dermal cortex can be regarded as 
of generic import. We shall discuss the true characteristics of 
Leucilla when dealing with that genus. 

The genus Leucandra as now defined may be regarded as 
derived from a Grantia-like ancestor by the conversion of 
the syconoid canal system into a leuconoid one, with the 
simultaneous replacement of the articulate tubar skeleton by 
an irregularly scattered skeleton of the chamber layer (compare 
Anamixilla). Indications of the syconoid ancestry can, however, 
frequently be detected in the skeleton (compare d/egapogon), 
while as regards canal system such species as L. australiensis 
Carter and ZL. infesta sp. n. (Leucilla intermedia Row [1909 )), 
which are of the so-called sylleibid type, form connecting links 
between Grantia and Leucandra. 


We recognise the following species as belonging to this 
genus :— 


Section A. With large, usually radially arranged oxea, 
but without microxea. 


1. L, ananas Montagu. 
Spongia ananas Montagu [1812]. 
Spongia pulverulenta Grant [1826], fide Haeckel [1872]. 
Scypha ovata 8. F. Gray [1821], fide Haeckel [1872]. 
Sycinula penicillata O. Schmidt [1870], fide Haeckel 
[1872]. 
Leucandra ananas Haeckel [1872]. 


. ANGUINEA Lidley. 
Leucortis anguinea Ridley [1884]. 


bo 
& 


3. L. anomMALA Haeckel. 
Leucetta pandora var. anomala Haeckel {1872}. 


4, L. anmata Urban. 
Leuconia armata Urban [1908]. 


5. L. aspera O. Schmidt. 
Sycon asperum O. Schmidt [1862]. 
? Spongia panicea Esper [¢], fide Haeckel [ 1872}. 
¢Spongia inflata Delle Chiaje [1828], fide Haeckel 
[1872]. 
Leucandra aspera Haeckel [1872]. 


6. L. AUSTRALIENSIS Carter. 
Leuconia fistulosa var. australiensis Carter | 1885-1886]. 
Leucandra austrahiensis Dendy [1892 B]. 


7. L. caminus Haeckel. 
Dyssyconella caminus Haeckel [1870]. 
Leucandra caminus Haeckel (1872). 


770 PROF. A. DENDY AND MR. R. W. He ROW ON 


8. L. capiniavra Poléjaeff. 
Leuconia multiformis var. capillata Poléjaeft [1883 ]. 


2 
eS 


. CIRRATA Jenkin. 
Leucandra cirrata Jenkin [1908 B]. 


10. L. crrrsosa Urban. 
Leuconia cirrhosa Urban [1908]. 


11. L. cuavirormis Schuffner. 
Leucandra claviformis Schuftner [1877]. 


12. L. compacta Carter. 
Leuconia compacta Carter [1885-1886]. 


13. L. cramBessa Haeckel. 
Leucandra crambessa Haeckel [1872]. 


14. L. crusracea Haeckel. 
Leucaltis crustacea Haeckel [1872]. 


15. L. cuUMBERLANDENSIS Lambe. 
Leucandra cumberlandensis Lambe | 1900 B). 


16. L. ponnant Dendy. 
Leucandra donnant Dendy [1905]. 


17. L. ecurnara Schuffner. 
Leucandra echinata Schuftner [1877]. 
Leuconia echinata Carter [1885-1886], fide Dendy 
[1913]. 
Leucandra echinata Dendy [1913]. 


18. L. ncepm O. Schmidt. Type species of the genus. 
Sycinula egedit O. Schmidt [1870]. 
Leucandra egedii Haeckel | 1872]. 


19. L. rancienra Schugfner. 
Leucandra falcigera Schuftner [1877]. 


20, L. risruLosa Johnston. 
Grantia fistwlosa Johnston [1842]. 
Leucandra fistulosa Haeckel [1872]. 


21. L. cemmrpara Thacker. 
Leucandra gemmipara Thacker [1908]. 


22. L. gosser Bowerbank. 
Leucogypsia gosset Bowerbank [1864-1882]. 
Leucandra gosset Haeckel [1872]. 


23. L. HIBERNA Jenkin. 
Leucandra hiberna Jenkin [1908 B}. 


24. L. uirsuta Topsent. 
Leucandra hirsuta Topsent [1907]. 


CALCAREOUS SPONGES. Con 


25. L. HisPipA Carter. 
Leuconia hispida Carter [1885-1886]. 


26. Li. INFESTA, sp. 7. 
Leucilla intermedia Row [1909]. 
The new name is given to this species as intermedia is already 
occupied, 


27. LL. KERGUELENSIS Urban. 
Leucandra kerguelensis Urban [1908]. 


28. L. LENDENFELDI Greitfuss. 
Leuconia lendenfeldi Breitfuss [1897]. 
Leucortis elegans von Lendenfeld, MS., fide Breitfuss 
[1897]. 


29. L. tunuuatTa Haeckel. 
Leucandra lunulata Haeckel [1872]. 


30. L. MASATIERRE Breitfuss. 
Leuconia masatierre Breitfuss [1898 E]. 


31. L. MEANDRINA von Lendenfeld. 
Leucandra meandrina von Lendenfeld [1885 B]. 


32. L. minima Low. 
Leucandra minima Row [1913 MS. }. 


33. L. mutrirormis Poléjaeff. 
Leuconia multiformis Poléjaeff | 1883}. 


34, L. pHItiirensis Dendy. 
Leucandra phillipensis Dendy [1892 B}. 


35. L. pauyipa Row. 
Leucandra pallida Row [1913 MS. ]. 


36. L. punyinar Haeckel. 
Sycolepis pulvinar Haeckel {1870}. 
Mlea dohrnii Michlucho-Maclay, MS8., jide Haeckel 
[1872]. 
Leucortis pulvinar Haeckel | 1872). 


37. L. rHULAKOMORPHA Low. 
Leucandra thulakomorpha Row [1913 MS.]. 


38. L. vactnata von Lendenfeld. 
Leucandra vaginata von Lendenfeld [1885 B]. 


39. L. vauipa Lambe. 
Leucandra valida Lambe [1900 B]. 


40, L. vituosa von Lendenfeld. 
Leucandra villosa von Lendenfeld {1885 B]. 


Proc. Zoou. Soc.—1913, No. UL. 5] 


772 PROF. A. DENDY AND MR. R. W. H. ROW ON 


41. L. wastnensis Jenkin. 
Leucilla wasinensis Jenkin [1908 A]. 
Leucandra wasinensis Dendy [1913]. 


Secrion B. With large, usually radially arranged oxea, 
and with microxea. 


42, L. amorpHa Poléjueff. 
Leuconia multiformis var. amorpha Polejaeff [1883]. 


43. L. anrracta Urban. 
Leuconia anfracta Urban [1908]. 


44, LL, aprcatis Urban. 
Leucandra apicalis Urban [1905]. 


45. L. BaLEARICA Lackschewitsch. 
Leuconia balearica Lackschewitsch [1886 }. 


46. L. BuLtBosa Hanttsch. 
Leucandra bulbosa Hanitsch [1895]. 


47. L. coImBr& Greitfuss. 
Leuconia coimbre Breitfuss [1898 C}. 


48. L. conica von Lendenfeld. 
Leucandra conica von Lendenfeld [1885 B]. 


49. L. catnHA Haeckel. 
Leucandra crambessa var. callea Haeckel [1872]. 


90. L. crossLanpi Thacker. 
Leucandra crosslandi Thacker [1908]. 


D1. L. cytinprica Fristedt. 
Leucandra cylindrica Fristedt [1887 ]. 


02. L. FERNANDENSIS Breitfuss. 
Leuconia fernandensis Breitfuss [1898 EK]. 


53. L. euapiator Dendy. 
Leucandra gladiator Dendy [1892 B]. 


54. L. neatuit Urban. 
Leucandra heathii Urban [1905]. 


55. L. sousrnt Topsent. 
Leucandra joubini Topsent [1907 |. 


56. L. toricata Polejaeff. 
Leuconia loricata Poléjaeff [1883]. 


57. L. minor Urban. 
Leucoma minor Urban [1908]. 


58. L. puater Breitfuss. 
Leuconia platei Breitfuss [1898 EF]. 


09. 


60. 


66. 


Ms. Ay, 


CALCAREOUS SPONGES. 173 


. PYRIFORMIS Lambe. 


Leuconia pyriformis Lambe [1893]. 


. RoDRIGUEZIL Lackschewittsch. 


Leuconia rodriguezti Lackschewitsch [1886]. 


. RUDIFERA Poléjaeff. 


Leuconia rudifera Polejaeft [1883]. 
Leucandra rudifera Thacker [1908]. 


spissA Urban. 
Leuconia spissa Urban [1908]. 


. TAYLORI Lambe. 


Leucandra taylori Lambe [1900 A}. 


. tyPicA Poléjaeff. 


Leuconia typica Poleéjaeff [1883]. 


. vitREA Urban. 


Leuconia vitrea Urban [1908]. 


Secrron C. With microxea, but without large oxea. 


. JOHNSTON Carter. 


Grantia nivea var., Johnston [1842]. 
Leuconia johnston Carter [1871 B}. 
Leucandra johnstonii Haeckel [1872]. 


. LOBATA Carter. 


Leuconia lobata Carter [1885-1886]. 


. MULTIFIDA Carter. 


Leuconia multifida Carter [1885-1886 ]. 


. NIVEA Grant. 


Spongia nivea Grant [1825-1826]. 
Leucandra nivea Haeckel [1872]. 


. ovata Poléjaeff. 


Leuconia ovata Polejaeff [1883]. 


. PRAVA Breitfuss. 


Leucona prava Breitfuss [1898 C}]. 


Secrion D. Without oxea of any kind. 


. BATHYBIA Haeckel. 


Dyssycum periminum Haeckel [1870], jfide Haeckel 
(1872). 

Leucaltis bathybia Haeckel [1872]. 

Grantia arabica Michlucho-Maclay, MS., fide Haeckel 
eas], 


The earlier of Haeckel’s names for this species, periminum, is 


a nomen nudum, as it never was accompanied by a diagnosis 


51% 


V74 PROF, A. DENDY AND MR. R. W. H. ROW ON 


73. L. BRUMALIS Jenkin. 
Leucandra brumalis Jenkin [1908 B]. 


74, L. curva Schuffner. 

Leucandra curva Schuftner [1877]. 
75. L. Fricipa Jenkin. 

Leucandra frigida Jenkin [1908 B]. 
76. L. aeuatinosa Jenkin. 

Lencandra gelatinosa Jenkin [1908 B]. 


77. L. HELENA von Lendenfeld. 
Leucaltis helena von Lendenfeld [1885 B]. 


78. L. impressa Hanitsch. 
Leucaltis impressa Hanitsch [1890]. 
79. IL. INNOMINATA, sp. 2. 
Leucilla crosslandi Row {1909}. 
The new name is necessitated by the fact that the name: 
crosslandi is already occupied in this genus. 


80. L. iwrermepIA Haeckel. 
Leucetta pandora var. intermedia Haeckel {1872}. 


81. L. Levis Poléjaeff’. 
Leuconia levis Poléjaeff [1883]. 
82. L. naustcaAm Schuffner. 
Leucaltis nausicae Schuftner [1877]. 
83. L. panporA Haeckel. 
Leucetta pandora Haeckel [1872]. 
84. L. pumita Dowerbank. 
Leuconia pumila Bowerbank | 1864-1882]. 
Leucaltis pumila Haeckel [1872]. 
85. L. saarrrara Haeckel. 
Leucetta sagittata Haeckel [1872]. 
86. L. sCHAUINSLANDI Prewwisch. 
Leucetta schawinslandi Preiwisch [1904]. 
87. L. rELuM von Lendenfeld. 
Polejna telwm von Lendenfeld [1891]. 
88. L. verpEnsiIs Zhacker. 


The following species are doubtfully assigned to this genus :— 


Leucandra verdensis Thacker | 1908}. 


89. L. mnFLata Delle Chiaje. 


Possibly identical with Lewcandra aspera, fide Haeckel 


Spongia inflata Delle Chiaje [1828]. 


[1872]. 


CALCAREOUS SPONGES. 175 


90. L. paANicEA Esper. 
Spongia panicea Esper, fide Haeckel [1872]. 
Ksper’s original reference to this species has not been 
found by us, but Haeckel [1872] states that the species 1s 
possibly identical with Leucandra aspera. 


Genus 36. Banria Michlucho-Maclay [1870] (emend.). 


Diagnosis. Canal system leuconoid. Skeleton of the chamber 
layer composed almost exclusively of irregularly scattered 
colossal quadriradiates. Microxea present in large numbers, 
and of very characteristic form, being almost always pierced 
with a small hole towards one end. 


For illustrations of this genus see Haeckel [1872]. 


The very characteristic ‘‘needle-eye ” spicules of this genus are 
really triradiates, in which two of the rays are very much reduced 
and have come to le approximately side by side, being actually 
fused at their distal ends. In this way we get a linear spicule 
very slightly swollen at one end, and in the centre of the swelling 
a small hole, the remnant of the space between the two originally 
separate rays. That this is the true explanation of these spicules 
was made abundantly clear from an examination by one of us 
(Row) of a microscopical preparation of the species preserved at 
Jena, for while most of the spicules were found to correspond 
exactly to the type described above, a few of them had the 
reduced rays not fused together but widely open, thus maintaining 
the triradiate condition. Exactly similar spicules occur in 
Kuarrhaphis cretacea (q. v.). 

It may perhaps be pointed out here that these spicules indicate 
a possible way in which all the calcareous monaxon spicules may 
have originated. At any rate their occurrence adds probability 
to the presumption that all calcareous oxea have been derived 
from triradiates In some way or other. 


The only known species of the genus 1s :— 


1. B. ocuorensis Wichlucho-Maclay. 
Baeria ochotensis Michlucho-Maclay [1870]. 
Leucandra ochotensis Haeckel [1872]. 


Genus 37. LEuUCOPSILA nov. 


Diagnosis. Canal system leuconoid. Skeleton of the chamber 
layer composed almost exclusively of irregularly scattered 
colossal quadriradiates. Gastral cortex well developed, but 
without any radiate spicules, the whole of the gastral skeleton 
being formed of a dense layer of microxea, 


776 PROF. A. DENDY AND MR. R. W. H. ROW ON 


For illustrations of this genus see Haeckel [1872]. 


The very peculiar character of the skeleton of the gastral cortex 
seems to justify the erection of a new genus for O. Schmidt's: 
Leuconia stylifera. 


The only known species is :— 


1. L. srynirera O. Schmid. 
Leuconia stylifera O. Schmidt [1870]. 
Leucandra stylifera Haeckel [1872]. 


Genus 38. APHROCERAS Gray [1858]. 


Diagnosis. Sponge usually a single person or a colony of such 
persons in which the component individuals are readily re- 
cognisable. Canal system sylleibid or leuconoid. Skeleton 
of the chamber layer more or less confused, but frequently 
with vestiges of an articulate tubar skeleton in the form of 
subgastral or other sagittal radiates. Dermal skeleton of 
tangentially placed triradiates supplemented by colossal oxea 
placed longitudinally and not projecting from the surface 
sufficiently to render it hispid. 


For illustrations of this genus see von Lendenfeld [1891], 
under Vosmaeria corticata. 


The genus Aphroceras was originally proposed by Gray [1858] 
to receive a sponge from Hong-Kong, whose chief characteristics,, 
at any rate from our point of view, were the leuconoid canal 
system and the colossal longitudinal oxea of the dermal cortex. 
Since then other species which combine these characters have 
been described, and, although recent authors have not seen fit to 
accept this genus, we feel that these species form a very natural 
eroup, and we consider that the characters distinguishing it are 
sufficiently well defined to render it possible to separate it from 
its nearest ally, Lewcandra. In fact, almost the only species that 
presents any difficulty is Haeckel’s Leweandra crambessa and its 
varieties, in which the colossal oxea are not arranged longi- 
tudinally, but lie scattered quite irregularly over the surface of 
the sponge. ‘This condition is somewhat intermediate between 
that of some Leucandras and that of Aphroceras, and we prefer 
to place this species in the genus Leucandra. 

It may be advisable to state that we do not consider Aphroceras 
to have been derived from U¢e or a Ute-like form, but directly 
from an ancestral Leucandra. 


We recognise the following species as belonging to this. 
genus :— 


CALCAREOUS SPONGES. COU 


Section A. Without microxea. 


1. A. auctcornis Gray. Type species of the genus. 
Aphroceras alcicornis Gray [1858]. 
Cyathiscus actinia, Haeckel [1870], fide Haeckel [1872]. 
Leucandra aleicornis Haeckel [1872]. 


2. A. carapHracta Haeckel. 
Leucandra cataphracta Haeckel [1872]. 


3. A. ELoNGATA Schuffner. 
Leucandra elongata Schuftner [1877]. 


Section B. With microxea. 


4. A. cusprrosaA Haeckel. 
Leucandra aleicornis var. cespitosa Haeckel [1872]. 


5. A. CLIARENSIS Stephens. 
Leucandra cliarensis Stephens [1912]. 


6. A. corricAta von Lendenfeld. 
Vosmaeria corticata von Lendenfeld [1891]. 


Genus 39. Leucerraca Haeckel [1872] (emend.). 


Diagnosis. Canal system leuconoid. Skeleton almost entirely 
composed of a confused mass of triradiates, which are mostly 
irregular and which form the dermal cortical skeleton as 
well as the skeleton of the chamber layer. Gastral cavity 
traversed by numerous endogastric septa, which possess a 
special skeleton of their own in the form of minute radiates. 


For illustrations of this genus see Haeckel [1872]. 


This genus is proposed for the reception of Haeckel’s Leucetta 
pandora var. loculifera, which is the only known species. It 
affords an example amongst leuconoid Calcarea of that remarkable 
development of endogastric septa which occurs also in certain 
Leucosolenias (e. g. ZL. wilsoni) among the Homoceelide, in 
Leucetiusa among the Leucaltide, and in Sycandra among the 
Sycettide. We have, however, only considered it necessary to 
attribute generic importance to this character when it is combined, 
as in the present instance, with the presence of an endogastric 
skeleton. 

We have revived this name, which was applied by Haeckel to 
one of the subgenera of his Lewcetta, for this genus. 


The only known species is :— 


1. L. nocunirera Haeckel. 
Leucetta pandora var. loculifera Haeckel [1872]. 


778 PROF, A. DENDY AND MR. R. W. H. ROW ON 


Genus 40. ParatEuctLia Dendy [1892 B]. 


Diagnosis. Canal system leuconoid. Skeleton of the chamber 
layer composed of confused triradiates. Subdermal cavities 
present, supported by an inner and an outer layer of quadri- 
radiates, whose apical rays cross each other in opposite 
directions. Dermal cortex with tangentially placed tri- 
radiates, between which lie large, longitudinally placed oxea. 


For illustrations of this genus see Haeckel [1872]. 


This genus was proposed by Dendy [1892 B] to receive Haeckel’s 
Leucandra cucumis, but abandoned by him in 1893 on the ground 
that it was not sufliciently distinct from Leucilla. He also 
pointed out that Poléjaeff had previously proposed the name 
Pericharax for the same sponge. Further consideration has, 
however, convinced us that the dermal: quadriradiates of 
Leucandra cucumis are not really comparable to the subdermal 
quadriradiates of Leweilla and other Amphoriscide, being related 
solely to the cortex and not to the chamber layer at all. We 
therefore transfer the species to the family Grantiide, with which 
it has much more in common than with any other family of 
Calearea. We know nothing, however, of the position of the 
nucleus of the collared cells. The first mentioned species of 
Pericharax, on the other hand, is P. cartert Poléjaeff [ 1883, p. 19} 
which we have now shown to belong to the Leucascid-Leucaltid 
line of descent. We cannot therefore associate Leucandra 
cucumis with either Zewcilla or Pericharax, and it therefore 
appears to us that as it is clearly distinguished from other 
Grantude by the presence of subdermal cavities with a special 
skeleton, it is necessary to revive the genus Paraleucilla for its 
reception. Haeckel, it is true, placed his Lewcandra cucwmis in 
the subgenus “ Leucogypsa,’ but Bowerbank’s name ‘ Leuco- 
gypsia,” if revived at all, which we think very undesirable, would 
have to be reserved for Leucandra. 


The only known species is :-— 


1. P. cucumis Haeckel. 
Leucandra cucumis Haeckel [1872]. 
Paraleucilla cucumis Dendy [1892 B}. 


Genus 41. Lamontia Kirk [1894]. 


Diagnosis. Sponge consisting of a single person with a specialised 
pore-zone below the terminal osculum. Canal system 
leuconoid. Skeleton of the chamber layer consisting of small 
scattered oxea. Dermal cortex with triradiates in addition 
to oxea. Gastral quadriradiates present. 


For illustrations of this genus see Kirk [1894]. 


CALCAREOUS SPONGES. 779 


The only species of this curious genus is Kirk’s Lamontia zona 
from New Zealand, which perhaps forms a transition from the 
genus Leucandra to the genus Leucyssa. 


1. L. zona Kirk. 
Lamontia zona Kirk {1894}. 


Genus 42, Luucyssa Haeckel [1872] (emend.). 


Diagnosis. Canal system leuconoid. Skeleton entirely composed 
of smooth oxea. 


¥or illustrations of this genus see Haeckel [1872]. 


We can only suppose that this genus, which has only been 
observed by Haeckel, owes its peculiar skeleton to the complete 
suppression of ancestral radiates. 


The only known species is :— 


1. L. sponemtia Haeckel. 
Leucyssa spongilla Haeckel [1872]. 


Genus 43. Tricnocypsi1A Carter [1871 B]. 


Diagnosis. Canal system leuconoid. Skeleton entirely composed 
of spined oxea. 


For illustrations of this genus see Haeckel [1872]. 


In 1871 Carter proposed the genus 77richogypsia for his species 
T. willosa. In 1872, in ‘* Die Kalkschwimme,” Haeckel regarded 
this species as a variety of his Leucyssa inerustans, the specific 
name incrustans having been given by him without description in 
1870 under the genus Sycolepis. The diagnosis of Sycolepis also 
contained no reference to the essential peculiarities of the species 
in question, and was subsequently abandoned by its author. 

The name Z'richogypsia has therefore priority in our opinion 
over both Sycolepis and Leucyssa, and we have retained it here 
for species which, like 7’. villosa, have spined oxea, while using 
Haeckel’s name Leucyssa for those with smooth oxea. 

We consider that Haeckel’s variety lichenoides, which is the 
first variety of his Leweyssa incrustans, 1s specifically distinct from 
Carter’s Trichogypsia villosa, and as the name villosa has priority 
over incrustans, we confine the latter to the form termed by 
Haeckel var. lichenoides. 


We recognise the following species of this genus :— 


1. T. rycrusrans Haeckel. 
Leucyssa incrustans var. lichenoides Haeckel [1872]. 


780 PROF. A. DENDY AND MR. R. W. H. ROW ON 


2. T. vittosa Carter. Type species of the genus. 
Trichogypsia:villosa Carter [1871 B]. 
Leucyssa incrustans var. villosa Haeckel [1872]. 


Tt is doubtful whether Haeckel’s Sycolepis incrustans is really 
a synonym of Zrichogypsia villosa or of 7’. incrustans. 


Genus 44. KUARRHAPHIS nov. 


Diagnosis. Canal system presumably leuconoid. Skeleton com- 
posed exclusively of perforated ‘ needle-eye ” spicules. 


For illustrations of this genus see Haeckel [1872]. 


We propose this name for Haeckel’s Lewcyssa cretacea, which 
differs from all the other species included by him in the genus 
Leucyssa in the remarkable perforation of the ‘“needle-eye” 
spicules. Spicules of practically identical form are found in the 
genus Baeria, and we must refer the reader to that genus fora 
diseussion of their nature and origin. In Baeria they are still 
associated with trivadiates and colossal quadriradiates. Whether 
Kuarrhaphis is to be regarded as derived from a Baeria-like 
ancestor by the complete suppression of the radiate spicules, 
or whether the remarkable ‘“ needle-eye” spicules have arisen 
independently in the two cases, it is impossible to decide. 


The only known species is :— 


1. K. creracea Haeckel. 
Leucyssa cretacea Haeckel [1872]. 


Genus 45. Hinnarptia Poléjaeff [1883]. 


Diagnosis. Sponge calyciform, with pores on the inner and 
oscula on the outer surface of the cup. Canal system leu- 
conoid. Skeleton of the chamber layer confused, composed 
of triradiates of various shapes and sizes, and of microxea. 
Cortex of inner surface with microxea and sagittal trivadiates,. 
cortex of the outer surface with large oxea and sagittal 
triradiates. 


For illustrations of this genus see Poléjaeff [1883]. 


The only known species of the genus is Poléjaeft’s Lilhardia. 
schulzet, a highly remarkable sponge in many ways, the distri- 
bution of the pores and oscula being exactly the reverse of the 
usual condition in cup-shaped sponges, and indicating that the 
sponge cannot be regarded as a single leuconoid person with 
expanded osculum, in the same way as the calyciform Pericharax 
pexiza. 


CALCAREOUS SPONGES. 781 


Although the genus Hilhardia was abandoned by Dendy 
[1892 5B}, we now consider that it is fully entitled to recog- 
nition. 


The only known species is :— 


1. HE. scnuuze1 Poléjaeff. 
Eilhardia schulzei Poléjaeff [1883]. 


Family 9. AMPHORISCID# Dendy [1892 B] (emend.). 


Diagnosis. Flagellate chambers ranging from elongated and 
radially arranged to small, spherical and irregularly scattered. 
With a distinct dermal cortex supported by a skeleton of 
tangentially placed radiates to which oxea may be added. 
Some or all of the dermal radiates with large apical rays, 
which project inwards through the chamber layer to a greater 
or less extent, and form the principal part of its skeleton. 
No articulate tubar skeleton, but sometimes, in the leuconoid 
forms, a confused skeleton of quadriradiates in the chamber 
layer. Nuclei of collared cells probably always apical. 


The most conspicuous feature of this family lies in the large 
dermal or subdermal quadriradiates with centripetally directed 
apical rays. Such spicules may indeed be present in certain 
species of Lewcandra, but in such cases they are always associated 
with a confused chamber-layer skeleton of scattered triradiates, 
which is never the case in the Amphoriscide. If there be a 
confused chamber-layer skeleton in this family it is found to be 
composed of quadriradiates, which presumably have been derived 
from the subdermal and subgastral quadriradiates themselves by 
immigration. 

The evidence seems to indicate that Zeucandra and Leucilla, 
though difficult to separate in practice, owe their resemblance 
largely to convergence, and that each has been independently 
evolved from some syconoid ancestor, in the one ease directly from 
some such form as Grantia, in the other through some such form 
as Amphoriscus. 


In some Amphoriscide large subgastral quadriradiates are 
present, and in others, or even in the same, subgastral sagittal 
triradiates (or quadriradiates) resembling the sagittal radiates 
of the first joint of an articulate tubar skeleton. Whether the 
centrifugally directed ray of the large subgastral quadriradiates 
is homologous with the basal ray of the subgastral sagittal tri- 
radiates, or whether it is an apical ray added to a tangential 
triradiate of the gastral cortex, is a question which we cannot 
decide without further evidence. 


782 PROF, A. DENDY AND MR. R. W. H. ROW ON 


Genus 46. AmpHoriscus Haeckel [1870] (emend.). 


Diagnosis. Canal system syconoid. Without any special root- 
tuft of anchoring spicules. 


For illustrations of this genus see Haeckel [1872] under 
Sycaltis and Sycilla, and Poléjaeff [1883]. 


We recognise the following species in this genus :— 


Section A. Without oxea. 


1, A. curysauis O. Schinidt. 
Ute chrysalis O. Schmidt [1864]. Type species of the 
genus. 


Sycilla chrysalis Haeckel [1872]. 


LS 
LS 


. cyaTuiscus Haeckel. 
Amphoriscus cyathiscus Haeckel [1872]. 
Sycilla cyathiscus Haeckel [1872 . 


3. A. cyLtinDrus Haeckel. 
Sycilla cylindrus Haeckel [1872]. 


4, A. Kryprorapuis Urban. 

Amphoriscus kryptoraphis Urban [1908]. 
5. A. oviparus Haeckel. 

Sycaltis ovipara Haeckel [1872]. 
6. A. semont Breitfuss. 

Amphoriscus semoni Breitfuss [1896 C]. 
7. A. TestipaRus Haeckel. 

Sycaltis testipara Haeckel [1872]. 


8. A. uRNA Haeckel. 
Amphoriscus urna Haeckel [1870]. 
Sycilla urna Haeckel [1872]. 


SEecTION B. With microxea, but without large oxea. 


9. A. BUCCICHIE von Hbner. 
Amphoriscus buccichii von Ebner [1887 ]. 


10. A. ELoncatus Poléjaef. 

Amphoriscus elongatus Poléjaeff [1883]. 
11. A. erucorit von Lendenfeld. 

Hbnerella gregorii von Lendenfeld [1891 }. 


12. A. opiatus Row. 
Amphoriscus oblatus Row [1918 MS8.]. 


CALCAREOUS SPONGES. 783 


Genus 47, Sycutmis Haeckel [1872] (emend.). 


Diagnosis. Canal system syconoid. With a root-tuft of oxea and 
anchoring quadriradiates. 


For illustrations of this genus see Haeckel [ 1872}. 


This is a highly specialised genus of a single species, but had it 
not already been proposed by Haeckel, we should hardly have felt 
justified in distinguishing a special genus on the characters 
available. 


The only known species is :— 


1. 8. synapra Haeckel. 
Syculmis synapta Haeckel [1872]. 


Genus 48, Leucinna Haeckel 1872 (emend.). 


Diagnosis. Canal system sylleibid or leuconoid. Skeleton of the 
chamber layer typically composed of the centripetally and 
centrifugally directed apical rays of subdermal and sub- 
gastral quadriradiates, but subgastral sagittal triradiates and 
confused chamber-layer quadriradiates may be present, while 
the subgastral quadriradiates may be absent. 


For illustrations of this genus see Haeckel [1872] and Dendy 
[1893 A]. 


The resemblance of this genus to some species of Leucandra 
has already been pointed out. It also resembles by convergence 
some species of the genus Leucetta, but may be distinguished by 
the fact that traces of syconoid ancestry are still to be met with 
in the skeleton (e. g., the presence in some species of subgastral 
sagittal triradiates), while the triradiates are not of the cha- 
racteristic regular type occurring in the Leucascide. The position 
of the nucleus of the collared cells, as determined in Leucilla 
australiensis and L. princeps, is, moreover, apical, instead of 
basal as in the Leucascide. 


We recognise the following species as belonging to this 
genus :— 
Section A. Without oxea. 


1. L. AmpHora Haeckel. Type species of the genus. 
Leucilla amphora Haeckel [1872]. 


2. L. AUSTRALIENSIS Carter. 
Leuconia johnston var. australiensis Carter [1885-1886]. 
Leucilla australiensis Dendy [1892 B]. 


784 PROF, A. DENDY AND MR. R. W. H. ROW ON 


3. L. carsuta Haeckel. 
Lipostomella capsula Haeckel [1870]. 
Leucilla capsula Haeckel [1872]. 


Section B. With large radially arranged oxea or 
trichoxea, but without microxea. 


4, L. ecuinus Haeckel. 
Leuculmis echinus Haeckel [1872]. 


5. L. OXEODRAGMIFERA Row. 
Leucilla oxeodragmifera Row [1913 MS.]. 


6. L. princers Row. 
Leucilla princeps Row [1913 MS.]. 


<1 
is 


. PROTEUS Dendy. 
Leucilla protews Dendy [1913]. 


8. L. urer Poléjaeff. 
Leucilla uter Poléjaett [1883]. 


Section C. Without large oxea, but with microxea. 


9. L. Nurriner Urban. 
Rhabdodermella nuttingi Urban [1902]. 


10. L. sAccHarata Haeckel. 
Leucandra saccharata Haeckel [1872]. 


The following species apparently also belongs to this genus, but 
was very inadequately diagnosed :— 


ll. L. Levconipes Bidder. 
ycaltis lewconides Bidder [1891]. 


Family 10. LELAPIIDA# nov. 
Didlytine Kirkpatrick [1911 A]. 


Diagnosis. Canal system presumably always leuconoid. Skeleton 
of the chamber layer containing fibres or bundles of modified 
sagittal triradiates placed side by side, but not cemented 
together. Nuclei of collared cells (presumably always) 
apical. 


This family appears to bea highly specialised offshoot from the 
leuconoid Grantiide. The presence in Lelapia of distinct sub- 
gastral sagittal triradiates and the apical position of the nuclei of 
the collared cells afford very strong evidence in support of this 


CALCAREOUS SPONGES. 785 


view, and necessitate, as we have already pointed out, its wide 
separation from the other so-called Pharetronid sponges. 

The presence of tuning-fork spicules in Lelapia cannot be taken 
as indicating close affinity with the latter, for, as is well known, 
such spicules occur in Haeckel’s Lewcandra (Leucortis) pulvinar 
and L. (Lewcetta) pandora, while they are replaced in Kebira, the 
only other known genus of Lelapiide, by radiates in which the 
oral rays have been practically suppressed. 


Genus 49, LeLapia Gray [1867]. 


Diagnosis. Skeleton of the chamber layer composed of large 
scattered oxea and loose fibres of tuning-fork spicules. 
Dermal skeleton of tangential triradiates and microxea. 
Gastral skeleton of tangential triradiates and quadriradiates. 


For illustrations of this genus see Dendy [1893 B]. 


This genus was originally based by Gray on figures published 
by Bowerbank of the characteristic tuning-fork spicules. Carter 
really first described the sponge, adopting Gray’s name, Lelapia 
australis, for the species which he studied. There is, of course, 
no guarantee that Carter’s species is either generically or 
specifically identical with that which furnished the spicules 
figured by Bowerbank. There is some probability, however, from 
the locality, and from the fact that the spicules are stated by 
Bowerbank to have been “loosely fasciculated,” that the two 
species are really identical, and in any case the genus and species 
may conveniently be retained for the sponge described by Carter. 


The only known species is :— 


1. L. austrauis Gray. 
‘A new species of sponge” Bowerbank [1858-1862]. 
Lelapia australis Gray [1867]. 
Lelapia australis Carter [1885-1886]. 
Lelapia australis Dendy | 1893 B). 


Genus 50. Kepira Row [1909]. 


Diagnosis. Skeleton of the chamber layer composed of large 
longitudinally arranged oxea, and of loose fibres of sagittal 
triradiates whose paired rays are vestigial. Dermal and 
gastral skeleton of tangential trivadiates. 


For illustrations of this genus see Row [1909]. 


The only known species is :— 


1. K. ureorpes Low. 
Kebira uteoides Row [1909]. 


786 PROF. A. DENDY AND MR. R. W. H. ROW ON 


GENERA AND SPECIES “INCERT# SEDIS.” 


Genus 51. Sycanris Haeckel [1872] (emend.). 


Diagnosis? Canal system syconoid (?). Skeleton of the chamber 
layer confused. 

The sense in which we employ this genus is obviously quite 
different from that in which it was used by Haeckel, who based 
it upon the presence of triradiate and quadriradiate spicules and 
the absence of oxea, Most of the species assigned to it by him 
have been relegated to other genera, but his description of the 
sundermentioned species appears to indicate that it possesses 
characteristics separating it widely from any syconoid sponge 
of normal structure, and therefore, although there seems to be 
some similarity between this sponge and Leucascus, we prefer 
to consider it as “‘incerte sedis” rather than to assign any 
definite position to it in our classification. It may be reiated to 
Ananiailla. 


The only known species is :-— 


1. S. pERForATA Haeckel. 
Sycaltis perforata Haeckel [1872]. 


The following species are so inadequately known as to render: 
all attempts to identify them ineffectual :— 


Ute viridis O. Schmidt [1868]. 
Medon barbata Duchassaing and Michelotti [1864]. 
Medon imberbis Duchassaing and Michelotti [1864]. 


The following species have been referred to without de- 
scription :— 


Grantia striatula Bowerbank MS8., referred to by Bowerbank 
[1864—1882, vol. i. p. 233]. 

Leucogypsia algoauensis Bowerbank MS., referred to by Bower- 
bank [1864-1882, vol. i. p. 166]. 

Ute papillosum O. Schmidt, referred to by Gray [1867]. 

(We have not been able to discover any reference to a 
species of this name in any of Schmidt’s papers, and think 
that Gray’s reference is probably a misprint for Ute 
capillosum O. 8.) 


LIST OF REJECTED GENERIC NAMES. 


The following list includes all the generic names that have, so 
far as we are aware, been applied to calcareous sponges, but which 
we have not made use of in this paper. Many of them have been 
used in various senses by different authors, and in such cases all 


- 


CALCAREOUS SPONGES. 187 


the instances which are of systematic importance have been 
referred to, though the list of references is by no means com- 
plete. Under each genus the date and type-species are given, in 
order to simplify the work of determining the true names of 
genera in accordance with the laws of priority, should any of 
these names be revived for future use. Wealso give (in brackets) 
the name under which the type species will be found in the present 


paper. 
ALCYONCELLUM Quoy et Gaimard [1833]. 


Type species, as regards calcareous sponges, 4. gelatinoswm 
de Blainville, (=Sycon gelatinosum). 


The name dAleyoncellum was originally proposed by Quoy and 
‘Gaimard for certain hexactinellid sponges, but was subsequently 
used by de Blainville [1834-1837] to include Sycon (Aleyoncellwm) 
gelatinosum. Gray [1867] has also used the name for a genus of 
calcareous sponges, and with the same type species. The name is 
now regarded as a synonym of both Huplectella and Sycon. 


AmpuHoripium Haeckel [1870]. 
Type species 4. viridis O Schmidt. 
Schmidt’s Ute viridis is absolutely unrecognisable, even 
generically. 


AmpuHorvuLa Haeckel [1870]. 
' Type species A. solida O. Schmidt, (= Leucandra solida). 


ArtyNAS Haeckel [1870]. 
Type species 4. compressus Fabricius, (=Grantia compressa). 


ARTYNELLA Haeckel [1870]. 

Type species 4. compressa Fabricius, (= Grantia compressa). 
ARTYNES Gray [1867]. 

Type species A. compressa Fabricius, (=Grantia compressa). 
Artynium Haeckel [1870]. 

Type species 4. compresswm Fabricius, (=Grantia compressa). 


ARTYNOPHYLLUM Haeckel [1870]. 
Type species 4. compresswm Fabricius, (=Grantia compressa). 


Ascautis Haeckel [1872]. 


Type species dA. canariensis Michlucho-Maclay, (= Leuco- 
solenia canariensis). 


Ascanpra Haeckel [1872]. 
Type species 4. cordata Haeckel, (= Leucosolenia cordata). 


The name Ascandra has been used in almost exactly Haeckel’s 
sense by several subsequent writers, notably von Lendenfeld, 
Breitfuss, Arnesen, and various systematists who have followed 

Proc. Zoo. LION. No. LIT. 52 


788 PROF. A. DENDY AND MR. R. W. H. ROW ON 


the modification of Haeckel’s original system propesed by vor 
Lendenfeld in 1891. The name has also been used by Minchin 
[1896, 1900, &c.] in an entirely different sense for a genus whose 
type, and only, species was A. falcata, placed by us in the genus 
Leucosolenia. In the latter sense it is the equivalent of von Len- 
denfeld's Homandra. 


Ascetta Haeckel [1872]. 
Type species A. primordialis Haeckel, (= Leucosolenia 
primordialis). 

As employed by Haeckel, this genus includes only species 
whose whole skeleton is composed of triradiates ; von Lendenfeld, 
however, has used it [1891] to include all those ascon sponges 
which do not possess oxea. 


AscitLa Haeckel [1872]. 
Type species A. gracilistHaeckel, (= Leucosolenia gracilis). 


AscometrRA Haeckel [1872]. 
Type species A. primordiale Haeckel, (= Leucosolenia 
primordialis). 

The name Ascometira was used by Haeckel, in the “ artificial 
system” given at the end of his ‘Monographie,’ to replace the: 
name Zhecometra used for exactly the same group in his earlier 
‘Prodromus, and for no apparent reason save nomenclatorial 
symmetry. He gave his new genus, however, a different type 
species. 


Ascortis Haeckel [1872]. 
Type species 4. horrida Haeckel, (= Leucosolenia horrida). 


Ascutmis Haeckel [1872]. 
Type species A. armata Haeckel, (= Leucosolenia armata). 


Ascuris Haeckel [1872]. 
Type species A. arrecife Haeckel, (= a variety of Leucoso- 
lenia canariensis). 
ASTROSCLERA Lister [1900]. 
Type species 4. willeyana Lister. 

Although originally described as a member of the Calcarea, this 
sponge is now known to be an aberrant member of the Non- 
ealearea (vide Kirkpatrick [1912)). 

AvLopLtecmMA Haeckel [1870]. 
Type species A. loculoswm Haeckel, (= Leucosolenia loculosa). 


Avtorniza Haeckel [1870]. 


Type species A. witestinalis Haeckel, (= Leucosolenia 
lamarckii). 


CALCAREOUS SPONGES. 789 


Caucisponeia de Blainville [1834-1837]. 

Type species C. compressa Fabricius, (=Grantia compressa). 

This genus was proposed by de Blainyille in exactly the same 

sense as Fleming’s Grantia [1828]. Since the latter genus takes 
priority, Calcispongia becomes merely a synonym of it. 
CiaTHRINA Gray [1867]. 

Type species C. clathrus O. Schmidt, (= Leucosolenia 

clathrus). 


This genus has been employed by Minchin [1896, 1900] as the 
type genus of one of his families of homoccel sponges. We have 
already (p. 718) given our reasons at length for not accepting his 
conclusions. 


CrystotyntHus Haeckel [1870]. 
Type species C. vesicula Haeckel, (= Lewcosolenia vesicula). 


CaNostoMELLA Haeckel [1870]. 
Type species C. caminus Haeckel, (= Leucandra caninus). 


Canostomium Haeckel [1872]. 


Type species C’. crambessa Haeckel, (= Leucandra crambessa). 


Canostomus Haeckel [1872]. 


Type species C. primigenius Haeckel, (= Leucetta primigenia). 


Cyatuiscus Haeckel [1870]. 
Type species C. actinia Haeckel, (= Aphroceras alcicornis). 


DERMATRETON Jenkin [1908 B]. 
Type species D. chartacewm Jenkin, (=Grantia chartacea). 
This genus was proposed by Jenkin for certain species of the 
family Grantiide, which possess ‘‘ linked” chambers. We do not 
consider that this character is of generic rank, and we have there- 
fore included both these species in the genus Grantia. 


Dseppra Michlucho-Maclay MS. (fide Haeckel [1872)]). 
Type species D. violacea Michlucho-Maclay MS., fide Haeckel 
[1872], (=Grantessa stawridea). 
The generic name Djeddew is quoted by Haeckel in the 


synonymy list attached to his Sycetta stawridea. This seems’ to 
be the only authority for the name. 


Dunstervitiia Bowerbank [1845]. 
Type species D. elegans Bowerbank, (=Sycon elegans). 
Bowerbank’s genus was adopted by Haeckel as one of the 
genera of his “artificial” system, but no other author seems to 


have made use of it save Gray. Itis now considered to be merely 
a synonym of Sycon. 


52* 


790 PROF. A. DENDY AND MR. R. W. H. ROW ON 


Dyssycartum Haeckel | 1872}. 
Type species D. egedii O. Schmidt, (= Leucandra egedii). 

In proposing this genus Haeckel states that it is equivalent to 
O. Schmidt’s Sycinula. Both these are considered by us to be 
synonyms of Leucandra, but the question is discussed in more 
detail under Sycinula. 


DyssyconELLta Haeckel [1870]. 
Type species D. pumila Bowerbank, (=Leucandra pumila). 


Dyssycum Haeckel [1870]. 
Type species D. fistuloswm Johnston, (= Leucandra fistulosa). 


Dyssyous Haeckel [1872]. 
Type species D. primigenius Haeckel, (= Leucetta primigenia). 
A genus of the artificial system, identical with Dyssycum of 
the ‘ Prodromus’ of 1870. There does not seem to be any reason 
for the change of spelling, but such changes, and sometimes more 
radical ones, were made by Haeckel in several cases in his later 
work. 


EBNERELLA von Lendenfeld [1891]. 
Type species £. buccichti von Ebner, (= Amphoriscus 
buccichit). 


GuancuHa IMichlucho- Maclay | 1868). 
Type species G. blanca Michlucho-Maclay, (= Leucosolenia 
blanca). 


HereropreeMa Poléjaeff [1883]. 
Type species H. nodus-gordit Poléjaeft, (= Leucaltis clathria). 
As we have shown when discussing Lewcaltis, Poléjaeff’s name 
is merely a synonym of the latter. 


Homanpra von Lendenfeld {1891}. 
Type species H. falcata von Lendenfeld, (= Leucosolenia 
falcata). 


This genus is the equivalent of Ascandra in Minchin’s sense 
[1896, 1900, &e.}], but must not be confused with Ascandra in 
the original sense of Haeckel. The latter genus was used by 
yon Lendenfeld in almost exactly Haeckel’s sense, so that, since 
he considered A. falcata to be worthy of generic separation, a 
new name became necessary. Ata later date Minchin revised 
the classification of the homoccel sponges, and relegated the 
name <Ascandra to A. falcata, so that the name Homandra 
was rendered unnecessary. (See, for details, Minchin [1896, 
1897].) Wehave given above (p. 720) our reasons for considering 
the separation of L. falcata from the genus Leucosolenia to be 
inadvisable. 


CALCAREOUS SPONGES. 791 


Homoperma von Lendenfeld [1885 A]. 
Type species H. sycandra von Lendenfeld, (=Sycon sycandra). 
This genus was proposed for a somewhat aberrant Sycon, and 
was made by its author the only genus of a new family of Ascones, 
the Homodermide. The question has been dealt with to some 
extent above (p. 716), and is fully investigated in Row’s report 


on the Calcarea of the Hamburg South-Western Australian 
Expedition of 1905 (see Row [1913 MS.]). 


Homerra von Lendenfeld {1891}. 


A genus proposed by von Lendenfeld on hypothetical characters, 
and without any species. 


Hypopicryon Jenkin [1908 B]. 
Type species H. longstaffi Jenkin, (=Sycon longstaffi). 

This genus was erected by Jenkin for the above species, and 
placed by him in his supposed family Chiphoride, on account of 
the presence of chiactines, being separated from Streptoconus on 
account of the “linking” of the chambers. Neither of these 
characters is considered by us as of generic value. 


Hypoerantia Carter [1885-1886 ]. 
Type species H. infrequens Carter, (=Grantiopsis infrequens). 
This genus was proposed by Carter for several species of diverse 
nature, and the diagnosis was extremely unsatisfactory. Further, 
the first (7.e. type) species was said by its author to be ‘“incertze 
sedis.” Under these circumstances we do not feel that we can 
allocate the name to any of our genera. 


LeucKkarteA Michlucho-Maclay MS., fide Haeckel [1872]. 
Type species ZL. natalensis Michlucho-Maclay MB8., jide 
Haeckel [1872], (=Sycon ramosum). 
The only warranty for this name seems to be that it is included 
in the synonymy list attached to Haeckel’s Sycandra ramosa as a 
MS. name of Michlucho-Maclay’s. 


Leucoeyrsia Bowerbank [1864-1882]. 
Type species LZ. gossei Bowerbank, (= Leucandra gosse?). 

One of the four genera of Calcareous sponges proposed by 
Bowerbank in his ‘ Monograph of British Sponges.’ It is now 
usually considered to be a synonym of Leucandra, and although 
perhaps it has right of priority over the latter, yet we feel that 
the name Leucandra is so well known that it should be preserved. 
Levucomerra Haeckel [1872]. 


Type species L. primigenia Haeckel, (= Leucetta primigenia). 


Levconia Grant [1841]. 


A genus proposed by Grant and used by many subsequent 
authors very nearly in the sense in which we use Lewcandra 


792 PROF. A. DENDY AND MR. R. W. H. ROW ON 


(e.g. Carter, Poléjaeff, Urban, etc.), but as shewn by Vosmaer 
[1887] and Dendy [1893 A], the name is not valid, as it had been 
previously applied to a genus of Montusca. 

Leucopsis von Lendenfeld [1885 B]. 


Type species L. pedunculata von Lendenfeld, (= Leucosolenia 
pedunculata). 


This species was supposed by von Lendenfeld to be a transition 
form between Haeckel’s Ascones and Leucones, but it seems to 
us that the structure described is far more probably that of a 
Clathrinoid Leucosolenia provided with a stalk. 

Lervucortis Haeckel [1872]. 
Type species Z. pulvinar Haeckel, (= Leucandra pulvinar). 


Leucutmis Haeckel [1872]. 
Type species LZ. echinus Haeckel, (=Leucilla echinus). 


LipostoMELLA Haeckel [1870]. 
Type species LZ. clausa Haeckel, (= Leucetta primigenia). 


Mepon Duchassaing et Michelotti [1864]. 


Type species VW. barbata Duchassaing et Michelotti (incerta 
sedis). 


A genus comprising two species, both of which are quite un- 
recognisable, and may even not be calcareous sponges. 
Merits Kirkpatrick | 1908}. 
Type species M. normani Kirkpatrick. 
This sponge was originally described as a member of the family 


Pharetronide, but recently Kirkpatrick has shown that its true 
place is among the Non-calcarea. 


Mura Michlucho- Maclay MS., fide Haeckel [1872]. 
Type species M/. dohrnii Michlucho-Maclay MS., fide Haeckel 
[1872], (=Leucandra pulvinar). 

The only authority for this name seems to be Haeckel, who 
published several MS. names in the synonymy lists attached to 
various species In his Monograph. 

Mosrusisponera Duncan [1880}. 
Type species JZ. parasitica Duncan. 

An organism originally described as a parasitic caleareous 
sponge, but it seems very doubtful whether it belongs to the 
sponges atall. We certainly do not feel inclined to recognise it, 
without further evidence, as a member of the Calcarea. 


Narpoa O. Schmidt [1862]. 


Type species WV. reticulum O. Schmidt, (= Lewcosolenia 
reticulum). 


Minchin [1896] has shown that this name was previously 


CALCAREOUS SPONGES. 793 


used for a genus of AsTERoIDEA, and is therefore permanently 
unavailable. 


Narpoma Haeckel [1872]. 
Type species WV. nitida Haeckel, (= Leucosolenia nitida). 


Narvorpsis Haeckel [1870]. 
Type species WV. gracilis Haeckel, (= Leucosolenia gracilis). 


Narporus Haeckel [1872]. 
Type species V. primordialis Haeckel, (=Leucosolenia pri- 
mordialis). 
OLYNTHELLA Haeckel [1872]. 
Type species O. coriacea Montagu, (= Lewcosolenia coriacea). 


OtynrHium Haeckel [1870]. 
Type species O. nitidum Haeckel, (= Leucosolenia nitida). 


Otyntuus Haeckel [1870]. 
Type species O. simplex Haeckel, (= Lewcosolenia pri- 
mordialis). 

This name, as an actual generic name, is considered by us to be 
merely a synonym of Leucosolenia, but we may point out that its 
author proposed it for what he considered to be the most primitive 
adult sponge known, and, although we now believe that he 
probably erred in considering specimens of this form to be adult, 
yet the name is still retained for a hypothetical genus of an- 
cestral Calcarea, and for an early stage in the ontogeny of the 
individual. Even should adult Olynthus-forms occur, however, 
we do not consider that they would be generically separable 
from Leucosolenia. 


Potesna von Lendenfeld [1891]. 
Type species P. uter Poléjaeff, (= Leucilla uter). 


One of the genera based by von Lendenfeld on the presence of 
a sylleibid canal system, a character which we do not consider of 
generic importance. 


Prosycum Haeckel [1870]. 
Type species P. simplicissimawm Haeckel, (= Lewcosolenra 
primordialis). 
RuHABDODERMELLA Urban [1902], 
Type species 2. nwttingi Urban, (= Leucilla nutting:). 
_ ScypHa S. J. Gray [1821]. 


Type species SX. coronata Ellis and Solander, (= Sycon 
coronatum). 


The genus Scypha actually has priority over Risso’s Sycon, but 


194 PROF. A. DENDY AND MR. Rk. W. H. ROW ON 


the latter has so long been in general use that it seems desirable: 
to retain it. Moreover, the earlier name was proposed by a 
botanist for organisms which he regarded as plants, and the- 
generic diagnosis was hopelessly erroneous and misleading. 


Sotenrpium Haeckel [1872]. 
Type species S. nitidum Haeckel, (= Leucosolenia nitida). 


Soteniscus Haeckel [1870]. 
Type species S’. loculosus Haeckel, (= Leucosolenia loculosa). 


SoLeNuLA Haeckel [1872]. 
Type species S. coriacea Montagu, (= Leucosolenia coriacea). 


SPHENOPHORINA Breitfuss [1898 B]. 
Type species S. singularis Breitfuss, (=Grantia ? singularis).. 
This name was originally proposed for a sponge showing certain 
apparently great peculiarities in its skeletal structure and in 
the form of its spicules. Possibly, if the structure described 
really represents that of the sponge, it merits a distinct genus, 
but the fact that only a fragment was found, and that that frag- 
ment had been preserved in spirit for no less than 33 years before 
it was examined by Breitfuss, led us to doubt whether there had 
not been some corrosion of the spicules during that time. This 
opinion was supported by an examination of type slides which one 
ot us (Row) was able to make when in Berlin recently, for the 
appearance of the specimens is just what might be produced by 
the very slow eating away of the terminal portions of the spicule- 
rays by very dilute acid. Under these circumstances we feel that 
the genus had better be abandoned for the present, at any rate 
until further material of the species assigned to 1t has been ob- 
tained, and we have accordingly placed the only described species 

provisionally in the genus Grantia. 


SPHENOPHORUS Lreitfuss [1898 B]. 
Type species S. singularis Breitfuss, (=Grantia % singularis). 
An earlier name for the previous genus, abandoned by its. 
author as being preoccupied. 


Sponera Linneus [1758-1759]. 
The name under which the earlier known species of Calcareous. 


sponges were, In common with non-calcareous forms, described, 
but now entirely abandoned. 


Srreproconus Jenkin [1908 B}. 
Type species S. australis Jenkin, (=Sycon australe). 


One of the genera of Jenkin’s family Chiphoride, which has. 
been abandoned by us for reasons given above. 


CALCAREOUS SPONGES. 795 


SycantHa von Lendenfeld {1891}. 

Type species S. tenella von Lendenfeld, (=Sycon ener), 

This genus was retained by Dendy [1892 B], but although the 

species placed in it by von Lendenteld is certainly aberrant, we 
agree with Jenkin [1908 B] that it does not merit generic 
recognition. 
Sycarium Haeckel [1870]. 

Type species S. ampulla Haeckel, (=Sycon ampulla). 


Sycriprum Haeckel [1870]. 

Type species S. gelatinosum de Blainville, (=Sycon gelati- 

NOsUnr). 

Sycitua Haeckel [1872]. 

Type species S. cyathiscus Haeckel, (= Amphoriscus cyathiscus). 
SYCINULA O. Schmidt [1868]. 

Type species S. aspera O. Schmidt, (= Lewcandra aspera). 

This name, strictly speaking, may have priority over Leucandra, 

but it was only mentioned casually by Schmidt, without diagnosis, 
although he indicated S. aspera as type of the genus, and it has 
never been accepted, except by Haeckel for one of his * artificial” 
genera, whereas Leucandra has come into fairly general use. 


And further, if the name Lewcandra were changed, it apparently 
should be changed to Leucogypsia rather than to Sycinula. 


Sycocystis Haeckel |1870]. 
Type species S. oviformis Haeckel, (=Sycon ciliatwim). 


Sycopenprum Haeckel [1870]. 
Type species S. ramoswm Haeckel, (=Sycon ramosui). 


SycotePis Haeckel [1870]. 
Type species S. inerustans Haeckel, (=either Zrichogypsi« 
incrustans or T’, villosa). 


Sycomerra Haeckel [1870]. 
Type species S. compressum Fabricius, (=Grantia compressa). 


Syconetta O Schmidt [1868]. 
Type species S. quadrangulata O. Schmidt, (= Sycon quad- 
rangulatun). 


SycopHyiuum Haeckel [1870]. 
Type species S. lobatum Haeckel, (= Grantia lobata). 


Sycorruiza Haeckel [1870]. 
Type species S. coriacea Montagu, (= Leucosolenia coriacea). 


Sycortis Haeckel [1872]. 
Type species S. levigata Haeckel, (=Grantia levigata). 


796 PROF. A. DENDY AND MR. R. W. H. ROW ON 


Sycortusa Haeckel | 1872). 
Type species S. levigata Haeckel, (=Grantia levigata). 
One of the subgenera into which Haeckel divided his genus 
Sycortis, raised by von Lendenfeld [1891] to generic rank. 


Sycornamnus Haeckel [1870]. 
Type species S. fruticosus Haeckel, (= Leucetta primigenia). 


Sycum Haeckel [1870]. 
Type species S. ciliatum Fabricius, (=Sycon ciliatwm). 


A modification of Risso’s generic name Sycon for which there 
does not seem to have been any adequate reason. 


Sycurus Haeckel {1872}. 
Type species S. primitivus Haeckel, (=Sycetta prinutiva). 


Tarroma Haeckel [1870]. 


Type species 7’. canariense Michlucho-Maclay, (= Leuco- 
solenia canariensis ). 


Tarropsis Haeckel [1872]. 
Type species 7’. coriacea Montagu, (= Leucosolenia coriacea). 


Tarrus Haeckel [1870]. 
Type species 7’, densus Haeckel, (= Leucosolenia densa). 


TEICHONELLA Carter [1878}. 
Type species 7’. prolifera Carter, (= Leucetta prolifera). 

A genus proposed by Carter for two species, whose slight 
similarity of external form misled him into thus associating them. 
One of these has now been placed by us in the genus Lewcetta, 
the other is the type of our genus Veichonopsis. For a criticism 
of the genus Teichonella and its species see Dendy [1891 B]. 


TENTHRENODES Jenkin [1908 B]. 
Type species Z’. antarcticus Jenkin, (=Sycon antarcticum). 
For a discussion of this genus and its species see under Sycon 
(p. 744). 
THecomerra Haeckel [1870]. 
Type species 7’. loculosa Haeckel, (= Leucosolenia loculosa). 


Uvetia Dendy [1892 B). 
Type species U. hystrix Haeckel, (=Sycodorus hystrix). 
We have found it necessary to substitute Haeckel’s name 
Sycodorus for the above. 
Vosmaeria von Lendenfeld [1885 B}. 
Type species V. gracilis von Lendenfeld, (=Grantia gracilis). 
One of the genera based by von Lendenfeld on the presence of 


CALCAREOUS SPONGES. 797 


a ‘“sylleibid ” canal system, a character which we do not consider 
to be of generic importance. 


WaGNnERELLA Merejkowshi [1878]! 
Type species W. borealis Merejkowski. 
An organism originally described as a calcareous sponge, but 
shewn by Mayer [1879] to be a Heliozoan. 


In addition to the above, an enormous number of subgeneric 
names, both ‘artificial’ and ‘natural,’ were proposed by 
Haeckel in his two works on the group, but the list is sufficiently 
swollen out with discarded generic names, without including 
subgeneric ones also. 


PHYLOGENY OF THE CALCAREA. 


Our views as to the phylogeny of the Calcarea, elaborated in 
the foregoing pages, may now be summarised as follows, and 
illustrated by the accompanying phylogenetic tree. This tree 
differs in certain important respects from that published by one 
of us twenty years ago [Dendy, 1893 A], which is only to be 
expected when we consider the great advances made in our 
knowledge of the group in the interval. All the families of the 
earlier scheme: and the general ideas of their relationships to 
one another in the main lines of descent, are, however, retained 
with but little alteration, but we recognise now four additional 
families of recent Calcarea, the Leucaltide, the Minchinellide, 
the Murrayonide and the Lelapiide, while several genera have 
had to be transferred from one family to another. 


One of the most important advances in the classification of 
the group was made by Minchin [1896], in his demonstration 
that even among the homoccel sponges two types of collared cells 
are met with, with apical and basal nuclei respectively; a 
suggestion which was followed up by Bidder [1898], who, it will 
be remembered, proposed to divide, not only the Calcarea, but 
the whole of the sponges into BAsINUCLEATA and APINUCLEATA 
accordingly, or, confining the suggestion to the Calcarea, to 
divide these into CALCARONEA with apical, and CancinEA with 
basal, nuclei. 

We think it quite likely that the latter of these two suggestions 
will ultimately prove to be thoroughly sound. With regard to 
the former, however, we consider it highly probable that several 
distinct types of collared cells will be shewn to exist in the non- 
calcareous sponges, though as yet we have very little information 
on this point. 


Continuing the observations of Minchin, we find that in the 


798 PROF. A. DENDY AND MR. RB. W. H. ROW ON 


genus Leucosolenia, which admittedly stands at the bottom of 
the line of evolution of the Calearea, both types of collared cells 
exist, but apparently not side by side in the same species; and 
we find further, that the two principal lines of descent, which 


Text-fig. 133. 
“PHARETRONES- 


© MURRAYONID/A 


PMINCHINELLIDA- 


LELAPIIDA- 


GRANTIID/AZ 


EILHARDIA 


neuen 7 


SS LEUCETTAGA 


ae LAMONTIA 


LEUCYSSA 


Sd ae 
APHIS 
Seed —TRICHOGYPSIA 
pmTRicy 


TEICHONOPS!S MEGAPOGON 
GRANTIA 


ANAMIXILLA 


GRANTIOPSIS 


SYCYSSA 


LEUCILLA 


[ee 
! i Aa 


ASCUTE 
Vi 
en 

LEUCOSOLENIA 


“OLYNTHUS” 


HOMOCOELIDAZ 
PHYLOGENY OF CALCAREOUS, SPONGES: 
appear, on quite other grounds, to have sprung from the homoccel 


sponges, are characterised respectively by the two types of 
collared cells. It is interesting to note that these two main lines. 


CALCAREOUS SPONGES. 799 


of descent were clearly indicated twenty years ago, as represented 
by the Leucascidee and Sycettide respectively. 

It must, of course, be remembered that the real difference 
between the two types of collared cells concerns, as Minchin has 
shewn [1909], the relation of the flagellum, with its basal granule, 
to the nucleus. “his relation has, of course, only been deter- 
mined ina very few cases. In ZLeucosolenia coriacea, for example, 
the basal granule is situated at the apex of the cell and the 
nucleus at the base, while in Z. complicata the flagellum appears 
to spring from the nucleus itself, which is apically situated. 
There can be no doubt that the actual position of the nucleus 
itself in the collared cell may vary temporarily under certain con- 
ditions, but in good spirit-preserved material it appears always 
to settle down into a characteristic position, which is either 
basal or apical, and which may be determined without resort to 
special methods of cytological investigation. We do not wish 
to lay undue stress upon this character at present, and we 
should not venture to use it were it not associated with other 
distinctive features, but we have been surprised, in view of our 
former opinion as to the systematic value of such a character, to 
find how constant the position of the nucleus is in the two lines 
of descent indicated. This will be sufficiently evident from 
reference to the table given in the Introduction. 


We have in vain attempted to split up the unwieldy genus 
Leucosolenia into smaller groups. The utmost we have been 
able to do has been to isolate from the main body of species three 
well-marked types, Ascyssa, Ascute and Dendya. We do not 
consider that Minchin’s proposal to divide the homoceel sponges 
into two families, Leucosoleniide and Clathrinide, is at all 
practicable in the present state of our knowledge, and if it be 
true, as he himself has pointed out [1909], following Goldschmidt, 
that the two types of relation of flagellum to collared cell may 
occur in the same genus of Protozoa (JZastigina), we see no reason 
for supposing that both may not occur in the genus Leucosolenia. 
According to our view, this is a large and heterogeneous group 
of primitive forms all closely related to one another and merging 
into one another to a large extent, from which the two lines of 
descent referred to have led the way to the evolution of the 
higher Calcarea. 

We will take the Dendya, or Leucascid-Leucaltid, line first, 
in which the nucleus of the collared cells is basal. The starting 
point of this lime seems to have been from some form closely 
related to Dendya. The radiate arrangement of the colony in 
this genus formerly misled Dendy [1893 A] into regarding it as 
on the line of evolution of the Sycettide, but there are several 
strong arguments against this view. The radiate arrangement 
appears to be but a modification of a reticulate ‘‘ Clathrinid ” 
character, and actual open anastomoses may occur between the 
radial tubes, which, in spite of what has been said by more than 


800 PROF. A. DENDY AND MR. R. W. H. ROW ON 


one author, appears rarely if ever to be the case in true Sycettide: 
or their descendants. To this must be added the primitive type 
of skeleton, composed exclusively of equiangular radiates, which 
do not exhibit the characteristic arrangement met with in the 
syconoid sponges, with their differentiated gastral cortex and 
articulate tubar skeleton. In this connection we may especially 
note the absence of subgastral sagittal triradiates (or quadri- 
radiates), which form such a constant feature of the Sycettide 
and their derivatives. 

The Dendya line seems to have given off two branches, repre- 
sented by the Leucascide and Leucaltide respectively. The Leucas- 
cide are undoubtedly the more primitive of the two. The genus 
Leucascus itself, indeed, might very easily be mistaken for a 
homoceel sponge were it not for the presence of a distinct and 
independent pore-bearimg dermal membrane; it retains the 
elongated, branched, and more or less radially arranged flagellate 
chambers of its Dendya-like ancestors. Within the family evolu- 
tion has led to the development of a more highly differentiated 
dermal cortex in Lewcetta and Pericharax, accompanied by great: 
reduction in the size of the flagellate chambers and complication 
of the inhalant and exhalant canal systems. In this way 
has arisen that remarkable convergence between Lewcetta and 
Pericharax on the one hand, and the leuconoid Grantiide on the 
other, which has for so long prevented the appreciation of the 
fundamental distinction which really exists between these forms. 
The remaining genus in the family, Leawcomalthe, is a highly 
specialised and aberrant type, which is only included here 
provisionally, until we know more of its minute anatomy and 
histology. 

In the Leucaltide the distinctive peculiarity has been the 
enormous development of the dermal cortex with its special 
skeleton, and the accompanying reduction of the skeleton of 
the chamber layer to a more or less vestigial condition, or even 
its complete disappearance. In this family, again, as regards 
canal system, we meet with the customary transition from the 
long chambers and radial arrangement of the more primitive 
forms (ZLeuwcaltis) to the spherical chambers and _ scattered 
arrangement of the highest (Lewcettwsa). 


To this line of descent must also be relegated two out of the 
three surviving families of ‘‘ Pharetrones,” namely, the Minchi- 
nellidee and the Murrayonide. We found this conclusion upon 
the basal position of the nucleus in the collared cells in 
Minchinella and Murrayona ; but it must be borne in mind that 
as regards their general organisation also the members of these 
two families differ very widely from Lelapia and Kebira, the 
only representatives of the Lelapiide, the third surviving family 
of ‘‘ Pharetrones.” 

We are therefore compelled to regard the so-called family 
Pharetronidx as of diphyletic origin, and the resemblance, such 


CALCAREOUS SPONGES. 801 


as it is, between the Lelapiide on the one hand, and the 
Minchinellide and Murrayonide on the other, as due to 
convergence. 

As to how many of the vast group of extinct ‘“‘ Pharetronid ” 
sponges should be associated with the Minchinellide and 
Murrayonide, and how many with the Lelapiidie, is a question 
which we cannot attempt to decide, but we think there is evidence 
to shew that the great majority belong to the basinucleate group, 
though this opinion, of course, rests only on skeletal characters. 
It seems highly probable that in past times the Dendya line of 
descent led to the evolution of the dominant Pharetronid group, 
while at the present day this group has dwindled away and has 
been replaced chiefly by the now dominant Grantiide on the 
apicinucleate line of descent. 

The great antiquity of the Pharetronid group, considering its 
high degree of organisation, is highly remarkable. It dates back 
far into the Paleozoic Epoch, perhaps even to Silurian times 
{| Ulrich, 1889], and almost certainly to Devonian [Zittel, 1878], 
so that it seems that the Calearea had already reached one 
of their highest states of evolution at the commencement of 
the Paleozoic Epoch. Throughout the Secondary Period the 
Pharetrones were dominant, and very numerous genera and 
species have been described, whereas at the present day they 
are almost extinct, though possibly a few more forms yet remain 
to be discovered. 

Of course it is quite possible that the predominance of the 
Pharetrones over other types of Calcarea in past times is apparent 
rather than real, owing to the fact that they alone, on account 
of their coherent skeleton, had much chance of being preserved 
in a fossil condition. Thus there may have been a kind of 
‘“‘ ceological selection” of these forms in a fossil condition, but 
it is indeed noteworthy that the apparently much more primitive 
groups should predominate over these ancient and _ highly 
specialised forms to such an extent as they do at the present day. 

It is possible that a fresh outburst of evolutionary vigour on 
the part of the more primitive persistent groups may have 
occurred in comparatively recent times. 


Turning now to the Sycettid line of descent, we must remind 
the reader, in the first instance, that this appears to have given 
rise to the vast majority of the recent Calcarea. 

The most primitive genus on this line appears undoubtedly to 
be Sycetta, with its radially arranged chambers standing entirely 
separate from one another, with no trace of dermal cortex, and 
without tufts of oxea at the distal ends of the radial chambers. 
This genus already possesses a well-developed articulate tubar 
skeleton, the first joint of which is composed of subgastral 
sagittal trivadiates, which appear never to have been developed 
along the Dendya line of descent, but which are remarkably 
constant throughout the whole of the Sycettid line, although 


802 PROF. A. DENDY AND MR. R. W. H. ROW ON 


absent by suppression in a few cases where the skeleton has 
undergone extreme modification. We are unable to indicate 
any intermediate forms between the genus Sycetéa and the 
Homocelide. It presumably arose from some homoccel ancestor 
which formed colonies by radial budding, not unlike those of 
Dendya, but the apical position of the nucleus of the collared 
cells and the much more advanced type of skeleton shew that 
the relationship to Dendya itself cannot be a close one, while the 
interval to be bridged over between the most primitive Sycetta 
and any Lewcosolenia is a very wide one. Moreover, Sycetta 
itself seems to be in the nature of a cul-de-sac, for the entire 
absence of the characteristic oxeote spicules of Sycon makes it 
doubtful whether we can derive the latter genus directly from 
the former, though both have probably sprung from some common 
ancestor. If, however, Sycon derives its oxea from an ancestral 
Leucosolenia, it is ditiicult to account for the absence of these 
spicules in Sycetta, but the distribution of oxea in the Calearea 
is an extremely difficult problem about which we have perhaps 
said enough in an earlier part of this paper. 

The fact that certain species of Sycon, for which von Lendenfeld 
[1885 A] proposed his genus Homoderma, retain the collared cells 
as a lining to at any rate a portion of the central gastral cavity 
throughout life, certainly shews that one can draw no hard and 
fast line of distimction between the Homoccelide and the old 
group Heteroccela in this respect, but the forms in question have 
such a highly specialised syconoid skeletal system that they 
hardly help us to bridge over the interval between the Homoceelide 
and the Sycettidee. 

The family Sycettide is a very small one, the typical genus 
being Sycon with a large number of species, while the only other 
known genera are S7 ‘ycetta and Sycandra, each with a very small 
number of species and each representing an offshoot which 
probably leads no further. From the Sycettide two lines of 
descent appear to lead to the Heteropiide and Grantiide 
respectively. 


In both these families the important step in further evolution, 
as in the Leucascide and the Leucaltidee, has been the develop- 
ment of a dermal cortex, but this cortex appears to have arisen 
somewhat differently in the two cases. In the Heteropiidee it is 
clearly associated with the out-turning of certain of the oral rays 
of the distal tubar trivadiates so as to arch over the entrances to 
the inhalant canals. We may assume that with these rays the 
dermal tissues of the sponge have spread over the intercanals 
and have given rise ultimately to the special cortical spicules 
developed in situ. The rotation of the distal tubar trivadiates 
in the manner indicated, and the preponderating development 
of the now centripetally directed oral rays, have finally converted 
these spicules into the ‘‘ pseudosagittal” triradiates which 


r\ ey 


CALCAREOUS SPONGES. 803 


constitute the outstanding feature of the Heteropiide. They 
are, so to speak, a new discovery, which the sponge utilizes to 
the utmost, until finally their strong centripetally directed oral 
rays, In association with the opposed basal rays of subgastral 
sagittal triradiates, give rise to an “ inarticulate” tubar skeleton, 
which replaces the articulate tubar skeleton of the ancestral Sycon. 

The evolution of the canal system within the family appears 
to have followed the usual lines up to a certain point. The 
known species of Vosmaeropsis exhibit the type of canal system 
described by von Lendenfeld as “ sylleibid,” intermediate between 
syconoid and leuconoid, but a Heteropiid with a typical leuconoid 
canal system has not yet been found. 


Tn the Grantiide, on the other hand, the development of a 
dermal cortex appears to have been inaugurated by the 
appearance of a thin pore-bearing dermal membrane over the 
ends of the inhalant canals in some syconoid ancestor, such as 
is known to occur in some species of the genus Sycon at the 
present day (e. g. S. boomerang). 

The Grantiide must be regarded as the dominant family of 
Calcarea at the present day, comprising, as they do, no less than 
23 genera, and exhibiting a very wide range of structure both 
as regards skeleton and canal system. It might be possible to 
divide these genera into syconoid and leuconoid subfamilies, but 
we should have no guarantee of the monophyletic origin of the 
latter from the former. Nevertheless, the scarcity of sylleibid 
forms, which might be regarded as connecting links between the 
two types, is somewhat remarkable, and suggestive of a natural 
cleavage. The known species of the genus Megapogon, however, 
are sylleibid, and although most of the Leucandras have small 
chambers, Z. australiensis Carter and L. infesta sp. n. (= Leucilla 
intermedia Row [1909]) have very large ones, and might also be 
considered as transitional forms. Also we must remember that 
in a considerable number of cases we have no accurate information 
as to the canal system. 

The arrangement of the genera within the family is a very 
difficult problem. They appear to group themselves around two 
central types, Grantia and Leucandra, but as we have just 
mentioned, our knowledge of the exact type of canal system in 
many forms is very imperfect, while in others the only evidence 
of their close relationship to Lewcandra is the canal system itself. 
The appearance of this part of our phylogenetic tree will pro- 
bably be greatly modified by subsequent investigations. Such 
aberrant genera as Leucopsila, Baeria, Kuarrhaphis, Leucyssa 
and Trichogypsia can only be included in the Grantiids pro- 
visionally. 

The difficulty of arranging the genera probably arises from the 
fact that great gaps exist in the family owing to extinction 
of intermediate forms. It might be argued that this family 

Proc. Zoou. Soc.—1913, No. LILI. 53 


804 PROF. A. DENDY AND MR. R. W. H. ROW ON 


cannot be very ancient, because hardly any of thei fossil remains 
have been discovered, but this may possibly be accounted for by 
the fact that they do not possess a coherent skeleton, and 
accordingly become disintegrated soon after death. 

It is, however, quite possible that some of the very imperfectly 
known extinct Pharetrones may really be offshoots from this 
family, as we believe to be the case with the recent Lelapiide. 
The customary association of the latter with the Pharetrones 
is based upon the presence of a fibrous skeleton composed of 
modified radiates, especially the tuning-fork spicule, and a 
fibrous skeleton of this type 1s perhaps present in some of the 
extinct genera. We have, however, purposely avoided discussing 
the latter in this paper, as we do not know enough about them 
to warrant us in drawing any but the most general conclusions. 
As we have already pointed out, the apical position of the nucleus 
of the collared cells in Lelapia, and the presence in it also of 
subgastral sagittal triradiates, render the Grantiid origin of the 
Lelapiide reasonably certain. 

Tn addition to the Lelapiide, one other family, the Amphoriscide, 
seems to have originated from the Grantiid stock. This family 
derives its distinctive character from the development of strong, 
centripetally directed apical rays on the tangential radiates of 
the dermal cortex. Such apical rays have undoubtedly appeared 
several times independently in the evolution of the Calcarea. 
We find them, for example, in some species of Leuwcetta, in 
Leucaltis, in some species of Leucettusa, in one species of 
Grantia (G. intermedia), in some species of Leucandra, and m 
Grantilla, as well as in this group. It may well seem doubtful 
whether, in view of these facts, the character in question ought 
to be regarded as of family significance in the Amphoriscide ; 
but inasmuch as the latter appear to us to comprise a natural 
assemblage of three closely related genera in which this character 
is no longer casual but has assumed great importance in the 
structure of the skeleton, we have decided to retain the group, 
at any rate for the present. We have, however, considerably 
reduced the size of the family by the removal of Lewcaltis 
(Heteropegma), together with some of the species formerly 
placed in Leucilla, but now divided between Leucetia, Leucettusa 
and Leucandra, to which they seem to be more closely affiliated 
by other characters, leaving in the genus Lewcilla a group of 
species which are, we believe, all directly descended from 
Amphoriscus. 

The most primitive Amphoriscide, belonging to the genus 
Amphoriscus, have a syconoid canal system and a somewhat 
feebly devel6ped dermal cortex, and we accordingly consider the 
family to be an offshoot from low down on the Grantiid stem. 
Finally, we may point out that in this family again the canal 
system has undergone its usual transformation from the syconoid 
to the leuconoid type, with intermediate sylleibid forms. 


CALCAREOUS SPONGES. 805 


BIBLIOGRAPHY. 


The following list of literature includes only those papers which 
are actually referred to in the text, as it has been found impossible 
to give anything like a complete list of the papers which deal with 
calcareous sponges in the space at our disposal. However, there 
will be found to be but few papers of real systematic importance 
which are not mentioned, and, for obvious reasons, all papers 
which describe new species are included, at any rate so far as we 
are aware, and we have no reason to fear that our list is not 
complete. 

When more than one paper has been published by an author in 
any year they have been distinguished by the letters A., B., C., &e. 


1901. ArRNeEsSEN, Emily. <‘‘Spongier fra den norske kyst. I. 
Calcarea. Systematisk katalog med bemerkninger og 
bestemmelsestabel.” Bergens Mus. Aarbog, 1900 (1901), 
No. 9. 

1891. Broper, G. P. “ Review.” (of) ‘A Monograph of 
Victorian Sponges, by Arthur Dendy. Part I.— 
The Organisation and Classification of the Calcarea 
Homoceela, with descriptions of the Victorian Species. 
Quart. Journ. Microse. Sci., (n. 8.) vol. xxxii. pp. 625- 
632. 

“The Skeleton and Classification of Calcareous 
Sponges.” Proc. Roy. Soc., London, vol. lxiv. No. 403, 
1898, pp. 61-76. 

1834-1837. Buatnvitixn, H. de. ‘ Manuel d’Actinologie et de 
Zoophytologie.” Paris. 

1845. BoweErsanx, J. 8. ‘Description of a new Genus of 
Calcareous Sponges (Dunstervillia).” Ann. Mag. Nat. 
Hist., (ser. 1), vol. xv. 1845, pp. 297-3800. 

1858-1862. “On the Anatomy and Physiology of the 
Sponeiade.” 

Part I. Phil. Trans. Roy. Soc., London, vol. exlviii. 
1858, pp. 279-332. 

Part Il. Phil. Trans. Roy. Soc., London, vol. clii. 
1862, pp. 747-836. 

Part III. “On the Generic Characters, the Specific 
Characters, and the Method of Examination.” Phil. 
Trans. Roy. Soc., London, vol. clii. 1862, pp. 1087— 
1135. 

1864-1882. —— “A Monograph of the British Spongiade.” 
Ray Soc., London, 4 vols. 

1872-1876. “Contributions to a general history of the 
Spongiadee.” Proc. Zool. Soc., London, 1872, pp. 115- 
129, 196--202, 626-634; 1873, pp.3-25, 319-333; 1874 
pp- 298-3805 ; 1875, pp. 281-296 ; 1876, pp. 768-775. 

53* 


1898. 


806 PROF. A. DENDY AND MR. R. W. H. ROW ON 


1896 A. Brurrruss, L. ‘ Kalkschwamme der Bremer-Expedition 
nach Ost-Spitzbergen im Jabre i889 (Prof. W. 
Kukenthal und Dr. A. Walter).” Zool. Anzeiger, Bd. xix. 
No. 514, pp. 426-432. 

* Ascandra Hermesi, ein neuer homocceler Kalk- 

schwamm aus der Adria.” Zeitschr. wiss. Zool., Bd. 1xii. 

Hft. 1, pp. 39-42. 

“ Amphoriscus semoni, ein neue Art heteroceeler 
Kalkschwimme.” Zool. Anzeiger, Bd. xix. pp. 435- 
436, 

1897. — ‘Catalog der Calcarea der zoologischen Sammlung 
des koniglichen Museums fir Naturkunde zu Berlin.” 
Arch. fiir Naturgesch., Jahrgang Ixii. Bd. 1, pp. 206- 
226. 


1896 Bb. 


1896 C, 


1898 A. 


“Amphoriscus semoni, ein neuer heteroceler Kalk- 
schwamm.” (Semon, Zoologische Forschungsreisen 
Australien malayischen Archipels, Bd. v. Lief. 4.) 
Denkschrift. med.-nat. Ges., Jena, Bd. vii. pp. 381- 
384. 

1898 B. —— “ Kalkschwammfauna des weissen Meeres und 
der Hismeerkiisten des europidischen Russlands mit 
Beriicksichtigung und Aufstellung der Kalkschwamm- 
fauna der arktischen Region.”  dMémoires de U Acad. 

' Impér. des Scvences, St. Pétersbourg, (ser. 8) vol. vi. 
No. 2. 

“ Kalkschwammfauna der Westkiste Portugals.” 
ZGoolog. Jahrbuch, Syst. Abth., Bd. xi. pp. 89-102. 

1898 D. —— “Die Kalkschwammfauna von Spitzbergen. Nach 
den Sammlungen der Bremer-Expedition nach Ost- 
Spitzbergen im Jahre 1889 (Prof. W. Kiikenthal und 
Dr. A. Walter).” Zoolog. Jahrbuch, Syst. Abth., Bd. xi. 
pp. L03-120, ‘ 

“ Die Kalkschwimme der Sammlung Plate. (Fauna 
Chilensis, Bd. 1.)” Zoolog. Jahrbuch, Suppl.-Bd. iv. 
pp. 455-470. 

1871 A. Carver, H. J. ‘On two undescribed Sponges and two 
Esperiade from the West Indies; also on the nomen- 
clature of the calcisponge Clathrina, Gray.” Ann. Mag. 
Nat. Hist., (ser. 4) vol. vii. pp. 268-283. 

“A description of two new Caleispongie (T'richo- 

gypsia, Leuconia), to which is added confirmation of 

Professor James-Clark’s discovery of the true form of 

the Sponge-cell (Animal) and an account of the Polype- 

like pore-area of Cliona corallinoides, contrasted with 

Prof. E. Haeckel’s view on the relationship of the 

Sponges to the Corals.” Ann. Mag. Nat. Hist., (ser. 4) 

vol. vill. pp. 1-28. 

“« On Teichonia, a new family of Calcareous Sponges, 

with descriptions of two Species.” Ann. Mag. Nat. 

Hist., (ser. 5) vol. 11. pp. 85-40. 


1898 C. 


1898 E. 


1871 B. 


CALCAREOUS SPONGES. EMS On 


1883. Carrer, H. J. ‘“ Further Observations on the so-called 
‘Farringdon Sponges’ (Calcispongie, Zittel), followed 
by a description of an existing species of a like kind 
(Leucetta elathrata, new sp.) Ann. Mag. Nat. Hist., 
(ser. 5) vol. xi. pp. 20-37. 

1885-1886. -—— “Descriptions of Sponges from the Neigh- 
bourhood of Port Phillip Heads, South Australia.’ 
Ann. Mag. Nat. Hist., (ser. 5) vols. xv., xvi., xvii. and 
xvi. (The calcareous sponges are described in the fol- 
lowing parts ;—vol. xvii. pp. 431-441, vol. xvii. pp. 502- 
516, vol. xvii. pp. 84-55, vol. xviii. pp. 126-149.) 

1886. —— “Description of a new species (Aphroceras 
ramosa).” (in Higein, T., Report on the Porifera of the 
L.M.B.C. district}. Proc. Lat. Phil. Soc., Liverpool, 
vol. xl. Appendix. 

1828. Deis Curasn, 8. “ Memoria sulla Storia e Notomia 
degli Animali senza Vertebre del Regno del Napoli.” 
Napoli, 1828. 

1891 A. Denpy, A. ‘A Monograph of the Victorian Sponges. 
Part I. The Organisation and Classification of the 
Calcarea Homoceela, with descriptions of the Victorian 
Species.” Trans. Roy. Soc. Victoria, vol. iii. No. 1, 
pp. 1-82. 

1891 B. —— “Studies on the Comparative Anatomy of Sponges. 
III. On the Anatomy of Granta labyrinthica, Carter, 
and the so-called family Teichonide.” Quart. Jowrn. 
Mrerose. Sct., (n.s.) vol. xxx. pp. 1-89. 

‘Preliminary Account of Synute pulchella, a new 

Genus and Species of Calcareous Sponges.” Proc. Roy. 

Soc. Victoria, (n.s.) vol. iv. pp. 1-6. 

‘Synopsis of the Australian Calcarea Heteroccela, 

with a proposed classification of the group, and 

descriptions of some new genera and species.” Proc. 

Roy. Soc. Victoria, (n.s.) vol. v. pp. 69-116. 

“On a new species of Leucosolenia from Port 
Phillip Heads.” Proc. Roy. Soc. Victoria, (n.s.) vol. v. 
pp. 178-180. 

“Studies on the Comparative Anatomy of Sponges. 
V. Observations on the Structure and Classification of 
the Calcarea Heterocela.” Quart. Journ. Microsc. Sci., 
(n.s.) vol. xxxv. pp. 159-257. 

1893 B. + “Studies on the Comparative Anatomy of Sponges. 
VI. On the Anatomy and Relationships of Lelapia 
australis, a living representative of the fossil Phare- 
trones.” Quart. Journ. Microsc. Sci., (n.s.) vol. xxxvi. 
pp- 127-142. 

‘Report on the Sponges collected by Professor 

Herdman at Ceylon in 1902.” Reports on the Pearl 

Oyster Fisheries of the Gulf of Manaar, vol. iti. pp. 59— 

246. (Royal Society, London.) 


1892 A. 


1892 B. 


1892 C. 


1893 A. 


1905. 


808 
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1897. 


1864. 


1887. 


1786. 


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Denby, A. ‘ Report on the Calcareous Sponges collected 
by the Sealark Expedition in the Indian Ocean.” Trans. 
Linnean Soc. London, Zool. vol. xvi. pp. 1-29. 

Dovertery, L. “ Description of Petrostroma schulzei, n.g. 
et sp. of Calearea, representing a new order Lithones.” 
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145. 


“Ueber die Lithonina, eine neue’ Gruppe von 
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Ducuassaine, P., et Micumnori, G. “Spongiaires de 
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xORlg 

Duncan, P. Martin. “On a parasitic Sponge of the 
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Kepner, V.von. ‘ Amphoriscus buccichii, n. sp.” Zoolog. 
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Euxis, J.,and Sonanper, D. “* Natural History of many 
curious and uncommon Zoophytes collected from various 
parts of the Globe.” London, 1786. 

Esper, EH. J.C. (In the synonymy list under Leucandra 
aspera, Haeckel gives a reference to Spongia panicea, 
Esper, ‘“ Spongien, Taf. 18, fig. 1, 2.” We have been 
unable to trace this reference, but the species is not 
referred to in Hsper’s “ Pflanzenthiere.”) 

Fasricius, O. ‘“ Fauna Groenlandiea.” Hafnie et 
Lipsie, 1780. 

Fremine, J. ‘A History of British Animals.” Edin- 
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Fristepr, K. “Sponges from the Atlantic and Arctic 
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pp- 365-367. 


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1826. 


1841. 


1861. 


1858. 


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1867. 


1821. 
1870. 
1872. 


1877. 


1908. 


1890. 


1894. 


1895. 


1886. 


1900. 


1869. 


1908 A. 


1908 B. 
1842. 


1893. 


1894; 


1895. 


CALCAREOUS SPONGES. 809 


Gray, J. E. ‘Notes on the Arrangement of Sponges, 
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Gray,S.F. “A natural arrangement of British Plants.” 
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1872. 


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JENKIN, C. F. “The Calcareous Sponges. (in) The 
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810 
1895. 


1900. 


1908. 


1910. 


TOMAS 


1911 B. 


1912. 


1864. 


1886. 
1893. 


1896. 


1900 A. 


1900 B. 


1885 A. 


1885 B. 


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pp. 177-179. 

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“Note on Astrosclera willeyana, Lister.” Proc. 
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Konimiker, A. von. ‘‘ lcones Histologice, oder Atlas der 
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‘* Description of a new Species of Caleareous 
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“A Monograph of the Australian Sponges. 

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1868. 


CALCAREOUS SPONGES. 811 


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Family of Sponges.” dA. Welley’s Zoological Results, 
Part IV. Cambridge. 

Lunpsick, W. “The Porifera of East Greenland.” 
Meddel. om Gronland, vol. xxix. 1909, pp. 423-464. 

Mayer, Paul. ‘“‘ Wagnerella borealis.”  Zoolog. Anzeiger, 
vol. 11. pp. 357-358. 

Mrresxkowsky, C. ‘On Wagnerella, a new genus of 
Sponge nearly allied to the Physemaria of Ernst 
Haeckel.” Ann. Mag. Nat. Hist. (ser. 5) vol. i. 
pp. 70-77 

Micunucno-Macnay, N. de. ‘“ Beitriige zur Kenntniss 
der Spongien. I. Ueber Guancha blanca, einen neuen 
Kalkschwamm.” Jenaische Zeitschrift, vol. iv. pp. 221— 
240, 


“Ueber einige Schwiimme des nordlichen stillen 
Oceans und des Hismeeres, welche im Zoologischen 
Museums der Kaiserlichen Akademie der Wissen- 
schaften in St. Petersburg aufgestellt sind ; ein Beitrag 
zur Morphologie und Verbreitung der Spongien.” 
Mémovres de? Académie des Scr. a St. Pétershourg, vol. xv. 
No. 3. 

Mincuin, H. A. “Suggestions for a Natural Classifica- 
tion of the Asconide.” Ann. Mag. Nat. Hist., (ser. 6) 
vol. xvi. pp. 349-362, 

‘* Ascandra or Homandra? A Test Case for the 

Rules of Zoological Nomenclature.” Zoolog. Anzeiger, 

Bd. xx. No. 524, pp. 49-50. 

“The Porifera.” Lankester’s Treatise on Zoology, 

Part 2, Chapter 3. 

‘The Characters and Synonymy of the British 

Species of Sponges of the Genus Leucosolenia.” Proc. 

Zool. Soc., London, 1904, vol. ii. pp. 349-396. 

‘““'The Relation of the Flagellum to the Nucleus 
in the Collar-cells of Calcareous Sponges.”  Zoolog. 
Anzeiger, Bd. xxxv. pp. 227-231. 

Montacu, G. “An Hssay on Sponges, with Descriptions 
of all the Species that have been discovered on the 
Coast of Great Britain.” Memoirs of the Wernerian 
Soc. Edinburgh, vol. ii. pp. 67-122. 


1788-1796. Mttnur, O. F. ‘“ Zoologica Danica.” (2nd Edition.) 


1888. 


1904. 


Hafnie, 1788-1796. 

Potrsanrr, N. de. ‘The Calearea.” Reports on the 
Scientific Results of the Voyage of H.M.S. ‘Challenger, 
Zoology, vol. vill. 

Prerwisco, J. ‘‘Kalkschwimme aus dem _ Pacific. 
Ergebnisse einer Reise nach dem Pacific, Schauinsland 
1896-1897.” Zoolog. Jahrbuch, Syst.-Abth., Bd. xix. 
pp. 9-26. 

Quoy, J. R.C.,et Garmarp, P. ‘“ Voyage de I’ Astrolabe.” 
Zoologie, vol. iv. Paris, 1&33. 


812 
1881. 


1884. 


1909. 


1913? 


1864. 


1864, 


1907. 


1869. 


ON CALCAREOUS SPONGES. 


Ripury, 8. O. ‘“Spongida collected during the Expe- 
dition of H.M.S. Alert in the Straits of Magellan and 
on the Coasts of Patagonia.” Proc. Zool. Soc., London, 
1881, pp. 107-139. 

‘““Spongida.” Reports on the Zoological Collections 
made in the Indo-Pacific Ocean during the Voyage of 
H.M.S. ‘ Alert,’ 1881-1882, pp. 366-482, 582-630. 
London, 1884. 

Risso, A.  ‘ Histoire Naturelle des principales Pro- 
ductions de |’Kurope Meéridionale, et particulicrement 
de celles des Environs de Nice, &.”’ Vol. v. Paris, 
1826. 

Row, Rk. W. H. ‘ Reports on the Marine Biology of the 
Sudanese Red Sea.—XIX. Report on the Sponges col- 
lected by Mr. Cyril Crossland in 1904-1905. Part I. 
Calcarea.” Journ. Linn. Soc., London, Zoology, vol. xxxi. 
pp. 182-214. 

‘‘ Report on the Calcarea obtained by the Hamburg 
South-Western Australian Expedition of 1905.” (Will 
shortly appear.) 

Scumipt, Oscar. ‘ Die Spongien des Adriatischen 
Meeres.” Leipzig, 1862. 

“Supplement der Spongien des Adriatischen 

Meeres, enthaltend die Histologie und systematische 

Ergainzungen.’ Leipzig, 1864. 

“ Die Spongien der Kiste von Algier, mit Nach- 

trigen zu den Spongien des Adriatischen Meeres. 

(Drittes Supplement.)” Leipzig, 1868. 

““Grundziige einer Spongien-Fauna des Atlant- 
ischen Gebietes.” Leipzig, 1870. 

Scuurrner, O. “ Beschreibung einiger neuer Kalk- 
schwamme.” Jenaische Zeitschr., vol. xi. pp. 403-483, 
Scuutze, f. KE. ‘“ Ueber den Bau und die Entwicklung 
von Sycandra raphanus Waeckel.” Zettschr. wiss. 

Zoologie, Suppl., vol. xxv. pp. 247-280. 

STEPHENS, (Miss) J. ‘‘Clare Island Survey. Part 59. 
Marine Porifera.” Proc. Roy. Irish Acad., vol. xxxi. 
No. 59. 

THacker, A. G. ‘On Collections of the Cape Verde 
Islands Fauna made by Cyril Crossland, M.A.(Cantab.), 
B.Se. (Lond.), F.Z.8. (ate of the Gatty Marine Labora- 
tory, St. Andrews University), from July to September 
1904. The Calcareous Sponges.” Proc. Zool. Soc., 
London, 1908, pp. 757-782. 

Toprsmnt, E. “ Eponges calcaires recueillis par le Francais 
dans l’Antaretique (Expédition du Dr. Charcot).” 
Bullet. Mus. Hist. Nat. Paris, 1907, pp. 539-544, 

Unricu, H. O. “ Preliminary Description of new Lower 
Silurian Sponges.” American Geologist, vol. ii. pp. 233— 


248, 


THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 813 


1902. Urpan, fF. “ Rhabdodermella nuttingi, nov. gen. et nov. 
spec.” Zeitschr. wiss. Zoologie, Bd. lxxi. pp. 268-275. 


1905. “ Kalifornische Kalkschwimme.” Archiv fiir 
Naturgesch., Jahrgang 72, Bd. i. pp. 33-76. 

1908. “Die Kalkschwimme der deutschen Tiefsee- 
Expedition.” Zoolog. Anzeiger, Bd. xxxili. pp. 247-252. 

1909. “Die Calcarea.” Wissensch. Ergebnisse der 


deutschen Tiefsee-Hapedition (Valdiia), Bd. xix. Jena, 
1909. 

1873. Verrinn, A. KE. ‘“ Exploration of Casco Bay by the U.S. 
Fish Commission in 1873.” Proc. American Assoc. 
Advane. Sci. 1873, Part 2, pp. 340-395. 

1880. Vosmarr,G. C.J. “Ueber Leucandra aspera H., nebst 
alleemeimen Bemerkungen ueber das Canalsystem der 
Spongien.” Leiden, 1880. (dn wtaugural Dis- 
sertation.) 

‘“Porifera.” Die Klassen und Ordnungen des 
Thierreichs, wissenschaftlich dargestellt in Wort und 
Bild. Von Dr. H. G. Bronn. Bd. i. Leipzig und 
Heidelberg, 1887. 

1878. Zinrnn, K. A. von. “Studien ther fossile Spongien. 
Ill. Monactinellide, Tetractinellide, und Calci- 
spongiv.” Abhandl. Akad. Wissensch. Miinchen. vol. xiii. 
part 2, pp. 1-48. 


1887. 


EXHIBITIONS AND NOTICES. 
May 20, 1913. 


Prof. HK. A. Mincnin, M.A., F.R.S., Vice-President, 
in the Chair. 


The Secretary read the following report on the Additions that 
had been made to the Society's Menagerie during the month of 
April, 1913 == 

The registered additions to the Society’s Menagerie during the 
month of April were 205 in number. Of these, 95 were acquired 
by presentation, 79 by purchase, 9 were received on deposit, | in 
exchange, and 21 were born in the Gardens. 

The number of departures during the same period, by death 
and removals, was 156. 

Amongst the additions special attention may be directed to :— 

1 White-handed Gibbon (Hylobates lar), from Penang, deposited 
on April 10th. 


814 ON DATES OF PUBLICATION OF TRANSACTIONS, 


1 Maholi Galago (Galago maholi), from 8. Africa, presented by 
Lady Yule, F.Z.8., on April 8th. 

1 Bangs’s Agouti (Dasyprocta colombiana), 1 Bonda’s Squirrel 
(Scturus saltuensis bonde), from Colombia, both new to the 
Collection, presented by W. K. Pomeroy, Esq., F.Z.S., on April 
21st. 

A Collection of Birds, from North-West India, presented by 
Major G. A. Perreau, F.Z.S., on April 1st, of which the following 
are new to the Collection :— 

1 Dark Grey Bush-Chat (Oreicola ferre), 1 Plumbeous Redstart 
(Lhyacornis fuliginosa), 1 Red-flanked Bush- Robin (Janthia 
rufilata), 2 White-tailed Blue Robins (WVotodela leucwra), 1 Great 
Niltava (Niltava grandis), 2 Short-billed Minivets (Pericrocotus 
brevirostris), 1 Larger Red - headed Crow - Tit (Scaorhynchus 
rujiceps), 1 Cinnamon-bellied Nuthatch (Sitta cinnamomeiventris), 
1 Grey-sided Laughing Thrush (Dryonastes cerulatus), 4 Rufous- 
necked Laughing Thrushes (Dryonastes rujicollis). 

1 Bushmaster (Lachesis mutus), from Trinidad, presented by 
the Baron K. G. K. Leijonhufvud, F.Z.8., on April 28th. 

1 large Anaconda (Hunectes murinus), from Trinidad, purchased 


on April 28th. 


The Rev. T. R. R. Srespsine, M.A., F.R.S., F.Z.8., drew 
attention to Prof. F. K. Schulze’s important work on zoological 
nomenclature, the ‘ Nomenclator Animalium,’ giving a_ brief 
description of its objects and scope, and urging its claims for 
assistance from British naturalists. 


The LiprariaAn submitted the following list of dates of publi- 
cation of the early parts of the Society’s “‘ Transactions,” which 
had been drawn up from the records kept by Messrs. Taylor and 
Francis, the Society’s printers. 

The date of issue of Parts subsequent to Vol. VII. Part 1 
appears at the foot of each sheet. 


VouumeE I. 

Part. Pages. Plates. Published. 
il, 1-90. I.-XIT. Aug. 14, 1833. 
2. 91-194. XITII.-XXVI. Apr. 23-25, 1834. 
3, 195-800. XXVIT.-XXXVITI. Mar. 20; 1835. 
4. 301-407. XXX VITI.-LIX. Dec. 3, 1835. 

Vouume IT. 
le 1-86. T-XVII. Oct. 2, 1836. 
2 87-164. XVITI.-XXTX. May 4, 1838. 
3: 165-248. XXX.-XLV. Dec. 5, 1838. 
4, 249-342. XLVI.-LVIII. Apr. 6, 1840. 
5. 343-420, LIX.-LXXT. May 15, 1841. 


Part. Pages. 
1-132. 


So trea ae 


1-30. 
31-74. 
75-88. 
89-148. 


GO IS OT G9 bo 


1-32. 
33-102. 


ee Sh 


1-48. 
49-86. 
87-124. 


G0 I OTR Ge bo 


I 1-144. 


133-234. 
235—276. 
277-344. 
345-380. 
381-431. 


149-182. 
183-268. 
269-352. 
303-382. 


103-242: 
243-336. 
337-421. 


125-226. 
227-308. 
309-376. 
377-494. 
495-537. 


ON GROWTH OF ANTLERS. 


Vouume III. 
Plates. 

IL Wl 
VIII.-X VIL. 
XVIII-XxXX. 
XXXI.-LI. 
LIT-LVIL. 
LVI1.-LXITITI. 


Vouume LV. 


I-VI. 

IX.-XXV. 
XXVI.-XXX. 
XXXJ.-XLII. 
XLIUI-LIT-~. 

LIV .—-LXITI. 
LXIV.-LX VII. 
LXVIla.—-LXXVIT. 


VoLuME V. 


T—-XITI. 

XIV .-XXVI. 
XXVIT.-XLUIT. 
XLIIL-LI. 
LITT.-LX VII. 


Vouume VI. 


T-XIV. 
XV.-XXIV. 
XXV.-XXX. 
XXXITL-XLI, 
XLIV.-LIV. 
LV.-LXII. 
LXIIT-LXXXVIT. 
UXXXVITT.-XCII. 


Vouume VII. 
= WIL, 


815 


Published. 


June 
Jan. 
June 
Dec. 


16, 1842. 
23, 1844, 
5, 1844. 
28, 1846. 


. 22, 1848. 


30, 1849. 


1, 1851. 
2, 1852. 


, 29. 1853. 
, 10, 1856. 
. 28, 1858. 
. 31, 1859. 
e. 23, 1861. 
. 15, 1862. 


11, 1862. 
97, 1863. 


. 28, 1864, 


5, 1865. 
6, 1866. 


. 15, 1866. 
lu, WG. 
, 28, 1867. 


5, 1867. 


>. 19, 1868. 


4, 1868. 


_ 15, 1868. 


1, 1869. 


. 20, 1869. 


Mr. R. EK. Honpine exhibited a number of antlers, skulls, and 
photographs illustrating variations in the growth of the Antlers of 
Deer, and stated that antlers were liable to considerable variation 
in form, due either to exuberance or proliferation of growth, or 
to congenital defect in the embryonic stage as in the case of the 
“Hummel” or Hornless Stag (text-fig. 134 A), which occasionally 
made its appearance in Scottish and continental Deer forests. 


816 MR. R. E. HOLDING ON 


Text-fig. 134. 


gat) P| 
ea “ ‘ 
E 553 


& \ 
yy i) Vy : 
ie ‘i 4, 


>I) 


REX det 


A. Upper portion of the skull of a “ Hummel” or Hornless Stag, aged 5 years. 

B. Upper portion of the skull of Red Deer (Cervus elaphus), with deficient right 
and arrest of left antler. 

Ci. First antlers of young Wapiti Stag (Cervus canadensis), showing unusual 
growth of nodules and “ burr.” 


GROWTH OF ANTLERS. 817 


Sometimes the arrest was on one side only, producing the Single- 
horned Stag as shown (B). Occasionally a small supernumerary 
horn might grow below the Burr or Coronel (6), as shown in the 
figures of Muntjac (BE) and Red Deer (D), probably due, as in the 
Four-horned Sheep, to accessory centres in the pedicle or horn 
support. Other variations, and by far the most common, were due 
to dichotomy, bifurcation, or segmentation of the beam or of 
the tines. That favourable environment and ample food promoted 
variation in the direction of exuberance or proliferation of points 
was well demonstrated by the photographs from Warnham Court 
and by illustrations in Mr. Winans’ book ‘“ Deer-breeding for 
Fine Heads,” published by Messrs. Rowland Ward. On the other 
hand, head-keepers and others in charge of Scottish Deer forests, 
had observed that scarcity of food, hard weather, and exposure 
have a marked effect on the antlers of Deer, causing deficiency 
in growth, bad form, and degeneration. 

It was not often, at least in Scotland, that a stag with a good 
head escaped the attention of the owner or lessee of the Forest, 
and therefore defects and variations due to extreme age were 
seldom seen in the Red Stag, but in the Park Fallow Deer a 
marked change took place in the form of the palmation of the 
antler due to advancing age. 

Malformations and variations due to accident, castration, or 
wounds, were not referred to, as being pathological. 

Of the specimens exhibited in illustration of his remarks, 
Mr. Holding drew special attention to the antler of a Sambur 
stag (J), showing excessive proliferation of points, and an 
exceptionally fine pair of antlers lent by Mr. C. J. Lucas, of 
Warnham Court. 


Explanation (continued) of Text-fig. 134 (see opposite). 


D. Head of Stag shot in Jura Forest, showing supernumerary right antler (a) growing 
from a separate pedicle on the frontal bone. 


Hd 


Muntjac (Cervulus muntjac), showing supernumerary horns (4) springing from 
the long pedicle. 


F. Head of Fallow-buck (Cervus dama), showing dichotomy or bifurcation of beam 
of right antler at the “burr.” 


G. Antler of Axis Deer (Cervus axis), the bifurcation occurring above the burr. 


H. Head of Red Stag from Warnham Park, having 42 points. From photograph by 
J. G. Millais, Esq., F.Z.S. 


I. Head of White-tailed Deer (Cariacus leucurus), showing excessive proliferation. 
Shown by Capt. Frewen at a recent Exhibition of American trophies. 


J. Antler of Sambur Deer (Cervus unicolor), showing profuse growth of “points ” 
obliterating the normal character of the antler. 


818 MR. R. LYDEKKER ON THE 


June 3, 1913. 
Prof. E. W. MacBrips, M.A., D.Sc., F.R.S., Vice-President, 


in the Chair. 


Mr. D. Sera-Suira, F.Z.S., Curator of Birds, exhibited the 
egg and young of the Mikado Pheasant (Calophasis mikado), a 
rare species, described first in 1906, from the mountains of 
Formosa. Some living specimens had been imported in 1912 by 
Mr. Walter Goodfellow, and the owners of these birds had 
entrusted the eggs to the Zoological Society, where they were 
being hatched. 

The egg was cream-coloured and very large compared with 
those of allied species of pheasants, measuring 57 x 41 mm. 

The incubation period proved to be twenty-eight days, instead 
of twenty-four as in the majority of pheasants, and the young 
when newly hatched were very large, and had the quil-feathers 
better developed than was the case in allied species. 


PAPERS. 


45. The ‘Transvaal Race of the Cape, or Khama, Hartebeest *. 
By R. LypKexesr, F.R.S., F.Z.S. f 
[Received April 1, 1913: Read April 8, 1913.] 
(Text-figure 135.) 
INDEX. 


Page 
Description of Bubalis caama selbornei ............ 819 


In their excellent volume, ‘The Sportsman in South Africa,’ 
Messrs. Nicolls and Eglington describe (p. 45) the Cape, or Rooi, 
Hartebeest (Lubalis caama), probably from fresh specimens, as 
follows :— 

“General colour reddish brown, with violet tinge throughout ; 
dark plum-coloured saddle-patch, commencing at point of shoul- 
der, extends over entire surface of back and ends in root of tail ; 
a similar patch extends over each shoulder, downwards as far as 
knees, and front of shin-bones of fore-legs; pale yellowish patch 
on cheeks of rump; front of face, which is very long, almost 
black, as is a stripe down back of neck.” 

In their figure of the head (pl. iv. fig. 13) the broad face-blaze 
is shown as extending uninterruptedly from the base of the horns 


* [The complete account of this new subspecies appears here, but since the name 
and a preliminary diagnosis were published in the ‘ Abstract,’ No. 119, 1918, it is 
distinguished by being underlined.—Eprror. | 

+ By permission of the Trustees of the British Museum. 


CAPE HARTEBEEST. 819 


to the muzzle; but, in many cases at any rate, this is divided by 
a narrow tawny line at the level of the eyes. They omit te 
mention a plum-coloured patch on the side of the thighs below 
the light area, extending to the hocks. 

In the ‘ Book of Antelopes,’ by Messrs. Sclater and Thomas, an 
obviously over-coloured plate (no. iv.) of the entire animal is 
given, in which the dark face-blaze, divided by an interocular 
hght bar, and the dark markings on the fore and hind limbs are 
clearly shown, although there is no sign of a plum-coloured 
saddle-patch. The authors describe the general colour as brown- 
ish fulvous, darker than in any other member of the genus. 

The only specimen of an adult male Cape Hartebeest from 
Cape Colony in the British Museum is one obtained by Sir 
Andrew Smith, which has recently been dismounted and con- 
verted into a flat skin. Although much faded by long exposure, 
it serves to show that the type of colouring was originally much 
the same as in the plate in the ‘ Book of Antelopes. 

A few years ago Lord Selborne was good enough to offer to 
endeavour to procure for the Museum specimens of such South 
African Antelopes as might be required for public exhibition ; 
and as a result of his lordship’s request the skin, skull, and horns 
of an adult male of the Transvaal representative of the Cape 
Hartebeest were received at the Museum in 1912, as a gift from 
the De Beers Mining Company. The specimen w as in due course 
set up by Rowland Ward, Ltd., and placed on exhibition in the 
galleries in lieu of the old and faded example from Cape Colony 
referred to above. At the time I was busy with other matters, 
and consequently did not pay any attention to the details of the 
new acquisition. Recently, however, I have had occasion to 
review all the Hartebeests in the collection; and this survey has 
left no doubt that the Transvaal Hartebeest, which was shot in 
the neighbourhood of Kimberley, represents a very distinct 
undescribed race of Bubalis caama. 

From the typical Khama this race differs by its much paler 
general colouring, which is yellowish fawn or tawny, not unlike 
that of B. cokei, by the minor development and intensity of the dark 
markings, and by the apparently less sharp definition of the white 
on the sides and back of the lower part of the rump. The general 
tawny tint tends to chestnut on the loins, as in B. cokei, but 
elsewhere the back is of much the same colour as the flanks. The 
nuchal stripe is indistinct ; and the face-blaze, instead of forming, 
with the exception of the narrow light band between the eyes, a 
continuous wholly black streak from the horns to the muzzle, is 
much broken up, and everywhere mingled with fawn- coloured 
hairs. It practically stops short of the horns, and is interrupted 
in the neighbourhood of the eyes by a fawn area, mingled with 
a few blackish hairs, for a length of about four inches, while the 
nasal portion does not nearly reach the muzzle. ‘Then, again, 
the dark patch on the shoulder and fore-leg is much less intense 
than in the typical race, and is everywhere mingled with fawn 


Proc. Zoo. Soc.—1913, No. LIV. 54 


820 ON THE CAPE HARTEBEEST. 


hairs, while it stops short of the knees, on which it forms a dark 
patch, and on the shanks is represented solely by a very small 
patch at about the middle of their length. The same lack of 
intensity characterises the dark area on the thighs, which 
appears to be smaller than in the typical southern race. 


Text-fig. 135. 


Transvaal Khama Hartebeest (Bubalis caama selbornei). 


The Transvaal race, which is typified by the aforesaid mounted 
buck, and may be named Dubalis caama selbornei [Abstract 


PeZoSo UGNGs [Pl LAKIN. 


G. Dollman, del. Henry Stone & Son, Ltd., Banbury 


1. SOREX GRANTI. 2. S. ARANEUS CASTANEUS. 


ON HEBRIDEAN MAMMALS. 821 


P. Z.8. 1913, p. 19 (April 15)]|, cannot, indeed, be described 
as a “ Rooi” (red) Hartebeest ; and is evidently a form 
which is either losing or acquiring—I cannot say which—the 
rufous body-colour and deep purplish-black markings of the 
typical southern Khama. The two races may be defined as 
follows :— 


a. General colour rich rufous-brown; face-blaze black and ex- 
tending, with the exception of a narrow fawn band between 
the eyes, from horns to muzzle; limb-markings plum-colour, 
and forming a continuous stripe on front of fore-legs. 

B. caama typica. 


&. General colour yellowish fawn; face-blaze mingled with tawny. 
stopping short of horns and muzzle, and interrupted by a 
long interval in region of eyes; limb-markings mingled with 
tawny, interrupted above knees, on which they form a cap, 
and represented by a small patch on front of shanks. 

B. caama selbornet. 


46. Ona Collection of Mammals from the Inner Hebrides. 
By G. E. H. Barrerr-Hamitton, F.Z.S., and Marrin 
A. C. Hinton. 


[Received and Read April 8, 1913.} 


(Plate LXXXITV.* and Text-figures 136-138.) 


INDEX. 

Systematic : Page 
SOPH GIRUMGU, SDs Ts 2 e-cosdcsonercsaa4ab0 000 seauvac0n venues tery 
JISOCOMGS GUSTO) Vo Vs sgnaceeadcaeceocopecncceccossuoce, SA 
Microtus agrestis macgillivrati, subsp. n. ......... 831 


The peculiar mammals of high interest yielded by investigation 
of the peripheral regions of the British Islands, led Barrett- 
Hamilton to believe that further valuable results might be 
obtained by a closer study of the mammals of the Scottish Islands. 
The despatch and management of collecters was, however, an 
undertaking beyond his means and available time, and the 
matter hung fire until taken up by Mr. W. R. Ogilvie-Grant, 
whose enthusiasm as manager and collector of funds has made 
it possible to amass the material on which this paper is based. 

Karly in 1912 Mr. R. W. Sheppard was sent to the Inner 
Hebrides, and commencing to collect at Bute, this young naturalist 
gradually worked his way northwards through the group as far as 
Coll, which he reached in July. As small mammals are difficult 


* For explanation of the Plate see p. 839. 


b4* 


822 MAJOR G. BE. H. BARRETT-HAMILTON AND 


to catch during the summer months, further investigation of the 
islands was then postponed. 

Among those who have assisted the work of exploration, either 
financially or with local assistance, we may specially mention 
the Duke of Argyll, the Marquis of Bute, Mr. A. R. Macgregor, 
Mrs. Hugh Morrison of Islay, Mr. Colin Campbell of Jura, and 
Mr. Harold Russell. 

The present collection has come well up to expectations, since 
it has resulted in the discovery of three quite distinct forms 
belonging to the genera Sorex, Hvotomys, and Microtus*. 

In working out the fauna of small islands, two distinct phe- 
nomena must be disentangled. A peculiar mammal found on 
a small island may represent a new development under insular 
conditions, it may represent the survival of an ancient type else- 
where extinct, and, as is often the case, it may combine both 
features. We believe that Sorex grantii is an entirely new de- 
velopment, and that Hvotomys alstonti and Microtus agrestis mac- 
gillivrati are instances of ‘“ relics” surviving with little alteration 
from the pleistocene epoch. The meaning of these three forms 
has been partly discussed under the description of each of them ; 
but it would be premature to enter upon an extended discussion of 
the cause of variation or survival in islands. The influence of a 
comparatively recent glacial period, and the extermination brought 
about by man, now proved to be an ancient inhabitant of Britain, 
are difficult to follow; and at all periods the British Islands have 
been the scene of constantly shifting environments and altering 
or variable climates, due principally to the proximity of a great 
ocean and the frequent oscillations of sea-level. Many of our 
common genera of mammals have inhabited the British Islands 
since the deposition of the late pliocene Forest-Bed, but no one 
has as yet ventured to guess how often they may have been 
exterminated, to return again in fresh vigour on the resumption 
of favourable conditions. The history of the British fauna 
resolves itself into that of the various invasions or ‘‘ migrations ” 
of mammals, which have been no less real, albeit more ancient 
and more difficult to trace, than those of Kelt, Roman, Saxon, 
Skandinavian, and Norman. ‘The severity of the struggle for 
existence has in our area undergone profound variation, accord- 
ing as the sea cut off or united our area with Europe, according 
as severe climatic conditions were combined with insularity or 
connection with the Continent. But, however frequently the 
fauna may have been exterminated, renewed junctions with 
the mainland have from time to time refreshed it with new 
stock, and the geological records of what seem from a distance 
almost kaleidoscopic changes are only now in process of 
disentanglement. 


* [The complete account of these new forms appear here, but since the names and 
preliminary diagnoses were published in the ‘ Abstract,’ No. 119, 1913, they are 
distinguished by being underlined.—Eprror. |] 


MR. M. A. C. HINTON ON HEBRIDEAN MAMMALS. | 823 


SOREX ARANEUS CASTANEUS Jenyns. (Pl. LX XXIV. fig. 2.) 


Three, GREAT CUMBRAE. Dimensions * : — 
Head & Hind 
body. Tail foot 
No. 34, male, 28 March 1912 ...... 70 30 12 
32, female, 27 e Feed ‘Nee 70 33 RS 
BS) wine Oe Be A midds Soa 67 32 11 
Average of 3 specimens of both sexes:— 69 32°6 12 


Hight, Bure. 


NowlSe males) 19eMaxeh:-[9i2) so. 70 Bie 12 
Gomme SWAY eam eae 00 33 12 
Osis Ona SOI a cat ee 70 32 12 
(Als ae 20 ienee PaMsoos re 70 36 12 
M4 temale, l9Miarch) 4) 222 65 32 ih; 
15) ee oe * See NR AN 65 32 12 
HORN ee.§ : Se wae iL ee 65 33 12 
GOyeane. bi ANCA O EGE Corn ee. 73 32 12 

Average of 8 specimens of both sexes:— 69°37 33°37 12 


Three, ARRAN. 


INowaSy malese NO esspril, LOZ es. Wi2 36 12 
ie hemealen Mees ea TEES 70 36 12 
CORO: one SEAT ee ( 36 12 

Average of 3 specimens of both sexes:— 72°33 36 12 


Four, Muu. 


INos 143, male, 29-June 1912 |... 78 35 12 
144, ,, SOM cs scin are 75 33 12 
1I38\temale, Zilkume ys. .-..-- 68 40 12 
1 ec Wee ep a ee 68 38 12 

Average of 4 specimens of both sexes:— 72°25 36:5 12 
Six, JURA. 

Nos OSS males la aMiay TOMO Sees. 70 36 12 
TON es Rid rid tee hegat at Boas 73 35 12 
TPs | gp Gee haved: Sarin ee 76 36 12 
11, temalesionys has! | tet ch 73 36 12 
LAO: tibiae WAMU bem s -eussy) | meee 70 33 12 
OU RONDE Wot Aas TAsiot Y3Ey 4 NO 

Average of 6 specimens of both sexes:— 72°5 35:3 12 


For cranial measurements see Table at p. 828. 


The specimens from Great Cumbrae, Bute, Arran, and Mull do 


* Collectors’ measurements: throughout hind feet without claws, tail without. 
terminal hairs. All dimensions are given in millimetres. 


824 MAJOR G. E. H. BARRETT-HAMILTON AND 


not differ appreciably from examples of castaneus from the neigh- 
bouring mainland. Skulls of castanews from South-eastern 
England seem to attain slightly larger dimensions than those 
from northern Britain, and this may indicate the existence of two 
forms. The Jura Shrew is interesting in having a pelage inter- 
mediate in character between that of castaneus and grantii de- 
scribed below; in the four skulls seen the dental characters are 
those of normal araneus, and therefore this form must be left 
with the latter species. 


Sorex Grant. (Pl. LX XXIV. fig. 1, & text-fig. 136.) 
Abstract P. Z.S. 1913, p. 18 (April 15). 


Twenty-two, Isnay. Dimensions :— 
Head & Hind 
body. Tail. foot 
INosiiommaleye 24 sApril 1912) eee 36 13 
(as 7 : Sig," tact 00 36 13 
(ET 26 fae Ie 79 Br 13 
SOs 95 BO on Pst bce 80 36 13 
SOR ie igen He gan Tl 2 80 36 13 
SB, 59 : ai a aa i ee 80 37 13 
Sau 5 Yel bi seth. ae 78 38 il 
OO. 2 Mays inn aaa 78 RD 12 
DB. as Bana ah CS C® 35 12 
OS os (ee eh irene: 75 35 12 
100, 55 Su ve Pe aie TA ie 75 35 12 
Ol,” 4 ae suites 78° 8B 12 
NOS; 55 HO: ayes achat Aes BRE 75 36 13 
i2mtemialles 3 Aspyr; tees naire 83 37 See 
Oman eieee Op) St Reg Ma 75 OM 13 Type. 
SH eee tate 3 (ii yy ba Sg SE pace 78 36 13) 
OB, bg 3 May Chim sdain’ a 35 ib 
OG Ra ee O: £5). oe leave 75 35 12 
157, female, juv., 3 Aug.1912 ... 75 38 12 
158, 99 op. 5) 9 oF NEKO) 34 12:5 
161, cf Ee es Dante reams AS) 36 12 
165, i ee Ot eamn Bae x Neen) 38 12 
Average of 18 adults of both sexes:—  77°3 36 12°6 


For cranial measurements see Table at p. 828. 


This is a very remarkable new Shrew, distinguishable at a 
glance from the common British S. araneus castaneus by its dusky 
upper side in strong contrast to the light flanks. The dentition is 
peculiar in the fact that more than half of the individuals have 
lost the posterior upper unicuspid, sometimes on one side only, 
but more frequently on both sides, the absence of this tooth in 
these cases being compensated for by enlargement of the four 
remaining unicuspids 


MR. M. A. GC. HINTON ON HEBRIDEAN MAMMALS. 825 


Description.—The size and proportions are about as in 
S. araneus castaneus, but average slightly larger than in north 
British specimens of the latter. 

Colowr.—Adults have the upper side deep blackish brown, 
perhaps darker than ‘‘clove-brown”*; slightly grizzled on the 
upper side of the head and neck with brown; the under side is 
silvery or smoky grey rather than yellowish or brownish as in 
castaneus, and this colour runs far up the flanks and shows up in 
contrast to the dark upper side. Between the colours of upper 
and under sides a narrow, inconspicuous, grey-brown flank-band 
intervenes; rarely it is ‘‘ wood-brown,” in which case it contrasts 
more sharply with the back. Nos. 72 & 76 (23 & 26 April) are 
moulting into a scarcely less dusky summer coat, but some speci- 
mens taken in May and Nos. 157, 161, & 165 (3, 5, & 8 August) 
are ina lighter brown coat, between “seal-brown” and ‘ clove- 
brown,” somewhat as in S. a. castaneus, with a flank-band near- 
‘““ wood-brown.” These specimens are no doubt immature and the 
brown pelage is that of immaturity. 

Cranial and dental characters—The skull agrees in form and 
size with that of S. a. castaneus, but attains rather larger dimen- 
sions than skulls of the latter species from northern Britain and 
the Inner Hebrides (vide Table, p. 828). The teeth agree in form 
with those of S. araneus; but the posterior unicuspid (p. 3) tends 
to disappear as shown by the following statistics :— 


p- 3 present on both sides in 7 individuals, 7. e. 43°7 per cent. 
. »  ononesideonly in 4 ee ss 25 g 
.. absent on both sides in 5 % SB 1LEB}" op 


Text-fig. 136. 


Right upper tooth-row of Sorex grantii. xX 8b. 


When p. 3 is absent the four remaining unicuspid teeth appear 
to be somewhat enlarged in compensation (text-fig. 136). There 
is no trace of pigment on the hypocones of the upper cheek-teeth 
nor on protocone of the last upper molar. 

By colour and dentition S. grantii is clearly differentiated from 
all Huropean members of the genus Sorex. Some of these, such 
as araneus araneus and araneus tetragonurus, have the back as 
dark, but none shows the conspicuously contrasted sides. 


* The names of colours in inverted commas are from Ridgway. 


826 MAJOR G. E. H. BARRETT-HAMILTON AND 


In our opinion S. grantii is to be regarded as an insular 
development of S. a. castaneus stock. It has maintained or 
acquired slightly larger dimensions than has the latter in the 
more northern parts of its range; it has developed a peculiar 
colour pattern, and is well on its way to reduce the number of its 
unicuspid teeth, perhaps because of a tendency to enlarge the 
anterior members of the series. The Jura Shrew is geo- 
graphically and morphologically the connecting link between 
S. a. castaneus and S. grantii: it has the small size and dental 
characters of the northern castanews; but its colour pattern is 
modified so as to approach that of S. grantii. 

The species is named in honour of Mr. W. R. Ogilvie-Grant of 
the British Museum, to whose enthusiasm the formation of the 
collection described in this paper is so largely due. 


SOREX MiINuTUS Linneeus. 


Nine, Bute. Dimensions :— 
Head & Hind 
body. Tail. foot. 
No. 18, male, adult; 20 March 1912... 52 35 10 
ts ay a Qt Bee Gade SOO 35 10 
FORA i 20 April Bate 8 OD 34 10 
2, female ,, 2s dMiarcluth ia |... 4 32 10 
fhe Sartore A 20y aes b Gdk TOD: 35 10 
OS eee aaa a 55 i OO) 33 10 
Dangers S e be Sua eo) O) 35 10 
Dona mes 8 * * seh een et 55) 35 10 
GOs gg gp AD eyoraail Pete aire). 35 10 
Average of 9 specimens of both sexes:— 53 aye 2 KW) 


All in dark adult pelage, between ‘“ seal-brown” and “ hai- 
brown.” 


Two, ARRAN. 


No, 59, male, adult, 11 April 1912 ... 58 35 10 
51, female, ,, Oo sf 59 35 leat 
Average of 2 specimens :-— 58° 35 10°5 


In dark adult pelage as above. 


Three, Istay. 


No, 75, male, adult, 26 April 1912 ... 60 40 10 
Sy 95 POO) betes iNet Oe) 37 10 

86, female, ,, “8 As Lee MeN 5574 Bel 10 
Average of 3 specimens of both sexes:— 586 38 10 


No. 85 is in dark adult pelage as above, Nos. 75 and 86 in 
juvenile or summer pelage. 


One, CoLu. 
No. 56, female, adult, 23 July 1912... 60 Ef 10 


MR. M. A. C. HINLON ON HEBRIDEAN MAMMALS. — 827 


Five, TrreEe. Dimensions :— 
Head & Hind 


i body. ail. foot. 
No. 152, male, juv., 16 July 1912... 50 34. 10 


Genie Se lal eli ia ahaa aan 1) 
154, ,, - Wider se Met | e508 TO) 34 10 
OO! Js a Whe ane ae le) 34 10 
151, female, fully | 

mae pe gen He 60! 38! sypi0 


The larger specimens from Coll and Tiree are in the light 
summer coat of young born in the same season; in this pelage the 
hairs are more plentifully grizzled or peppered with tawny than 
in adults. The very young specimens show an interesting early 
pelage, in which the upper side is lighter than in adults, the under 
side very clear and nearly white. 

For cranial measurements see Table at p. 828. 


Elsewhere in the Hebrides S. minutus is known to inhabit Skye, 
Higg, Sanda, Great Cumbrae, Lewis, North and South Uist, 
Benbecula, and Barra. (Barrett-Hamilton, ‘ History of British 
Mammals,’ 11. pt. 8, pp. 111-115.) 


EyoroMYs GLAREOLUS BRITANNICUS Miller. 


Two, Butr. Dimensions :— 


Head & Hind 
bodys. Mails toot. aHars 
No. 6, male, adolescent, 13 March 1912. 90 42 17 10 


Zils eee echullit: PA soaker 5 Smo Bir. IS, yO 
For cranial measurements see Table at p. 830. 


With this material we are unable to distinguish the Hvotomys 
of Bute from the form inhabiting the mainland. In No. 6 the 
cement-spaces of the cheek-teeth are just closed; in No. 27 the 
roots of the cheek-teeth are moderately developed. In each 
specimen the last upper molar has a third inner fold. The 
skull agrees in size and general form with that of britannicus, 
but the external measurements indicate a rather smaller animal 
with slightly larger hind feet than the latter. 


Hyoromys atstont. (Text-fig. 137.) 
Abstract P. Z.S. 1913, p. 18 (April 15). 


Five, MULL. Dimensions :— 
Head and Hind 
bodyzu stan toot.) Har: 
No. 134, male, aged, 18 Junel1912 108 44 18 11 Type 
See, Cr ee pape lel O emme Aan: Oreille 
SOs iss aR OW Gis a LOSE aa Ss Ae 
128} female) (ABiaw;,, cpp OO eA On) i Bro neleliga 
LSKOS Se ie ales eee 55 BMOO BOA i aS (0) 
Average of 5 specimens of both sexes: 105°2 43°38 184 11 


For cranial measurements see Table at p. 830. 


828 


MAJOR G. E. H. BARRE'TT-HAMILTO 


N AND 


Cranial and Dental measurements of Sorex. 


: 


Ke Fas a 
ni g le 
— a 8 o & | | LES I 
be =| | g z fe 2 | 2 | * Be a 
Aya | S| al elena ee BP Vise 12 Si 2 
rai | Sl || oS | oat ose | oe Er @ Ss || ey le Si) 
S| = |. & co Sh SU A lic |= |e ey 8 
Ee aee| ee | — | Sula Aue ey evel 
SUE ES OMS | 5 | EONS 8) ee caloN ais ica Ponte 
S.aranus |>|\/2|e\|e)5/2)|4 | 4/3/\/4|2) 2 Bel 2 
castaneus. |) 12) 2) P/E |S )8) SB) Sie) = le sies 
S|) 2 2 Seles | Sq Sue ie lok | = 
URA. oS a ee aes a eae 
IN@s dL opocee 18°7 5°06 3°5 9:0 47 | 8°2 | 4:4 | 6:93) 9°3 (5-17 | 3°52) 4°51) 2°8 | 1°32 
Prin 2 Olen 18:3: 5°17) 3°5 | 9°2 | 5:0 | 7-7 ize 66 9:3 5:17 | 3°52) 4:29] 2°75) 1°43 
JLL. | | | | | | 
Now14 ona 18°7/ 4°84) 3:5 |9°0 | ... |8:1 | 451] 6:93) 9-4 [5-28 |3°63) 4-4 | 2°86) 1°32 
NAS: s56be 19:0) 5°28) 3°9 |9°6 | 5:1 | 81 | 44 | 7°04 9°8 [5°28 | 3°57) 4°51) 3:08) 1°32 
ee 17°7, 484/33 89 4:9 | 7-6 4:18] 6-49 9°0 [4:95 | 3°52) 4-4. | 2°86 1°32 
4), | | | 
IN@ BB sesso 18°4 5°06 3°5 19-2 | 4:8 | 8-0 | 4:4 | 6°93 9°3 5°17 | 3°63) 4-4 | 2°86) 1°32 
MARR ee): 18°4) 4:95 3°5 |9°0 | 4°8 | 7°9 | 44 | 6°82: 9°3 (5°06 | 3°52) 4rd. | 2°97) 1°32 
Syria 17°9| 5:28 3°5 | 9:1 | 5:0 | 7°8 | 429] 6°71 9:0 [5°06 | 3°52) 4-45) 2°86) 1°32 
Gir “Sosseo 18°5| 5:12 3°5 |9°2 49 7:9 | 4:29) 6°82 9°2 |5°17 | 3°63) 4-4 | 2°86) 1°43 
68 118: H 5:06 3:5 | 92 52 | 7-7 | 434/665 91 5:06 | 3°52 4-29] 2°91) 1:32 
CUMBRAE, | | | acid mea ie 
iNowoO ee | 187) 5:06 3°7 | 91 7°8 | 4:29] 6°71/9:4 5:06 | 352) 44 | 2°91) 1°37 
34 18°9) 5°28 3:7 |... | ... | 7:9 | 4°29] 6:9319°7 15°17 | 3°62)4°51) 2:97) 1:32 
SON Rees 18° 3) 5:06 3°53 | 89 48 | 8:0 | 4:29) 6°71'9°5 4:95 | 3°46) 4-4 | 2°86) 1°32 
ARRAN. | | aes 
IN@s BY soo o05 18-2) 506136 |89 ... | 7°8 | 4:29) 6°71 5:06 | 3°46) 4°51) 2°97) 1:43 
GIS) sop ave 18°6) 5°22, 3°6 | 9°4 |5°0 | 8:0 | 44 | 6°93 9°3 5:11 | 3'57| 4°62) 2:97) 1:37 
S. granti. 
ISLAY. | 
No. 73 .| 1971 5:28 3°7 |9°3 | ... | 8:1 | 4:4 | 6°93) 9°6 5°17 | 3°52) 4°51) 2°75 1-48 
Ty eee 19°83) 5°39 3°8 | 9°6 5:0 | 8°4 | 4°62) 7:15 9°9 5:28 | 3°63 4°55) 2°8 1:43 
MN TWO” 7B © sasooe 19°0 5°28 3°8 (91 52 | 83 | 4°62) 6:93 9:9 517 | 3°57) 4°51) 2:97) 1:32 
Ml 18°9) 5°39) 3°8 | 9°3 | 5:1 | 8:2 | 44 | 682 9°7 5°17 | 3°63) 4°56) 2°86 1esy7/ 
80 18:8) 5:28) 3°8 | 9:4 | 5:2 | 81 | 44 | 6°82, 9°6 5°17 | 3°63) 4°62| 2°97 1:43 
Sou raat 19:0|5°5 | 3°8 |9°2 51 | 84 | 4°62) 7-2 | 9°9 5:39 | 3°63) 4°62) 2°97 1:43 
SA: 191 5:28 3:8 |9'2 52 82 | 4:51) 6:93 98 5:28 | 3°63) 462] 2°97 1:43 
95 18°6| 5°17 3°8 |} $9 5:0 | 81 | 4:4 | 6:82 9:4 5-17 | 3°63) 4°51] 2°86 1°32 
HOW scsccc 18°2,6:17 3°5 |90 5:0 | 7°8 | 4°29) 6°71) 9:4 817 | 3°57 4:4 | 2°75 1:37 
OMe eeeee 18°5 5°33: 3°7 | 9°2 5:0 | 7:9 | 4:29) 6°71 9:6 \5°06| 3°46 4°67|2°5 | 1°43 
NOS. cceoce 18°4 5°39 3°8 | 9°4 |50 | 7:9 | 429/66 9°6 4°06 | 3°52) 4-62) 2°86 1°32 
NOS soacoe 18°6| 5°17 3°7 | 9:1 |5:1 |.7:9 | 44 | 6°71|9°3 (5°17 | 3:57| 4-4 | 2°86 1°32 
S. minutus. LA WHE. I 
BUTE. el 
Nowr2) gine 15:2) 407) 2:7 | 7°3 | 35 | 65 [3525-4 |7°3 4°07 | 2°97) 2:97 214 088 
¢/ 15°5| 4°18) 2°7 | 7:2 | 4:0 | 6:6 | 3°74! 6°61) 7°7 |4°29 | 3°08! 3:08) 2-2 0°88 
24 15°2| 3°96 2°7 |72 | 4:0 | 63 | 3°52) 5°39 7:4. 4:07 | 2°86) 2°97| 2°14 0°88 
DASA ewe 15:2] 3:96 2°8 | 7:3 | 4:0 | 6:3 | 3°52] 5°39) 7°4 4°07 | 2:97 2:97) 2°14 0°88 
26 15:0) 3°96) 2°8 |'7°3 | 4:0 | 6:2 | 3°47| 6°8 | 7°3 4:07 | 2°86) 2:97) 2°2 0°88 
CORP ae 15°3] 3°96] 2°7 | 7:1 | 4:0 | 6°83 | 3°52) 5°5 | 7-4 4:07 | 2:97} 3°08) L 98 O'77 
ARRAN. | | | 
No. 51 159) 418 2°8 | 7-1 4:0 | 68 | 3°85 5°83 7°8 4:18) 3:08) 3°08 2:2 | 0°88 
59 .| 15:9] 4°18) 2°8 | 7-2 | 4:0 | 6:6 | 3°74) 6°72) 7-8 4°29 | 3°14) 3:08) 2:2 | 0°88 
ISLAY. | 
No. 85 | 15:4) ... 127 |7-2 | 4:0 | 65 | 8°74) 6°61) 7°6 4:23 | 3:08) 3:08) 2:25, 0-94 
| SGN ieee 15°6) 4:18 2°8 73 (40 | 65 (3°74 561,76 418 | 3°08 3°08) 2°31) 0°88 


* These measurements were made with an eyepiece micrometer and 5-inch objective, 
vide Hinton, “ British Fossil Shrews,” Geol. Mag. n. 8., dec. 5, vol. vil. pp. 537 & 538. 


MR. M. A. C. HINTON ON HEBRIDEAN MAMMALS. 829 


2 


General characters.—This mouse is a little larger than LZ. 
glareolus, approaching EL’. norvegicus, but the tail is shorter than in 
the latter, and about as in glareolus; smaller than H. skomerensis 
or cesarius. ‘The ears are about as in #, glareolus. The hind 
feet are as in norvegicus and skomerensis. 

Cranial and dental characters.—In size the skull is larger than in 
H. glareolus and agrees with that of norvegicus. As in the latter 
form, the jugals are heavy, but the curvature of the zygomatic 
arches agrees with giareolus. The brain-case is very broad and 


Text-fig. 137. 


Dorsal view and profile of skull of Evotomys alstoni. Natural size. 


smoothly convex, the temporal ridges being but faintly indicated 
even in aged skulls: the parietal region is convex in dorsal profile. 
with the highest point a little behind the middle of the parietals, 
instead of flattened ; these features impart an appearance of 
relatively greater cranial capacity than is seen in any of the 
other European species of Hvotomys. The postorbital (squamosal) 
processes are not conspicuous. The interorbital region is broad, 
with a wide shallow median sulcus. The nasals are rounded 
or slightly and narrowly emarginate behind, ending flush with 
or slightly behind the ends of the premaxille; they are slightly 
longer than the diastema, expanded in front, their lateral borders 
slightly but distinctly concave. The rostrum is shallow, as in 
EL. norvegicus, its least depth behind the incisors not exceeding 
the anterior width. The bulle are nearly as in norvegicus. The 
last upper molar has a third inner fold and fourth inner salient 
angle—the latter being usually well developed. 

The colowr is much darker than in /. skomerensis, but 1s very 
similar to that of #. glareolus, being deep russet above ; the under 
side is richly washed with yellowish or buff. 

This is an interesting and quite distinct species, recognizable 
externally by its generally large size combined with small ears 
and short tail, while in addition its skull characters are quite 
distinctive. On the whole it makes a nearer approach to H. nor- 
vegicus than to any other species of Hvotomys. 


830 MAJOR G. E. H. BARRETT-HAMILTON AND 


€ 


Cranial measurements of Hvotomys. 


eee | x 
ele | are | 2 \2 
oD ae | dc) 5 = 4 
2 | *S | a a | = | | at : 5 
eeu t| SSlch ea te Ome iS aha e ere 
ese een ea ol Sess P=surs l= 2 
|e ps sees a |>S| 6 a= 
eee |e era ene 
S| See Se eS he |e Is 
6 5 lS (Se EAE h eae Seale 
peGlarmoritaniicss ===. 2a |2s ee a a a 
| | | | | 
Chingford, EssEx.........) 24°2 14:0) 3°9/ 11:2) 60 68 65) 55 145 51 Aged. | 
Bure, No. 27, male ...... | 23°5| 12°8) 3°8 11-2) 6:0) 6:7 | 64 5°4/14°6 5:4) -Adult. | 
| | | | | 
| | 
. | | | | | | 
E. alstoni. -———|—— — ny S| Site 
Mutt, No. 134, male ...| 25°3) 18°9| 8°8)11°5 64) 7°5| 7-1) 56/156 5-4) Aged. | 
ss S60 (2573/1473) 3:9)| 12-1, 6:61) 7-81) 7271) bez) e671 4) ee 
ee 136, ,, _.../ 249145 41/11°8 63) 7:3) 68) 56 15:3 51) 
ye 130, female .| 24"1) 135) 3°9| 11-4) 6:0) 6'8| 63| 52|1571| 54) 
lie icone 
EH. norvegicus. fe | a ne | ee eee 
Hardanger, 5.8.5.2 ...... 24°5, 134) 3°8/ 11-1] 6:0! 6:9) 7:0| 5:°4| 15:3) 51) Aged. 
Bergen, max. (Miller) .... 26°2 14°6 4°0| 12:0 6°6 80) 7°8| 58160 54) Adult. 
MIcROTUS AGRESTIS EXSUL Miller. (Text-fig. 138 a.) 
Four, Muu. Dimensions :— 
Head & Hind 
bod\zylbarlearetootameeLan: 
No. 139, male, ad., HY wine WOU, MAD. (BG UD 12 
NL se ie oy OTe aa ike a LOO “SO . Me 
132, female, subad., 15 _,, us Iss NasA By ee 
145, a UNE 3 July as OEY GO hss) 
Hight, ARRAN. 
No. 41, male, ad., Pyare We = [PAS aI IN) Le 
AOS Tae ae sVULV See % NOG wage UO sl 
AG eemesuioaGs. se4 o ey BBM) oclkss 
DOL ee 3 Dy, ae : WAS 2 Way AS) LE 
S25 ks 3 Ongar = Greta Gs The 
G25 Seems peel yee 5 NAGY SG) IG 4s 
AS Lemaleweiives | oO)», r NOB BR. WO Ll 
AU. Tees ACen. GA.” ee ; Ud Leagan lle WLS): 9 hel 
(damaged) 
One, GIGHA. 
No.127, male, adult, 23 May 1912. 125 33 18 12 


MR. M. A. C. HINTON ON HEBRIDEAN MAMMALS. 831 


Hight, Jura. Dimensions :— 
Head & Hind 
body. Tail. foot. Bar. 
No l00S male ssubad Lo May 1912) 15s "30 19 1s 
TESS . +5 lO >; i AS ee MOR we 2 


116, 3 “A if so i TUS BY OAD). cess 
ina Stenralesre Ori: ss IPAQ) SVE co ME A 
Sea BA aan aS Ss TIL a4 TE) 112 
EAD tee lie. Pee . IAG BOQ Wes We 


Tite Tenemaene eter ts oak Mi (N95), (TLOMAMiTS 
124 oo LPs bed asl Sa ew est Hunts alls Dl aOMeaAS 


For cranial measurements see Table at p. 834. 


M. agrestis exsul Miller, of which, in addition to the material 
in the British Museum, we have examined several specimens 
from North and South Uist, kindly lent by Mr. Eagle Clarke, 
of the Royal Scottish Museum, Edinburgh, is a thick-coated form 
with light upper side and flanks, and under side heavily washed 
with buff. The fur of the back in winter reaches a length of 
about 12 mm., including the hght tips of about 2-3 mm.; the 
longer hairs reach about 15 mm. Weare not able to distinguish 
the specimens from Mull, Arran, Gigha, and Jura from this sub- 
species. Those from Mull are practically quite typical. 

The following Table shows the character of the anterior upper 
cheek-tooth in the skulls examined :— 


Mutu. ARRAN. GicHa. JURA. 


Ath inner angle quite absent... 1 
represented by microscopic ‘ 

; VESEICO aed as eo Porcine 2 yee 12 : ae 

5p RCTEMIL LONG GUIS BUOVCE 5 sconecoae 1 4 3 8 

Specimens examined ...... 5) 9 1 8 21 


Some features of the skull are discussed below in connection 
with the form living in Islay. 


MIcROTUS AGRESTIS MACGILLIVRAI. (Text-fig. 138 B.) 
Abstract P. Z.8. 1913, p. 18 (April 15). 


Fourteen, Isuay. Dimensions :— 


Head & Hind 
body. Tail. foot Ear. 
No. 78, male, subad., 26 April 1912. 115 34 19 13 
OA siesshs | sae iamles 3 i OB ellis “anal 


Hy Ween exohribin’ SON, LAOS A Ride P12 
BOC vig cls nel barca te, el PELE SOM ee 
OOo Me, 2 May LB SYP IG) 1 


vlna 25), 1132) edi yaeertlyspe 


832 MAJOR G. E. H. BARRETT-HAMILTON AND 


Head & Hind 
body. Tail. foot. Har. 


No.103, male, juv., @) Jileng TESA IO aah MS) TL 
MOST as excholke, IMO) 8 Ff Wy ase) Ns) 
159, ,, subad., 3 August ,, WO, aie IS) 
NOS Am 5s a3 (ee me . ie 8H IS le 

87, female, , 30 April - ay Say UO 
I 4,0 adulty 2 May. re IPAQ By IDS 
160. |e subbadk., 3 August 4 Itsy ee IG) 
G2 es Fan 46 e a WO aa It 


For cranial measurements see Table at p. 854. 


This mouse resembles J/. agrestis exswl in size and general 
proportions, but may be recognized at all seasons by its much 
thinner coat and the much lesser amount of yellowish wash on 
the under side, resulting in the slaty bases of the hairs being 
visible and taking part in the general coloration; the under side 
thus resembles that of immature exsul. The upper side is slightly 
less brightly coloured than in easwl; the hairs of the back reach 
a length of about 8 mm. 


Text-fig. 138. 


Dorsal view and profile of skulls of (A) Microtus agrestis exsul and 
(B) UM. a. macgillivraii. Natural size. 


A quite young male (No. 79) is in dusky pelage, with very 
slight development of the ight hair-tips. The hair-tips of the 
under side are of a very ight tint, increasing in depth through 
the flanks to the back and poll, where they are deepest but never 
approach the rich buff of adults. The rump of this specimen is 
more dusky than the back. 

In each of the eight skulls examined the anterior upper cheek- 


MR. M. A. C. HINTON ON HEBRIDEAN MAMMALS, 833 


tooth has a distinct and often rather large fourth inner angle. As 
is usual in microtine subspecies, the subspecific cranial characters 
first become obvious in old age, although, if care be taken to 
compare only skulls of equal age, close observation will find the 
beginnings of such characters in younger stages too. The oldest 
skull, the type (No. 102), is fairly comparable with that of No. 139, 
a M.a. exsul from Mull, from which it is distinguished by its deeper 
rostrum, lighter jugals, and more nearly vertical occiput; because 
the occiput is less sloping the interparietal is a little less 
reduced,—its posterior border is straight instead of sinuous, 
and it is rather longer antero-posteriorly. The most striking 
difference is seen, however, in the form of the ‘“ shield” or 
flattened dorsal area delimited by the temporal ridges upon the 
sides of the braincase. In ewsul (text-fig. 138 a) that part of the: 
temporal muscle which arises between the interorbital region 
and the glenoid articulation continues apparently throughout 
life to extend its area of origin by creeping up the sides of the 
braincase and so pushing the temporal ridge backward ; the result 
is that in old age the anterior width of the “shield,” taken 
between its antero-external angles, is considerably less than its 
glenoid width: in the Islay mouse the upward creeping of the. 
temporal muscle ceases at a relatively early stage of growth, and 
the result is that in old age the ‘ shield” presents a form which 
is found only in relatively early stages of development in exswl. 
The oldest skull of ewswd from South Uist (Edinburgh, No. 300) 
is younger than the skulls from Islay and Mull just described, 
the temporal ridges not yet being quite fused in the interorbital 
region, but the form of the “shield” is already more modified 
than in the aged type of the present subspecies. 


MICROTUS AGRESTIS NEGLECTUS Jenyns. 


Three, Bute. : Dimensions :— 
Head & Hind 
body. Tail. foot. Lar. 
INOmOomemalemanuve OPA pri onic Ne BO lly NB 
Scena, 4, IP Wileigela By soc ICO "2G eh 
6 a ea SeAsorilbh” io atey cs. PEL NG alates: 


For cranial measurements see Table at p. 834. 


The short tails are, no doubt, due to immaturity. 

Making allowance for the youth of these specimens, we 
ean detect no character by which the skins or skulls can be 
differentiated from those of J/. agrestis neglectus of the mainland. 

In his recently published ‘Catalogue of the Mammals of 
Western Europe’ (1912), Mr. Miller treats the various agrestis- 
like forms as subspecies. We think it preferable, somewhat 
as proposed by Barrett-Hamilton in 1896 (P.Z.8. 1896, 19th May, 
pp. 599-608), to regard these forms as belonging to two species, 


834 MAJOR G. BE. H. BARRETL-HAMILTON AND 


a larger northern with isolated southern colonies, now known to 
be older in western Europe, a smaller southern, now known to be 
a recent comer. Such an arrangement shows the relationships 
most clearly. The earliest known member of the group in Europe 
is a species of which the fossil and fragmentary remains have 
been found in the early middle pleistocene brickearth of the 
Thames at Grays in Essex; whether this form has any close 
affinity with any living now it is impossible to say. In the 
late pleistocene deposits of Britain, such as that fillmg the 
Ightham Fissures, and in many of the caves, remains of a 
large form not certainly distinguishable, with the available 
materials, from J/. agrestis neglectus occur. The Skandinavian 
agrestis, the Hebridean exsul and macgillivrai, the pleistocene 
and recent British neglectus, together probably with the southern 
French and Swiss levernedii, are all intimately connected forms 
and may be regarded as subspecies of J/. agrestis, and of thera 
macgillivrati may be counted as the most primitive. At some 
time since the close of the pleistocene period J/. agrestis 
neglectus has been succeeded in England and the lowlands of 
Scotland by a smaller, brighter form, which being a newer 
immigrant, may well be granted specific rank as J/. hirtus 
Bellamy, with Jf. hirtus baillona de Sélys of France as a 
subspecies. 


Cranial measurements of Jicrotus agrestis. 


es : = | [3 
ey il) cel > | |= 
& = a ee es | 3 
= |wiBeall & Saplomeialveame \ 2. 
heen ie Be et ehh SS | es sf SS 
M. a. exsul. Ss fia a So 4 Sys 
Murz. No. 139, male. | 286 | 162 | 34 | 195 | 71 | 84 | 81) 69 
ARRAN. 43, juv. f. | 26:8 | 15:2 374 | 116 6:2 68 7-4 69 
52, 5, m.| 27:0. | 15:8 316 | 125 6°4. 75 TA 70 
JURA. 119, female | 27°3 | 16°5 vA | 12°5 6°7 WS || BPS 70 
N. Uist. CY op | 26°) 14°6 33 | 116 | 64 Vs NW YB 68 
S. Uist. R00, 4 |) Br3 | 168 | 33 || Tp, 68 79 | 78 65 
BN sath | 
M. a. macgillivran. | | 
i | | 
Istay. No. 89, male. 264 | 15°3 38 | 12:0 67 79 | 7:2 66 
TO eel 26:5 | 16:3. |) 3:6 1 d1s8) |, G2, 16 726. Ives) eee 
OS See al e2OrOn oA: 37 12°0 62 | 76 7-4, 68 
87, female .| 27°4 | 16:0 37 | 127 1 OB 2) 7 68 
102, Type .:.| 287 | 164 | 37 | 196 | 67 | 87 | 78 | 71 
| 
M. a. neglectus. | | | | 
| | 
Bure. No. 67,juv. m. | 243 | 140 36 | 115 | 60 | 68 69 61 
Bh SUS paley | Gl Ge Ge) ay 


MR. M. A. C. HINTON ON HEBRIDEAN MAMMALS 


APODEMUS SYLYATICUS SyLyATICUS Linneus. 
Seventeen, Buts. 


Head & 
body. Tail. 

IN@ Il, imeiley @ Wiesel OR Say ahs} 

Ae dess gos eee ; VO SO 
ead eo Loud ese Ms 90 80 
10, 29 16 7) ? 90 (es 
11D), dane rg 90 77 
Deo Be WE 100"! 86 
23, br) bh) 9) 9 87 76 
21 Ah eT A aN NSE 
3], 2? 23 2? 9? 95 75 
5, female, 12 Be 80 78 
Sia es ga etek aes 59 S879 
9, ”? 14 ” ” 90 75 
gg OT HD 
A, gg AW Ne ; 90 90 
22, ” ” ” ’ 83 67 
DORR RUM ae , 80 80 
3 tne 4 ett, 22) A) a 807-73 
Average of 17 specimens of AEE 
both sexes : sCachsterae wach | Ber Oe 

Six, GREAT CUMBRAE. 

No. 33, male, 27 March 1912. 95 92 
SD) el Maer Deke aa OH OO 
SOR POM: ae 5 95a Oe 
37, ” ” ” ” 90 88 
38, female, ,, 3 be 93 90 

Ona: vO. os. 3 JO 90 
Average of 6 specimens of | 93 90-3 


A rather rufous series, perhaps due 


worn. 


both sexes . 


Nine, ARRAN. 


No. 44, 


Average of 9 spec 


towards the large foot and short ear of hebridensis. 


male, 3 


9? 


3 


99 
female, 


39 


April 1912. 


ye) 


both sexes 
This series shows a rather large reddish mouse, with a tendency 


>)%) 
imens of 


* Trapped on moss-covered bank. 


t Trapped in middle of rough pasture. 


Dimensions :— 


100 «88 
10S es 
105-86 
100. 85 
LOO Wess 
SORES 
ea) to) 
JOM RE Ts: 
wa) fen) 


98°3 84°59 


SH SCN 


OU 


bol po bh po by pp 
WNMDNWPH bd w& 


kD bD DO bO BO bo 
Gs Oo CO to KD CO 


Skull 
Kar. Coaldy Ve 
TLegeey 
115)%15) 
15 22D 
Loui yy 2289 
15 
15 23°2 
14 M95) 
15 Sal 
Dee ae 2) 
14:5 21:5 
14:7 22-2 
S25) Die 
16 Hil 
15 22°8 
14°5 20-4 
14°5 21°8 
16 21°6 
14-76 
5 23°4 
14:5 22°8 
16 
16 
15 Q32, 
16 22°4 


22°8 15:4 


14:5 
15 
14 
14 
15 
14 
15:5 
14 
14:5 § 


bo 
i 
=) 


bo 
wo) 
=r) 


bo bo bo bo 


02 He 02 19 
OH Oo 


+ Trapped in grass near sea-shore. 
§ 'Frapped in heather on side of cliff. 


Proc. Zoou. Soc.—1913, No. LV. 


DD 


Or 


to late winter coat being 


836 MAJOR G. BE. H. BARRETT-HAMILTON AND 


One, GIGHA. Dimensions :— 
Head &. Hind Skull 
body. Tail. foot. Kar. c.eb: I 


No. 126, female,22 May 1912. 100 85 22°5 15 
A rather large nursing specimen with dimensions tending 
towards those of hebridensis. 


Ten, Istay. (8, Sheppard ; 2, Royal Scottish Museum.) 
No. 97, male, 3 May 1912. ee 4b 283 13} 


99, eee mY Gee Ms Vo ie ses Ns Ba) 
93, female, 2 ,, 5 SSS Ome 2 melee 22:1 
4 a. Re e LOO. sh 23 115) 24°] 
10405, Ome ss 5S {3 Bo 23°7 
GAs aes ger AKWe® JOM SO nee 22 onal 3 
GBs 55 She ee) Go OO tom a oem 
NGA eRUVGS: ye ae Co CLG 8 
Royal Scottish Museum :— 
INO, BOG HemNAes ocwanbesodsonsces OIL Baie BR. abo 
SO Ot emEey PES Li Sia ee Pec 15 OO 2a 1G 


Average of 9 adults of | 


Inatilh SEEElosesyese: 93°3 81:6 22°5 14 


A moderately large mouse, with back heavily washed with 
black, probably indicating the new summer coat; stomach clear 


white, and shght trace of yellow collar. 


Three, JURA. 


iINoswl22; male, 18!) May W912] 9884 24) a4. 5 
123, 9 - HOOe 5. 722) EL oy 23-9 
Les, Resanenlley 16 - a On me 1 en) 

Average of 3 specimens aa : 

of both sexes......... | HY GSO 2a) Mew 


Five, Mutt. 
Non 29) male, 13 June 19125 9a 922 2:oe It 


1th, 115 Gy Oh toe) foi) 24 tlie ea 
TSS ce tlO: Lt) OE NOOw) Ging e22nn gl eeu eS 
Sie ” 20 ” 39 92 88 24 14:5 

140, Ne Te ak KOO IO Eb NEG) Bdlom 


Average oe WINES, so0ose Vi Sie 2322) ASA 


Four, TIRE. 


No. 146, male, 8 July 1912. 105 88  23°5/13°5 
148, PO es, at as, 105 88 24 (13 
149, a cla tag ee TOOW S73 ete guage 

iLO), Fennel. Layeng t LOOM RS Sire tego 


Aver age of + specimens of 102° 84:25 23:1 13:5 
both SOMES manatee: 


| 
| 
| 
| 


MR. M. A. C. HINTON ON HEBRIDEAN MAMMALS. 837 


Six from Tiree, in Royal Scottish Museum :— 
Head & Hind 
bodys Tail; foot. Har: 
No. 344, male, 19 Nov. 1906. 80 85 23:5 14 
Be), og, lB) IPGlo SOV, OB SR WS} i eyes 
342, female, 19 Nov. 1906. S68 OO BF la 
Oe ess |, Maat tse 4s SHA fe BB A 
Ato). RR eet oe oA VS (ee Bil 1S 
BAG Re ae Wee 5 S277) 2805 ono 
Average of 6 specimens of |... : : 
both sexes ........ esl Boe BO le 
Average of 10 ara 92-9 84:5 21-1 13-8 


The specimens from Tiree and Mull are, when adult, apparently 
above the average size, and tend to have the large hind feet and 
small ears of hebridensis. They also approach hebridensis in a 
tendency to have the under side washed with yellow. 


Cranial measurements of Apodemus sylvaticus sylvaticus. 


We are unable to distinguish the specimens from the islands 
from A. sylvaticus sylvaticus of the mainland. The dimensions 
recorded above afford an idea of the variability encountered in this 
form: even allowing for errors on the part of the collector, and 
for the differing ages of individuals, there seems to be an inherent 
tendency to vary. On all the islands, except Bute and Great 
Cumbrae, there is a dimly seen tendency to acquire greater size, 
larger hind feet, and shorter ears, 7. ¢.,a tendency to vary in 
the same direction as hebridensis, as further shown by the speci- 
mens from Tiree and Mull, in which the belly tends to be 
washed with yellowish tints. 


DD* 


ener aR 7 | 

| | os : a Ge rage 

| 3 oS og ape 28 

= 5 S| “ | rece 38 

2 @ ices Py Bey ae | oe | aS 
ee | eS eS) iy cS om | oo Be] se SS 
Se S 25 aS) S : B Sa | 2a | 
Bute, 21, male ...... 23:2 | 13:0 40 | 112 | 80 | 102 66 39 | 142 3°6 Aged. | 
POO Napea Theat: 23°1 | 12:8 Lomeli? 80 | 95 67 40 | 143 38 jAduli.| 
| Cumbrae, 38,female.| 23°2 | 13°2 38 | 11:8 82 | 92 | 68 39 | 145 36 | Aced. | 
| Arran, 48, male......] 24:0 | 136 | 4:0 | 12-0 83 | 68 4°2 | 149 3°7 |Adult.| 
| Islay, 94 female ...... 241 | 140 | 41 | 114 HS Mes ew a ZeO 37 | 149 37 | Aged. | 
Jura, 123, male......| 23°9 | 13:0 |. 40 | 11:6 | 82 9°4. 7-0 4:0 | 144 3:8 |Adult.| 
Wind, WO Gy earn 24°2 | 13:0 | A) | wale, | SL) OA PS 4-1 | 14:7 | 38 | Aged. | 


838 MAJOR G. E. H. BARRETY-HAMILYTON AND 


Mus muscuuus Linnzeus. 


One, TIREE. Dimensions :— 

Head & Hind 

body. Tail. foot. Ear. 
No. 147, male, 10 July 1912... 80 Co 18% 11? 


An example in the yellowish outdoor pelage. It was trapped 
in sand-hills. 


In conclusion we have to sum up the distributional evidence 
and offer a few remarks, of a quite tentative character, upon 
its meaning as it appeals to us at present. Of the shrews, 
S. minutus oceurs throughout the Outer and Inner Hebrides, 
and in the Orkneys, Ireland, and other islands; S. araneus, 
on the other hand, is confined to the mainland, the Inner 
Hebrides and such islands as Anglesea and Wight; it is 
represented in Islay by the peculiar S. grantii connected 
morphologically and geographically with the parent form by 
means of the partially differentiated Jura form. We may 
therefore perhaps conclude that S. minutus was the first form to 
arrive in this region; but this point is of little importance, 
because both species have had representatives in Britain from an 
extremely early period, and the problem before them in the past 
may have simply resolved itself into a contest of endurance to 
decide which of them could survive the rigours of the glacial 
period in the outlying districts. Be this as it may, one fact 
stands out clearly, that Islay has been separated from the 
mainland longer than have the other islands of the Inner 
Hebrides, with Jura approaching it in age. The microtine 
species lead us to much more definite conclusions. We have in 
the Inner Hebrides two genera, Microtus and Hvotomys ; the 
former alone, so far as is known, inhabits the Outer Hebrides as 
well. As is well known, neither genus occurs in Ireland, and 
Microtus alone is met with in the Orkneys. There is a great 
difference, however, between the Orcadian and the Hebridean 
species of Microtus: the Orcadian forms are members of the 
M. arvalis group, while those of the Hebrides are members of 
the WM. agrestis group. Now, at first sight, having regard to the 
fact that J. oreadensis and WW. sandayensis have a near ally in 
the Guernsey J/. sarnius and another in the late pleistocene 
English J. corneri, as well as to the fact that this group is no 
longer represented on the mainland of Britain, one might feel 
inclined to look upon the Orcadian fauna as an older one than 
that of the Hebrides. On the other hand, we have to remember 
that both the MW. agrestis group and the genus Hvotomys appeared 
in Britain long before any of the other modern groups of Voles. 
Each is represented in such an early horizon as that marked by 
the Grays brickearth by a form much like the living ones; while 


MR. M. A. C. HINTON ON HEBRIDEAN MAMMALS. ——— 839 


the J. arvalis group in a modern guise does not appear until the 
late pleistocene era. Secondly, we have to face the remarkable 
fact that both #. alstoni and M/. agrestis exsul, with its ally 
macgillivrait from Islay, are most closely related to the Skandi- 
navian species. Moreover, MW. agrestis macgillivrati appears to be 
a little more primitive in its organization than ewsul; the latter 
form in turn a little more primitive than true agrestis. The 
interpretation of these peculiar facts which we suggest is that 
the Hebrides have formed a refuge for some very old mammals ; 
that from the former continuous land area, of which they are 
now the disconnected remnants, certain of these old mammals 
were dispersed along former land bridges to Scandinavia. The 
proximity of the Atlantic would ensure the existence of much 
milder conditions in the Hebridean area during the glacial 
period, at whatever moment in pleistocene time that remarkable 
event transpired, than would be present elsewhere to the east. 
It will be seen therefore that we are practically in full accord 
with the views put forward by Mr. L. Stejneger in his very able 
and suggestive paper “ On the origin of the so-called Atlantic 
Animals and Plants of Western Norway ” (Smithsonian Misc. 
Coll. vol. xlviii. p. 458, 1907). Just as the presence of S. granti 
implies that Islay was separated from the mainland at an earlier 
date than were the other islands of the Outer Hebrides, so the 
existence upon it of J/. agrestis macgillivrati and of a distinct 
subspecies of Stoat, Mustela erminea ricine (found in Jura as 
well), may be taken as evidence that it was detached from the 
common Hebridean land surface at a relatively early moment. 

The early detachment of Islay and its peculiar mammals has 
an important bearing on the origin of the Irish Fauna. It 
indicates that Islay and Ireland have not been connected since 
the genus M/icrotus and Sorex araneus reached the Inner Hebrides. 
And if that were so, the existence of the deep North Channel 
between Ireland and South Scotland must at the same time have 
interrupted communications in that direction also. We are 
thus unable to accept E. R. Alston’s suggestion (Fauna of 
Scotland, 1880, p. 5) that Ireland received its mammals from 
Southern Scotland, and are forced to derive the Irish Fauna from 
England and Wales (see Barrett-Hamilton, Clare Island Survey, 
Mammalia, Proc. Roy. Irish Acad., March 1912), 


EXPLANATION OF PLATE LXXXIV. 


Fig. 1. Sorex grantii. Natural size. 
2. Sorex araneus castaneus. Natural size. 


840 MR. T. H. WITHERS ON 


47, Some Miocene Cirripedes of the Genera Hexelasma 
and. Scalpellum from New Zealand. By Tnomas H. 
Wiruers, F.G.8.* 


[Received May 6, 1913: Read June 3, 1913. } 


(Plates LXXXV. & LXXXVI.7 & Text-figures 139, 140.) 


INDEX. 

Page 
Introduction ........ AR La cde Ae SOCK. ct OM0 Leet ner nA) 
Hexelasma Ron iandicuit (Hector VOM b ardaeeE cadaunaeeeeneradcumely tor all 
Structure and Affinities of H. aucklandicum .................. 846 
TE LCROCUGISTIAGD See | wade Bea bone Pee Hedoai can mps OprinbiepaeoRePoRaS dss haehonoas: ts! 
Scalpellum Bispianuns Sie seeeeeceee Hae re ee CAS 
Scalpellum Fae angulation, Oo INS saves 850 
Distributions (Geological) yeas) eeeeseheaseeaes B41, 848, 851 


This paper contains the results of a study of the remains of 
the ‘gigantic Cirripede” (= Hexelasma aucklandicum) from 
New Zealand, as well as some notes on a smaller species of 
Hexelasma, and descriptions of two new species of Scalpellum. 
One of the latter is founded on some valves in the Geological 
Department of the British Museum, and the remaining species 
of Scalpellum and the small Hexelasma were found associated in 
the matrix with the remains of Heaelasma aucklandicum. 

Remains of a gigantic Cirripede have long been known to 
oceur in the Waitemata Beds (Miocene) of Motutapu Island, 
Auckland Harbour, New Zealand. These remains have been 
considered by Sir James Hector (1887) and Prof. W. Blaxland 
Benham (1903) as belonging to a pedunculate Cirripede; but 
while the former referred them to the genus Scalpellum, the 
latter thought that they approached more closely to the genus 
Pollicipes. 

On learning of my wish to see some of these remains, 
Prof. James Park was good enough to write to Dr. J. Allan 
Thomson, Paleontologist to the Geological Survey, Dominion 
Museum, Wellington, who most kindly sent me the actual 
specimens collected by Prof. Park in 1887. Prof. Park wrote 
also to Prof. Benham, who sent me plaster-casts of the specimens 
figured by him in 1903; these casts are now in the Geological 
Department of the British Museum. My thanks are therefore 
due to Professors Benham and Park and Dr. J. Allan Thomson, 
and I have also to acknowledge the kindness of Dr. A. Smith 
Woodward in allowing me to describe the new species of 
Scalpellum in the Geological Department of the British Museum. 


* Communicated by Dr. W. T. Catman, F.Z.S. 
+ For explanation of the Plates, see p. 854. j 


WARS MNS) ENE ILYOCQVi 


i 


Huth imp. 


del, et lith, 


AH. Searle 


HEXELASMA AUCKLANDICUM, Hector sp. 


A Meena 
RT EU RUE 
as 


Ne eat 
ey 


Ate 


i oh 


IPAS) hee gel ILAO.ORVAL 


Y 

‘ 
4 
1 
; 
I 
1 


Huth imp. 


t ith. 


AH Searle dele 


+6.SCALPELLUM SUBPLANUM. sp.n: 


3 


Fig 


TI8.SARCOSG ALPELLUM) UNGULATUM. sp.n. 


i) 


Fis 


FOSSTL CIRRIPEDES FROM NEW ZEALAND. 84] 


BALANIDA. 


Genus HEXELASMA. 


1913. Hewelasma P, P. C. Hoek, Siboga-Expeditie, Cirripedia 
Sessilia, p. 244. 

“Compartments six; carina, carino-lateral, and lateral compart- 
ments with ale, but without radii, the rostrum having neither 
radii nor ale. Parietes not porose and without longitudinal 
ribs on their inner surfaces; basis membranous. Opercular 
Vvalives sulo-uanoular. eee by ky 0. bloel: 


HEXELASMA AUCKLANDICUM Hector sp. (Pl. LXXXYV.) 


1888. Scalpellum aucklandicum Uector, Trans. N.Z. Institute, 
vol. xx. (1887) p. 440. 

1903. Pollicipes ? aucklandicus Hector sp.: W. B. Benham, “ On 
some Remains of a Gigantic Fossil Cirripede from the 
Tertiary Rocks of New Zealand,” Geol. Mag. London, 
dec. 4, vol. x. p. 111, pls. 9, 10 (non figs. 8, 9). 

1905. Pollicipes aucklandicus Benham: E. Clarke, ‘‘'The Fossils 
of the Waitemata and Papakura Series,” Trans. N.Z. 
Institute, vol. xxxvi. (1904) p. 419. 

1910. Pollicipes ? aucklandicus Hector sp.: J. Park, ‘ Geology 
of New Zealand,” p. 115 (pl. 7), pp. 1138, 134. 

Diagnosis. Compartments attaining a length of at least 187 mm., 
carinal, carino-lateral, and lateral compartments with simple ale 
(i.e., there is no distinct upturned extension at the margin). 
Sheath feebly developed, almost absent, and with no suturai edge 
to abut against the longitudinal ridge formed on the inner 
surface. Opercular valves unknown (except for probably a single 
tergum). 

Material. 7 rostral compartments, 9 carinal, 13 lateral (8 right 
and 5 left), 5 carino-lateral (3 right and 2 left), together with a 
small tergum (Pl. LXXXYV. figs. 13 a, 6), which may or may not 
belong to the species; all these are more or less imperfect. The 
specimens are in the collection of the Geol. Surv. New Zealand, 
and are marked with the locality-number 695. They are pre- 
sumably the syntypes of Hector. In addition to these specimens 
I have examined plaster-casts of the specimens figured by 
Prof. Benham. 

Holotype. From among the syntypes of Hector I select as 
holotype the rostrum here figured on Pl. LXXXY. fig. 1. 

Horizon and Locality. Miocene, Oamaruian, Base of Waitemata 
Beds: Motutapu Island, Auckland Harbour, New Zealand. 

General Remarks. Sir James Hector (1887) first called at- 
tention to this fossil, and at a meeting of the Wellington 
Philosophical Institute he remarked on some remains of it there 
exhibited :— 

“Specimens of a large fossil stalked Cirripede, recently 


842, MR. T. H. WITHERS ON 


collected by Mr. Park, at Motatapu Island, Auckland. A careful 
restoration will have to be made before definitely determining 
this fossil, but it will probably be found to belong to the genus. 
Scalpellum and is distinguished provisionally under the name 
S. aucklandicum. In size, this fossil Cirripede greatly exceeds 
any previously known, in 8. magnum the capitulum being only 
13 inches in length, while in the Auckland specimen it is at 
least 8 inches. These fossils oceur in a breccia, marking the old 
shore line of the upper part of the W aitemata Series, similar 
to the Cape Rodney beds. The associated fossils are Corals, 
Brachiopods, and Hchinoderms. Among the latter are two 
specimens having plates of a Cidaris of enormous size.” 

Attention was again called to this fossil in 1903, when 
Prot. Benham described and figured certain valves. He con- 
sidered that they showed closer resemblance to the capitular 
valves of certain species of Pollicipes, and doubtfully referred 
them to that genus. 

After an examination of the present material I am convinced 
that the valves belong to a sessile Cirripede allied to Balanus. 
There are six compartments—a rostrum, carina, right and left 
lateral, and right and left carino-lateral. Prof. Benham figured 
only four valves, namely, “a carina, left scutum, ?rostrum, and 
upper latus.” The carina figured by him is the same as that 
now considered as a carina, and the scutum and ? upper latus. 
correspond to the right and left lateral compartments respec- 
tively; but the valve figured (Benham, 1903, pl. 10, figs. 8, 9); 
as a “?yrostrum” is really a carina of Scalpellum subplanwm, 
sp. n. (see p. 848). The valves considered here as rostral and 
right and left carino-lateral compartments were not figured 
by Benham, and it has now been possible to give figures of the 
inner surface of each different compartment. 

Description of Valves. Valves with solid walls of variable 
thickness, apparently not more than 2°25 mm.; externally 
marked with prominent, more or less regular, transverse growth- 
ridges, rather more strongly marked on the carina; sometimes 
ridged longitudinally, and in one or two cases the valves are 
distorted by linear depressions; but all the valves are more 
or less irregular in shape, and this is obviously caused by the 
surface of attachment; inner surface not longitudinally ribbed 
near the base as in Balanws, for the smooth imner surface slopes 
gradually to meet the outer surface and forms a definite, more 
or less smooth edge. 

Rostral compartment (PI. LXXXV. figs. 1-3) without radi, 
almost symmetrical, moderately convex transversely and slightly 
convex longitudinally, bluntly angular at the apex, and either 
rounded or slightly concave at the basal margin; triangular in 
shape when young, but in the older and consequently longer 
valves the lateral margins for the greater part are almost 
parallel to each other. On the inner surface two more or jess 
prominent ridges extend from the apex, and die out at a point. 
halfway from the apex in the young valve (fig. 2) and at a 


POSSIL CIRRIPEDES FROM NEW ZEALAND. 843 


Text-fig. 139. 


OUTER VIEWS. 


Text-fig. 140. 


INNER VIEWS. 


Hexelasma aucklandicum Hector sp. 
Restored compartments. 
r., rostrum ; 7., lateral; ¢./., carino-lateral; c., carina. 


844 MR. T. H. WITHERS ON 


point about one-third the length of the valve from the broken 
apical portions in the larger valves (figs. 1, 3). Obviously these 
ridges serve for the reception of the angle of the ale on the 
adjacent lateral valves, and the space between the ridges is 
marked, in the older specimens, with transverse lines. The 
portion of the valve enclosed by the ridges is half as wide as the 
adjacent parts of the valve in the specimen figured (fig. 3 6), but 
in the valves figured (figs. 1 6, 26) it is wider. At the base of 
the two ridges extending from the apex, and only in the largest 
valve (fig. 16), a slight transverse ridge is formed by the 
thickening of that part of the valve enclosed by the ridges. 
From a point about one-third from the base of the valve, a small 
extent of the inner surface, parallel to the outer margins, is 
marked with lines which extend upward, and bend abruptly 
inwards towards the base of the ridge on either side ; these lines 
are made by the ale of the lateral compartments which are 
overlapped by this part of the valve. 

Lateral compartment (figs. 4-6) with an ala on the rostral 
side, moderately convex transversely, and longitudinally almost 
flat, irregularly convex, and in one case bent in an elongately 
S-shaped curve; the whole valve is usually strongly bent 
towards the rostrum, but one valve is strongly bent away from 
the rostrum. Parietal portion very much wider than in the 
carino-lateral compartment, as much as three times as wide as 
the widest part of the ala in one valve, and in others from two 
to under one and a half times as wide. The two margins of the 
ala form an obtuse angle, the upper margin is practically straight 
and the lower somewhat concave, but their shape is influenced by 
the degree of curvature of the valve; the growth-lines on the ala 
are closely set and extend obliquely upwards from the base, and 
on reaching almost to the upper margin curve downwards to the 
angle of the ala; on the lower margin, near the angle, a small 
smooth portion is left just below where the growth-lines bend 
downwards. On the inner surface, almost at the middle of the 
parietal portion, a more or less prominent ridge extends from 
the apex and dies out at a point opposite the angle of the ala; 
near the parietal margin the inner surface is marked with lines 
which extend upwards, and on reaching a point just above the 
base of the longitudinal ridge bend sharply inwards and down- 
wards to meet its lower extremity; these lines are obviously 
made by the ala of the carino-laterai compartment, the angle of 
which abuts against the longitudinal ridge. Between the longi- 
tudinal ridge and the upper margin of the ala the inner surface 
is marked with indistinct and irregular transverse lines. 

Carino-lateral compartment (figs. 7-9) obtusely triangular in 
general outline, with an ala on the rostral side. The whole valve 
bent, especially in its apical half, towards the rostrum, is almost 
flat transversely, and the parietal portion is much narrower than 
that of the lateral compartment; the two margins of the ala 


FOSSIL CIRRIPEDES FROM NEW ZEALAND. 845 


form an obtuse angle, the upper margin of which is straight and 
obliquely inclined towards the apex, and the lower margin, which 
is concave, emerges from just above the base of the valve, and 
curves gently upwards and then sweeps sharply outwards to meet 
the upper margin. Parietal portion of valve extremely narrow, 
the widest part being about one-fourth the width of the widest 
part of the ala. On the ala the growth-lines are closely set, and 
follow a similar course to those on the lateral compartment. 
Beneath the upper half of the ala, a portion of the valve on the 
inner surface is somewhat thickened for about one-third the 
width of the valve; its inner margin is steep-sided, and forms a 
ridge, against which abuts the angle of the ale of the carinal 
department; the thickened portion of the valve widens gradually 
from the apex, dies out at a point opposite the angle of the ala, 
and is marked with fine, regular, closely-set, transverse lines. 
Near and parallel to the lower half of the parietal margin, and 
for about one-third the width of the valve, the inner surface is 
marked with lines which extend upwards, and, on reaching about 
half the length of the valve, ave angularly bent downwards and 
inwards, and the lowest of them meet the base of the steep-sided 
ridge near the upper margin of the ala; these lines correspond 
to, and are obviously made by, the ale of the carina, which are 
overlapped by the portion of the valve thus marked. 

Carinal compartment (figs. 10-12) gently curved longitudinally, 
with an ala on each side, and these are bent at a sharp angle 
from the parietal portion. Parietal portion narrow, transversely 
convex, especially near the apex, much narrower than the parietal 
portion of the lateral compartment, but wider than that of the 
carino-lateral compartment ; ale about one and a half times as 
wide as the widest part of the parietal portion, and in one young 
valve about the same width as the parietal portion. The ale 
emerge from near the base of the valve, widen gradually upward 
until about two-thirds the distance from the base, and here they 
bend further outwards and then sharply upwards to the apex ; 
the two margins, therefore, roughly form an obtuse angle, the 
basal margin of which is somewhat concave, and the upper 
margin, which is the shorter, is straight. The growth-lines on 
the alee extend obliquely upwards from the base and, on reaching 
a point more than halfway across the ale, bend sharply and 
angularly downwards to the margin; a smooth triangular portion 
of the valve is left beneath the angularly bent growth-lines. 
The inner surface is quite smooth except for some transverse 
lines, which mark the surface above the angle of the ale and 
which are more.prominent at this point. 

Measurements. The largest valve in the present series (a carina, 
fig. 10) would measure, if complete, circa 90 mm., and the smallest 
valve (a lateral compartment, fig. 6) cirea 12mm. One of the 
compartments (a lateral) figured by Prof. Benham as a scutum 
measures 187 mm. 


MR. T. H. WITHERS ON 


Structure and Affinities. It is evident from the structure of 
the inner surface of the compartments of this Cirripede, and 
from the modification of the side wall of the compartments to 
form ale, that the elements combined to form a shell something 
like the compartments in the genus Salanus. This is proved by 
the fact that the lines on the inner lateral portions of the rostrum 
correspond to, and are obviously made by, the growth-lines of 
the ala of each lateral compartment, which was overlapped by the 
lateral portions of the rostrum; similarly shaped lines on the 
lateral compartment correspond to those on the ala of each 
carino-lateral compartment which was overlapped by the lateral 
compartment; the more angularly bent lines on the carimo- 
lateral compartments correspond to those on the ale of the 
carinal compartment which were overlapped by the carino-lateral 
compartment. In addition, more or less prominent longitudinal 
ridges are developed in the upper portions of the rostral, lateral, 
and carino-lateral compartments against which the angle of the 
ala of the adjacent valves abutted. The transverse lines on 
the portions of the compartments not covered by the ale are 
similar to those in Balanws, in which they are caused by the 
successive exuviation of the opercular membrane. To a similar 
cause may be attributed those in the present Cirripede. 

That this form is a sessile Cirripede is, | think, beyond doubt, 
and the irregularity in shape of the several compartments, as 
well as the fact that some are externally and irregularly ridged 
longitudinally and others distorted with linear depressions, 1s. 
further evidence in support of this conclusion, for these features 
could be caused only by the irregularity in the surface of 
attachment. In accordance with the above interpretation, six 
compartments would complete the wall of the shell, as in 
Balanus; and it is important to note that every fragment in 
the present collection can be allocated to one of the six valves 
figured. (Pl. LXXXV. & Text-figs. 139, 140.) 

The shell of this form could not have been very strong, for, in 
comparison with the great length attained by the compartments, 
the walls aré relatively quite thin. The compartments could 
have been only weakly attached, and are in consequence always 
found quite apart from one another. Moreover, the absence of 
radii, together with the absence of sutural edges to the alee and 
the comparatively feeble ridges or shoulders developed on the 
inner surface, and against which only the angles of the ale could 
have abutted, show « quite clearly the great ‘structural frailty of 
the shell. It could hardly have been a littoral barnacle. 

Although this form agrees with the typical species of the genus 
Balanus in the number of compartments forming the walls of the 
shell, it differs markedly in the structure of these compartments. 
These differences are (1) the absence of radii, (2) the simpler 
structure of the ale, (3) the absence of longitudinal ribs on the 


5 
inner surface, (4) the feebly developed sheath, as well as the 


FOSSIL CIRRIPEDES FROM NEW ZEALAND. 847 


absence of a sutural edge to abut against the longitudinal ridge 
on the inner surface. All these are primitive characters, and 
show that in this Cirripede we have a Balanid more primitive 
than Balanus. 

In my preliminary consideration of this form I regarded it as 
being related, mainly in the absence of radii, to the recent species 
Bataan Honaunms and BL. corolliformis, winich were included by 
Dr. Hoek* in a new section (G) of Balanus, and B. hoekianus 
and B. callistoderma, which were referred by Dr. Pilsbry f to the 
same section, and I intended to found a new genus to include 
these species. On seeing a proof of Dr. Hoek’s work (1913, 
‘ Siboga-Expeditie, Cirripedia-Sessilia,’ pp. 244-246), however, 
I found that he had included these recent species, together with 
two new species (/7. velutinwm and H. arafurae), in a new genus 
Hexelasma. 

Dr. Hoek kindly sent me drawings of the type-species HZ. velu- 
tinwm, and from these it could be seen that while Hewxelasma 
differs from Balanws in the absence of radii and the absence of 
longitudinal ribs on the mmner surface, it agrees in having a well- 
developed sheath, and in the carino-lateral compartments having 
an upturned extension of the ale as well as a well-developed 
sutural edge. 

Since ‘ Pollicipes (¢) aucklandicus” agrees with Hexelasma in 
the absence of radii and of longitudinal ribs on the inner surface, 
it seems advisable to refer it to that genus ; but in some respects 
it appears to be somewhat more pr imitive than the typical species 
of Hexelasma, especially in the feeble development of the sheath, 
in the absence of a distinct upward extension to the ale of the 
earino-lateral compartments, and of a sutural edge to the ale of 
the carino-lateral compartments. 

The species included in Hewelasma are all deep-sea forms, and 
occur at depths varying from about 100 to 900m. In length 
the shell of the largest species, HY. corolliforme Hoek, measures 
nearly 45 mm., and since the largest-known compartment of the 
fossil H. aucklandicum measures about 190 mm., the great 
difference in size is apparent. 

Except for Balanus psittacus Molina sp., which has been known 
to attain a length of 9 inches (circa 225 mm.), Heaxelasma auck- 
landicum 1s the largest-known Cirripede. Salanus evermanni 
Pilsbry, another large barnacle, is recorded as measuring 150 mm. 


HEXELASMA sp. 


A large number of the disconnected compartments of a small 
Balanid are to be seen scattered about in the matrix containing 


* 1883. P. P. C. Hoek, ‘Challenger’ Report, Zoology, vol. viii. pp. 155-160. 
a Ugh, Jets Jeiilsjony, Barnacles of Japan and “Bering Sea,’ Bull. Bureau 
Fisheries, Washington, vol. xxix. 1909, pp. 76-80. 


3848 MR. '. H. WITHERS ON 


the compartments of Heaxelasma aucklandicum and the valves of 
Scalpellum subplanwm, sp. nu. Those that I have been able to 
extract and clean appear to be somewhat worn, and the largest: 
of them does not measure more than 5 ov 6 mm. in length; their 
outer walls are thrown into comparatively wide longitudinal 
folds. Owing to their worn appearance the finer characters are 
not well shown, but since they do not appear to possess radit 
they must be referred to the genus Hewelasma. They differ, 
however, from the typical species of Hewelasma, as well as from 
the fossil H. wucklandicum, in having a well-developed sheath, 
and in the presence of strong ribs on their inner surface. Balanus : 
hoekianus Pilsbry (1911, ‘ Barnacles of Japan and Bering Sea, 
Bull. Bureau TDi Werate, Washington, vol. xxix. 1909, p. 2 
text-fig. 8), which has now been referred by Dr. Hoek to his 
genus Hexelasma, agrees with the present compartments in the 
presence of ribs on the inner surface, but it would be rash to 
say that they are related specifically. In view of the unsatis- 
factory preservation of these compartments, and in the absence 
of the opercular valves, I do not think it advisable to institute a. 
new species. 

Horizon and Locality. Miocene, Oamaruian, Base of Waitemata 
Beds; Motutapu Island, Auckland Harbour, New Zealand. 

Collection. Geol. Surv. New Zealand. 


POLLICIPEDID4. 


Genus SCALPELLUM. 


1817. Scalpellum Leach, Journ. de Physique, &e., Ixxxv. p. 68. 


SCALPELLUM SUBPLANUM, sp.n. (PI. LXXXVI. figs. 1-6.) 


1903. Pollicipes (?) aucklandieus Hector sp.: W. B. Benham, 
Geol. Mag. dec. 4, vol. x. p. 114, pl. 10, figs. 8-9. 


Diagnosis. Carina not separated into tectum, parietes, or 
intraparietes, flatly arched transversely, basal margin bluntly 
angular; tergum with the upper carinal margin unusually short, 
and making with the occludent margin an obtuse angle ; rostrum 
with a wide, flat median keel extending from the apex to the 
basal margin. 

Material. 2 carine, 2 scuta, 4 terga, | rostrum, and 1 sub- 
carina; most of these are incomplete, and they were all extracted 
from the matrix containing the valves of Hexelasma aucklandicum. 

Holotype. The carina figured on Pl, LXXXVI. fig. 2. 

Collection. Geol. Surv. New Zealand. 

Horizon and Locality. Miocene, Oamaruian, Base of Waitemata 
Beds: Motutapu Island, Auckland Harbour, New Zealand. 

Carina not separated into tectum, parietes, and intraparietes, 
bowed moderately either inwards or outwards, widening gradually 


FOSSIL CIRRIPFEDES FROM NEW ZEALAND. 849: 


from the apex to the basal margin, which is more or less bluntly 
angular; flatly arched transversely and indistinctly carinate. 
The apical portion of the smaller specimen is thickened to quite 
a third of its extent, the inner portion forming almost a flat 
surface extending from each side of the carina. 

Length (fig. 1, basal half of valve) 16-5 mm. ; breadth 10 mm. 

Length (fig. 2, basal 2 of valve) 16°6 mm.; breadth 6-4 mm. 

Scutum trapezoidal, nearly twice as long as wide, divided 
almost equally by an indistinct, flat, wide ridge extending from 
the apex to the basi-lateral angle. Oceludent margin convex ; 
basal margin almost straight. “Tere ral margin slightly coneave, 
of about the same length as the convex lateral margin, with 
which it forms an obtuse angle. Along the tergal margin the 
valve is rounded towards the inner surface. The inner occludent 
edge is broad, flat, of about the same width throughout, marked 
with growth-lines, and overhangs the depression for the adductor 
Eoutor um opposite the tergo-laterai angle; the inwardly rounded 
tergal edge is marked with growth- Hines, but the extent thus 
marked narrows rapidly towar -ds the ter go-lateral angle. 

Length (fig. 3, incomplete valve) 28: 7 mm.; breadth 17 mm. 

Tergum subrhomboidal, with an obscure ridge extending in a 
straight line from the apex to the basal angle, and dividing the 
valve unequally, the occludent portion being in its widest part 
almost twice as wide as that of the carinal portion ; lower carinal 
margin weakly convex, somewhat longer than the scutal margin, 
and forming with it an acute angle; upper carinal margin 
slightly convex, unusually short, and forming an obtuse angle 
with the slightly convex occludent margin, which is almost twice 
its length. The occludent margin forms a somewhat raised 
border, and this is followed by a wide, shallow depression 
bounded by an obscure ridge extending from the apex to a point 
on the scutal margin, which is slightly produced about one-third 
the distance from the basal angle; from this ridge the valve 
slopes upwards towards the apico-basal ridge, and slopes rapidly 
down to the carinal margin. On the inner surface a narrow 
portion of the valve along the occludent and upper carinal 
margins is marked with growth-lines, and the extent thus marked 
is wider beneath the apex. 

Length (fig. 4) 20 mm. ; breadth 12:4 mm. 

Rostrum subtrianglar, strongly convex transversely, bowed 
inwards, basal margin convex ; lateral margins slightly concave ; 
a flat submedian keel extends from the apex to the basal mar Sn, 
and this on the right side is followed by a further longitudinal 
ridge near the lateral margin, but on the wider left side this 
ridge, if present, is extremely obseure. 

Length (fig. 5) 10°5 mm. ; breadth (when complete) circa 8 mm. 

Subearina triangular, moderately convex transversely, bowed 
inwards; apex rounded ; basal margin concave ; lateral margins. 
slightly concave. On the inner surface a slight, but well-defined 


850 MR. T. H. WITHERS ON 


ridge extends from the apex to about the middle of the valve, 
and is there met by two further ridges extending from each basal 
angle; the valve is thus divided into three almost equal portions, 
the basal one being smooth and doubtless covered at one time by 
the corium, the two upper portions being marked with growth- 
lines and most probably overlapping the adjoining carino-lateral 
valve on either side. 

Length (fig. 6) 5 mm.; breadth 5°2 mm. 

Comparison with other Species. This species is referred to the 
genus Scalpellum (sensu lato), a course that is advisable until we 
can find the remaining valves of the capitulum, which may enable 
us to refer the species to one of the subgenera into which 
Scalpellum has been divided. Judging from the known valves 
(carina, scutum, tergum, rostrum, and subcarina), this species is 
related to S. zancleanwm Seguenza* from the Pliocene of Messina, 
Sicily. The carina of S. zancleanum differs in having a strong 
median keel, from which the sides of the valve slope steeply, and 
in the less angular growth-lines; the scuta differs in the rounded 
basal. margin and in the usually less acute tergo-lateral angle ; 
the terga differ in being much narrower in proportion to width, 
in the much more acute apical portion, and in the carinal margin 
not being divided into an upper and a lower portion; the rostrum 
differs in the absence of a wide, flat, median keel. 


Subgenus ARCOSCALPELLUM. 


1907 (Oct.). Arcoscalpellum P. P. C. Hoek, Siboga-Expeditie, 
Cirripedia Pedunculata, p. 59. 

1907 (Nov.). Holoscalpellum H. A. Pilsbry, Bull. No. 60, U.S. 
Nat. Mus. p. 25. 

1908. Arcoscalpellum Hoek; H. A. Pilsbry, Proc. Acad. Nat. 
Sci. Philadelphia, p. 109. 

1912. Arcoscalpellum Hoek; T. H. Withers, Proc. Zool. Soe. 

London, p. 538. 


SCALPELLUM (ARCOSCALPELLUM) UNGULATUM, Sp. n. (Pl, LXXXVI. 
figs. 7-13.) 


Diagnosis. Carina with its tectum almost flat, parietes more 
than half the width of the tectum, intraparietes narrow and bent 
abruptly inwards, basal margin rounded; upper latera sub- 
triangular, with rounded basal margin and wide lateral portions 
obliquely inclined towards the umbo; rostral latus with sub- 
parallel scutal and basal margins, and about one-third of its 
apical end, which is much thickened, projecting freely beyond the 
scuta, 


* 1876. G. Seguenza, Atti Accad. Pontaniana, vol. x, p. 386, pl. vii, figs. 1-13. 


FOSSIL CIRRIPEDES FROM NEW ZEALAND. B51 


Material. 6 carine, 4 scuta, 2 terga, 1 upper latus, and 1 rostral 
latus. 

Holotype. The carina, I. 15409 (fig. 8). 

Collection. Brit. Mus. (Nat. Hist.). 

Horizon and Locality. Miocene, Upper Oamaruian. Blue 
Clay: Pareora, South Canterbury, New Zealand. Jimestone : 
Takiroa, Oamaru District, N. Otago, New Zealand. 

In the Geological Department of the British Museum there is 
a series of 14 valves of a species of Sealpellwm (sensu lato), 
registered I. 15409-I. 15422. Although these come from different 
localities, they are of interest, since they appear to represent a 
single new species and enable us to gain some idea of the form 
of the capitulum (see restoration, fig. 13). 

Three of the valves (a carina, scutum, and rostral latus) are 
labelled “ Scalpellam | (Blue Clay) | Parimoa, New Zealand | Mr. 
C. Mantell.” The remaining eleven valves (5 carine, 3 scuta, 
2 terga, and 1 upper latus) are labelled “Scalpellwm | Takiroa, New 
Zealand | Mr. C. Mantell.” With the latter series is a MS. label 
(probably written by the collector) bearing the words “‘ Lime- 
stone | Takiroa—Crinoline Qu. 1852.” 

It is fairly obvious that the “ Mr. C. Mantell” was intended 
for the late Hon. Walter Mantell, from whom a collection of 
New Zealand fossils was obtained- by the British Museum in 
1856. On failing to trace the above-mentioned localities, I wrote 
to Prof. James Park, who kindly informed me that ‘‘ Parimoa ” 
should be Pareora, South Canterbury, and Takiroa is in the 
Oamaru District, N. Otago; he was of the opinion that the 
Cirripede valves came from beds of Miocene (Upper Oamaruian) 
age. ‘Crinoline Qu.” may mean Crinoline Quarry. 

Description of Valves. Valves thick, ornamented exteriorly 
with numerous exceedingly fine ridges radiating from their 
apices. 

Carina narrow, widening very gradually from the apex, 
moderately bowed inwards, basal margin rounded. ‘Tectum 
almost flat transversely, not carinate. Parietes more than half 
the width of the tectum (in one old specimen nearly as wide), 
bent nearly at right angles to the tectum. Intraparietes very 
narrow, bent abruptly inwards and meeting just below the apex, 
so that a small portion of the valve projected freely. Umbo 
apical. 

Length (fig. 7, incomplete valve) 19:2 mm. ; breadth 6°5 mm. 

Length (fig. 8, small complete valve) 16 mm.; breadth 3°7 mm. 

Scutum trapezoidal, almost twice as long as wide; basal and 
lateral margins slightly concave, nearly at right angles to each 
other; tergo-lateral angle reaching a point a little above the 
middle of the valve; tergal margin straight and its outer edge 
flat ; tergo-lateral portion of valve slightly concave; occludent 
margin moderately convex and very obliquely inclined towards 
the terga. Umbo apical. 

Length (fig. 11) 27-4 mm.; breadth 15 mm. 

Proc. Zoou. Soc.—1913, No. LVI. 56 


852 MR. IT. H. WITHERS ON 


Tergum subtriangular, convex transversely, thick, elongated, 
with a prominent, nearly straight ridge, extending from the apex 
to the basal angle. On both sides of this ridge the valve slopes 
rapidly to the outer margins. Carinal margin gently convex ; 
occludent margin gently convex, with a raised border, followed 
by a wide depr ession, extending ‘from the apex to about half the 
width of the scutal margin, whichis “ncleniter! to that extent. 

Length (fig. 9, incomplete valve) 21°4 mm.; breadth 10°8 mm. 

Upper latus subtri angular, thick, convex transversely and 
longitudinally ; umbo a little below the apex, owing to a thick 
ledge formed beneath it, which extends from the lateral angles to 
just beyond the umbo. Scutal margin slightly convex, its upper 
surface forming a prominent ridge, followed by a wide depression, 
which is bounded by an indistinct ridge, and between these two 

ridges the growth-lines are abr uptly upturned; a second in- 

distinct ridge runs almost parallel to the slightly convex tergal 
margin, and between these the growth-lines are also abruptly 
upturned. 

Length (fig. 10) 15-4 mm.; breadth 13°5 mm. 

Rostral latus about four times as wide as long, with a promi- 
nent, rounded, wide ridge extending its whole length, from which 
the surface of the valve slopes rapidly ; outer (rostral) extremity 
bluntly angular, much thickened, and this must have projected 
freely more than one-third the extent of the valve; inner 
(lateral) extremity obliquely truncated. Scutal margin concave, 
sloping upwards to the inner margin of the thickened portion, 
and then descending rapidly in a curve to the umbo; basal 
margin straight and almost parallel to the inner two-thirds of 
the scutal margin, 

Length (fig. 12) 4-4 mm.; breadth 15:1 mm. 

Remarks and Comparison with other Species. In a paper by 
Dr. G. A. Mantell* (1850, p. 329) a Cirripede from the Ototara 
Limestone (Miocene, Upper Oamaruian) is recorded as ‘“ Pollicipes, 
resembling a Cretaceous species.” There is nothing to indicate 
whether this is the present species or not. ‘The two scuta 
figured by Zittel T as “ Cirrhipedenschalen,” and recorded from 
Whaingora and Aotea, North Island, New Zealand, are certainly 
distinct from the present species.  Zittel’s figures were sub- 
sequently reproduced and recorded as Scalpellum sp. by Prof. 
James Park, who considered the specimens to be of Miocene 
(Upper Oamaruian) age. 


* 1850. G. A. Mantell, ‘* Notice of the Remains of the Dinornis and other Birds 
and of Fossils and Rock-Specimens, recently collected by Mr. Walter Mantell in 
the Middle Island of New Zealand; with Additional Notes on the Northern 
Island. WithsNote on Fossiliferous Deposits in the Middle Island of New Zealand, 
by Prof. E. Forbes.” Quart. Journ. Geol. Soc. London, vol. vi. pp. 319-343, 
pls. xxvill., xxix. 

+ 1865. K. A. Zittel, “ Fossile Mollusken und Echinodermen aus Weuscelandig 
Palaontolozgie von Neuseeland (Novara-Exped.), Geol. Teil., Bd. 1. Abth. 2, pl. ix. 
figs. 12 a, b. 

£1910. J. Park, ‘ Geology of New Zealand, p. 141 (pl. xu1.), figs. 9a, 6. 


FOSSI“L CIRRIPEDES FROM NEW ZEALAND. 853 


From a study of the isolated valves here described, it seems 
reasonable to suppose that they formed a capitulum just as is 
shown in the restoration (fig. 13). No examples of the carinal 
and inframedian latera have yet been found, but doubtless these 
will prove to be somewhat like the valves diagrammatically 
indicated in the restoration. The capitulum, as_ restored, 
possesses 13 valves, but this number may have been increased 
to either 14 or 15 by the addition of a rostrum or subcarina, or 
both. In any case, it is apparent from the structure and dis- 
position of the valves, that the species falls into the group now 
included in the subgenus Arcoscalpellum Hoek. 

Scalpellum (A.) ungulatwm may be compared with S. michelot- 
tianum Seguenza*, from the Pliocene of Messina, Sicily, and 
S. quadratum Dixon t sp., from the Eocene of Bognor, Sussex, 
but both these species differ in the valves being appreciably 
thinner. 

Although size is often of little account, none of the numerous 
valves of S. quadratum that I have seen are more than half the 
size of the largest valves of S.(A.) wnguwlatum. The carina of 
S. quadratum difters in having inwardly bent intraparietes, the 
scutum in having a raised border to the tergal margin, the tergum 
in the carinal margin being elongately §-shaped and the apical 
half bent towards the carina, the upper latus in being more 
symmetrical and proportionally wider, and the rostral latus 
differs in the absence of the prominent, transverse, rounded 
ridge or fold and in the outer extremity of the valve being 
thickened only to a small extent. 

In S. michelottianum the carina differs in having broad, 
rounded, longitudinal ridges on the outer margins of the tectum, 
in the tectum being proportionally much wider and the basal 
margin less convex, the scutum in having the tergo-lateral 
portion of the valve markedly convex and rounded inwards at 
the tergal margin, the tergum in being much thinner and very 
much flatter transversely and the median longitudinal ridge 
hardly perceptible, the upper latus in being more triangular in 
shape and in the absence of a thick ledge formed beneath and 
beyond the umbo, the rostral latus in having a much smaller 
extent of the valve thickened at the outer extremity and in the 
inner extremity not being obliquely truncated. 

S. nuchelottianwm var. gassinensis Alessandyvi ~ (Pliocene to 
Miocene) agrees more closely: with S. (A.) wngulatum in having 
the basal margin of the carina more strongly convex, but the 
other characters of this and the remaining valves differ quite as 
much as the valves of S. michelottianum Seguenza. 


* 1876. G. Seguenza, Atti Accad. Pontaniana, vol. x, p. 381, pl. vi. figs. 15-25 ; 
p. 464; pl. x. fig. 26. 

+ 1846. F. Dixon, in J. de C. Sowerby, Min. Conch. vol. vii. pl. 648; 1851. 
C. Darwin, Pal. Soc. Monogr. Foss. Lepadide, p. 22, pl. i. fig. 3. 

£ 1906. G. de Alessandri, Palwontogr. Ital. vol. xii. p. 252, pl. xiii. figs. 10-15, 


854 ON FOSSIL CIRRIPEDES FROM NEW ZEALAND. 


EXPLANATION OF THE PLATES. 
Prare LXXXV. 


Hewxelasma aucklandicum Hector sp. 


Miocene, Oamaruian, Base of Waitemata Beds: Motutapu Island, 
Auckland Harbour, New Zealand. 


Fig. 1. Rostrum. a, outer view ; 6, inner view of apical portion of same. 

2. Id. a, outer view of small example; 6, inner view of same. 
3. Id. a, outer view; 0, inner view of same. : 
4. Lateral compartment (right). a@, outer view; 6, inner view of same. 
5. Id. (left). Outer view. 
3. Id. (right). Outer view of small example. 
7. Carino-lateral compartment (right). Outer view. 
8. Id. (right). Inner view. 
9. Id. (left). Outer view of incomplete example. 

10. Carina. a, outer view; 4, side view of same. 

11. Id. Outer view of small complete example. 

12. Id. Inner view of incomplete example. 

13. Tergum. Probably of the same species. a, outer view; 6, inner view of 

same. 


All figures reduced to } nat. size, except figs. 2, 6, and 9, which are nat. size, and 
figs. 13 a, 6, which are X 4 diam. 


Prate LXXXVI. 


Sealpellum subplanum, sp. n. 


Miocene, Oamaruian, Base of Waitemata Beds: Motutapu Island, 
Auckland Harbour, New Zealand. 


Fig. 1. Carina. a, outer view of basal half of valve; 6, side view; c, transverse 
section. 
2. Id. a, outer view of basal two-thirds of a smaller valve; 6, side view ; 


c, transverse section. 
3. Scutum. a, outer view of incomplete valve; 4, inner view of sane. 


4. Tereum. a, outer view; 6, inner view of same. 
5. Rostrum. Outer view. 
6. Subearina. a, outer view; 6, inner view of same. 


Scalpellum (Arcoscalpellum) ungulatum, sp. n. 


Miocene, Upper Oamaruian, “ Limestone”: Takiroa, Oamaru District, 
; N. Otago, New Zealand. 
Fig. 7. Carma. a, outer view of basal half of valve; 0, inner view; c, transverse 
section. 
8. Id. a, outer view of small complete valve; 6, side view. 
9. Tergum. a, outer view; 6, inner view of same. 
10. Upper latus. a, outer view ; 0, inner view of same. 


Miocene, Upper Oamaruian, “ Blue Clay”: Pareora, South Canterbury, 
New Zealand. 


11. Scutum. a, outer view; 5, inner view of same. 
12. Rostral latus. a, outer view; 6, inner view of same. 


13. Restoration of capitulum, based on the disconnect®d valves here figured. 
The inframedian and carinal latera are not known, but these valves 
: have been diagrammatically indicated in the restoration. 


All figures of nat. size, except figs. 5, 6, 8, and 12, which are twice nat. size. 


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OSTEOMALACIA IN A BABOON. 


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OSTEOMALACIA IN A BABOON, 


PROF. T. WINGATE TODD ON OSTEOMALACTA. 855 


Observations on Osteomalacia in the Zoological Collec- 
tions of Manchester and Cleveland. By T. WinGate 
Topp, M.B., F.R.C.S., Professor of ent orie Western 
Reserve University, Cleveland, Ohio * 


[Received May 5, 1913: Read June 3, 1913. ] 
(Plates LXXXVII.-LXXXIX.7) 


It is the object of this communication to call attention to the 
widespread appearance of osteomalacia among animals in captivity, 
and to point out the opportunity afforded by the disease for the 
study of changes in formed bone-tissue. 

Through the kindness of Messrs. Jennison of Manchester and 
Director Springborn of Cleveland, I have been enabled to study 
the disease as it exists in the Zoological Gardens of both these 
cities. 

In Manchester, while the cercopitheques, macaques and small 
baboons were kept in a large cage in a moderately heated 
monkey-house, the animals sickened and died of miliary tubercle 
within a few months after their arrival. When, however, the 
glass was removed from the windows of the monkey-house and a 
free current of air permitted to pass through the building, the 
animals remained healthy for as longas three years. Eventually, 
however, most of them begin to show a disinclination to move 
about, crouching and walking with the aid of the fore-limbs—the 
hind ones being curled up beneath the body in squatting attitude. 
When attacked by healthy animals they seem unable to defend 
themselves, or, if they try to run away, they do so groaning with 
apparent pain. ‘This is more marked if they are compelled to use 
their hind legs, as, for instance, in climbing, Left to themselves 
they are very much addicted to masturbation. Indeed, in Cleve- 
land, my attention was first drawn to the condition by my being 
asked to inspect some macaques of both sexes which had developed 
this objectionable habit. As the disease progresses the joints 
become stiff and enlarged, although post-mortem joint-changes 
have not been present in the cases which I have examined. The 
disease is not of itself fatal and may last two years, although it 
is usually found necessary to destroy the animals after twelve 
months. If the animals die, there are frequently indications of 
broncho-pneumonia found at the autopsy. The food on which all 
the apes, healthy and diseased, are fed consists of rice-pudding, 
carrots, onions, potatoes, greens, apples, and bananas, with a little 
meat occasionally. 

While suffering from the disease, the hair may become dry, 
ruffled, or fall out. This is more frequently the case in rabbits 


* Communicated by the SecRETARY. 
+ For explanation of the Plates see pp. 859-860. 


856 PROF. T. WINGATE TODD ON OSTEOMALACTA. 


and rats than in monkeys. The disease seems to localize itself 
in different sites in different animals. In the macaques and 
baboons it usually affects the hind limbs, which are in con- 
sequence badly bent and twisted, but that the whole skeleton is 
affected is shown by the brittle character of all the bones. The 
ability of the bones to unite after fracture does not seem to be 
impaired. Fig, 4 (Pl. LX XXVIII.) represents the tibia of a 
young Anubis Baboon (Cynocephalus anubis) after healing of 
a fracture near the upper extremity. This bone was broken 
by a fall from a height of some three feet while the animal was 
attempting to escape from a pursuer. The animals may become 
affected by the disease while still young and before the epiphyses 
have joined, so that the disease is readily confounded with rickets. 
The lesions, however, are the same at whatever age the disease 
occurs. In some instances the bone-changes are found most 
marked in the fore limbs. This was the case in a peceary from the 
Manchester Collection. The animal struggled about with its fore 
limbs bent backwards beneath the body. In other animals the 
bones of the face are affected. The maxille are much enlarged 
and give a puffed-out appearance to the cheeks. This condition 
occurred in a horse, a leopard, a rabbit, and a chimpanzee. In 
some cases the necks of the teeth are decayed and the teeth may 
fall out. This was observed in the leopard and rabbit just 
referred to, and also in a Californian sea-lion from Cleveland. 
Uleeration and falling of the teeth has not occurred among 
the apes in Manchester since the glass was taken out of the 
windows of the monkey-house. The bones of the chest are 
frequently softened and deformed, which would seem to pre- 
dispose the animal to respiratory disorders. The lungs of the 
sea-lion showed emphysema and patches of broncho-pneumonia, 
the latter being confirmed by histological examination. 

Post mortem, the viscera show no lesion whatever in animals 
which have been intentionally killed. Nor are there any atrophic 
changes in the muscles of the disabled limbs. 

The ductless glands are apparently normal. Pathological 
appearances are to be found only in the nervous system and 
the bones. As regards the former opinion is very varied. Gayet 
and Bonnet describe an increase in volume of the nerves with 
an overgrowth of the fibrous tissue between the nerve-bundles 
and disappearance of myelin in certain cases. The blood-vessels 
of the nerves exhibit endarteritis (1). They found no cellular 
lesion nor any alteration in nerve-fibres in the spinal cord. 
Morpurgo, on the other hand, describes diffuse chromatolysis 
in the cells of the anterior horns of the cord, but denies any 
change in the cells of the spinal ganglia (2). These changes 
occur very early in the course of the disease, but there would 
appear to be no definite evidence to show that they initiated the 
disease (8). 

It may be that these different histological pictures were 


PROF. T. WINGATE TODD ON OSTEOMALACIA. __ 857 


produced by differences in the etiology of the disease. For the 
bone-changes can only be a symptom, caused, perhaps, in many 
different ways. 

The bone-marrow in osteomalacia is much modified. The 
marrow-cavities of the long bones are enlarged and filled with a 
gelatinous fatty tissue and by bright red marrow. Marrow is 
divided microscopically by Ziegler into splenoid and fine-fibred 
constituent factors (4). Of these the latter form is much in- 
creased, partially filling up the marrow-cavity and penetrating 
the bony tissue. 

In early cases the compact bone is invaded by this vascular 
fibrous tissue, which has already replaced to a large extent the 
spongy bone and which is not preceded by osteoclasts. The same 
observation has been recorded by Morpurgo (8). 

In later stages the whole of the compact tissue of the shaft has 
disappeared and is replaced by fibrous tissue, in which are to be 
found discreet areas or islets of osseous material (see Pl. LX X XIX. 
figs. 5, 6). There is meanwhile no subperiosteal deposit of new 
bone. The histological picture of the bone closely resembles that 
seen in ostitis fibrosa. The remaining islets of bone lose their 
ossein and become transformed into a tissue which, from its 
staining properties, appears to be hyaline in character. But, at 
the same time, in other situations the regressing bony substance 
exhibits a fibrillar change of the ground-substance similar to that 
described by Retterer as the basis of normal bone (5). 

Cells with similar staining reactions to osteoblasts may still be 
seen bordering the islets of bony tissue (see Pl. LX X XIX. fig. 6). 
In many places so-called ‘“ osteoclasts” are observed. But, on 
examining several slides one is struck by the comparative infre- 
quency of these cell-masses. As Gayet and Bonnet remark (1) 
it is difficult to believe that osteoclasts can play more than a very 
subsidiary part in the destruction of bone, because of their scarcity 
when compared with the extent of the process. It would appear 
that osteoclasts are not at all necessary for the production of 
change in bone-substance. 

As the bony tissue becomes transformed or replaced by the 
fibrous material, the Haversian systems disappear. The peri- 
osteum becomes intimately united to the mass of fibrous tissue 
which remains in place of the true bone. I have not observed 
hemorrhages in subperiosteal or other localities. The joints were 
unaffected. At the diaphyso-epipyseal junction, changes similar 
to those in rickets were observed. 

The theories of causation of osteomalacia have been well 
reviewed by Morpurgo (2), who, among others, has succeeded in 
producing the disease by inoculation of a micro-organism (6). 

The detailed histological changes have been discussed by 
Basset (7), whose description is amply borne out by my own 
shdes. 

The disease is one which may be secondary to some other 


858 PROF. T, WINGATE TODD ON OSTEOMALACIA. 


lesion, and therefore localised, as in the variety appearing after 
trauma. 

It may develop in connection with giant-celled sarcoma and be 
more generalised in type (Schonenberger, 8). 

It may occur in the so-called spontaneous form, such as is seen 
in animals in captivity. 

The last-mentioned variety is certainly not a simple decalcifi- 
cation. There is absorption of osseous substance with rarefaction 
of the tissue. The process starts from the marrow-cavity and 
involves first spongy and later compact bone. The Haversian 
systems disappear and the bone becomes fibrillar in character and 
later is transformed into fibrous tissue. 

An intermediate hyaline change is shown in places. All these 
changes point to a revolution in the constitution of bone as a 
whole, which is accompanied by changes in the marrow and 
periosteum, as already described. Similar changes in bony 
tissue are to be observed in ostitis fibrosa and leprosy. For it 
has been my good fortune to be able to investigate all three 
diseases at the same time. The clinical symptoms in these cases 
of generalised bone-softening, which I have described as osteo- 
malacia, suggest a nervous origin. The obvious inference to be 
drawn from the histological picture in leprosy is that in the last- 
named disease the bone-changes are certainly trophic in character. 
I would emphasise the fact that we are as yet ill-acquainted with 
the symptoms consequent on lesions to the sympathetic nervous 
system. But there would seem to be ample confirmation of nerve~ 
lesion in the histological changes found in the nerve-bundles by 
Gayet and Bonnet (1). Moreover, the intimal proliferation de- 
scribed by these authors in the vessels of the nerve-trunks may 
be produced by a lesion in the sympathetic nerves, as I have 
recently been enabled to show (9). 

If the disease is infectious, the incubation-period must be con- 
siderable, for it seems to appear spontaneously in animals which 
have been isolated for a long while. After inoculation, Morpurgo 
found the animal became ill in a week or two. Such evidence as 
we have points to the nervous system as the seat of primary 
disease, whether it be infectious or not, and suggests that the 
bone-changes are consequent on nervous lesion. Treatment is 
unsatisfactory. Dr. Fox, of Philadelphia, has recently adminis- 
tered calcium lacto-phosphate and adrenalin, separately and in 
combination, to animals suffering from the disorder in the 
Philadelphia collection, but without success (10). 

In making observations on the living animals Messrs. Antliff 
and Readinger, keepers of the monkey-houses at Manchester and 
Cleveland respectively, have rendered generous assistance. The 
histological sections are the work of Mr. Gooding, of the Ana- 
tomical Department of Manchester. Mr. J. C. Miller, of the 
Laboratory here in Cleveland, has assisted me in gathering and 
abstracting the literature. To all these gentlemen, I would 
therefore express my obligation. 


PROF, T. WINGATE TODD ON OSTEOMALACIA.. - 859 


Summary. 


1. Osteomalacia, or so-called spontaneous generalised bone- 
softening, is not a simple decalcification of bone, but a re- 
organisation of bone as an organ, in which the loss of osseous 
tissue is not brought about by osteoclasts. 

2. From this it is evident that the cell-masses known by the 
name of “ osteoclasts” are not indispensable in the transformation 
of bone to less specialised tissue. 

3. The disease may be infectious in origin. If so, the evidence 
at our disposal points to the nervous system as the site of infec- 
tion. The bone-changes appear to be the symptoms consequent 
on the nervous lesion. 


REFERENCES. 


(1) Gaver et Bonner.—“‘ Contribution 4 l'étude des ostéo- 
malacies.” Revue de Chir. vol. xxiii. pp. 44, 228, 1901. 

(2) Morpurco.—‘ Ueber eine infectidse Form der Knochen- 
briichigkeit bei weissen Ratten.” Verhandl. d. Deutschen 
Path. Ges. vol. iii. p. 40, 1901. 

(3) Morpurco.—‘ Durch Infection hervorgerufene malacische 
und rachitische Skeletveranderungen an jungen weissen 
Ratten.” Cent. f. Allg. Path. u. Path. Anat. Bd. xiii. 
p. 113, 1902. 

(4) Zrecter.— Ueber Osteotabes infantum u. Rachitis.” Cent. 
f. Allg. Path. u. Path. Anat. Bd. xii. p. 865, 1902. 

(5) Rerrerer.—‘ Structure et histogénése de Vos.” Journ. de 
VAnat. et de la Physiol. p. 561 eé seg., 1905. 

(6) Morpurco.—‘ Ueber die infectidse Osteomalacie und 
Rachitis der weissen Ratten.” Verhandl. d. Deutschen 
Path. Ges. vol. xi. p. 282, 1907. 

(7) Basser,—“ Anatomie pathologique de ’Ostéomalacie spon- 
tanée et expérimentale.” Arch. de Med. Expérimentale, 
Meant, joy (sy IQOG. 

(8) SCHONENBERGER.—‘ Ueber Osteomalacie mit multiplen 
Riesensarkomen und multiplen Fracturen.” Virchow’s 
Archiv, Bd. clxv. p. 189, 1901. 

(9) Topp.—“ The Arterial Lesion in Cases of Cervical Rib.” 
Journ. Anat. & Physiol. vol. xlvii., 1913. 

(10) Fox.—The 40th Annual Report of the Board of Directors 
of the Zoological Society of Philadelphia, p. 55, 1912. 


EXPLANATION OF THE PLATES. 
Prats LXXXVII. 


Fig. 1. Right Radius from young Anubis Baboon (Cynocephalus anubis). Advanced 
stage of osteomalacia. 

Note large size and dark colour of marrow cavity. ‘The dark appearance is 
due to the large quantity of red marrow. The rarefaction of bone and the 
progression of the osteoporosis from marrow-cavity outwards is well shown 
in this photograph. The compact tissue can be seen to have been replaced 
by spongy bone. 


Proc. Zoou. Soc.—1913, No. LVII. 57 


Fig. 3. 


Fig. 5. 


PROF. T. WINGATE TODD ON OSTEOMALACTA. 


. Right Ulna from same animal. Here and there are districts of gelatinous 


fatty marrow giving rise to lighter areas between the dark-coloured sites 
of red marrow. The thickened periosteum and its intimate union with 
the tissue composing the bone can be seen well, especially near the centre 
of the shaft. In this situation the bone was accidentally broken while 
stripping off the muscles. 


Prats LXXXVIII. 


Right Humerus of same animal. 

Note the irregular diaphyso-epiphyseal junction at the upper end of the 
shaft. The extreme brittleness of the bones resulted in the fracture of the 
surgical neck of this humerus while the muscles were being removed. 
There is no sharp line between the articular cartilage of the condyles and 
the underlying spongy bone. This photograph shows clearly the articular 
cartilage being destroyed on its bony aspect. 


. Right Tibia and Fibula from same case. 


Note extreme recurved upper portion of tibia. The bone had been fractured 
some time previously and had united in this position. The union is com- 
plete. The fibula was likewise bent, its lower extremity is cut obliquely 
and is shown behind the lower portion of the tibia. 


Prate LXXXIX. 


Longitudinal section of shaft of Left Ulna from the same animal. X40. 

Note the entire disappearance of Haversian systems from the bone and the 
replacement of osseous tissue by fibrous material. The bony substance is 
here shown broken up into islets. The section represents what once was 
the compact bone layer of the shaft. 


. Longitudinal section of shaft of Left Ulna from the same animal. 206. 


A bone-islet is shown surrounded by non-osseous tissue. Note the gradual 
transition of the bone into fibrous tissue. There is no sharp line of 
demarcation, and no osteoclasts are to be observed. The decalcification 
progresses from the centre of the illustration to the periphery. Inter- 
mediate between bone and fibrous tissue is a zone where the nuclei are 
surrounded by considerable non-calcitied protoplasm. This corresponds 
in appearance to the pre-osseous stage in Retterer’s description of 


bone (5). 


No. 122. 


ABSTRACT OF THE PROCEEDINGS 


OF THE 


ZOOLOGICAL SOCIETY OF LONDON.* 
May 20th, 1913. 


Prof. EH. A, Mincuin, M.A., F.R.S., Vice-President, 
in the Chair, 


The Minutes of the last Scientific Meeting were confirmed. 


The Srcrerary read a Report on the Additions that had been 
made to the Society's Menagerie during the month of April 
1913. 


The Lrprarian submitted a list of dates of publication of the 
early parts of the Society’s ‘Transactions,’ drawn up from the 
records kept by Messrs. Taylor & Francis, printers to the Society. 


Mr. R. E. Hotpine exhibited a large number of specimens and 
photographs illustrating variations in the growth of the Antlers 
of Deer. . 


The Rev. T. R. R. Srespine, M.A., F.R.S., F.Z.S., drew 
attention to Prof. F, E. Schulze’s important work on zoological 
nomenclature, the ‘Nomenclator Animalium,’ giving a brief 
description of its objects and scope, and urging its claims for 
assistance from British naturalists. 


Dr. R. Broom, C.M.Z.S., read a paper “On the South African 
Pseudosuchian Reptile Huparkeria and allied Genera.” Besides 


* This Abstraet is published by the Society at its offices, Zoological Gardens, 
Regent’s Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge, 
to all Fellows who subscribe to the Publications ; but it may be obtained on the 
day of publication at the price of Sixpence, or, if desired, sent post-free for 
the sum of Sic Shillings per annum, payable im adyance. 


30 


giving an account of the very completely known South African 
form, he also discussed the structure of the Elgin allied forms, 
Ornithosuchus and others. The group of Pseudosuchians he 
regarded as an extremely important primitive reptilian order, 
as there is good reason to believe that not only does it eontain the 
ancestor of the Dinosaurs, but also the aneestors of the Ptero- 
dactyles and Birds. Huparkeria and Ornithosuchus are, in 
structure, almost Dinosaurs, and it is held that when the bipedal 
habit was more fully acquired the few characters not quite 
Dinosaurian would become Dinosaurian. Birds are held to 
have originated from a Pseudosuchian which, by a bipedal habit, 
had acquired a Dinosaur-like hind limb, and had then become 
arboreal in habit and acquired the peculiar power of flight. : 


Mr. E. G. Bouneneer, F.Z.S., Curator cf Reptiles, read a paper 
giving an account of the experiments which he had, for the past 
year, been conducting on the Metamorphosis of the Mexican 
Axolotl (Amblystoma tigrinum), and gave a detailed description 
of the changes that take place in the course of transformation. 
He also exhibited a number of specimens in the perfect or 
Amblystome condition. The conclusions arrived at by the 
author, as a result of his experiments, were that, in accordance 
with Mlle. de Chauvin’s experiments, and contrary to those of 
Dr. Powers, the Axolotl will, with a few exceptions, transform 
if placed under special conditions which force it to breathe air 
more frequently than usual; that starvation, irregular feeding, 
and temperature have no influence on the metamorphosis; that 
the elimination of oxygen from the water has likewise no bearing 
on the point, as the animal will not, under the circumstances, 
rise to the surface and make use of its lungs at more frequent 
intervals than animals placed under normal conditions. 


Mr. G. E. BuLien contributed a short paper, communicated by 
Mr. John Hopkinson, F.Z.8., “On some Cases of Blindness in 
Marine Fishes.” Work hitherto performed, e.g. that of Hofer, 
de Drouin de Bouville, and others, upon the pathology of fishes 
has been directed largely upon species of freshwater habitat. 
The present author has found, in certain specific cases of blindness 
in marine fishes, pathological conditions similar to those described, 
and others with slight modifications, in several freshwater species. 
The examples dealt with in detail are traumatic corneal opacitis 
in a Conger-Hel, corneal opacitis, etc., in a Greater Weaver, 
and corneal opacitis and cataract in a Pollack. 


Dr. R. W. Suurevpt, C.M.Z.8., sent a paper dealing with the 
Patella in the Phalacrocoracidee. From a study of the patella in 
a number of species of this family, he had found that in adult 
individuals that bone was composed of the true patella solidly 
fused with the proximal portion of the cnemial process of the 


31 


tibio-tarsus, which became dissociated from the latter early in 
the life of the bird. Late in life this fusion obliterated the 
tendon of the ambiens muscle, which heretofore had been 
described as passing through the patella and persisting through 
life, 


The next Meeting of the Society for Scientific Business (closing 
the Session 1912-1913) will be held on Tuesday, June 3rd, 1913, 
at half-past Hight o’clock p.m., when the following communications 
will be made :-— 


EXHIBITIONS AND NOTICES. 


Sir ArrHor H, Cuurcu, K.C.V.O., M.A., D.Sc., F.R.S., F.S.A. 


Notes on Turaecin and Turacin-bearers. 


-P. Coatmers Mitcuent, M.A., D.Sc., LL.D., F.R.S., F.Z.8. 
Observations on the Anatomy of the Shoe-bill (Baleniceps 
Tex). 
T. H. WitHers, F.G.S, 
Some Miocene Cirripedes of the Genera Hexelasma and 
Scalpellum from New Zealand. 
Prof. ArtHuR Denpy, D.Sc., F.R.S., F.Z.8., and R. W. H. Row, 
B. Se. 
The Classification and Phylogeny of the Calcareous Sponges, 


with a Reference List of all the described Species, systematically 
arranged. 


Surgeon JosepH C. THomeson, U.S.N. 


Contributions to the Anatomy of the Ophidia. 
Prof. T. WineatE Topp, M.B., F.R.C.S. 


Observations on Gsteomalacia in the Zoological Collections 
of Manchester and Cleveland. 


32 
The following paper has been received :— 
§ pap 


D. M.S. Watson, M.Se. 


Batrachiderpeton lineatum Hancock & Atthey, a Coal Measure 
Stegocephalian. : 


Communications intended for the Scientific Meetings should 
be addressed to 
P. CHALMERS MITCHELL, 
Secretary. 
ZOOLOGICAL Society or Lonpon, 
Re@ent’s Park, Lonpon, N.W. 
May 27th, 1913, 


No. 123. 


ABSTRACT OF THE PROCEEDINGS 


OF THE 


ZOOLOGICAL SOCIETY OF LONDON.* 
June 3rd, 1918. 


Prof. EK. W. MacBripz, M.A., D.S8c., F.R.S., Vice-President, 
in the Chair. 


The Minutes of the last Scientific Meeting were confirmed. 


Mr. D. Sersa-Smitu, F.Z.S., Curator of Birds, exhibited the 
ege and young of the Mikado Pheasant (Calophasis mikado), a 
rare species, described first in 1906, from the mountains of 
Formosa. Some living specimens had been imported in 1912 by 
Mr. Walter Goodfellow, and the owners of these birds had 
entrusted the eggs to the Zoological Society, where they were 
being hatched. 

The egg was cream coloured and very large compared with 
those of allied species of pheasants, being 57 x 41 mm. 

The incubation period proved to be twenty-eight days, instead 
of twenty-four as in the majority of pheasants, and the young 
when newly hatched were very large, and had the quill-feathers 
better developed than was the case in allied species, 


Sir Artuur H. Caurcu, K.C.V.O., F.R.S., read a paper, com- 
municated by the Secretary, entitled ‘‘ Notes on Turacin and the 
'Turacin-bearers.” 

This paper contains a summary of the chief facts as to the 
composition, properties, and occurrence of turacin, the soluble 
crimson pigment of the Musophagide. Special stress is laid upon 
its constancy of composition, the limitation of its occurrence to 


* This Abstract is published by the Society at its offices, Zoological Gardens, 
Regent’s Park, N.W., on the Tuesday following the date of Meeting to which 
‘it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge, 
to all Fellows who subscribe to the Publications ; but it may be obtained on the 
day of publication at the price of Sixpence, or, if desired, sent post-free for 
the sum of Six Shillings per annum, payable in adyance, 


34 


certain plantain-eaters, and the relation of its spectrum to the 
spectra of hemoglobin and chlorophyll. Some current errors 
concerning turacin are corrected. 


The Secretary, Dr. P. Coatmers Mrircuet, F.R.S., gave an 
account of his observations on the Anatomy of the Shoe-bill 
(Baleniceps rex), illustrating his remarks with lantern-slides. 
He showed that Baleniceps and Scopus shared so many anatom- 
ical characters, and of these so many occurred also in Storks, that 
if the reasoning generally followed by anatomical ornithologists 
were adopted, Baleniceps and Scopus must be placed with Storks 
rather than with Herons. He submitted, however, that such a 
method was irrational, unless it were accompanied by a much 
closer scrutiny of the value of the characters than had hitherto 
been made or was yet possible, and that for the present Baleniceps 
must be regarded as the representative of a division equivalent to 
Storks and Herons. He thought also that the relation of the 
Steganopods to these three groups required reconsideration. 


A paper on ‘‘Some Miocene Cirripedes of the genera Hexelasma 
and Scalpellum from New Zealand,” communicated by Dr. W. 
T. Carman, F.Z.8., was read by Mr. T. H. Wrrumrs, F.G.S. 
An account is therein given of the ‘gigantic Cirripede’ of New 
Zealand, originally deseribed as Scalpellum aucklandicum, of 
which remains have long been known to occur in the Waitemata 
Beds (Miocene) of Motutapu Island, Auckland Harbour. The 
valves of this Cirripede attain a length of 8 inches, and have 
been previously supposed to belong to a pedunculate form, but 
while Sir James Hector (1887) referred them to the genus 
Scalpellum, Prof. W. Blaxland Benham (1903) thought that they 
approached more closely to the genus Pollicipes. From a study 
of the criginal material collected by Prof. James Park (1887), 
it is now shown that this Cirripede is a sessile form allied to 
Balanus, and it is referred to Dr. P. P. C. Hoek’s recently 
instituted genus MHexelasma (1913). A smaller undetermined 
species of Hexelasma, and a new species of Scalpellum (sensu lato), 
are also described. These are in the collection of the Geological 
Survey, New Zealand, and occur in the same beds as the 
‘ vigantic Cirripede.’ 

A second new species of Scalpellum is founded on some valves 
from New Zealand, and a restoration is given, the remains 
being sufficient to justify their reference to the sub-genus 
Arcoscalpellum Hoek. 


A paper on “ The Classification and Phylogeny of the Calcareous 
Sponges, with a Reference List of all the known Species, sys- 
tematically arranged,” was received from Prof. Artaur Denby, 
D.Sc., F.R.S., F.Z.8., and Mr. R. W. Haroip Row, B.Sc., F.L.S. 
This memoir aims at a complete revision of the genera of 


35 


Caleareous Sponges. Fifty recent genera are recognised and 
diaynosed, and all the described species, amounting to 433, are 
arranged under these genera. The rejected generic names, which 
are listed separately, amount to 97. The fifty accepted genera 
are grouped in 10 families, and Poléjaeff’s subdivision into 
Homocaita and Hursrocata is abandoned. The systematic part 
of the werk is prefaced by a discussion on the principles of 
classification, and followed by a discussion on the phylogeny of 
the group, accompanied by a genealogical tree. The suggestions 
of Minchin and Bidder as to the systematic value of the position 
of the nucleus of the collared cell is followed up, and this position 
has been determined in 75 species. It appears from the evidence 
thus secured, taken, of course, in connection with other characters, 
that in the primitive family Homoceelide the position of the 
nucleus is basal in some species and apical in others, but that it 
is not correlated with other characters so as to justify a sub- 
division of the family accordingly. From the Homoceelide, 
however, two lines of descent have sprung, the one basinucleate 
and the other apicinucleate; the former including the families 
Leucascide, Leucaltide, Minchinellide and Murrayonide, and 
the latter the Sycettide, Heteropiide, Grantiide, Amphoriscide 
and Lelapiidee. 


Surgeon J. C. Tuompson, U.S.N., sent a paper, communicated 
by Dr. F. E. Bepparp, M.A., F.R.S., F.Z.S., containing eon- 
tributions to the Anatomy of the Ophidia, 


The SzcreraRyY communicated a paper by Prof. T. WincATE 
Topp, M.B., F.R.C.S., entitled ‘“ Observations on Osteomalacia in 
the Zoological Collections of Manchester and Cleveland.” 


——* 


This Meeting closes the Session 1912-1913. The next Meeting 
of the Society for Scientific Business will be held on Tuesday, 
October 28th, 1913, at half-past Hight o’clock P.M. 


The following papers have been received :— 


1. D. M.S. Warson, M.Sc. 


Batrachiderpeton lineatum Hancock & Atthey, a Coal-Measure 
Stegocephalian, 


36 
2. R. W. Parmer, B.Sc. 


The Brain and Brain-case of a fossil Ungulate of the Genus 
Anoplotherium. 


3. KREDERIC WILMET. 


Notice sur l!’Okapia johnstoni, dont Vespece s'est retrouvée 
intacte dans une forét commune a |’Ituri et 4 1 Uelle. 


_ Communications intended for the Scientific Meetings should 
be addressed to ; 
P, CHALMERS MITCHELL, 


Secretary. 


ZOOLOGICAL Society oF LonpDoN, 
Recurnt’s Park, Lonpon, N.W. 
June 10th, 1913. 


30, 


3l, 


32, 


33. 


34. 


35. 


36. 


37. 


38, 
39. 


40. 


“ill 


42, 


43. 


44, 


45: 


46, 


Papers (continued). 
Page. 
On the Patella in the Phalacrocoracide. By Dr. R. W. Suursupt, C.M.Z.S. 
(Pl, LXI.) Bese eceeecseeeeer set HOF se se sett SPFFso®F es ee ee re eee FH s7 Hse 289899 OD 393 


Experiments on the Metamorphosis of the Mexican Axolotl (Amblystoma tigrinum), 
conducted in the Society's Gardens. By EH. G. EOUUENS ¥.Z.S., Ourator of 
Reptiles. (Text-figs. 75 & 76.) ...... eel ateraie stares sfoie wists eres Bes wegicarets msttaliece's 9 400 
Oontributions to the Anatomy of the Ophidia. By Josrru C. Tuompson, Surgeon, 
United States Navy. (Text-figs. 77 & 78.) ........++0+.00. mis cletets, Sioielo reise Se faite, 414 
The Polyzoa of Waterworks. By Sipney Bo Harmer, S8c.D., F.RS., F.ZS. 
(ELS SEGRE aks IDG NTE Siem iichno anion ne on dace cocCU moan AMC ne OOO ED secre emia 4 


The Marine Fauna of British East Africa and Zanzibar, fom Collections made by Cyril 
Crossland, M.A., B.Sc., F.Z.S., in the Years 1901-1902. Bryozoa—Cheilostomata. 
By Artnur Wu. Waters, F.L.S., F.G.8. (Pls. LXIV.-LXXIII. and Text-figs, 79-82.) 458 


Notes on Albinism in the Common Reedbuck (Cervicapra arundinum), and on the 
Habits and Geographical Distribution of Sharpe’s Steenbuck (Raphiceros sharpet). 
By Major J. Srnvenson-Hamitton, C.M.Z.S., Game-Warden of the Transvaal ........ . 537 


Contributions to the Anatomy and Systematic Arrangement of the Cestoidea,—X. On 
Two Species of Tapeworms from Genetta dongolana, By Frank H, Bepparp, M.A,, 
D.Sc., F.R.S., F.Z.S., Prosector to the Society. (Text-figs. 85-94.) -.........s2e000 549 


Pacific Salmon: An Attempt to evolve something of their History from an Examin- 
ation of their Scales. By Jonn Apam Mitnu, of Ardmiddle, Turriff, Aberdeenshire. 
Uber trap O STONE som gee scars moe caress oh ierotai oisr ae ctecaisinGr Stee uae win slaice @ ease e accra srure tie 572 


Notes on Peripatoides woodwardi Bouvier. By Karnuuun Happon ......00+-eeeees 611 


Field-Observations on the Enemies of Butterflies in Ceylon. By J. C. F. Fryur, M.A, 
F.E.S., Fellow of Gonville and Caius College, Balfour Student in the University of 


Cambridge ..... Nstvecionshiortu veins ioha ist eaeepart By etre orovslentt atatetera orca a oat ora Miibsises eo. 613 
On the South African Pseudosuchian Huparkeria and Allied Genera, By R. Broom, 
MEDS DESes OC, NME Ais. (Pls, TyXeXV SE XOWENG i ck ieee coo see © APTOS Haine Spo ORS 619 
On some Cases of Blindness in Marine Fishes. By G. E. Butusn, the Hertfordshire 
AVES SUITING SEMAN DATS ce Gisece cenclretat nS mesic oadeaeee a aienrs oe Gc opeUN rear Shae alawte Slee orate e ciinTowacn olateTece . 634 


Notes on Turacin and the Turacin-Bearers. By Sir ArtHur H. Onurcn, K.C.V.O., F.R.S. 639 


Observations on the Anatomy of the Shoe-bill (Baleniceps rex) and allied Birds. By 
P. Cuaumers Mircnent, M.A., D.Sc., LL.D., F.RS., ey of the Society. 
(Pls. LXXX.-LXXXIII, and Text-figs. 119-132.) perder siete stere ores Sa isrelaereles sais ce 644 


The Olassification and Phylogeny of the Calcareous Sponges, with a Reference List of 
all the described Species, systematically arranged. By Artuur Denpy, D.S8c., F.R.S., 
F.Z.S., Professor of Zoology in the University of London, and R. W. Haroup Row, 
B.Sc., F.L.8., Assistant Lecturer and Demonstrator in Zoology at King’s College. 


(CRextafe Gos) ~unbes ss etereraitine se sary wey = 5 mray oboe ofer eta ceatshsTOMh tera =ce cyefaietes vast ciel atets 704 
The Transvaal Race of the Cape, or Khama, Hartebeest. By R. Lypzxxnr, F.R.S., 
InolAds bes (USS ¢Failanel BIR Boca. ire rin Capi St. iad Gn ano eid OE ene Bint ceo oe 818 


On a Collection of Mammals from the Inner Hebrides. By G. EH. H. Barrent- 


- Haminton, F.Z.8., and Martin A.C, Hinron. (Pl. LXXXIY. and Text-figs. 186-138.) 821 


47. 


48. 


Some Miocene Cirripedes of the Genera Hexelasma and Scalpellum from New Zealand. 
By Tuomas H. Wirunrs, F.G.8, (Pls. LXXXV. & LXXXVI. and Text-figs. 189 & 140.) 840 


Observations on Osteomalacia in the Zoological Collections of Manchester and Cleve- 
land. By T. Wincatx Topp, M.B., F.R.C.S., Professor of Anatomy, Western Reserve 
University, Cleveland, Ohio. (Pls, LXXXVII.-LXXXIX.) ...... SGonUOhHOnoe acre Leow 


wee Page 
L. \ : 
LI. | 
LIT. } Costa Rican Butterflies ............¢ Bae APIA RAS _ 839 
LIII. | 
LIV. ) 
LV. ° Acanthistius fuscus ...cccesccree Que moomoO Ne ooodDS ) 
NaS Kahiiavmiutabunda 6.) eres seen cin inicio ts 
LVIL. ~Girellops nebulosus 2. ce ce oes s cco uc uceusecuceses 
LVIIL. 1. Pomacentrus inornatus. 2: Labr ichthys big + 368 
3. Anampses pulcher....c+..ccrecess AE OTONU Dog Ho 
LIX: Anampses pulcher . 0. cocvccce sees en seas on,te © Haetinod 
LX. Pseudomonacanthus paschalis .....0..eccersces-vecees ) 
LXI. Patella of Phalacrocoracidz .........+2+cccceseees -. 898. 
PRU Pa idicella Greiculacd, ame mate vies ce a cn eleake s+ | 496 
LXIII. Statoblasts of Phylactolemata ........c..cseeeoecees \ 
LXIV.) 
LXV. | 
LXVI. 
LXVII. 
ea (eago7es from Zanzibar .....+++e06 NAIA D RSIS ernie e 458 
LXx. 
LXXI. | 
LXXxIi. 
LXXIIt. ) 
LXXIV. Zonurus giganteus. Rone as oar quaamoo as “sities eke 548° 
ene \ Huparkeria capensis ....-- Sher inatietavate eters canine aRecCtene Te i. 
LXXVII. 9,10. Ornithosuchus woodwardi. 11. O, taylori.. ee 
LXXVILI. Mesosuchus browns and others.....0+..cessecccescees 
LXXIX. Browniella africana and others .....-....-- sie eee ae ae 
POO TAMIUGHSOS REO ebo co ogo pon pO OU Oo CUO ONO OUbndOGaCdr: 
LXXXI, Skulls of 1. Baleniceps. 2. Scopus. 3. enoriynchis 
LXXXII. Skulls of 1. Dissura. 2. Tantalus. 3. Cancroma.... 
UXXXIIT. Skulls of 1. Ardea. 2. Pelecanus. 3. Plotus......2-.. 
LXXXIV. 1. Sorex grantii. 2. S. araneus castaneus.. 
LXXXV. Hexelasma auchlandicum Hector sp. ..++-...-ses.seeee 
LXXXVI. 1-6. Scalpellum subplanum. 7-13. 8S. (Arcoscalpellum) 
UMGULACUTU : ons si vinin® = onto) dere eis = nidietelale reheat an 
LXXXVII. 
LXXXVIII. | Osteomalacia ina Baboon ......-- eS elaec ere eaeeareicia eee 
LXXXIx. 


LIST OF PLATES. 


1918, Part III. (pp. 339-860). 


NOTICE. 


The ‘ Proceedings’ for the year are issued in four parts, paged consecutively, 
so that the complete reference is now P. Z. 8.1913, p.... The Distribution 
is as follows:— 

Part I. issued in March. 
SLs a June. 
Poise [El Beeler, September. 
Peseta AZ December. 


‘ Proceedings,’ 1913, Part II. (pp. 153-337), were published on 
May 30th, 1913. 


The Abstracts of the ‘ Proceedings,’ Nos. 122 & 123, are 
contained in this Part. 


SE DS DEP 


PROCEEDINGS 


OF THE 


GENERAL MEETINGS FOR SCIENTIFIC BUSINESS 


OF THE 


ZOOLOGICAL SOCIETY 
OF LONDON. 


(1913. 


PART IV. 


- CONTAINING Paces 861 to 1104, witH 24 PLaTEs 


xf eS ae ae ee at 0". — — 
J . eae es eee our 
3 ’ : 5 ay 
; : . 4 Ste 
SS 
ZL 
* ° 
as . 


AND 55 TEXT-FIGURES. 


DECEMBER 1913. 


PRINTED FOR THE SOCIETY, 
SOLD AT ITS HOUSE IN REGENT’S PARK. 
LONDON: 
MESSRS. LONGMANS, GREEN, AND CO,, 


PATERNOSTER ROW. 


ee a 


[Price Twelve Shillings.) 


LISET SOF CONT ENS: 


1913, Part IV. (pp. 861-1104). 


EXHIBITIONS AND NOTICKS. 
Page 
The Szcrerary. Report on Additions to the Society’s ee during the months of 
Miayationsepbenmtbar LOU. ls tc. cicinic © ater cp ate ele whetn «cin cons te Geceieinge Suerte oe tenia tate eee 1090 


“Tue Secrnrary, Exhibition of the Alimentary Tract of a young Elliot's Pheasant 
G@Phastanusielliote) ae caccies sae oles bs Sule See Ee ee toe LEE Gee 1094 


Mr. RB. I. Pococr, F.R.S., F.Z8. Exhibition of young Woolly Opossums (Philander 
laniger) and young Degus (Octodon degus) .. 0... .cccer sere oe ce nc ress cdeeeeoers 1094 


Mr. D. Srtu-Suiru, F.Z.8. Heel-Pads in a young Toucanet (Selentdera macuitrostris). 
(Moxt Roe mlOd\cctek suis as inctseis ser pi\ p weciee yummie en gana ior noe aeT: Sona ods: +. 1095 


Mr. CO. Tarn Recan, M.A., F.Z.S. A Deep-Sea Angler-Fish, Melanocetus johnsonii .... 1096 


Mr. E. G. Bouncer, F.Z.S. Exhibition of a young specimen of the Matamata Terrapin 
(QSL HED EOE HRT OC pe oc ies oro iitiow ee hee nee cone ene 1097 


Mr. D. Setu-Ssuru, F.Z.8. Exhibition of Hybrid Birds. (Text-fig. 192.) ............ 1098 
Mr. R. I. Pocock, F.R.S., F.Z.S. Dorsal Glands in Armadillos. (Text-figs. 193-195.) .. 1099 


The SecreTary. Report on Additions to the Society's Menagerie during the month of 
October NOlSr 2. .5.< a. SAO NGG Ga ote Oa aN OU SO ‘Gaielletabeis acetate eee 1103 


Prof. H. Maxweti Lerroy, M.A., F.Z.8. Remarks upon the New Insect House..... o.- 1104 


Mr. R. H. Borns, M.A., F.Z.8. Exhibition of Foetal Skeletons prepared by the Beale- 
Selulizepme tod yejr-couetey civ enelasereve's. ors ote 5 ots vs oteseupinlnte oy tecwions cite eet ao hole ees ene eee 1104 


PAPERS. 


49. Contributions to the Anatomy and Systematic Arrangement of the Cestoidea,—XI. On 
a new Tapeworm from (dicnemus. By Frank HK. Bupparp, M.A., D.Sc., F.BS., 
F.Z.S., Prosector to the Society. (Text=figs. 141149.) oe os en's ot niin sm sips ea &61 


50. The Brain and Brain-Case of a Fossil Ungulate of the Genus Anoplotherium. By 
R. W. Paumer, M.Sc. (Manchester), Research Fellow in Zoology, University College, 
Reading; (Dext-feeS0= Laie): wih csistls w nlas ae oma Setar ae Maric eo bb eae 878 


Contents continued on page 3 of Wrapper. 


OF LONDON. 


a 


ZOOLOGICAL SOCIETY 


Tars Society was founded in 1826 by Sir Sramrorp Rarries, 
Mr. J. Sasrne, Mr. N. A. Vigors, and other eminent Naturalists, 
for the advancement of Zoology and Animal Physiology, and for the 
introduction of new and curious subjects of the Auimal Kingdom, 


and was incorporated by Royal Charter in 1829, 


Patron. 


HIS MAJESTY THE KING, 


COUNCIL. 
HIS GRACE THE DUKE OF BEDFORD, K.G., F.RS, President. 


Str Jonn Rost BrapForp, Str Watters Roper Lawrence, 


K.C.M.G., M.D., D.Sc., F.RB.S., 
Vice-President. 

Ricuarp H. Burns, Ese., M.A. 

Sure.-Genu. Srr R. Havenock 
Cuargtes, G.C.V.O., M.D. 

Tue Rr. Hon. Toe Eart or 
Cromer, P.C., G.C.B., 
G.C.M.G., K.C.8.1., F.R.S., 
Vice-President. 

F, G. Dawrrey Drewitt, Ese., 
M.A., M.D. 

Caartes DrumMonp, 
Treasurer. 


Kse., 


Str Epwarp Duranp, Br., C.B. 

Freperick Gittett, Ese. 

Tue Lorp GLENCONNER, 

F, Du Canz Gopman, Ese.,D.C.L., 
F.B.S. 


Br., G.C.LE. 


Str Epwunp G. Loper, Br., Vice- 
President. 


Ernest W. MacBripr, Ksa., 
M.A., D.Sc., F.R.S., Vice- 
President. 


Proressor Epwarp A. Mrycurn, 
M.A., F.R.S., Vice-President. 


P. Caatmers Mrrcnert, Ese., 
Wises IDES, IA aID) 5 JB oIey sie. 
Secretary. 


W. R. Ogitviz-Grant, Ese. 
Aprian D. W. Pottocg, Ese. 
OtprieLtp Tuomas, Kse., F.R.S. 
AntHony H. WineFrrezp, Hse. 


Henry Woopwarp, Ksa., LL.D., 
E.R.S., Vice-President. 


2 
The Society consists of Fellows, and Honorary, Foreign, and 
Corresponding Members, elected according to the By-Laws. It 


carries out the objects of its foundation by means of the collection 
of living animals, by its Library, and by its Scientific Publications. 


The Office of the Society, Regent’s Park, N.W., where all com- 
munications should be sent, addressed to “The Secretary,” is open 
from Ten till Five, except on Saturdays, when it closes at One p.m. 


The Library, under the superintendence of Mr. Henry G. J. Peavot, 
is open daily (except Sunday) from Ten till Five, except on 
Saturdays, when it closes at Two p.m. The Library is closed from 
Good Friday to Easter Monday, and upon all other Bank Holidays. 
It is also closed annually for cleaning purposes during the whole 
month of September. 


The Meetings of the Society for General Business are held in the 
Meeting Room at the Society’s Office on the third Wednesday in 
every month of the year, except in September and Octeber, at half- 
past Four o’clock p.m. 

The Meetings for Scientific Business are held in the Meeting 
Room at the Society’s Office fortnightly on Tuesdays, except in 
July, August, September, and December and January, at half-past 
Hight o’clock p.m. 

The Anniversary Meeting is held on the 29th. of April, or the 
nearest convenient day, at Four p.m. 

The Society's Gardens are open daily from Nine o’clock until 
Sunset. Mr. R. I. Pocock, F.R.S., F.L.S., is the resident Super- 
intendent and Curator of Mammals, Mr. D. Seth-Smith is Curator 
ot Birds and Inspector of Works, Mr. E. G. Boulenger is Curator 
ot Reptiles, and Prof. H. M. Lefroy is Honorary Curator of Insects. 
The Prosectorium for Anatomical and Pathological work is under 
the charge of Mr. Frank E. Beddard, M.A., D.Sc., F.R.S., Prosector, 
assisted by Mr. H. G. Plimmer, F.R.S., M.R.C.S., Pathologist to 
the Society. 


‘TERMS FOR THE ADMISSION OF FELLOWS. 

Frrrows pay an Admission Fee of £5, and an Annual Contri- 
bution of £3, due on the Ist. of January, and payable in advance, 
or a Composition of £45 in lieu thereof; the whole payment, 
including the Admission Fee, being £50. 

No person can become a Frtxow until the Admission Fee and 
first Annual Subscription have been paid, or the annual payments 
have been compounded for. 


Frtiows elected in November and December are not liable for 
the Subseription for the vear in which they are elected. 


PRIVILEGES OF FELLOWS. 


Fettows have Personal Admission to the Gardens upon signing 
their names in the book at the entrance gate, and may introduce 
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The Wire or Hussanp of a Fattow can exercise these privileges 
in the absence of the Fellow. 


Until further notice, Frtnows will receive 40 undated Green 
Cards, available on any Sunday or week-day up to the end of 
February of the year following the year of issue, and 20 White 
Cards available on any week-day up to the same date. Twenty 
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available for any day except Sunday in the week ending with the 
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It is particularly requested that Fellows will sign every ticket before 
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Frettows are not allowed to pass in friends on their written 
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Frertows may obtain a TransreraBLe Ivory Ticker admitting 
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LISTS OF THE ANIMALS IN THE SOCIETY’S GARDENS. 


List of the Vertebrated Animals now or lately Living in the Gardens 
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THE OFFICIAL ILLUSTRATED GARDEN GUIDE—11th Edition 
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animals in the Collection (now containing over 3000 spe- 
cimeus), together with 50 Photographic Illustrations and 
Index. Price 6d. in Stiff Paper Cover, postage 14d., or in 
Green Cloth Cover price 1s. 2d. post free. 


P. CHALMERS MITCHELL, 
Secretary. 
Regent's Park, London, N.W. 
December, 1913. 


These publications may be obtained at the Socinry’s OFFIce, 
or through any bookseller. 


ON AVIAN CESTODES. 861 


49. Contributions to the Anatomy and Systematic Arrange- 
ment of the Cestoidea. By Frank HE. Bepparp, M.A., 
D.Sc., F.R.S., F.Z.S., Prosector to the Society. 


[Received August 27, 1913: Read October 28, 1913. } 
XI. On A New TAPEWORM FROM GQ/DICNEMUS. 


(Text-figures 141-149.) 


INDEX. 
Page 
EHugonodeum edicnemi, gen. eb sp. mn. ............ Saw cette eS Ol 
Characters of Hugonodeuwm ...... A is NE as een eae Sal eS) 7(7 
[DiREOMVMMES Ce OOCMOPIRIIAE  sononucansasetcnteectsboseecacdseuedcanooscs  ov/4t 
(GSMS OQ OSAAAMMONS oocaccéscnceasopsccocss sos ssbaenoaseesmssbonssercoovecca. ot 


In Prof. Fuhrmann’s exhaustive list of Tapeworms * (of the 
Cyclophyllidea only) which occur in birds, only one parasite 
of this order is recorded from an Mdicnemus. The species in 
question is Choanotenia coronata, and it occurs in the Kuropean 
“dicnemus edicnemus. 1 was, therefore, particularly interested 
to find in a New World species of the genus, viz. Gdicnemus 
bistriatus, a number of examples of a tapeworm which does not 
belong to the genus Choanotewnia, though it should probably be 
referred to the same family and even subfamily. 

I shall, however, defer the consideration of the systematic 
position of this Cestode until an account has been given of its 
anatomical characters. 

The species dealt with in the present communication was 
obtained from a Thick-knee of the species mentioned above, which 
died in April of the present year, and had lived in the Society’s 
Gardens fora year. It is not unlikely, therefore, that the tape- 
worms had infected the bird hefore its arrival. But no fact 
of importance can be at present deduced as to locality, since the 
very parasite of Gdicnemus owdicnemus mentioned above has 
been also discovered in Wgialitis nivosa, which is an American 
bird. 

The @dicnemus bistriatus contained no other tapeworms in its 
intestine, except that which forms the subject of the present 
communication to the Society. JI am not quite certain how 
many individuals there were; but there were at any rate six or 
seven. The worms are somewhat slender and measure at least 
120 mm. I infer this from the fact that one of the largest 
fragments, consisting of both ripe and unripe proglottids, measured 
110 mm. ; but, as it had no scolex nor trace of the thin neck, at 
least 10 mm. may be added safely. The greatest breadth of the 


* “ Die Cestoden der Vogel,” Zool. Jahrb. Suppl.-Bd. x., 1908. 
Proc. Zoou. Soc.—1913, No. LVIII. 58 


862 DR. F. E. BEDDARD ON 


ripe protoglottids is 2 mm. The accompanying figure (text- 
fig. 141) shows the scolex and anterior end of an example of this 
worm, as well as a portion of the posterior region of the same or 
of another example. The anterior part of the body shows the 


Text-fig. 141. 


Kugonodeum cedicnemi. 


Portions of two examples about twice nat. size. 


The left-hand figure shows the rather large scolex and whiplash-like anterior 
part of strobila. he right-hand figure consists of mature segment». 


rather long and oval scolex followed by a slender neck which 
widens out rather suddenly. Only one specimen out of the six 
or seven which I have examined showed a gradual increase in 


AVIAN CESTODES. 863 


thickness from the neck onwards. This characteristic of the 
species is not unfrequently met with in tapeworms; it is, 
however, perhaps unusually strongly marked in this worm from 
“dicnemus bistriatus. 

The scolew is long and oval, and provided with four large 
suckers which possess no armature of hooks. 

When the scolex and the rostellum are extended the suckers are 
quite in contact laterally. They are then oval in form. The 
rostellum is long and retractile; when fully extended it is as long 
as the rest of the scolex. The extremity is entirely muscular 
with several layers of stout fibres running both transversely and 
longitudinally. But in spite of its great muscularity there are 
no hooks. It has been recorded, and I myself have had the 
opportunity of observing a case, that among tapeworms which 
possess an armed rostellum, the hooks are occasionally absent. 
But in the present species I have never found them in a single 
individual. J am thus disposed to think that they are really 
absent, and to rely upon this as one of the distinguishing 
characters of the genus, which will be duly set forth later *. 
Behind the scolex there is quite a distinct neck, in which there 
is no trace of segmentation to be observed. The segments of 
the body overlap in the usual fashion and are never greatly 
elongated. The most posterior are only slightly longer than broad. 
The last external character to which I draw attention is the 
position of the genital pores, which are unilateral. In the ripe 
and more elongated segments they lie towards the hinder margin 
of each proglottid. These orifices are strictly lateral. In the 
narrow anterior segments the genital pores have a peculiar dis- 
position. In horizontal sections through a consecutive series of 
such proglottids the edges of the proglottids stand out for a con- 
siderable distance laterally. The genital pores open on these 
extensions, but on to the anterior margin of each where it runs 
parallel with the posterior margin of the lateral extension of the 
proglottid in front. The orifices are thus completely concealed on 
a lateral view. In more mature segments these lateral extensions 
do not stand out in the same straight line with the rest of the 
segment, but become bent backwards; so that the anterior margin 
becomes lateral. 

The structure of a proglottid is illustrated in text-fig. 142 
(p. 864). The cortical layer is about as thick as the medullary 
layer. The former contains two rows of bundles of longitudinal 
muscular fibres; the number of fibres constituting each of the inner- 
most bundles is greater than that of those of the outer. There 
appear to be a larger number of bundles also in the inner row. 
These longitudinal muscle-bundles do not extend outside of the 
nerve-cord ; they are not at all closely arranged. The water- 
vascular system consists of two lateral tubes on each side, of which 
the outermost, or ventral, is several times larger in cross-section 


* Vide infra, }. 877. 


C1 


864 DR. F. E. BEDDARD ON 


than the innermost, which I take to be the dorsal vessel. The 
tubes run nearly parallel to each other; but the dorsal vessel is 
slightly to the dorsal side of the proglottid. The ventral vessels 
are connected in each proglottid by a transverse vessel which is 
shown (in part) in text-fig. 143. 


Text-fig. 142. 


Part of a transverse section through a proglottid of Hugonodeum cedicnemi, to show 
the arrangement of the water-vascular tubes and cf the longitudinal muscles. 


d. Dorsal vessel. m. Longitudinal muscles. #. Nerve-cord. v. Ventral vessel. 


The ovary of this tapeworm is single and not divisible into two 
lobes, as is so often the case. It lies distinctly to the pore side of 
each proglottid median of the large (ventral) water-vascular tube 
and partly ventral, as well as extending to both sides of the 
dorsal water-vascular tube. It les in each case anteriorly in the 
proglottid, and not very far behind the transverse water-vessel 
of the proglottid in front. In front of the ovary, however, lie 
some of the coils of the sperm-duct; it is not, therefore, quite at 
the anterior margin of the segment. The vitelline gland lies 
behind the ovary and towards its inner (median) side. The 
relationship is shown in the accompanying drawing (text-fig. 143). 


AVIAN CESTODES. 865 


Tt will also be seen, on an inspection of that figure, that the 
fully grown ova ave largely grouped together in cavities, and that 
the ovary is more or less hollow. J am not, however, disposed 
to think that this fact has any morphological significance. Two 
interpretations are, of course, conceivable. Firstly, that the 
Space is celomic, the eggs having been freed from its walls into 
its cavity. The occurrence of a similar cavity in other tape- 
worms may perhaps be an argument in favour of this view, which 
is by no means impossible (and, indeed, has been theoretically 
demanded by Bergh and others) ; yet I am disposed to regard the 


Text-fig. 143. 


tv 


\ es Vv. 


Part of a horizontal section through a proglottid of Hugonodeum cedicnemi. 


e.s. Egg-sacs. J.v. Ventral water-vascular tube. sp.d. A part of ccil of sperm-duct 
lying in front of ovary. ¢. Testes. ¢.v. Transverse water-vascular tube. 
v. Vitelline gland ; in front of this is the ovary, of which the darkly stained 
mature ova are partly received within cavities of the parenchyma. 


cavities as merely due to shrinkage. These hollows might also 
be considered as the commencement of the uterus. The position, 
however, would be rather abnormal; and, moreover, as again 
is insisted upon later in this paper, ripe ova are already scattered 
through the parenchyma, the cavities surrounding which can 
therefore have nothing to do with the cavity in the ovary, what- 
ever may be its nature. In younger stages than that which has 
just been described the ovary forms a quite solid mass. Later 
on it seems to disappear as a definite structure. 


866 DR. F. E. BEDDARD ON 


S Genital Ducts. 


This worm is characterized by the great depth of the genital 
atrium, which is much more developed than in most tapeworms 
and quite as much so asin any. ‘The orifices of these follow in 
successive proglottids upon the same side of the body. In the 
ripe proglottids they are situated quite posteriorly and, therefore, 
the genital ducts which open into them run forwards towards the 
anterior region of the proglottid. The genital atrium, or genital 


Text-fig. 144. 


Part of a horizontal section through a proglottid of Eugonodeum cedicnemi, 
showing a generative aperture. 


c. Cirrus extruded from cirrus-sac (shown lying in front of it) and received within 
vagina (v;). e.g. Very deep cloaca genitalis. sp.d. A coil of sperm-duct. 


cloaca, is lined by a prolongation of the outer cuticle; outside of 
this is a very thick layer of rather stout muscle-fibres which, 
when the genital cloaca is cut through transversely to its axis, 
are seen to have a radial arrangement. The cavity is circular in 
transverse and oblong (more or less) in longitudinal section. At 
the base, where the genital ducts join it, it widens out and forms 


AVIAN CESTODES. 867 


a circular cavity projecting beyond the rest of the genital cloaca. 
The general shape is suggestive of a silk hat, if the brim of 
the hat were hollow. From the centre of the lumen of the 
genital cloaca a slender tube, which from its structure appears to 
be morphologicaliy a part of the genital cloaca, leads to the 
junction of the male and female ducts. This is shown in text- 
fig. 144. It should be noted that the muscular layer of the 
cloaca genitalis 1s covered externally by a layer of subcuticular 
cells continuous with those which underlie the body cuticle. It 
is, therefore, perhaps to be presumed that the muscular layer in 
question is a local thickening of the delicate layer of fibres which 
underlies the body cuticle. Occasionally I have observed the 
cloaca genitalis to be slightly protruded ; in such cases the cavity 
was more cup-like than cylindrical owing, of course, to the gaping 
of the external orifice. 

The sperm-duct and cirrus-sac, as already mentioned, pass 
obliquely forwards in the mature segments and are parallel to the 
vagina, which follows an identical course. ‘The cirrus-sac in this 
worm is large and directed in a straight line towards its opening 
into a chamber in common with the vagina. This latter chamber 
has nothing to do with the terminal cloaca genitalis from which 
it is sharply marked off by the muscular walls of the cloaca, 
which have a narrow. tube of intercommunication, doubtless 
capable of being widened. In all the individuals which I ex- 
amined the cirrus of the fully mature proglottids was largely 
protruded from the cirrus-sac, but not through the cloaca 
genitalis to the exterior of the body. I found the cirrus, in fact, 
to be invariably inserted into the neighbouring vagina, which 
latter was as invariably filled with spermatozoa. here is, of 
course, nothing new in this record of self-fertilization, which is 
well known to occur among Cestodes, and has been recorded by 
van Beneden, Leuckart, and others. A large number of instances 
are given in Bronn’s ‘ Klassen und Ordnungen des Thierreichs ’ 
by Prof. Max Braun*. I have not, however, noticed it myself 
in the considerable number of species which I have examined, 
excepting in the present species. This auto-copulation is there- 
fore far from being universal in occurrence. It appears to me, 
furthermore, that the structure of the efferent apparatus in the 
Cestode, which forms the subject of the present communication, 
may at times necessitate this auto-copulation. A closure of the 
cloaca genitalis coinciding with the eversion and protrusion of the 
cirrus would force the latter into the vagina, which is widened at 
its extremity, and does not project into the common chamber 
into which both efferent ducts open t. The passage for the cirrus 
is therefore not in any way hampered. 

The cirrus-sac has thickened walls, as in so many species, and 


* Bd. iv. Abth. 1, p. 1462. 

+ v. Janicki in describing Schizotenia haymanni (Zeitschr. wiss. Zool. 1xxxi. 
1906, p. 585), where there is also a very deep cloaca genitalis, comments on the 
mechanical necessity for auto-copulation on rather different grounds. 


868 DR. F. E. BEDDARD ON 


is somewhat bottle-shaped. ‘The narrow region, however,—the 
neck of the bottle—is not anterior where the cirrus-sac opens into 
the common genital atrium, but posteriorly where the sperm- 
duet opens into it. I have not always seen this narrow backward 
prolongation of the cirrus-sac ; but I have seen it in more than one 
case. There is no difference in its structure from that of the 
main part of the cirrus-sac*. The cirrus, as usual, lies coiled 


Text-fig. 145. 


e. m. ue 


. 


Part of a horizontal section through a proglottid of Eugonodeum cedienemi, 
to show further course of vagina. 


e. Egg-sacs. m. Longitudinal muscle-fibres. 2. Nerve-cord. v. Vagina, full of 
sperm and widening above to form a receptaculum seminis, thence bending 
hack again to divide into the two usual branches o. & V7it., which are arranged 
in the same straight line with each other. 


within the sac and when protruded, as shown in text-fig. 144, is 
broader at the protruded end. This is inserted into the vagina, 
the proximal dilatation of which it completely fills, as is shown in 
the same figure. The sperm-duct, after leaving the cirrus-sac, 
runs in a straight or slightly sinuous course, at first quite parallel 


* Tt is possibly temporary, and due to nnequal contraction. 


AVIAN GESTODES. 869 


to the vagina on the anterior side of it. It does not expand any- 
where into a vesicula seminalis, but forms a large coil. This coil 
lies in the neighbourhood of the two water- vascular vessels. 

The vagina opens behind the cirrus-sae and is wider, and with 
more muscular walls in that section which lies close to the ex- 
ternal pore than elsewhere. It is this region which receives the 
cirrus during the auto-copulation referred to above. The tube 
then becomes finer and passes parallel to the sperm-duct, as shown 
in text-fig. 144. At about the level of the larger, ventral, water- 
vascular tube it expands into a not very large receptaculum seminis 
(text-fig. 145), which in fully mature proglottids is gorged with 
sperm. From this point in fully mature segments the vagina 
then bends back again and divides into two tubes, one running 
anteriorly and the other posteriorly. In less fully mature pro- 
glottids the course of the whole vagina is straight and but slightly 
oblique, being nearly parallel to the transverse axis of the body. 
The shell-gland in such segments is very plain. 


§ Egg Sacs. 


The egg-containing cavities (which do not, as I think, collec- 
tively represent a uterus) occur in segments which have already 
begun to lengthen slightly, although they are still much broader 
than long. In such segments the ovary and testes are fully 
mature , but show no signs of degeneration. Several of these egg- 
holding cavities are dis splayed i in text-fig. 143 (p. 865). They are 
all small, but not of uniform size; their shape is quite uniformly 
spherical. There is no regularity of arrangement among them 
that I can detect; they lie everywhere in the proglottids, even 
among the bundles of longitudinal muscle-fibres; they thus 
axe into the cortex, a position which is, however, not unknown 
in other tapeworms, though it is not common. There is no 
connection to be observed between adjacent ege-cavities, though 
they may he in actual contact: that is to say, there is no question 
of a network—the cavities are totally independent in fact and, as 
I shall point out, probably so in origin. 

These egg- containing cavities, he largest of which are smaller 
than the teste es, are definitely marked off from the parenchyma, in 
which they lie, by a thickish wall. The cavity which lodges the 
ege@* and the vitelline cells is naturally more apparent in the case 
of the larger sacs. In the smaller ones the egg and accompanying 
eells fill up the available space completely or nearly completely. 
The inference appears to me to be that the cavity is formed later 
perhaps by the exudation of fluid as well as growth of the peri- 
phery in a way similar to that of the mammalian Graaftian follicle. 
On the other hand, the structure of the walls of these egg- 
containing cavities suggests another interpretation. As already 
mentioned, they are rather thick and thus very conspicuous: in 


* Occasionally two eggs are found in the same cavity. 


870, DR. F. E. BEDDARD ON 


them occur numerous nuclei sometimes close together and some- 
times more sparsely arranged. These nuclei are more numerous 
in the larger egg-holding spheres, and would thus seem to have 
multiplied during a growth. But it must be admitted that the 
existence of sacs with the lining apparently epithelial is highly 
suggestive of a state of affairs like that shown by Dipylidium 
caninum. 


Text-fig. 146. 


A horizontal section through mature proglottid of Hugonodeum ceedicnemi. 


A, Advanced embryo in egg-holding cavity. B. Less advanced embryo in sinaller 
cavity. v. Ventral water-vascular vessel giving off transverse vessel. 


In this worm, as is well known*, the uterus finally breaks up 
into numerous quite separate cavities, 1n which he many ripe 
ova, and which are lined by a continuous epithelium. The 
arguments against this supposition, however, outweigh those in 


* Of. e.g. Beddard, P. Z.S. 1918, p. 555 text-fig. 88. 


AYIAN CESTODES. 871 


its favour. In the first place there is no trace that I can discover 
of a pre-existent uterus*, whose subsequent fragmentation might 
produce the result described above. Furthermore, the actual 
cavities of the egg-holding apparatus are a later development, or 
at any rate they increase in extent as the eggs within them grow. 
Indeed it may, I think, be safely asserted that in the youngest 


Text-fig. 147. 


i 


Part of the section illustrated in text-fig. 146 more highly magnified and showing 
younger egg-sacs in parenchyma (e.s.) and testes (¢.), which are very much 
larger. 


groups of egg and surrounding cells there is no free space at all. 
Nevertheless it might be said that all this was due to precocious 
development, that in fact the rapid protrusion of ripe eggs from 
the ovary had outstripped the growth of the uterus, which in 
consequence appeared subsequently in point of time, and that 


* Of., however, p. 865 under description of ovary. 


872 DR. F. E. BEDDARD ON 


the uterus for this latter reason appeared in an already divided 
condition. 

It is to be noted that egg-sacs of the character just described 
occur in the younger proglottids, and that they are also to be 
found (text-fig. 145, p. 865) in quite young proglottids, in which 
the ovary and vitelline gland are at their full development and 
have not commenced to degenerate. Without having any positive 
proof, | am disposed to think that the cells which encircle the 
ovum are ovarian or vitelline gland-cells*, which come to be dis- 
posed round the ovum after the fashion of a follicular epithelium 
in higher animals. It looks to me, in fact, very much as if a ripe 
ovum with a few adherent cells moved out into the adjacent 
parenchyma directly from the ovary. There is here clearly a 
similarity with the Graattian follicle of Vertebrates, a similarity 
which is even increased by the later appearance of a space (perhaps 
containing fluid) which surrounds the growing ovum. 

It does not always follow that a proglottid which has attained 
the dimensions and acquired the appearance of a fully mature 
proglottid should contain only embryos. On the contrary, as 
be readily seen from an inspection of text-fig. 146 (p. 870), 
a large flat proglottid, quite as large as it will grow, may have 
egg-sacs 1n many stages of growth. In this figure, which is 
dvawn from a section seen under a low power, the size of the 

egg-sacs 18 a rough measure of their relative stages of growth. 
The largest are of course the oldest, and contain fully developed 
embryos surrounded by a thick shell and enclosed in rather a 
large space. Intermediate conditions are to be found between 
this stage and that in which an undivided ovum occupies all the 
space of the cavity of the body-parenchyma in which it lies. I 
have also cut sections of apparently fully mature proglottids, in 
which there were no advanced embryos. 

This state of affairs is not unlike what has been described in 
some other Tapeworms by others as well as by myself. We may 
exclude Monopylidiwm and Dipylidium, which only show an 
apparent likeness to Hugonodeum. In the former genus, and in 
certain species belonging to the latter genus, the inclusion of the 
ripe ova singly in compartments of the medullary parenchyma 
is preceded by a stage in which a functional uterus exists or 
(Dipylidium, various spp.) at any rate a cavity which—full of 
eggs——is ultimately broken up into: single compartments. On 
the other hand, there is a much greater likeness to Hugonodeum 
in a genus which I have lately described as new under the name 
of Diplopylidium*. Here we have, as it would appear, no trace 
of a uterus; but the eggs are found scattered widely through 
the medullary parenchyma enclosed in a cavity singly ; the cavity 
grows in correspondence with the growth of the contained 


* (Of. description of ovary and vitelline gland above. 
+ P.Z.S. 1913, p. 565, text-fig. 93, p. 566, text-fig. 94. 


AVIAN CESTODES. 873 


embryo. There is no great difference in size between the egg- 
holding cavities in the two genera. But there is an important 
difference in the structure of the wall of these cavities in Diplo- 
pylidium, and in the presently described genus Hugonodeum. In 
Diplopylidiwm 1 was unable to find any nuclei lining the cavities, 
which are simple excavations in the medullary tissue. This was 


Text-fig. 148. 


Another part of the same section of Hugonodewn cedicnemi showing older embryos 
surrounded by a greater egg-holding cavity in the medullary parenchyma. 


A, B. Egg surrounded by thick shell and lying in cavity from which nutritive 
cells (?), such as are shown at es. in text-figure 147, have disappeared. 
C. Older embryo with larger space surrounding it. D. A degenerating (?) 
egg-holding cavity and embryo. 


the case, not only with the fully mature embryos, but also in com- 
paratively newly formed cavities. In Hugonodewm, on the other 
hand, the egg-holding spaces possess a lining of cells. It is true 
that in fully mature embryo-containing cavities these cells are 


874 DR. F. E. BEDDARD ON 


often, perhaps generally, indistinguishable. But a granular 
detritus often visible seems to me to indicate that their tendency 
to disappear is due to the fact that they serve as nutriment for 
the growing ovum. It may be noted, furthermore, that the egg- 
shell in Lugonodewm is not nearly so broad as in Diplopylidium. 

A reseinblance exists between the disposition of the ripe ova 
in the present genus and that which characterizes Oochoristica. 
I have re-examined examples of a species of Oochoristica which I 
described lately to the Society as a near ally of, if not identical 
with, Oochoristica wageneri of Janicki*. In the course of my 
account of that species I figured the scattered condition of the 
ripe eggst, well known from the investigations of others to he 
one of the characters of this genus. I did not, however, attempt 
any details, merely showing the inclusion of the ripe embryos 
eich im a separate chamber, and these scattered through the 
general medullary parenchyma. I have now some details to add 
to that account. The accompanying figure (text-tig. 149) shows a 
part of a section through one of the last two proglottids of 
specimen of this species. It will be seen that the eggs are more 
closely packed than in Mugonodeum. Vhey are not, however, in 
contact or enclosed in one cavity. This much concerning the 
ege-sacs of Oochoristica 1s well known from the observations of 
several writers previously to myself. e. gy. Zschokke and v. Janick1. 
But neither Zschokke{ nor v. Janicki§ give any detailed figures 
in their memoirs dealing with Oochoristica of the structure of the 
scattered eggs or oncospheres, or of the spaces which they oceupy 
in the parenchyma. Zschokke describes the three membranes 
which surround the hexacanth embryos, and observes of them 
(the embryos) that they ‘“ liegen je einzeln in dichtgedringten, 
rundlichen, kapselartigen Follikeln des Parenchyms.” Goin) 
figures the ‘crowded embryos with their three surrounding mem- 
branes entirely filling the proglottids at the end of the body ; 
but he also does not enter into various histological points which 
are, as I think, of interest in comparing the scattered egg-capsules 
of Oochoristica with those of other tapeworms. 

In longitudinal sections of the entire ripe proglottid these egg- 
capsules are seen to be circular or more oval in contour, and to 
be larger or smaller in size with no regularity of arrangement. 
The differences of size correspond as I think partly at least to 
differences of age. The more or less circular contour may have 
a relation to the plane of the sections. Contrary to the obser- 
vations of Zschokke and Cohn upon the species studied by 
themselves, I can see but one egg-membrane which occupies all 
the available space in the egg-follicle. V. Janicki gives no 


Zeitschr. wiss. Zool. lxxxi. 1906, p. 533. 

P. Z.S. 1911, p. 633, text-fig. 150. 

“Das Genus Oochoristica Lithe,”’ Zeitschy. wiss. Zool. Ixxxiu. 1906, p. 53. 
Loe. cit. 

Arch. f. Naturg. 1903. 


tt & 


AVIAN CESTODES. 875 


description of the eggs of Oochoristica wagenert, which is nearest 
to, if not identical with, my species, except to state that they are 
provided with a shell. The sharply defined edge of the shell 
differentiates the ovum or the cells of the embryo, if it has deve- 
loped so far, which lie within it from a layer of cells which forms 
a follicle, and is so far like that which has just been described in 
Eugonodeum. There are, however, differences to be noted which 
will be apparent from the two figures (text-figs. 148, 149) which 
illustrate the two species, and which are drawn practically to the 
same scale. The cells of the egg-follicle in Oochoristica have 


Text-fig. 149. 


A section through a portion of the medullary parenchyma of Oochoristica, showing 
five eggs or developing embryos. p. One pole of the usually elliptical 
embryo-sac. 


nuclei of a different appearance and form a thicker layer in that 
the nuclei are more abundant and closer together. Furthermore, 
this cellular layer is just as well developed in the largest as 
in the smallest follicle. The cells do not disappear during the 
growth of the contained embryo, as would appear to be the case 
with Hugonodeum. It is easy to believe, therefore, in accordance 
with the opinion of the authors quoted above, that the egg-sacs 
in Oochoristica are portions of a subdivided uterus. I have, 
however, myself no facts to offer in confirmation of this view. 


876 DR. F. E. BEDDARD ON 


In many of the ege-follicles the form of the entire follicle is, as 
already stated, oval. In these cases, which are illustrated in text- 
fig. 149, the layer of cells extends at the two poles of the ellipse 
for a considerable distance outwards, and thus a heap of cells is 
formed. hese masses of cells generally enclose a distinct cavity, 
the function of which arrangement may be to protect the egg 
from the effect of pressure due to contortions of the worm’s body. 
In any case we have here a state of affairs very unlike that of 
Kugonodeum. It appears to me to be very possible to make a 
comparison between the egg-sacs of Oochoristica and the cor- 

responding structures in two species of Davainea studied by 
v. Janicki®. Referring, for example, to fig. 9 of his memoir, the 
space lying between the two “ Embryonalhiille” at the poles 
of the elongated embryo might well correspond to the cavity 
which I describe and figure here in Oochoristica. But this com- 
parison, which may be carried a good deal further, is not germane 
to the object of my present communication, which is to compare 
Hugonodeum with other allied genera. 


S General Observations. 


On the whole the characters of the present species ally it most 
nearly to the genus Monopylidium, the alliance being very largely 
due to one species only of that genus, viz. Monopylidiwm rostel- 
latum. The genus Monopylidium is thus defined by Ransom }:— 


(1) Rostellum armed with a double or single crown of hooks. 

(2) A single set of reproductive organs in “each segment. 

(3) Genital pores irregularly alternate, rarely unilateral: 

(4) Genital canals pass between the longitudinal excretory 
vessels and dorsal of the longitudinal nerve, or dorsal of 
both excretory vessels. 

(5) Testicles numerous (20 to 40 or more), behind the femaie 
glands or, also, on both sides of the latter. 

(6) Vas deferens coiled, seminal vesicle absent. 

(7) Uterus breaks down into egg-capsules, each containing one 
or several eggs. 


The worm which forms the subject of the present paper differs 
from Monopylidium in Nos. (1) and (7), and agrees with the 
genus in the other characters set forth. These ditlerences appear 
to be quite enough for generic separation were it not for the 
structure of Monopylidium rostellatum. This species has no 
rostellar hooks, and of the uterus or rather the disposition of the 
ripe eggs Prof. Fuhrmann writes: “ Les oncosphéres se trouvent 
réparties dans tout le parenchyme: chacune delles est entourée 


* “Ueber zwei neue Arten des Genus Davainea,” Arch. de Parasitologie, vi. 1902, 
p. 257. 

+ Bull. U.S. Nat. Mus. No. 69, 1909, p. 76. 

{¢ “ Nouveaux taenias @Oiseaux,”’ Rev. Suisse Zool. xvi. 1908, p. 65. 


AVIAN CESTODES. 877 


par un parenchyme vacuolaire, qui forme autour de Poncospheére 
des cellules hexagonales d’un diametre d’environ 0". 17. Les 
proglottis murs ressemblent ainsi beaucoup a ceux de certains 
Davainea & capsules utérines ne contenant quun ceuf.” The 
difficultyin comparing my genus with this particular Wonopylidium 
is that Fuhrmann says very little of a pre-existing uterus. 

He merely observes that quite early in the body ‘“ Vutérus 
commence déja 4 se former,” and that further back ‘“‘ les segments 
sont remplis Veeufs.” Iam disposed, however, to think that this 
implies a likeness to the more typical species—such as, for example, 
M. unicoronata, which Fuhrmann describes immediately after 
Healing with WV. rostellatum. Moreover, in his résumé of Avian 

lestodes, Furhmann™* defines the genus Monopylidium by, inter 
ae the following character :-—“ Uterus stark verzweigt, lost sich 
in Parenchymkapseln auf,” etc. As I. rostellatum had been already 
described by him it is to be inferred that this species agrees with 
the others, and is, therefore, so far not like the Teeniid which 
forms the subject of my present communication to the Society. 
We may, therefore, I think, exclude my species from the genus 
Monopylidium. Nor does it appear to me that we can refer it 
to any other known species. I propose, therefore, to name and 
characterize a new genus as follows :— 


Eugonodeum, gen. nov. 


Rostellum very muscular, long and retractile, without hooks. 
Suckers unarmed. Ripe proglottids never much longer than broad. 
Dorsal excretory tube much smaller than ventral, lying more or 
less laterally to it ; ventral vessels connected by transverse vessels in 
each proglottid. Longitudinal muscles in two rows of bundles. 
Genital pores unilateral. Genital ducts pass between excretory 
tubes. Testes chiefly behind ovary, and then not very numerous. 
Ovary to pore side, in front of vitelline gland. Genital atriwm 
very deep, with radiating muscles. Cirrus-suc large and muscular, 
lying in front of vagina. Sperm-duct with coil. Cuwrus unarmed. 
Receptaculum seminis present, but not strongly marked. Uterus 
not present. Ova imbedded singly in parenchyma accompanied by 
other cells; round each ovum a cavity is formed later which is 
lined by cells. 

I name as the type Hugonodeum edicnemi, sp. n., with the 
characters of the genus. 


* “Die Cestoden der Vogel,” Zool. Jahrb. Suppl.-Bd. x. 1908, p. 65. 


Proc. Zoou. Soc.—1913, No. LIX. 59 


878 MR. R. W. PALMER ON THE 


50. The Brain and Brain-Case of a Fossil Ungulate of the 
Genus Anoplotherium. By R. W. Paumer, M.Sc. 
(Manchester), Research Fellow in Zoology, University 
College, Reading *. 


[Received May 29, 1913: Read October 28, 1913. ] 


(Text-figures 150-157.) 


INDEX. 
MorruoLoey. Page 
Anoplotherium : Skull—description of ..................... 878 
s Brain— A FY isan aan es auce tetshs) 
SYSTEMATIC. 
Anoplotherium related to Orycteropus ............... 886, 892 


For some time a cranium from the Phosphorites of Quercy, 
together with an exceptionally perfect and well-marked brain- 
cast obtained from it, has been awaiting description in the British 
Museum. The cast was submitted to Prof. G. Elliot Smith, 
F.R.S., of Manchester, who intended to describe it, but he was 
prevented from so doing by the pressure of other work, and very 
generously passed it, together with the notes he had made on it, 
to myself. By the courtesy of Drs. Smith Woodward and C. W. : 
Andrews, of the British Museum, I have been allowed to describe 
both the skull and brain-cast. 

The whole of the facial region of the specimen is missing, and 
since the dentition cf mammals forms such an excellent guide to 
affinities that in original descriptions cranial features are for the 
most part entirely overlooked, the identification of a toothless 
skull such as this is a matter of difficulty. A study of the 
literature of the subject and of such specimens as are available, 
however, leaves little doubt as to the genus of the creature. 

The skull is broken away at the back of the orbits, and the 
paroccipital, zygomatic, and pterygoid processes are incomplete. 
In dorsal view (text-fig. 150) it is seen to be narrow, elongate, and 
somewhat fusiform. Its greatest width in front of the zygo- 
matic processes is 49 mm., but behind the orbits it is greatly 
constricted and its width reduced to less than one half, or 23 mm. 
At the back of the zygomata, not including the lambdoid ridge, 
it measures 40 mm. across. The marked postorbital constriction 
is by no means uncommon in early Tertiary mammals, and 
indicates at once a poor development of the frontal lobes of the 
hemispheres. 

The most striking feature of the dorsal surface of the cranium, 
however, is the great strength of the sagittal and lambdoid crests. 
The former arises in front by the union of the two low postorbital 
ridges on the frontals, and traverses the whole length of the brain- 


.* Communicated by Dr, C. W. Anprews, F.R.S., F.Z.S. 


BRAIN OF A FOSSIL UNGULATE, 879 


ease as a knife-like ridge which rises as high as 16 mm. above the 
general surface. As it nears the occiput the crest somewhat 
thickens and meets the equally highly developed and more massive 
lambdoid crest at right angles. 

The root of the zygomatic process is very extensive and slightly 
convex antero-posteriorly. Its widely concave form in a trans- 
verse direction indicates a temporal arcade standing well out from 


the skull. 


Text-fig. 150. 


Dorsal view of skull of Anoplotheriwm. 


fr., frontal; par., parietal; sq., squamosal. 


In lateral view (text-fig. 151) the great height of the sagittal 
and the true form of the lambdoid crests are displayed. This 
latter strikes off at right angles to the sagittal for about 2 ems., 
slightly dropping in elevation as it does so, then falling very 
sharply downwards, forwards, and slightly inwards, is con- 
tinued as a much less pronounced ridge to the posterior root 
of the zygoma. Here it bifurcates, the anterior branch being an 
extension of the vertical flange of the zygomatic process, while 
the posterior runs down, and slightly back, on to the huge par- 
occipital process, on which it dies out. The aperture of the 
external auditory meatus betweer. the paroccipital and zygomatic 
processes is remarkable in being formed above by a groove in the 
squamosal, while the tympanic forms its floor alone, there being 
no ordinary bony tube. 

In occipital view (text-fig. 152) the foramen magnum is seen to 
be nearly circular and the condyles massive. Above, the lamb- 
doidal crest forms a large shield-shaped area much roughened for 

59* 


880 MR. R. W. PALMER ON THE 


Text-fig. 151. 


SS SS == 


PAS 
il) eam pg 


Lateral view of skull of Anoplotherium. 


ais., alisphenoid; e.a.m., external auditory meatus; j:ov., foramen ovale; gl.s., 
glenoid surface of the squamosal; m,f., mastoid foramen; m.per., mastoid 
portion of periotic ; 0.s., orbitosphenoid; par., parietal; par.p., paroceipital 
process ; p.g,f., post-glenoid foramen; p.g.p-, post-glenoid process; p.orb.p., 
post-orbital process; IT., optic foramen; III.—VL., united sphenoidal fissure 
and foramen rotundum ; AIT., condylar foramen. 


Text-fig. 152. 


=> 


( i poy) 


Occipital view of skull of Anoplotheriwn. 


Lettering as in text-fig. 161, 


BRAIN OF A FOSSIL UNGULATE. 881 


muscular attachments. The paroccipital processes are seen to 
diverge slightly below. 

The palate (text-fig. 153) is very long and narrow. The glenoid 
surfaces of the squamosals are large and flat with strong post- 
glenoil processes. On the left side the tympanic has been 
preserved, while on the right this is conveniently missing, leaving 


Text-fig. 153. 


Palatal view of skull of Anoplotheriwm. 


Lettering as in text-fig. 151, with addition of 6o., basioccipital ; f£, venous foramen ? ; 
flm., foramen lacerum medium; f/.p., foramen lacerum posterius ; per, 
periotic; p.s., presphenoid; sy., squamosal; séin.f., position of stylomastoid 
foramen; é.hy., pomt of attachment of tympano-hyal; f.y., tympanic; wf, 
canals for vessels of the diploé. 


882 MR. R, W. PALMER ON THE 


the lower surface of the periotic exposed. The secondary palate 
is completely lost. 

In a longitudinal section of the skull (text-fig. 154) the great 
thickness of the bones and the general development of diploé are 
striking features. A skull which supported such massive tem- 
poral muscles as this beast possessed had of necessity to be of a 
strong build. The extreme shallowness of the pituitary fossa and 
the practical absence of clinoid processes are features of interest, 
while the mode of attachment of the cribriform mass to a groove 
in the pre-sphenoid is curious. The pear-shaped cerebral fossa is 
partly separated from the cerebellar by the heavy tentorial ridge, 


Text-fig. 154. 


ae Sie 
eS Sis 


Ya mieat 
bo ue fim fou bs vi. bs, 


Median section of skull of Anoplotherium. 


Lettering as in preceding figures, with addition of 6.s., basisphenoid ; er.p., cribri- 
form plate; f.fo., floccular fossa; v.d., canal for vessel of diploé. 


most of which belongs to the squamosal. The spacious cavity for 
the transverse snus opens immediately behind this ridge above 
the periotic. It goes straight down to the post-glenoid foramen 
and is joined by Bioanal for a vein which ran in a deep groove on 
the side-wall of the skull, and was formed by the eonlinenes of 
two vessels which ran in the substance of the alisphenoid. 

The main features of the individual bones of the skull will now 
be outlined. 

The bones of the occipital segment are indistinguishably fused 
together, but their anterior sutures are for the most part distinct. 
The thick supraoccipital is of great extent owing to the extreme 
development of the lambdoid crest, of which it forms the entire 


BRAIN OF A FOSSIL UNGULATE. 883 


upper portion. It articulates with the parietals by a jagged 
transverse suture anterior to this crest. The exoccipitals were 
also large bones, including, as they no doubt did, the large con- 
dyles and practically the whole of the paroccipital processes. 
They take no share in the lambdoid ridge, for their suture with 
the mastoid and squamosal lies just posterior to this ridge. It is 
a Jagged line, starting at the base of the steep drop in the crest, 
and travelling behind it, runs down on to the paroccipital process. 
The basvoccipital is clearly marked off from the basisphenoid by a 
slightly irregular groove. It is very wide behind, between the 
paroccipital processes, and notched by the condylar foramina 
which must pierce the skull between it and the exoccipitals (text- 
fig. 153, p. 881). In front it is narrowed by the lacerate foramina 
and by the gap in which the periotic is set. Its lower surface 
is here heavily marked for muscular attachments. In section, 
the bone is seen to be thin and dense posteriorly, but in front it 
is thicker and more cancellous in structure (text-fig. 154). 

The parietals are remarkably extensive, and externally they 
appear to form the entire cranial roof. Anteriorly, they con- 
siderably overlap the frontals, and the extent of this overlap can 
be seen in section (text-fig. 154), where the junction of the bones 
is indicated by a break in the structure of the roof (at “a” in the 
figure). The parietals do not share in the formation of the lamb- 
doid crest, but meet the supraoccipital anterior to it. There is no 
trace of a distinct interparietal, and the strength of the sagittal 
ridge has necessitated a firm union of the parietals. The suture 
with the frontals is first seen as a jagged line on the postorbital 
ridge. It travels forwards along this for about 12 mm., then 
turns suddenly backward and curves down on the side of the 
skull till it strikes the alisphenoid. Below, the parietal is 
suturally connected with the alisphenoid along a short horizontal 
line, and posteriorly it is considerably narrowed by its bow-shaped 
suture with the squamosal (text-fig. 150, p. 879). Inside the 
skull, the fronto-parietal suture 1s seen as a very irregular line 
running vertically on the wall of the cerebral fossa, and the 
squamoso-parietal suture as an even more irregular line in front 
of the tentorium. Neither suture can be completely traced. 

The sguamosal forms a considerable part of the side-wall of the 
skull. Posteriorly it overgrows the mastoid, so that only a small 
isolated surface of this bone is exposed. The suture with the 
mastoid is anterior to the lambdoid crest, and as this bone gets 
submerged the suture crosses the crest to form the junction with 
the exoccipital. The squamosal sends a small vertical thin flange 
on to the root of the paroccipital process. It is partly anchylosed 
with the tympanic, and in front of the meatus is bored by the 
very large post-glenoid foramen. Almost the whole length of the 
bone is concerned in forming the zygomatic process, which is con- 
cave from side to side above, has a vertical flange to the exterior, 
and a flat glenoid surface below. The flange is roughened on its 
upper edge for attachment of the temporal fascia. ‘T'o the inside 


884 MR. R. W. PALMER ON THE 


of the glenoid fossa, where the squamosal meets the alisphenoid, 
there is a narrow ridge which both bones help to form, and which 
runs parallel to the middle line to separate the glenoid and 
tympanic cavities. Where this ridge meets the post-glenoid 
process there is a depression which may represent a venous 
foramen (text-fig. 153, /,, p. 881). 

Seen from the inside the periotic is quite an extensive bone. 
The double openings of the internal auditory meatus and of the 
aqueductus Fallopii are seen in the centre of the lower triangular 
part. Above this, the floccular fossa (/,fo.) is excavated, and 
from this region the large mastoid portion inclines backwards. 
The mastoid part, as mentioned, 1s exposed on the outside of the 
skull only as a small island between the exoccipital and the 
squamosal, forming the lambdoid crest for about 1-5 ems. Through 
its suture with the exoccipital runs the small mastoid foramen 
(mf.). he inferior surface of the petrosal portion is seen as a 
subovate mass inclined steeply outwards and jammed in between 
the basioccipital and squamosal. It separates the median and 
posterior lacerate foramina. Posteriorly it is pierced by the 
fenestre rotunda and ovalis. 

The tympanic is preserved on the left side. It is partly 
anchylosed to the squamosal, but not to the periotic. It consists 
of a small, very thick, uninflated bulla which does not meet the 
basioccipital, but projects freely under the petrosal. A cylin- 
drical hollow on the bone represents the attachment of the 
tympano-hyal (¢.hy.), and on the right side the position of the 
stylomastoid foramen (sém.f:) is seen to be immediately behind 
this. The rest of the bone forms an irregular mass below the 
meatus and presents no points of interest. 

The alisphenoids ave firmly fused with the basisphenoid, and 
the three bones must be taken together. The basisphenoidal part 
is very elongated —behind, it 1s wide and notched by the median 
lacerate foramina. In front, it narrows very considerably, and 
at the junction with the alisphenoidal part the vertical pterygoid 
wings arise. The alisphenoidal part is a hatchet- or L-shaped 
bone, having a narrow limb confined to the base of the skull 
between the squamosal and the basisphenoid and a square limb 
in front of this, forming part of the side wall of the skull and 
meeting the parietal above and the frontal in front. ‘The long 
limb is deeply grooved for the Kustachian tube, and in its middle 
is perforated by the foramen ovale (fiov.). Anteriorly, below 
the pterygoid flange, the bone is tunnelled by a canal which 
represents the united sphenoidal fissure and foramen rotundum 
(III.-VI.). A couple of venous foramina (v,f.) appear externally 
on the alisphenoid. These were traversed by the vessels which 
united inside the skull to form the large vein already mentioned 
as running into the transverse sinus. On its upper surface the 
alisphenoid is very deeply grooved to shelter the cranial nerves 
lying on it. These grooves are better seen from the cast. They 
will be mentioned later. In section, the basisphenoid is seen to 


BRAIN OF A FOSSIL UNGULATE. 885 


be a thick bone, and it is scarcely thinned by the shallow 
pituitary fossa. 

On the dorsal surface of the skull the frontals form a flat 
arrowhead-shaped area between the post-orbital ridges, Laterally 
they are concave. Below, they unite by suture with the ali- and 
orbito-sphenoids, and they extend well down on the skull since 
the palatines cannot be traced below them. Within the skull 
they form the division between the olfactory and cerebral 
cavities. In front and above they are hollowed out by the 
frontal air-sinuses. 

The orbitosphenoid is quite small. Its suture with the ali- 
sphenoid is not plain, but it appears to completely surround the 
optic foramen. Itis indistinguishably fused with the presphenoid, 
which is a very long and thin bone with a marked median palatal 
ridge. It is very thick, and its upper surface, as seen in section, 
is Shaped like the roof of a house, that is, its surface is very high 
in its middle length and slopes down both forwards and backwards 
(text-fig. 154, p. 882). The posterior slope shares in the tunnel for 
the optic nerve, while the anterior one is excavated by a rounded, 
undercut hollow which gives a firm support to the cribriform 
plate (cr.p.). This last bone i is a solid mass of perforated tissue 
and presents no special points of interest. 

Comparison of the specimen with the figures given by 

Juvier [1] and Blainville [2], as well as with the casts available at 
the British Museum, leaves little doubt but that the skull has been 
correctly identified. In Cuvier’s figures the general form of the 
skull with the postorbital constriction, the form of the condyles, 
the immense paroccipital processes, the peculiar formation of the 
meatus, the post-glenoid process, its foramen, and a host of other 
details are identical. Fig. 1, pl. xlvi. of Cuvier’s 1822 edition 
shows the upper surface of the base of the skull of an Anoplo- 
theriwm determined from the teeth, and the arrangement of the 
foramina, of the grooves for the nerves, and of the transverse 
sinus agree precisely with this specimen. The agreement with 
Blainville’s figures is no less conclusive, and though the crests on 
the skull are considerably stronger than those in any specimen 
figured by these writers, they are of the same form, and their 
degree of development can hardly be of greater than specific 
value when other features agree so markedly. 

Support for the identification also comes from the brain-cast, 
for it agrees essentially with the natural one figured by Blainville 
on plate u., but not with the one given by Cuvier (plate lv., 
1822 edit.), which is incorrectly identified as an Anoplothere. 


Brain-cast. 


The narrow, elongate form of the skull is naturally also a 
feature of the brain. Its total length from the front of the 
olfactory bulbs to the caudal end of the cerebellum is a fraction 
under 10 ems. Of this total, the cerebrum accounts for 5°6 cms., 


886 MR. R. W. PALMER ON THE 


the cerebellum for 3°35 ems., and the olfactory bulbs for the 
remaining | em. The eranial capacity, as measured with sand 
from the cast, is 86 cc. 

The parts of the cerebral hemispheres chiefly concerned with 
the sense of smell, are, as usual in early Tertiary mammals, 
highly developed. This is seen in the large size of the olfactory 
bulbs and of the rest of the “smell-brain,” and in the small size 
of the neopallium. The greatest width of the cerebrum, which 
is near its caudal extremity, is 45 mm. 

The cerebellum is separated from the hemispheres, but there is 
no evidence that the corpora quadrigemina were exposed. Its 
width cannot be measured accurately, owing to the casts of the 
cavities for the transverse sinuses being continuous with its mass 
on each side. Hstimating the extent of these sinuses, its trans- 
verse is not much more than its antero-posterior diameter—that 
is, about 37 mm. 


Text-fig. 155. 


/ 7 
1 
( 
/ 


afb lef’ 


mm finv © xa “Yh lb, 
Lateral view of brain-cast of Anoplotherium. 


fic f., floccular lobe; f-l.m., cast of foramen lacerum medium ; f.l.p., cast of foramen 
lacerum posterius; f-rh., rhinal fissure; o1.b., olfactory bulb; s.coz., coronal 
sulcus; s.orb., orbital sulcus; s.ss/., suprasylvian sulcus; v., vessel; IT. 
IIT-V1., V3, VIL.-VILIL., casts of foramina of cranial nerves. 


Looked at in side view (text-fig. 155), the cerebellum stands as 
high above the axis of the brain as does the cerebrum. This is 
no doubt a primitive feature, for in recent Ungulates the hemi- 
spheres are distinctly more elevated than the cerebellum. The 
Middle Kocene Palwosyops agrees with Anoplotherium in this 
particular [3]. 

On the base of the brain the courses of the cranial nerves and 
the casts of the lacerate foramina are conspicuous, while below 
the floccular lobe on each side the periotic has left a ragged scar. 

The brain, in general form, arrangement of parts, and furrows, 
is remarkably similar to that of Orycterepus. ‘This is seen in side 
view, and especially from below. If the figure of the ventral 


BRAIN OF A FOSSIL UNGULATE. 887 


surface of the brain of the Aard-Vark given by Elliot Smith 
[4] be compared with text-fig. 156, the similarity will be seen to 
be very striking. The detailed resemblances will be noticed below, 
and they undoubtedly help to strengthen the evidence in favour 
of aprimitive Ungulate origin for the Tubulidentata—a view held 


by Elliot Smith, Max Weber, and Loinnberg. 


Text-fig. 156. 


fa! 
4 Mi 
\ 
\\ 4 
\ y 
AN nf 
\\ y Gh 
\W Wy? it / 
iy \ 


Me 


\ 


Ventral view of brain-cast of Anoplotherium. 


Lettering as in text-fig. 155, with addition of m.f., mastoid foramen ; 
ol.tb., olfactory tubercle. 


Ventral Surface of Cerebrum. 


The whole of this region is well developed. This can be clearly 
seen from text-fig. 156, for, seen from below, the pyriform lobes 
almost completely conceal the neopallium—a part of which is 


888 MR. R. W. PALMER ON THE 


visible only to the outside of the anterior parts of the pyriform 
lobes. <Anoplotheriwm, then, was far more highly macrosmatic 
than any living Ungulate, and not much less so than the bur- 
rowing Aard-Vark, 

The olfactory bulbs at their widest part are together 21 mm. 
across. The olfactory peduncles are distinct thick stalks, swelling 
out behind into the anterior ends of the pyriform lobes. 

The olfactory tubercles are well shown as smooth circular 
elevations placed rather further back than in Orycteropus, and 
sufficiently prominent to be noticeable in a side view of the 
cast. 

Between these tubercles the cast shows the course of the optic 
nerves. These tunnelled the base of the skull very near the 
middle line. The position of the chiasma is indicated by the 
sinking of these tubular casts on to the general level of the brain- 
surface. Diverging outwards from the chiasma, the vallecule 
Sylvii are distinct, marking off the globular posterior parts of the 
pyriform lobes. Running back along the lower surface of each 
of these lobes is the cast of the vein described as running on the 
side of the cranium. This vessel does not seem to be cerebral, 
but rather to collect blood from the diploé of the bones. The 
only other feature on the pyriform lobe worthy of note is a short 
straight fissure running along it, above the vessel just mentioned, 
and below and parallel to the rhinal fissure. A similar fissure 
occurs in the pig and the tapir. In text-fig. 155 (p. 886) it 
has been marked with the letter ‘‘ x.” 


Lateral Surface of Cerebrum. 


The olfactory bulbs having been described from below need no 
further mention. 

The neopallium is clearly marked off from the rest of the hemi- 
spheres by the great horizontal rhinal fisswre. In text-fig. 155 
there is a break in this fissure at about one-third of its total length 
from the front. This break does not occur on the other side, and 
seems to be due to damage of the skull-wall. In side view the 
rhinal fissure 1s seen to divide the hemisphere horizontally into 
two equal parts, and the primitive character of the brain is 
emphasized by the relatively small size of the upper part (neo- 
paliium) in comparison with the rest of the cerebrum. Its great 
development in the higher mammals, which led to the degenera- 
tion of the “smell-brain” and to its enwrapping by the 
neopallium, has scarcely begun. 

About 17 mm. from the anterior end of the hemisphere, there 
is a vertical sulcus running into the rhinal fissure below, and 
connected, on the right side only, by a shallow furrow with the 
lateral sulcus. This sulcus can be no other than the orbital *. Its 

* T am using the same terms as were employed in the descriptions of the brain of 
Orycteropus [4], to which I have referred, although Dr. Elliot Smith tells me the 


whole question of nomenclature of cerebral sulci urgently needs revising in the light 
of recent research on anatomical localization of the cerebral cortex. 


BRAIN OF A FOSSIL UNGULATE, 889 


position, relation to the rhinal fissure, and its possible connection 
on the right side to the lateral suleus ave exactly like the con- 
ditions in the brain of Orycteropus. 

Behind the orbital, the side of the hemisphere is marked by a 
clearly defined sulcus, rather under 2 cms. in length, and running 
obliquely from behind forwards and downwards to join the orbital. 
This sulcus is nearly certainly the coronal. Its relations on the 
two sides are similar, but while on the left it is simple, on the 
right it forks slightly at its superior or dorsal end, and in its 
middle length it is connected with avery short horizontal sulcus 
(4), the identity of which is not clear. 

There is no trace of a pseudosylvian sulcus on either side of 
the brain. 

Dorso-laterally the neopalltum is marked by a suleus which, 
starting at the back of the hemisphere, runs straight forward, 
parallel to the middle line, for about 2 cms., and then, curving 
down towards the rhinal fissure, ends on the right side, about 
7 mm. from that fissure, but on the left actually joins it, or very 
nearly so. This must be the suprasylvian sulcus. Though small 
and not so strongly arcuate as in the higher Ungulates, it is of 
the typical Ungulate form in the strength of its posterior branch. 
Near its caudal end the posterior limb of the suprasylvian is 
joined by a very short sulcus (c), which lies external to it. This 
sulcus, though minute, is.constant on both sides. 

Midway between the suprasylvian sulcus and the rhinal fissure, 
and within the curve of the former, there is a short deep hori- 
zontal sulcus about 5 mm. long on the right side, and about twice 
that length on the left. This sulcus is exactly analogous in 
position to a similar short one in Orycteropus. For purposes of 
reference it will be spoken of as sulcus “ d.” 

On the dorsal surface of the neopallium (text-fig. 157, p. 890), 
running in general parallel to, and on an average about 7 mm. from, 
the middle Tine. 4 is a deep sullen, which extends from very near the 
olfactory peduncle to the posterior surface of the hemisphere. 
This long furrow bears several branches and is really compound. 
On the left side of the brain, the most anterior part of it, about 
1 em. long, lies further from the middle line than the rest of the 
furrow, and was probably not directly continuous with it, though 
in the cast there is a connection by means of a shallow depression, 
This short suleus must be the prorean. On the right side it is 
directly continuous with the rest of the sulcus, which is clearly 
the lateral. The relation between the prorean ‘and lateral sulci 
is exactly as in Orycteropus. 

The large lateral sulcus, though roughly parallel to the middle 
line, is by no means straight, and pursues a sinuous course along 
the hemisphere. On the right side it has two very short branches 
going outwards from it, at distances respectively of 1:5 and 
3°3 ems. from its caudal extremity. 

At the junction of the prorean and lateral sulci on each side 
there is another sulcus, a few millimetres in length and running 


890 MR. R. W. PALMER ON THE 


straight forward. What exactly this represents is not clear. 
It may possibly be a rudiment of the crucial. 


Text-fig. 157. 


‘i My ZB Via 
) WV, 

in, 24. 

aN WY os 


ACAN gr 


$I 


Dorsal view of brain-cast of Anoplotherium. 


Lettering as in text-fig. 155, with addition of a.md., area medullaris ; fsp., fissura 
suprapyramis; s.crw.?, crucial sulcus(?); s.la¢., lateral sulcus; sp., supra- 
pyramis ; s.p70., prorean sulcus; wv., uvula; v., v’., vessels. 


At this point a couple of blood-vessels, marked on the dorsal 
surface of the cast, may be mentioned. On the right side, a 
small transverse ridge in the middle of the hemisphere and near 
the middle line may represent the course of one of the meningeal 
vessels or of a branch of the internal carotid artery. Branches 
of this latter artery probably account for some of the obscurity of 


BRAIN OF A FOSSIL UNGULATE. 891 


the brain-surface behind the orbital fissure, where the cast shows 
the marks of the fronto-parietal suture. 

Posteriorly, between the lateral sulcus and the middle line, 
traces of a spacious paired vessel (v’) are seen. ‘This is formed by 
two branches, one of which strikes out from the median line, while 
the other comes inwards and backwards from the surface os the 
brain. The main trunk, from its direction, seems to have gone 
round the tentorium and to have joined the transverse sinus. It 
was probably a vein collecting blood from the superior surface of 
the hemisphere. 


Cerebellum. 


The general proportions of this part of the brain have already 
been given, ‘The cast does not, of course, show the minute folding 
of the cortex characteristic of a fresh specimen, but the main 
areas and the chief fissures are distinctly marked. 

The most striking feature of this organ in Anoplotherium lies 
in the small size of the lateral lobes. This is probably a primitive 
feature, since, in a comparative series, evolution in the cerebellum 
is chiefly marked by the development of these parts [4]. 

The three fundamental divisions of the cerebellum are indicated 
by transverse fissures marked on the vermis. 

The fissura prima is not very clearly shown, but there is a 
suggestion of a groove in the place where it nearly certainly must 
have been situated, on the dorsal part of the anterior face of the 
vermis. 

The fissura secunda is clearly shown on the posterior surface 
of the vermis, marking off from the rest of the cerebellum a 
smooth hemispherical area, which faces posteriorly and is un- 
doubtedly the wvusa—the anterior division of the lobus posticus. 

The whole of the dorsal part of the vermis belongs to the lobus 
medius. It is divided into two almost exactly equal parts by a 
transverse fissure, which can be nothing but the fissura supra- 
pyramis. The vermis of the median lobe shows a twist such as 
is seen in a great number of mammals of different groups, and 
which gives a marked asymmetry to the organ. 

To the sides of the suprapyramis—that part of the vermis 
of the median lobe anterior to the suprapyramidal fissure—lie 
triangular depressed areas indicating the exposure of white 
matter such as is found in many small. mammals including most 
Marsupials, Rodents, Bats, etc. [5]. For this region Elliot Smith 
has suggested the term ‘ area medullaris.” 

To the sides of the pyramis he the indefinite masses of the 
lateral lobes, the small size of which has been commented on above. 
Their irregular form adds greatly to the asymmetry of the whole 
organ. 

_ To the outside of the lateral lobes he the probably relatively 
large jfloccular lobes. ‘These can be traced as being connected 
behind, in the usual manner, to the pyramis. Below, they rest 
on the periotics and fill in the floceular fosse. Their exact extent 


892 MR. R. W. PALMER ON THE 


cannot be determined, owing to the continuity of their casts with 
those of the transverse sinuses. 

Such a primitive form of cerebellum in an organ of this size 1s 
unknown, except in Marsupialia (acropus and Thylacinus) {6}. 


Brain-stem. 


In cranial casts this region is never represented with any 
accuracy, owing to the fact that the nerves and vascular spaces, 
which lie below the brain, obscure the outlines of the latter. 

Conspicuously shown (text-fig. 156, p. 887) to the outside of the 
pituitary region are the courses of the nerves escaping by the 
sphenoidal fissure (which, as mentioned above, is confluent with 
the foramen rotundum) and by the foramen ovale. 

The course of the trigeminal nerve can be followed from the 
outside of the region of the Pons Varolii, along a groove in: the 
alisphenoid to the foramen ovale, through which it gave off its 
mandibular branch, and then, still lying in a groove on the bone, 
the second and first branches ran forward to pierce the skull with 
the third, fourth, and sixth nerves through the sphenoidal fissure. 
Occupying the centre of the figure formed by these two pairs of 
foraminal casts, the pituitary body is represented, but the extreme 
shallowness of its fossa makes it very inconspicuous. The pos- 
terior clinoid processes leave a scarcely noticeable impression on 
the plaster. 

As to the divisions of the rest of the brain-stem nothing can 
be distinguished. It only remains to describe the casts of the 
periotic region, of the foramina for the veins of the sinus, and 
of the last cranial nerves. 

The peviotic bone has left an irregular crater-like scar on the side 
of the cast (text-fig. 155, p. 886). This hollow is walled by various 
nodose elevations, w hich are chiefly casts of parts of the transverse 
sinus. In the Ponte of the “crater” a tiny ridge represents the 
passage through the internal auditory meatus and the Aqueductus 
Fallopii of the ViIlIth and VIIth nerves. The upper boundary 
of the crater is formed by the cast of the floccular fossa. In 
front and below is the cast of the foramen lacerum medium, 
from which a slight ridge runs forward on the base of the brain 
to meet its fellow of the other side below the pituitary fossa. 
This ridge may have some connection with the course of the 
internal carotid. Below and at the back of the periotic depres- 
sion, an elevation represents the foramen lacerum posterius. At 
the back of the floccular lobe of the cerebellum, a tiny knob marks 
the position of the mastoid foramen (text-fig. 156, m.f.). 

On the side of the medulla oblongata, in line with the jugular 
foramen, the exit of the XIIth nerve is represented by a tubercle 
which is the last feature to be seen on the cast. 

At this pot a summary of the resemblances between the 
brains of Orycteropus and Anoplotherium may not be out of 
place. The Aard-Vark’s brain has been described as differing 
from a primitive Ungulate type only in a higher development of 
the olfactory parts. The only other important contrast with the 


BRAIN OF A FOSSIL UNGULATE. 893 


Ungulate brain lies in the insignificance of the suprasylvian 
sulcus compared with the lateral, which is the reverse of the 
usual condition found in that group [4]. 

The brain of Anoplotheriwm has the general proportions of 
that of Orycteropus, and the “smell-brain” is nearly as well 
developed. The horizontal continuous rhinal fissure in the brain 
of the first agrees with the usual condition in the second. ‘The 
relations of the orbital, prorean, and lateral sulci agree in 
both. The suprasylvian suleus of the Anoplothere is much more 
developed than in Orycteropus, but its size in relation to the 
lateral sulcus is intermediate between the usual Ungulate con- 
dition and the state of affairs in Orycteropus. The brain of the 
Aard-Vark has long been known to approach the Ungulate type, 
and in Anoplotherium we have an Ungulate which somewhat 
approaches the Tubulidentate type. 

On the ventral surface, the differences between the two brains 
ave practically all accounted for by the different degrees of macros- 
matism. In the cerebella there are no striking similarities, but 
there is nothing to contradict a relationship. 

If cerebral anatomy be of any systematic value, the view 
that Anoplotherium and Orycteropus arose from a common, 
though necessarily remote, ancestry can hardly be doubted. 

Tn conclusion, | must emphasize my indebtedness to Prof. Elhot 
Smith, whose notes on the brain-cast I have freely used, and who 
has given me a great deal of help in many other ways. My 
thanks are also due to Dr. Andrews and Mr. D. M.S. Watson 
for kindnesses received at the British Museum. 


LITERATURE. 


1. G. Cuvrer.—‘ Recherches sur les ossemens fossiles”... 
Vol. ii. 1822. 

2. H. M. D. pe Buarnvitte.—“ Ostéographie.” 1839-1864. 

3. C. Harte.—‘‘A Memoir upon the Genus Palewosyops Leidy 
and its Allies.” Journ. Acad. Nat. Sci. Philadephia, 
2nd ser. vol. ix. pp. 267-388. 1892. 

4. G. Exiior Smiru.—‘ The Brain in the Kdentata.” Trans. 
Linn. Soc. Lond. (Zool.) vol. vil. pp. 277-394. 1899. 

5. G. Huxtior Suira.—‘ Further Observations on the Natural 
Mode of Subdivision of the Mammalian Cerebellum.” 
Anatomischer Anzeiger, Bd. xxii. p. 369. 1903. 

6. Descriptive and Illustrated Catalogue of the Physiological 
Series of Comparative Anatomy contained in the 
Museum of the Royal College of Surgeons of England. 
Vol. 1. 2nd ed. p. 168. 1902. 


General. 


H. Fitnor.—< Recherches sur les Phosphorites du Quercy.” 
1876-77. 
G. Extior Smiru.— Notes upon the Natural Subdivision of the 
Cerebral Hemisphere.” Journ. Anat, & Phys. vol. xxxv. 
p. 431. 1901. 
Proc. Zoou. Soc.—1913, No. LX. 60 


894 DR. F. A. BATHER ON 


D1. The Fossil Crinoids referred to Hypocrinus Beyrich. 
By Hea. BATEER, MAC aDESc:aheitas heen 


[Received September 8, 1913: Read October 28, 1913. ] 


(Plate XC. and Text-figures 158-160.) 


ConTENTS. 
Page 
PrebanGe sh iil ane Mea GR AG SOR STR ti aR veh aria al nes MESS 4 
Previous Historyscie sania ccm tenta tase ccahe ees EEE 
Suborder Graniocrmonies BRO eC AAPE RO ne aan caahcadnauonccocbes ole) 
Themis CEICAROGOWMICEA sco casocesococo sna vvo acoso mngcaccasacocceose SS 
TEFANAPDUS, CUMENONS, \ so6:ssc 000429003206 000009 095 295 960 a2ebon000 200800 899 
JEANS PUOUD SOP DCOUIP, csc 408 s29008 06406080000 S6H0300G0 can esanoseas GOW) 
“ Hypocrinus”’ pir aS Batiste Ue n cay aren aN onMes MocnnGlighecobeg neko) 
Gener Cidia on OS18/0\7 sora sass cce eee resem en nce seen cee er Rey een eed 
SUMMA YE ose cece shee ese el nea ence eeban tte ohim aac teeleh ae eee eae LS 
Be olenacion of Plate Beene tba aionan socnnaniatoedands uonanusacianons woo ue) 
INDEX. 

Hypocrinus, history of Opimion .....5.....-2....- ences cee eee se) 890 
5 rediagOs dil aa ce tecannan Hea e eRe ERASE EE eRe OO, 
Hypocrinus schneideri vedescribed saan . 900 

‘ Hypocrinus”’ piriformis PidescrinedI and ened Ao the 
Taxocrinide as a new genus, diagnosed but not named ... 905 
@cenocystist GintyAmotiaa Cystidia eee eee eee ee Een eee ere SON] 
Myrtillocrinus Sandb., not a Costexoannidl . SEEN OOD, 
Lecythiocrinus adamsi compared with Flapnoeray WNUS) Bostcoree 899 
Cydonocrinus compared with “ Hypocrinus”’ piriformis ... 911 
Gasterocomide diagnosed and discussed ............2....2.00..--.. 898 

PREFACE. 


Early in 1908 a crinoid cup from the Carboniferous rocks of 
Yorkshire, elsewhere described as C'ydonocrinus parvulus ¥, was 
submitted to me by Mr. D. M.S. Watson. Some curious features 
in it suggested a comparison with Hypocrinus, which had oceupied 
my pen ‘ab intervals duri ing the previous twenty years, but of which 
IT had not seen actual specimens, Without delay all the known 
fossils that had been referred to the genus were borrowed for 
study. ‘This proved a lengthy business, involving patient pre- 
paration and microscopic examination. By the end of 1910, 
however, their description was drawn up and a series of illustra- 
tions prepared, when another specimen was reported as obtained 
from Timor by the Elbert-Sunda Expedition. Correspondence 
ensued with Dr. J. Elbert and Dr. J. Wanner, but the latter 


* Publishea by permission of the Trustees of the British Museum. 
+ F. A. Bather, 1913, “ British Fossil Crinoids, [X.,” Ann. Mag. Nat. Hist. ser. 8, 
vo}. xll. pp. 588-394, 


ID ZSis HON Z), IPL, 2G, 


H. G. Herring. Phot. London Stereoscopic Co. imp. 


1-6. HYPOCRINUS SCHNEIDERI. 7-10. H. PIRIFORMIS. 


THE CRINOID HYPOCRINUS, 895 


was away in Timor, and access to the specimen proved for the 
time impossible. 

My paper, therefore, was laid aside and other work taken up. 
Until the publication of that work in July of this year (1913) I 
could not return to Hypocrinus ; then, just as [ was about to do 
so, I was informed that Professor Wanner wanted to see the 
specimens that were in my hands, because he intended to re- 
describe the species in his account of the Permian Echinoderms 
collected by himself in Timor. Naturally I resumed at once the 
communications with Professor Wanner, but, friendly though 
our exchange of views and of information has been, I regret that 
it has proved impossible for us to come to any arrangement more 
convenient to our colleagues and to future workers than this: 
that I shall publish my discussion of the original specimens, 
and that the description of the new material shall be left to 
Dr. Wanner. 

This paper, then, is admittedly a fragment. References to one 
or two recent papers have been introduced, and my observations 
have in places been checked or corroborated by the information 
courteously sent me for this purpose by Prof. Wanner. But in 
its main lines the paper stands as it was written more than 
three years ago. It will, [ hope, be accepted as Prolegomena to 
Prof. Wanner’s memoir, relieving him of much needful drudgery, 
but not forestalling his more important results. 


Previous History. 


The genus was founded by Beyrich (1862, Zeitschr. deutsch. 
geol. Gesell. Bd. xiv. p. 537) to receive the single species Hypo- 
crinus schneidert, based on a unique cup from the bed of a brook, 
Kali Mati or Ajer Mati (the dead water), about half a kilometre 
south of Kupang in Southern Timor. Beyrich (1865, Phys. 
Abhandl. Akad. Wiss. Berlin, Jahrg. 1864, p. 83), when publishing 
a fuller description, was doubtful whether his new genus was a 
Crinoid or a Cystid, being inclined to the latter view by the 
curious infra-radial position of the anal opening. 

The first author to notice the new genus was Quenstedt, who, 
in his ‘ Handbuch der Petrefaktenkunde’ (ed. 2, 1867, p. 751), 
definitely referred it to the Cystidea, placing it with Hchino- 
encrinites, but separating it from Cryptocrinus. In ‘ Petre- 
faktenkunde Deutschlands’ (1876, Bd. iv. p. 687), Quenstedt 
put Mypocrinus, together with Cryptocrinus, near Hchino- 
encrinites and its allies, although he considered that the probable 
presence of five large arms brought it near to the true Crinoidea. 
In the Atlas to that work, pl. 113, fig. 94, Quenstedt gave a view 
of the posterior side, and another of the base. These views show 
the relation of the basals to the posterior side, and thus indicate 
the position of the small basal, a piece of information not given 
by Beyrich. Unfortunately, Quenstedt’s interpretation of the 
base proves incorrect. In the third edition of the ‘ Handbuch 

60* 


896 DR. F. A. BATHER ON 


der Petrefaktenkunde’ (1885, p. 967) Hypocrinus was still placed 
with Hchinoencrinites and its allies. 

Zittel (1875, ‘Handbuch der Paleozoologie, Bd. i. p. 413), 
who established the Cystidean group Aporitide on the lines 
indicated by Miiller (1854), placed Hypocrinus therein next to 
Cryptocrinus. KR. Hoernes (1884, Elem. d. Pal. pp. 123, 124), 
remarking that both genera possessed pores, especially Hypo- 
crinus, transferred them to the Caryocrinide. Hypocrinus was 
also associated with Cryptocrinus by M. Neumayr (1889, ‘ Die 
Stiimme des Thierreiches, p. 403), who, however, made no 
particular remarks on the genus. 

Hypocrinus was not mentioned by Wachsmuth and Springer 
in their ‘Revision of the Paleocrinoidea,’ although the name 
appears within round brackets in their privately issued index. 

Before 1890 no one had placed the genus anywhere except 
with the Cystidea ; in that year, however, in the second of my 
papers on “ British Fossil Crimoids” (Ann. Mag. Nat. Hist. 
ser. 6, vol. v. p. 324), [ associated it with Achradocrinus, stating 
that it appeared to differ from that genus only in having three 
infrabasals instead of five. In the concluding part of the same 
paper (tom. cit. p. 885) Achradocrinus and Hypocrinus formed 
Series 3, Achradocrinites, in the family Cyathocrinide, Series 2 
being Codiacrinites, containing Codiacrinus and Lecythiocrinus. 

The view that Hypocrinus was a crinoid was accepted by 
P. Herbert Carpenter (1891, ‘On certain Points in the Mor- 
phology of the Cystidea,” Journ. Linn. Soc., Zool. vol. xxiv. 
pp. 14-16). “ Hypocrinus,” he said, “is certainly a very singular 
form and one would like to know more about it.” Carpenter 
considered it to be allied to Lecythiocrinus White, of the Coal 
Measures of Central North America, but also mentioned its 
affinity to Codiacrinus. 

Our knowledge of the genus was increased through two cups 
collected by Dr. A. Wichmann in 1888-9 at the same locality 
of Ajer Mati, and described by Professor A. Rothpletz (1892, 
‘ Paleeontographica,’ Bd. xxxix. p. 74) under the names Hypo- 
crinus milleri Beyr. (a lapsus calami, as Prof. Rothpletz tells me, 
for H. schneidert) and Hypocrinus(?) pyriformis (melius pari- 
formis), n. sp. Since Dr. Rothpletz stated that Crinoidea were 
represented in the Wichmann Collection by columnars alone, and 
since between their description and that of Hypocrinus he placed 
the account of an echinoid radiole, he must be supposed to have 
regarded Hypocrinus as a Cystid. 

Whether the views of Herbert Carpenter and myself were 
unknown to foreign writers or whether we were merely thought 
unworthy of attention, subsequent authorities continued to refer 
Hypocrinus to the Cystidea, associating it as before with Crypto- 
crinus. Among text-books one notes Félix Bernard (1893, Elém. 
de Paléont. p. 159), Von Zittel (1895, ‘Grundziige der Pala- 
ontologie,’ p. 155), and Koken (1896, ‘ Die Leitfossilien,’ p. 290).* 
In 1895, Dr. O. Jaekel was describing from the Devonian 


THE CRINOID HYPOCRINUS. 897 


of Germany several crinoids which I regard as close allies of 
Hypocrinus, but he did not mention that genus for comparison. 
Haeckel (1896, ‘‘ Die Amphorideen und Cystoideen,” Festschr. 
fiir Gegenbaur, Bd. i. p. 147) dealt with the genus between Crypto- 
crinus and Lichenoer ystis, but concluded his ‘deser iption by saying : 
“wegen ihrer sonstigen nahen Beziehungen zu _ einfachsten 
Crinoideen kénnte man sie auch fur reduzirte oder verkummerte 
Formen dieser Klasse ansehen.” This statement, while perfectly 
true as regards Hypocrinus, is not equally applicable to Crypto- 
crinus. 

Undismayed by this weight of authority, in a “ Phylogenetic 
Classification of the Pelmatozoa ” (1899, Rep. Brit. Assoc. 1898, 
p- 923) I placed G'asterocoma, Seaeeninie. Achradocrinus, aad 
Hypocrinus in the family Gasterocomide, which came in the 
suborder Cyathocrinoidea, among the Dicyclica Inadunata. 
This family was maintained, with the addition of Manocrinus, 
on pp. 177, 178 of my contribution to Lankester’s ‘ Treatise on 
Zoology’ (Part III. Echinoderma, 1900). 

In the second edition of the ‘Grundziige der Paliontologie’ 
(1903, p. 141) Von Zittel included in the family Gasterocomidee 
the same genera as I had placed in it. His example was followed 


by Potta (1904, ‘ Rukovet Palaeozoologie,’ I.), who, however, 
quoted Hypocrinus as “ peemelserbon rusky.” 

Delage and Heérouard (1904, ‘Traité de Zoologie concrete,’ 
Tome iil. pp. 379, 380) reduced my suborder Cyathocrinoidea to 
the rank of a family Cyathocrinusine, but retained Hypocrinus 
and the other genera in the same relative position. 

The latest writer to maintain Hypocrinus in the Cystids under 
a family Cr yptocrinids appears to be Dr. G. H. Girty (1908, 
“Guadalupian Fauna,” Professional Paper 58, U.S. Geol. Surv. 

p. 108). This he has done in connection with ‘the description of 
Coenocystis richardsoni, an alleged new genus and species of this 
Family, from the Delaware Mountain Formation of uppermost 
Carboniferous age in Texas. Whatever Coenocystis may be, I 
find no reason for supposing it to be a Cystid. But even if it 
were, this would not affect the position of Hypocrinus, which, as 
Dr. Girty says, 1s evidently distinct. 

The preceding account shows that the position now occupied by 
Hypocrinus in our leading text-books depends chiefly upon the 
opinions expressed by Herbert Carpenter and myself. It is, 
therefore, advisable to point out that, when we wrote, neither 
of us had examined any specimens of Hypocrinus. For us it 
remained, as Carpenter expressed 1t, a very singular form which 
one would like to know more about. At last, the great courtesy 
and kindness of Professors Dr. W. Branca of the Museum fiir 
Naturkunde, Berlin, and Dr. C. K. A. Wichmann of Utrecht 
University, have enabled me to study all the described specimens 
of this genus. The following pages contain redescriptions of that 
material and fresh diagnoses. 


898 DR. F. A, BATHER ON 


CRINOIDEA DICYCLICA INADUNATA. 
Suborder CY ATHOCRINOIDEA. 


Dr. Frank Springer, in his paper on ‘“ Some new American 
Fossil Crinoids” (July 1911, Mem. Mus. Comp. Zool. Harvard, 
vol. xxv. no. 3), has published a very interesting discussion of 
my Suborders Cyathocrinoidea and Dendrocrinoidea. While 
differing as to the delimitation of these divisions and the 
principles upon which they are based, he readily agrees in 
recognizing their existence. He prefers to regard them only as 
large Families, to which he applies the names Cyathocrinide and 
Poteriocrinidee, but admits that this is ‘“‘a matter of detail, 
depending upon the general plan of treatment.” It was necessary 
for me to treat them either as Suborders or Superfamilies, because 
J split them up into Families, several of which are again divided 
into Subfamilies. Iam, therefore, emboldened to retain for the 
present the Suborder Cyathocrinoidea. 


Family Gasterocomide. 


Cyathocrinoidea with anal opening in the side of the dorsal 
cup, below the level of the arm-bases. Radial facets of horse- 
shoe shape, with distinct axial canal. Infrabasals small, often 
fused into three plates or one. Orals, so far as known, for the 
most part concealed by the cover-plates of the subvective system ; 
posterior oral a large madreporite. Stem frequently with peri- 
pheral canals. 

Dr. Springer, in the above-quoted memoir (July 1911, p. 122), 
has given a full account of the Gasterocomide, although he has not 
provided any fresh diagnosis. The characters which he mentions 
as distinetive are included in the above diagnosis, which differs 
only verbally from that published by me in Lankester’s ‘ Treatise 
on Zoology’ (Part 11. 1900, p. 177). Dr. Springer further agrees 
with Jaekel and with me in referring to the Family the genera 
Gasterocoma, Nanocrinus, Scoliocrinus, and Achradocrinus. To 
these headds a new genus, Schultzicrinus, found in the Onondaga 
Group of the Middle Devonian in Livingston County, New York. 
This genus has the infrabasals coalesced and five simple uniserial 
arms. Further, Dr. Springer adds Arachnocrinus Meek and 
Worthen, of which he has made a complete study, showing that 
our previous views as to the structure, and therefore as to its 
systematic position, were erroneous. ‘This genus differs from 
Schultzicrimus mainly in having dichotomous arms. I am, of 
course, prepared to accept its reference to the Gasterocomide. 

Apparently, Dr. Springer would also place in the Gasterocomide 
the genus Myrtillocrinus Sandberger, of which he considers 7’77- 
pleurocrinus HK. Wood to be a synonym. In this he follows 
Wachsmuth and Springer (‘ Revision of the Paleeocrinoidea,’ 1885), 
as well as some other writers therein quoted. It may, however, 


THE CRINOID HYPOCRINUS. 899 


be pointed out that the genotype, Wyrtillocrinus elongatus Sand- 
berger, is not very well known, and that Dr. Springer here bases 
most of his argument on J/. americanus Hall, which possibly has 
not the same structure. If, however, we accept the generic 
identity of the two species, then it follows that Myrtillocrinus is 
devoid of the most characteristic feature of the Gasterocomide, 
namely, the passage of the anus through the dorsal cup. It 
seems to me, therefore, as it did to Prof. O. Jaekel (1895, Pal. 
Abhandl. Bd. vii. p. 75), that the Family Gasterocomide is far 
more homogeneous if JMyrtillocrinus be removed from it. 
Whether I was right in placing it with the Cupressocrinide is 
another matter, not worth discussion until Sandberger’s type- 
specimen has been restudied. 

Dr. Springer, on p. 138, somewhat casually mentions Hypo- 
crinus, but leaves its systematic position uncertain. On the 
grounds of its structure Hypocrinus is naturally referred to this 
Family, and there does not appear to be any objection to that 
course, except the fact that all the other genera of the Family are 
restricted to the Middle Devonian, whereas Hypocrinws is not 
known below the top of the Carboniferous. This difficulty is 
scarcely lessened if the species Lecythiocrinus adamsi Worthen be 
proved to belong to Hypocrinus, since that species is similarly 
removed by a large interval of time from the rest of the Gastero- 
comidee, being found about the horizon of Coal No. 8 of the Lower 
Coal Measures in Peoria County, Illinois. 

According to Worthen’s description and figure of Lecythio- 
crinus adamsi (1882, Bull. Hlinois State Mus. Nat. Hist. no. 1, 
p- 87; and 1883, Geol. Surv. Illinois, vol. vil. p. 317), that species 
has five infrabasals, and therefore could not be referred to Hypo- 
crinus. This fact, however, would not prevent its being placed 
with the Gasterocomide, since Achradocrinus also has five infra- 
basals. The reason for placing this species with the Gastero- 
comide is that ‘Just above the summit of one of the subradials 
basals], and in the lateral angles of two of the radial plates, there 
appears to have been a circular opening, rather larger than the 
arm-facets, which is now filled with stony matter”(Worthen). This 
no doubt is an anal opening, as suggested by Worthen himself. 

Although, for reasons of structure, Lecythiocrinus adamsi and 
the species hitherto referred to Hypocrinus ave naturally placed 
in the Gasterocomide, it should not be forgotten that this peculiar 
disposition of the anus may have arisen independently in the late 
Carboniferous Cyathocrinoidea. In that case, the forms in 
question, wherever they might be referred, would not belong to 
the Gasterocomide. 


HYPocrinus. 


1862. Hypocrinus H. KE. Beyrich, Zeitschr. deutsch. geol. 
Gesell. Bd. xiv. p. 537. 

For other references, see ‘‘ Previous History.” 

Diagnosis.— A Gasterocomid with anal opening at the summit 


900 DR. F. A. BATHER ON 


of the posterior basal, and at the lower angles of the right and 
left posterior radials, which meet above. Two pairs of infrabasals 
fused ; the remaining small infrabasal being the anterior. Radial 
facets small. (Arms unknown.) Stem-facet from +13 to -2 
diameter of cup. (Stem unknown.) 

The diagnosis is based on the characters of the genotype, since 
H., piriformis must be removed from the genus. 'The somewhat 
hexagonal shape of the anal opening suggests that it was closed 
by six valvular plates. 

Genotype.—ZH. schneidert. 

Distribution.—This species, as well as “ Hypocrinus” piri- 
formis, has been found only in Timor, in beds which are either 
Permian or Carvoniferous. Recent workers have regarded them 
as Permian, but the Echinoderms at present described would 
rather favour a Carboniferous age. JI have already discussed 
this in my account of Schizoblastus from Timor (1908, N. Jahrb. 
f. Mineral., Beil.-Bd. xxv. p. 318). The affinities of Hypocrinus 
also, as shown above, are closer to the Upper Carboniferous 
‘“« Lecythiocrinus” adamsi, 1f not to the Devonian Gasterocomide, 
than they are to anything as yet known from undoubted Permian 
rocks. It is of no avail to discuss the question further in this 
place, for the abundant materials collected by Dr. J. Wanner 
and others will probably settle it when they have been described. 
I would only suggest that the possibility of some of the fossils 
being derived from older beds should be borne in mind. Some 
of the few Echinoderms that I have studied had certainly been 
much worn before they were included in their present matrix. 


HYPocRINUS SCHNEIDERI. (Pl. XC. figs. 1-6 ; text-figs. 158, 159.) 

1862. Hypocrinus schneideri H. K. Beyrich. 

1865. Hypocrinus schneidert Beyr.; H. E. Beyrich. 

1892. Hypocrinus millert [err. pro schneideri| A. Rothpletz. 

For references, see ‘“‘ Previous History.” 

Diagnosis.—This being the only undoubted species of Hypo- 
crinus known to me, no specific diagnosis can be given. 

Holotype.—The original of Beyrich’s figures (1565) preserved 
in Museum fiir Naturkunde, Berlin (our Pl. XC. figs. 1-4). 

The only other specimen described is that of Rothpletz (1892). 

Locality.— Both specimens come from Ajer Mati, 8S. of Kupang, 
Southern Timor, 

Horizon.— Permian 2 (vide supra). 

Description of Holotype.—A dorsal eup, with no trace of stem, 
anal plates, brachials, or tegmen. Surface of stereom stained 
pinkish. Infilling matrix, soft micro-erystalline calcite with 
small dark specks. 

General shape squat, pyriform. Cross-section subpentagonal, 
owing to projection of umbones of basals, which is especially 
marked in the posterior basal. Height 20°8 mm.; greatest 
sagittal diameter 20°38 mm., greatest transverse diameter 
19°9 mm., these being at about 14°5 mm, from basal plane. 


THE CRINOID HYPOCRINUS. 901 


Stem-facet (Pl. XC. fig. 4) circular, markedly concave, bounded 
by a slight rounded rim; external diameter 2°7 mm.; internal 
diameter 1:75 mm. Lumen circular, about -3 mm. diameter. 
There are exceedingly faint traces of about 15 or 16 radiating 
grooves. 

Infrabasals (Pl. XC. fig. 4) closely united. The posterior 
suture is most distinct, chiefly because a’ crack is partially co- 
incident with it. When the specimen reached me the right 
anterior suture could scarcely be detected, only being indicated 
by a faint band of lighter colour. Various processes of prepa- 
ration have now rendered it perfectly visible. The left anterior 
suture at first seemed conspicuous, but the appearance was due to 
a vather irregular crack continuous with that neticed along the 
posterior suture. The left anterior suture itself could not be 
traced with certainty; it now appears to be only in part co- 
incident with the crack. In the concavity of the base, after 
careful preparation, there are dimly seen two dark streaks 
running from the lumen in the direction of the posterior and 
right anterior sutures respectively. There appears to be a third 
such streak a little on the anterior side of the above-mentioned 
erack ; and this streak probably indicates the position of the left 
anterior suture. 

Tt follows from the preceding account that Beyrich’s description 
and figures of the base,.as composed of two large plates and one 
small plate, though unsupported by valid evidence, were in the 
main correct. His figures are little more than inexact diagrams, 
and neither they nor the text indicate the position of the small 
infrabasal ; it is anterior (text-fig. 158). 


Text-fig. 158. 


RR GS ce (en) e @ 


Hypocrinus schneideri. 


IBB 


Analysis of the cup of the holotype. 


The periproctal outline is dotted in. Nat. size. 


Heights of IBB :—anterior 8-7 mm. ; right large 1B 7-3 mm. 
along median line, 9°3 to posterior angle; length of posterior 
suture 7°3 mm. 

Basals bounded by fairly distinct sutures, emphasized by the 


902, DR. F. A. BATHER ON 


bands of lighter colour that accompany them (Pl. XC. fig. 3). 
The measurements in millimetres are :— 


ey ere ile gy maa. 
posta eee ee LOS alee lais 118 13°7 
THs DOS 1B coe cee eepecn ae A TOG ee ates | 5G |) Rs 
HPs, BUI: 183i ganado eos eeeonsane 8:7 | Tale) 9°3 | 17 
Rar eB edhe seen Viieccvel S774 il weelO; Olea ei | | 518hs 
Ost Neen 9°2 | 11:7 | 103 | ca. 139 


The greatest width of each basal is on a level with its umbo, 
at a little more than half its height. The umbo is most 
prominent in the posterior basal, which is also the widest of the 
basals and would be the highest were it not truncated by the 
periproct. 

Radials sloping from top of basals to the peristome rather 
suddenly. Height of each from the lower angle to the inner 
margin of the peristome about 8 mm. Width of each below 
about 8°3 mm.; width above about 2 mm. 

The upper margins of the radials form a distinctly pentagonal 
border to the dorsal cup (Pl. XC. figs. 1, 2). Each side of the 
pentagon forms a smooth slightly raised rim, and is about 2°3 mm. 
long. At each angle of the pentagon les a brachial facet. 

Facets rather worn, so that their precise constitution cannot 
be detected. A ventral groove leads downwards into the cavity 
of the cup. From this to the outer edge of the facet is about 
15mm. Width of facet about 13mm. Neither muscle-fossve 
nor an axial nerve-canal can be detected. 

Within the pentagonal rim, the edges of the radials slope 
steeply downwards into the cavity of the cup (cf Pl. XC. fig. 5). 
The downward extension appears to be greater near the facets 
than near the interradial sutures, and may have served for the 
attachment of muscles. There is no trace of any tegminal or 
oral plates, or of any sutural surface on which they might have 
abutted. This fact and the contraction of the cup at its upper 
end lead one to regard the pentagonal opening as a peristome, 
covered in life only by oral folds of a flexible and thinly-plated 
or naked integument. Below the left posterior facet, in the 
peristomial cavity, 1s a fragment of a plate, but its nature is not 
clear. 

The Periproct (Pl. XC. fig. 1) truncates the uppermost angle 
of the posterior basal with an obtuse re-entrant angle, 2°8 mm. 
across, and cuts into the adjacent lower angles of the right and 
left posterior radials with a curve of approximately the same 
chord (2°8 mm.), also somewhat angular, so that the outline 
is approximately an irregular hexagon nearly 3 mm. high. Of 


THE CRINOID HYPOCRINUS. | 903 


anal plates no trace remains, but the relatively large size of the 
opening suggests that some probably existed. 

Oimament:—Beyrich says ‘‘Sammtliche Platten sind aussen 
glatt und scheinen zum Theil von pordsem Gefiige zu sein.” 
Rothpletz remarks that his smaller specimen ‘ die pordse Structur 
der Platten, welche Beyrich an seinen stark abgeschliffenen 
Exemplare nur vermuthen konnte, sehr deutlich zeigt.” As a 
matter of fact, the holotype is only ‘“ strongly worn down” in one 
or two places, and it shows the texture of the plates (Pl. XC. 
fig. 6) no less clearly than the specimen described by Rothpletz. 
The latter author continues :—‘‘ Die Porositiit der Platten wird 
evkannt an den abwechselnden kleineren und grésseren rundlichen 
Vertiefungen, welche die Oberfliche bedecken. Die kleinen 
Poren stehen regelmissig und dicht gedrangt, die grosseren 
liegen vereinzelt dazwischen und hiufen sich auf den Radialia 
nahe den Anzatzstellen und auf den Parabasalia in der Gegend 
der Buckel.” This account applies equally well to the holotype ; 
but I must add that in neither specimen can I detect any 
regularity of arrangement in the smaller pores. 

The diameter of the smaller pores is about ‘075 mm. or more. 
The diameter of the larger pores is*3 min. or less. There is 
every gradation between these sizes. 

This micro-structure of the test is so unusual, if not unique, in 
its absence of regularity and in the sporadic occurrence of larger 
pores, that Beyrich’s use of the word ‘“ scheinen” may have been 
due toa hesitation as to the true nature of the appearance. ‘The 
irregular surface, especially in the umbonal region of a basal, 
markedly resembles some encrusting hydrozoon with autopores 
and dactylopores. Were the appearances really due to some such 
encrustation, this would not have been the only occasion on 
which an echinoderm plate has been described as porous on 
similar evidence. But in the present instance, where the rough 
surface has been worn down, the same porous structure is still 
seen penetrating the plates. The pores are rendered distinct by 
the infiltration of a reddish iron colouring. 

Since the matrix itself is of a granular micro-crystalline com- 
position, with darker specks, it is very difficult to discriminate 
between the matrix and the rough surface of the plates. Beyrich 
may have been uncertain whether he was looking at the actual 
stereom or at an imperfectly cleaned surface. The porous 
substance is of a saccharoidal appearance, and is easily scraped or 
broken with a needle, even with a brass pin. It does not cleave 
with a smooth fracture, as does the denser stereom near the 
sutures. 

This apparent porosity of the test doubtless weighed with 
those authors who referred Hypocrinus to the Cystidea, but 
the structure is really nothing but an exaggeration of that seen 
in the cup-plates of the larval Antedon before the connective 
strands of stroma have impressed on the stereom a radiate or 
fasciate arrangement. The exaggeration consists partly in the 
greater size of the pores, which, as figured by W. B. Carpenter 


904 DR. F. A. BATHER ON 


in his classical memoir (1866, Phil. Trans. B, vol. 156, pl. 41), 
have, in the * eribriform films” of Antedon, a maximum diameter 
of ‘02 mm., as contrasted with -38 mm. in Hypocrinus ; partly in 
the greater relative thickness of the plates. But, so far as I 
can make out, in a place where the stereom was already worn, 
the inner layer, or hypostereom, was much denser, so that the 
pores nowhere pass right through the test. 

Description of the Specimen figured by Rothpletz. (Pl. XC. 
fig. 5 & text-fig. 159.)—In its main features this agrees with the 
holotype, and only the points of difference need here be 
mentioned. i 


Text-fig. 159. 


a Os OL eae 


Oe es 
es eo ae 


Hypocrinus schneideri. 


Analysis of the cup in the specimen figured by Rothpletz. 
The periproctal outline is dotted in. Nat. size. 


Height 13°5 mm. Posterior basal not so prominent, partly, 
perhaps, because worn down. Greatest sagittal diameter 
13°2mm. Greatest transverse diameter 13°3 mm. These both 
at about 8 mm. from basal plane. 

Stem-facet obscure, but appears irregularly triangular, the 
angles directed towards the sutures between the infrabasals. 
Greatest external diameter (in direction of |. ant. suture) circa 
27mm. No trace of radiating grooves. 

Infrabasals. The left larger one is considerably cracked, and 
some of the cracks in part coincide with the sutures ; otherwise 
all the sutures are very hard to trace. Heights of IBB: 
anterior 6°6 mm.; right large IB 5:1 mm. along median line, 
5°8 mm. to posterior angle; length of posterior suture 4°7 mm. 

Measurements of Basals in millimetres :— 


Nar Re ere eee) ete 
POSHWB had. welee ene 66 70 6°8 84 
if OS tal} eee ee aes | 57 7-0 61 86 
TMT ber kan egeeone eee ee 61 85 74 90 
OEE Vii BHM Ge atekce Seccauae 66 71 67 87 
[PE OSES FB ascent oc conesnns 61 7 6:1 7-8 


THE GRINOID HYPOCRINUS, 905 


Tt will be noticed that the width is here on the anterior, rather 
than on the posterior, side. 

The Radials have a height of circa 5-4 mm.,a width below of 
6-6 (1. ant.), 6°3 (ant.), 5-7 (1. ant.), and a width above of circa 
1-7 mm. 

The Facets (Pl. XC. fig. 5), with their ventral grooves, are 
rather clearer than in the holotype, but still no muscle-scars can be 
detected. From the cavity of the cup to the outer edge of each 
facet is circa 1 mm., and this is also the width of each facet. 

The Periproct has a rather more regular shape than in the 
holotype, and its upper angle is slightly produced adorally. The 
height of the opening is 2°4 mm., the width 2 mm. 


‘“‘ Hypocrinus” prrirormis. (Pl. XC. figs. 7-10; text-fig. 160.) 


1892. Hypocrinus (?) pyriformis A. Rothpletz, 1892, p. 75, 
jOlls 3x6 ths Aa). 


As will appear from the description, this species must be placed 
in anew genus, So long as the imperfect holotype was the only 
available specimen, the curious features of its structure might be 
interpreted as abnormalities, and the proposal of a fresh name 
was scarcely justified. The material obtained by Dr, Wanner 
and the facts which he has very kindly communicated to me 
render a more precise interpretation possible and necessitate a 
new genus, - That Dr. Wanner may not be deprived of the 
fruits of his own discoveries, I leave to him the privilege of giving 
it a name and of completing my imperfect descr iption. 

Holotype.—The unique specimen figured by Rothpletz and 
preserved in the Mineralogisch- Geologisch Instituut te Utrecht. 

Locality.—Ajer Mati, S. of Kupang, Southern Timor. Dr. J. 
Wanner has recorded this [or possibly an allied] species from 
Fatu Bitaoni (or Bitatnu) in Insana district, Timor (Centrlbl. f. 
Mineral. 1910, p. 737). 

Description of Holotype.—A dorsal cup with no trace of stem, 
anal plates, or tegmen. Stained pinkish. Matrix fragmentary 
organic limestone with crystalline cement. General shape 
elongate- -pyriform ; somewhat irregular owing to peculiarities in 
the radial circlet ; with a tendency - to be flattened along the per- 
radial meridians, or, conversely, to be broadly ridged ‘along the 
interradial anemia, from the top of the basals downwards. 
This produces a sub-pentagonal cross-section. Height circa 
43 mm. Greatest sagittal diameter 28 mm. Greatest trans- 
verse diameter 31°50 mm, ‘These diameters are measured by 
callipers with the jaws parallel to the main axis of the cup; 
but the diameters are not at right angles to the main axis. 
On the left-hand side the greatest projection is 26°7 mm. 
above the basal plane, on the right-hand side it is 24°5 mm. 

The Stem-facet was broken away, and the proximal end of the 
cup was therefore slightly ground down by Dr. Rothpletz (Pl. XC. 
fig. 7), who says that it Lisst in der Mitte einen Centralcanal 
erkennen, von welchem die drei Niihte der Basalia [7, e. IBB| 


906 DR. F. A. BATHER ON 


ausstrahlen. Nahe der Peripherie liegen fiinf rundliche Poren 
in den letzteren, von denen eine auf das kleinere, je zwei auf 
die grdsseren [Infra-|Basalia fallen. Dies diifte fir die 
urspriingliche Anlage von fiinf [Infra-|Basalia sprechen, von 
denen je zwel nachtriiglich erst verwachsen sind.” The cross- 
section of the worn end is not regularly circular: the sagittal 
diameter is 7°4 mm., the transverse diameter 6°6 mm. The 
diameter of the stem-facet may therefore be taken as about -25 
that of the cup. The supposed * Centralcanal” is of irregular 
outline and is far from ‘“‘in der Mitte,” since its centre lies at 
2-7 mm. from the posterior side and 4:7 from the anterior side. 
Proceeding from this irregular area are three cracks, which, 
however, do not appear to reach the periphery precisely at the 
points where it is cut by the sutures between the three infra- 
basals. Under careful lighting one can detect traces of sutures 
continuing those on the posterior side, and they seem directed 
towards a point more strictly central than the supposed axial 
eanal. The “ fiinf rundliche Poren,” which represent the axial 
nerve-canals, are not quite so obvious as might be supposed. 
They are irregular in shape, size, and position, and that in the 
small infrabasal lies at the outer edge of a long depression, 
which seems to have been regarded by the draughtsman of 
Dr. Rothpletz (Taf. x. f. 236) as formed of two pores. That 
figure has the small infrabasal towards the observer. 

Infrabasals closely united. By removing a little more of the 
firm adherent matrix, I have been able to make the sutures 
clearer, so that the proof of their position no longer rests on the 
doubtful appearances of the proximal end. The axis of the small 
infrabasal is in a line with the periproct. Heights of IBB:— 
small IB 15 mm.; right large IB 14°4 mm. along median line, 
15-1 to upper angle; length of suture between the large IBB 
12°3 mm. The infrabasal cirelet seems to bear indications of 
growth-lines parallel with its upper margin and crossing the 
vertical sutures (Pl. XC. fig. 10). 

Basals (Pl. XC. figs. 8, 10) bounded by distinct sutures. 
While all rest symmetrically on the infrabasals, none is quite 
symmetrical above. The lack of symmetry corresponds with 
abnormalities in the radials, and will be described in connection 
therewith. The measurements in millimetres are :—— 


Width Greatest Width Hecht | 
below. width. above. ea 
ey OCK\ine Bip peenpaseelsoanoc scans 12:7 171 116 26°8 
Ye POstoUBin.  daencseeaesecees 13°7 168 iLleil ca. 25°6 | 
Pian B ie, ee ceed myet2so8 lull 70 ca. 120 248 | 
lant. B i ee 145 | 176 ca. 11-0 22°7 + | 
L.gpost Bb vx. yeusveners 12:0 16°6 12°5 25°3 


THE CRINOID HYPOGRINUS. 907 


The greatest width of each basal is at about two-thirds of its 
height, and here also the sides slope equably to a prominent 
umbo, above which the plates bend towards the oral pole, the 
bend being more sudden on the left than on the right. 

The Radials (Pl. XC. fig. 9 ; text-fig. 160) continue the slope of 
the upper part of the basals. They present several abnormalities 
due to growth, and are further obscured by crushing. Only four 
were distinguished by Dr. Rothpletz, but I believe I can see 
traces of a fifth, though much atrophied [and this is fully con- 
firmed by Dr. Wanner in litt.|. Ant. R and 1. ant. R were 


Text-fig. 160. 


RR 


BB 


IBB 


“ Hypocrinus” piriformis. 


Analysis of the cup of the holotype. 


The sutures obscured by wear or fracture are represented in dotted line, 
as is also the outline of the periproct. 


v.p.R denotes the minute plates, of rather uncertain number and shape, one or all of 
which probably represent this radial. On l.p.R the outline of the cleaved 
surface is represented. Tbe outline of a.R is modified by a slight shear. 
Nat. size. 


apparently of less modified outline than the others, though in 
this specimen weathering and crushing have rendered that out- 
line incomplete. The l. ant. R rests on |. ant. B by a longer 
side than that which rests on |. post. B. Similarly, the side of 
ant. R, resting on the r. ant. B seems to have been longer than 
that on |. ant. B. Ant. R is broken, and part of it, still united 
to l. ant. R has been pushed inwards and under the remaining 
portion. Thus it is difficult to compare the interradial sutures 
bounding these two radials, but there is nothing to show that 
they were not of equal length. The upper margin of each of 
these radials was excavated by a concave curve, and the edge 


908 DR. F. A. BATHER ON’ 


was slightly bevelled so as to give this the appearance of a very 
obtuse angle, with its apex about in the middle line of the radial. 
This appears to represent a greatly simplified brachial facet, but 
there are no articular markings, unless it be a slight depression 
near the middle. 

The r. ant. R was similar in its genera] outline to the ant. and 
l. ant. RR, having, like them, the basiradial suture longer on the 
posterior than on the anterior side, It differs, however, in that 
its left or posterior side is divided into two tracts: one a very 
short suture connecting the upper angle of the plate with what 
seems to be part of the r. post. R; the other an excavate margin 
to the periproct. The curve of this margin throws the left upper 
portion of the radial away from the median line, so that the sub- 
angular concavity of the facet is longer on the left than on the 
right. 

The r. post. R is probably represented partly by a small 
fragment still attached to r. ant. R and partly by a fragment 
attached to |. post. R. The right-hand fragment has been pushed 
slightly inwards. Whether these fragments all represent the 
r. post. R, whether a part or the whole may not represent anal or 
perisomic plates, are questions that cannot be settled on the 
evidence of this specimen. [Dr. Wanner iw Jitt. confirms and 
extends my observation of some small piate or plates in this 
position, one of which, at any rate, he identifies with r. post. R.] 
The really important fact is that r.post.R has been so far 
atrophied that we may safely infer the entire absence of any 
r.’ post. arm. 

The 1. post. R is, in every direction, larger than the others. 
It rests, almost symmetrically, on the shoulders of the 1. post. and 
post. BB and abuts normally on |. ant. R. On its right (7. e. 
posterior) side, however, the radial in its lower part bounds the 
peviproct, and in its upper part has a short sutural union with 
the supposed r. post. R. The upper, or peristomial, margin is 
regularly concave, with a slightly more marked curvature than 
that of the other radials, but shows no bevelling, groove, or 
depression, such as might suggest an articular facet. Outside 
this margin, however, the highest part of the plate rises into a 
mass of calcite (possibly stereom), which has been truncated by a 
cleavage-fracture, sloping slightly downwards and outwards from 
the oral pole. ‘The evidence of the holotype leaves it uncertain 
whether this protuberance formed the base of a mere hump, of an 
unjointed spine, or of an arm; but Dr. Wanner tells me that 
some of his specimens permit “die sichere Beobachtung, dass 
l. p. R eine Gelenkflache fiir einen Arm zeigt,” and his diagrams 
show that this joint-face corresponds with the cleavage-surface 
just described. It will be interesting to learn the exact consti- 
tution of this facet, whether it has a fulcral ridge and axial canal, 
and whether there is a ventral groove leading down to the 
peristome. Of such structures the holotype presents no trace 
whatever. 


THE CRINOID HYPOCRINUS. 909 


The projection of this facet is due not merely to the protu- 
berance on the radial, but also to the greater height of that 
radial and of the subjacent posterior basal. The sum of these 
heights is again relatively exaggerated by the nearer approach of 
' the left posterior side of the theca to the vertical as compared 
with the right anterior side, which bulges outward. ‘Thus the 
oral surface of the theca slopes downwards from the left posterior 
perradius towards the right anterior interradius. If the erect 
position of the stem be admitted, then all these departures of the 
theca from the normal symmetry result in raising the supposed 
large |. post. arm further and further from the sea-floor. 

The evidence thus far does not absolutely warrant the con- 
clusion that the r. ant., ant.,and ]. ant.arms had become atrophied 
out of existence, or even that they were so reduced as to be use- 
less as food-collectors ; but it does lead us to infer that the task 
of sending a food-stream to the mouth was mainly, if not entirely, 
thrown on the I. post. arm. 

The following are measurements of the radials in millimetres :— 


Left Right 


: : | 
Height. aedeh ee | lower lower 
| : a | margin. | margin. 
a, Cialis 1 Na ee 75 9°4 Be |) Gal 45 
CNG Gh gene ee ? 2 58) | ? Bb yl 
preserved. | | 
Wants oh een evenvea| ka geo. TIN azo B4or| 54 | 
to edge of | AA 
| periproct. | | 
| | | | | 
Ik, OSS IB, searesdencatwecc| ian CeLeen lip hy wlIEC/ AN Ratssyeal Note 83 | 


Peristome (Pl. XC. fig. 9).—The measurement “ width above” 
indicates how much of each radial enters into the peristomial 
margin. The opening is “ elliptische,” as Rothpletz describes 
it, only because the plates are shifted; probably it was of 
irregular outline, with an angle about the middle of each of 
the four large radials, and a longer side in the posterior region. 

One would like to know how this peristome was closed: by 
oral plates, by other plates of interradial position, by enlarged 
cover-plates of the left posterior arm, or, asseems highly probable, 
by the brachials of the reduced arms of the trivium. 

Periproct (Pl. XC. fig. 9)—Dr. Rothpletz regarded the 
subcircular or subhexagonal opening between the post. and 
r. post. BB, and the 1. post., r. post., and r. ant. RR as the 
periproct. He did not observe all the difficulties to which this 
interpretation led, such as the apparent absence or almost com- 
plete atrophy of a radial, or, still more, the unusual position thus 
assigned to the small infrabasal, or even the necessary conclusion 


Proc., Zoou. Soc.—1913, No. LXI. 61 


910 DR. F. A. BATHER ON 


that his species could not be a Hypocrinus. So long as there was 
only one specimen known, it might have been legitimate to evade 
some of those difficulties by a vague hypothesis of abnormality, 
or, more precisely, to have supposed that the anus occupied its 
usual position opposite the small basal (which region is worn 
away in the holotype}, but that it had become choked, and that a 
new outlet had been formed by the partial resorption of that 
radial which in the present paper is called r. post. R, but which, 
on that interpretation, would have been ant. R. When, however, 
Dr. Wanner informs me that all his new specimens agree in this 
curious position of the opening, and that none of them shows any 
trace of an opening on the opposite side of the theca, then J am 
bound first to accept the identification by Rothpletz, and after- 
wards to face the difficulties involved. 

The measurements of the periproct in millimetres are: height 5; 
width below, 5:4; left lower margin 3°9; right lower margin 2°7. 

There is no trace of any periproctal plates, unless they are to be 
sought in the minute plates between the left posterior and right 
anterior radials. 

Ornament.—In addition to the growth-lines already mentioned, 
there are to be seen on the better-preserved infrabasals, 7. e. on 
the posterior side, traces of pustules, apparently coinciding with 
the growth-lines. There also seem to be similar pustules on a 
small tract of the posterior basal. The rest of the surface is too 
badly preserved for the ornament to be detected. Dr. Rothpletz 
says that “die durchweg abgeriebenen Platten lassen ihre pordse 
Beschaffenheit zwar noch erkennen.” ‘They are, however, far 
from presenting the curious appearance of H. schneideri, and on 
the better-preserved portions no pores can be observed. Most of 
the surface is irregularly worn and contains numerous holes. 
These latter, however, are either due to some boring organism or 
to the impression of hard particles in the matrix. Often the 
hard particles, which may be grains of a dark mineral or frag- 
ments of other fossil organisms, are still seen closely adherent to 
the cup-plates, or even forced into them. 

Affinities of “ H.” piriformis.—Comparison of the analyses 
shows at once that this species is not a Hypocrinus. The position 
of the small infrabasal, the relations of the periproct to the pos- 
terior basal and to the superjacent radials, and the modifications 
of all the radials, are considerable distinctions. 

It is even doubtful whether the species should be referred to 
the Gasterocomide, although it agrees with the diagnosis of 
that Family, as given above, in all the known essential features 
except in the shape of the radial facets. No other Gasterocomid, 
however, has the infrabasals arranged in quite the same way. 
These two points demand closer scrutiny. 

The position of the small infrabasal in the right posterior 
radius is characteristic of Flexibilia Impinnata rather than of 
Dicyclica Inadunata. When the latter forms have a tripartite 


THE CRINOID HYPOCRINUS, 911 


base, the small IB is usually anterior, as in Hypocrinus. A few 
other positions for it have been recorded, but not, so far as I can 
find, the same as it occupies here, unless it be in the little- 
known Ampheristocrinus Hall (1881) from the Niagara group of 
Indiana. 

The radial facets of the Gasterocomide are of the horse-shoe 
shape so common in Cyathocrinoidea. In “H.” piriformis the 
facets are so greatly modified that it is perhaps unsafe to base con- 
clusions onthem. The arm-bearing facet seems to have occupied 
almost the full width of the peristomial margin of the left pos- 
terior radial in the holotype; but the diagram Dr. Wanner has 
sent me shows it as occupying about half the width. ‘There is, 
in the holotype at any rate, no trace of the axial canal. The 
other facets are atrophied, it is true, but such traces of them as 
there are seem to indicate a pre-existing facet as wide as the 
radial rather than one of horse-shoe outline. Wide facets are, 
of course, almost universal in the Flexibilia Impinnata, though 
not confined to that group. 

If the reference of “/.” piriformis to the Flexibilia be taken 
as a possible hypothesis, we have next to inquire whether in that 
Order the periproct ever emerges below the summit of the radials. 
I am unable to find that such an arrangement has ever been 
deseribed in any known genus, but that is no valid argument 
against the possibility. Among Dicyclica Inadunata this peculiar 
position first appears suddenly in the Devonian, and I have already 
hinted at the possibility of its independent reappearance towards 
the close of the Carboniferous. Why, then, should it not have 
appeared with equal suddenness in some other group, especially 
when the form displaying it is peculiarly specialized in other 
respects ¢ 

Here, moreover, some actual corroboration of the hypothesis - 
is afforded by the fossil which impelled me to make this first- 
hand examination of Hypocrinus. That is a small cup or patina 
from the uppermost bed of the Yoredale series, in Nidderdale, 
Yorkshire. On it is based the new genus and species Cydono- 
crinus parvulus (Bather, Oct. 1913). 

Cydonocrinus belongs to that group of Flexibilia in which, to 
use Dr. Springer’s words, ‘“‘the rays and their divisions are 
rounded exteriorly, and the interbrachial spaces relatively de- 
pressed ” (Journ. Geol. vol. xiv. p. 510, Oct. 1906), This character 
affects also the anal plates, which are ‘not united by suture with 
adjacent rays, but 1 In arm- -like series, more or less separated from 
them by perisome ” (Springer, op. ai. p. 919). Such forms con- 
stitute the Taxocrinidz as defined by Dr. Springer, and it is clear 
that, of the included genera, Cydonocrinus is most nearly allied 
to Taxocrinus. Thus it has the same fundamental plan of the 
cup as has “4.” piriformis. 

Now the features in which Cydonocrinus differs from Tazxo- 
crinus all bring it nearer to “H.” piriformis. They are the 

ole 


912 DR. F. A. BATHER ON 


subglobular shape of the cup, the small diameter of the stem- facet, 
the relatively large size of the infrabasals, which in the Flexibilia 
are rarely visible, the rise of the radials towards their facets 
curiously resembling the hump of the arm-bearing radial in 
“H.” piriformis, the crowding of the facets round the peristome 
which thus becomes relatively small, and, finally, of greatest 
interest in this connection, the outward projection of the posterior 
basal, which, combined with the constriction of the upper part 
of the cup, almost entirely separates the rectal channel from the 
general thecal cavity. In the actual fossil the anal plates, which 
once rested on the posterior basal, have been broken away, so 
that the appearance is that of a periproct, opening at the top of 
the posterior basal and at the lower corners of the adjacent 
racials. 

Given, then, the existence of such a form in Middle Car- 
boniferous times, it is easy to see how the accentuation of the 
left posterior radial with its arm would have pushed the periproct 
further to the right, and crowded out the right posterior radial. 
This and the diminution of the other arms would have produced 
a form agreeing with “/7.” piriformis. On this hypothesis some of 
the minute plates between the left posterior and right anterior 
radials may be relics of the perisomic plates. 

Physiologically considered, a modification bringing the food- 
intake so close to the vent would be curious in any group of 
erinoids; but that such a modification would not be altogether 
out of character with the Flexibilia may be gathered from a 
perusal of two suggestive paragraphs by Dr. Springer (Journ. 
Geol. vol. xiv. pp. 496, 497, 1906). Speaking of “that strange 
influence which has modified the bilateral symmetry of almost 
every genus in this entire group,” he points out that ‘‘if the arms 
have an asymmetrical distortion, it is to the right, never to the 
left.” If the anals are shifted, it is likewise to the right, so that 
the excavation of the posterior basal for their reception is “fon the 
right shoulder of the plate.” 

Timor is a long way from Britain, and I am not going to say 
that the genus which must be established for “//.” piriformis is 
necessarily descended from the genus to which our Yorkshire 
fossil belongs; but I do claim to have shown some reason for 
thinking that “ H.” piriformis may belong to the Taxocrinide, 
and that, peculiarly modified though it is, its very modifications 
are exaggerations of a tendency natural to that Family. 

Accepting this systematic position, at all events until it is 
proved erroneous, we may draw up the following :— 


Generic Diagnosis.—A Taxocrinid with no radianal, with large 
IBB forming a conspicuous part of the cup, with left post. R and 
arm enlarged and all others reduced in size, the right post. R 
being almost entirely atrophied, so that the rectum passes between 
post. and r. post. BB below and 1. post. and r. ant. RR above. 


THE CRINOID HYPOCRINUS. 913 


SUMMARY. 


The two specimens of Hypocrinus schneidert Beyr., described by 
Beyrich and Rothpletz respectively, are redescribed and refigured. 
The structure of the genus is shown to agree with that of the 
Devonian family Gasterocomide, but it is suggested that in this 
case and in that of “ Lecythiocrinus” adamsi the distinctive 
features may have been independently acquired. 

The holotype of “ Hypocrinus” piriformis Rothpletz is ve- 
described and refigured, and proved to be no Hypocrinus. It is 
thought to be a highly modified descendant of the Taxocrinide, 
by way of such a genus as C'ydonocrinus. The left posterior 
radial appears to have borne a large arm, but the other arms were 
more or less atrophied, and the right posterior radial has almost 
disappeared. 


EXPLANATION OF PLATE XC, 
Hypocrinus schneideri Beyr. 


Figs. 1-4 are photographs of the holotype, enlarged slightly 
less than 2 diameters. 


Fig. 1. View to show the relations of the periproct, which is the opening nearest 
the observer. 
2. View trom above the oral pole, showing the peristome and radial facets. 
hig. 3. The cup from the right side: o., oral pole ; a., anal opening or periproct. 
Note the light colour of the dense stereom at the sutures, contrasted 
with the staining of the porous stereom of the general surtace. 
ig. 4. The cup trom below, with the posterior imterradius away from the observer. 
Note the sutures between the infrabasals. 
Fig. 5. A photograph os the peristomial area of the specimen figured by Rothpletz. 
The periproct is at the lower corner of the figure. X 5 diam. 
Fig. 6. Micro-photograph of the upper part of one of the basals in the holotype. 
x 7 diam. 


“HH ypocrinus” piriformis Rothpletz. 


Fig. 7. The ground-down base of the holotype, photographed under water, so as to 
bring out the sutures between the intrabasals and the five axial nerve- 
canals in those plates (see detailed description on pp. 905, 906). The 
orientation is as in fig. 4, and approximately opposite to that of Roth- 
pletz, pl. x. f.236. X 4 diam. < 


Figs. 8-10. Photographs of the holotype, nat. size. 


Fig. 8. Shows the posterior basal with the protuberant left posterior radial (1. p. R.) 
above it and the periproct at its right upper corner. 

Fig. 9. View from above the oral pole, showing the distorted peristome with the 
large but obscure facet of left post. R. to the left, and the anal opening 
towards the observer. 

Fig. 10. The basal facing the observer is the right anterior. Above it is the peri- 
stome (0.), with the left post. R. rising up behind it, and with the anal 
opening (a.) to the left. Note the ornament of growth-lines in the lower 
part of the cup. 


All the photographs were taken by Mr. H. G. Herring, and represent a 
selection from many attempts upon these dithcult subjects. 


[| Note. The following corrections were received too late for insertion in previous 
sheet :— 
Page 896, line 15 from bottom ‘ For Hypocrinus miilleri. 
Pee X00, = I) 4 ve Read H. mnlleri. 
EpITor. | 


914 DR. W. T. CALMAN ON 


52. On Freshwater Decapod Crustacea (Families Potamonidee 
and Paleemonide) collected in Madagascar by the Hon. 


Paul A. Methuen. By W.T. Caiman, D.Sc., F.Z.8.T 
[Received October 8, 19138: Read November 11, 1913. } 
(Plates XCI. & XCII.¢ and Text-figure 161.) 


INDEX. Page 
Imtroductioniande lis trot Mocalitvesseemereeeeeeaiteeeree ee eee eee eee em Le 
Part I. Family Poramontp™. 
1. Description of the Material .. 915 
Potamon (Potamon) madagascaric iense A. Milne- ‘Edwards, 
and its Varietal Forms ...... cet cee OG 
Potamon (Potamon) goudoti H. Milne-Edwards ............... 920 
Potamon (Potamon) methueni, sp. n. SSomahr escent aoe OOO) 
Hydrothelphusa agilis A. Milne- Edwards. Sioa O22 
2. Systematic Afhinities of the Madagascar River! ior abe besa eas an Ce 
3. Geographical Relations of the Madagascar River-Crabs ......... 925 
Part Il. Family PanmMonip 2. 
Palemon (Macrobrachium) lepidactylus Hilgendorf ............... 926 
Palemon (Parapalemon) dolichodactylus Hilgendort ............ 926 
Palemnon (Hupalemon) ritseme de Man ..... .......2.-0.000ee0-00-.. 927 
Palemon hildebrandti Hilgendorf, and its Varietal Forms ...... 928 
INTRODUCTION, 


The collection described in this paper was obtained in Mada- 
gascar in 1911 by the Hon. Paul A. Methuen, to whom I am 
indebted for the opportunity of examining it. The specimens 
are the property of the Transvaal Museum, Pretoria, but, by 
permission of the Acting Director of that institution, a selection 
from among them, including the holotype of the new species, has 
been retained for the British Museum. 

For the purpose of comparison with Mr. Methuen’s material 
the specimens of Potamonidze and Palemonide from Madagascar 
already in the British Museum collection have been re-examined 
and the results have been incorporated in the paper. 

The following is a list of the localities whence Mr. Methuen’s 
specimens were obtained. Those from the localities marked with 
an asterisk were collected for Mr. Methuen by M. Herschell- 
Chauvin of Tamatave :— 


Hastern Region of Madagascar. 


Manambato, streams running into Lake Rasoabé. July, 


SNL 
Ambilo, streams running into lagoon. July, 1911. 
* Rapides de l’Ifotry, a deux joursde Tamatave. September, 
OE 


*Ivondro. August, 1911. 


+ Published by permission of the Trustees of the British Museum. 
{ For explanation of the Plates sce p. 932. 


1 ZS MONS) IP COI, 


G,M, Woodward delet lith. fith amp. 
POTAMON METHUENI. 


Ie Wigs), WS) WS Ie CIO, 


G.M.Woodward ta et hth. Huth imp. 
PALA MON HILDEBRANDTI. 


DECAPOD CRUSTACEA FROM MADAGASCAR. 915 


Mountains leading to the Mangoro Terrace. 
Ambatonharanana, near Ampasimpotsy. July, 1911. 


Mangoro Terrace. 
Imerimandrosa, northern end of Lake Alaotra, altitude 
800-900 metres. July 1911. 


Mountains leading directly to the Plateau. 
Ambohidratrimo. June, 1911. 


South-western Region. 


Andranolaho and Tongoroby, Onilahy River, Dist. of 
Betsioky. October, 1911. 


Part I.—Family PoramMonip& (River-Crabs). 
1. Description of the Material. 


The River-Crabs of Mi. Methuen’s collection are referred to 
the following species :— 


Potamon (Potamon) madagascariense A. Milne-Edwards. 
* i goudoti H. Milne-Edwards. 
+ methuent, sp. n. 


Hydrothelphusa agilis A. Milne- Edwards. 


As will be explained below, the specimens included under the 
first of these names differ considerably among themselves and 
represent at least three distinct forms which may possibly deserve 
to rank as varieties or even as species. 

In the specific descriptions which follow, regard has been 
given chiefly to the characters of the carapace, to the exclusion of 
those of the appendages, since the latter seem to afford no useful 
points for separating the species here dealt with. Attention may 
perhaps be called to the difficulty of deciding whether any single 
specimen is to be regarded as adult. In the case of females the 
broadening of the abdomen seems to afford a fairly definite 
indication, and it is usually easy to sort the specimens into 
“adults” and ‘‘immature.” In the males, however, the genital 
appendages of the first and second abdominal somites are often 
found apparently well developed in very small individuals, which 
differ considerably in the outline of the carapace from larger 
males. 

In the lists of specimens the numbers preceded by the letters 
‘“« BLM.” refer to the British Museum Register of Crustacea. The 
measurements given in the tables are expressed in millimetres 
and were taken by means of a scale graduated to°5mm. No 
very high degree of accuracy can be claimed for them, but I 
believe that they are fairly comparable among themselves; it is 
not in all cases certain that they can be directly compared with 


916 DR. W. T. CALMAN ON 


measurements given by other writers, owing to the frequent lack 
of precise indication of the points between which measurements 
have been taken. In those given here ‘‘length” is measured 
from the hinder edge of the carapace to the front in the median 
line, 7. e., to the bottom of the notch between the frontal lobes ; 
‘“‘ breadth” is the greatest breadth of the carapace wherever it 
may be; ‘‘exorbital width” is the distance between the tips of 
the exorbital teeth, and is therefore least accurate where these 
points are blunt and indefinite; “frontal width” is the most 
difficult of all to find fixed points for, owing to the obliquity of 
the sides of the front ; the measurements here given have been 
taken between the innermost points of the orbits, 7. e., just above 
the antennal flagella, when the carapace is viewed from in front. 

Complete references to the earlier literature relating to the 
species dealt with will be found in Miss Rathbun’s monograph, to 
which I am glad to acknowledge my great indebtedness. 


PoraMon (POTAMON) MADAGASCARIENSE A. Milne-Edwards. 


Potamon (Potamon) madagascariensis A. M.-E., Rathbun, Nouv. 


Arch. Mus. Paris, (4) vi. 1904, p. 264, pl. ix. fig. 7. 


I have ventured to group under this name a number of speci- 
mens which differ considerably among themselves, and of which 
some at least would probably be regarded by other students as 
representing distinct species. I have e so grouped them, partly as 
a confession of failure, for I feel no great confidence that they 
really belong to a single species, although I have found no 
satisfactory characters by which to distinguish them, and partly 
in order to emphasize the fact that the various forms can be 
arranged, in a way that seems to me significant, around the: 
typical form of the species. They radiate outwards from this 
typical form towards the other species of Potamonide recorded 
from Madagascar, and while in some cases there remains enough 
of a gap to justify the retention of the names given to these 
other species, the relations are clear enough to suggest for the 
Madagascar River-Crabs an autochthonous origin from some 
form resembling the typical P. madagascariense. Miss Rathbun, 
in her monograph, attributes a certain range of variation to 
this species, and notes that one of the forms shows a transition 
towards P. hwmbloti. If the view here put forward be correct, 
the range of variation must be considerably extended, and 
P. humbloti, P. grandidierit, and P. bombetokense must be 
included as varietal forms. 

The specimens examined appear to fall naturally into five 
series as shown on the accompanying table of measurements. ‘To 
avoid prejudging the question of their systematic rank, I designate 
these series by letters. 


DECAPOD CRUSTACEA FROM MADAGASCAR, 


Measurements of Potamon madagascariense. 


oii 


Locality, &c. 


Series A. 


Madagascar, B.M.81.9 .. 
3 B.M. 85.18 ... 
BY BICTOOO ss 


Betsileo, SMSO Che aise 


39 32 


39 


32 


SpErRizs B. 


| Ifotry, Methuen, No. 90............ 
9] 


SERIEs C. 


| Ambatonharanana, Methuen, 
No. 25... 
Oe 


Surtes D. 
| Madagascar, B.M. 88.5 


29 33 


SERIES EH. 


Imerimandrosa, Methuen, 


TB, COL copsprootessee 


No. 35 


O4 40404040 Oy OY +0+0 044004 05 +0 


O303+0+0 


OsQ3 00404 400440054005 


Sex.| 


4005 


Length. | Breadth. 
50 66 
30°25 39 
P55) 21:25 
11 14°5 
10 12°5 
16°75 22 
13°5 17-25 
13 16°75 
12°75 16 
25°25 33°5 
Zi 27-5 
20°26 26 
28°5 36 
20°35 26 
19°75 25°75 
37°5 46°5 
32°5 385 
22°25 26°5 
19°5 23°25 

| 
165 21°5 
16°25 21 
16 22 
1575 | 205 — | 
15°5 20°5 
14:5 19 
14°5 19 
NEP || Tes 
13°25 17 
11 13°5 
10°25 13 
34 45°5 
35 45°5 
AT 575 
46°5 58°75 
45°5 56 
38°5 AT 5 


Exorbital 


width. 


Frontal | Breadth 
width. | ratio. 
20 1°32 
12 1:28 

ut 1eBy/ 
5°25 131 
4:75 1:25 
7-25 1°31 
6 OT 
5'd 1:28 
515 1°25 
105 1:32 
9 11583 
85 1:28 
11 1:26 
Sion E26 
Sion melee) 
14°75 1-24, 
13 118 
9°25 1:19 
8 119 
i I 
7B || 128 
CAS | SI 
7 | 133 
7 | 16 
65 1:31 
65 131 
6 1:27 
6 1:28 
4:75 1:22 
4-75 1:26 
14 1:33 
14 1°33 
17°25 1°22 
ile 1:26 
16°5 1:23 
14°75 1:13 


SERIES A.—These specimens are all, without much doubt, 
referable to the typical form of P. madagascariense as described 
and figured by A. Milne-Edwards and by Miss Rathbun. 
characters in which they differ among themselves are relatively 
unimportant, with two exceptions, viz. :—(1) the distance between 
the epigastric and the postorbital crests, which, in the largest 


The 


918 DR. W. T. CALMAN ON 


Specimen (81.9), isa trifle less than half the distance between the 
former crest and the frontal margin, is about 2/5ths of that 
distance in the largest specimen from 85.18, 1/3rd_ in the largest 
from 82.6, and only about 1/4th in an adult female from 80.27, 
while in the smallest specimen from 85.18 the two crests are all 
but continuous ; (2) the roughness of the surface of the carapace, 
which, in most of the specimens, is in accordance with the 
published descriptions and figures, becomes notably reduced in 
the specimens from Betsileo registered under the numbers 82.6 
and 80.27 ; in these, the whole carapace is much smoother, with 
only a few oblique granules or raised lines near the antero-lateral 
margins and on the posterior branchial regions, and one or two 
obsolescent granules on the front; it is to be noted that the 
eranulation or serration of the antero-lateral margins themselves 
is not correlated with the general roughness or smoothness of the 
surface, being stronger in the Betsileo specimens than in the much 
rougher specimens from 85.18. 

In the smaller specimens of this series, especially in the two 
smallest from 85.18, the external orbital notch becomes very 
shallow or is altogether obliterated, and the outline of the carapace 
is more quadrilateral than in larger specimens. 


Serres B.—The four specimens of this series differ from those 
included in Series A in the relative flatness and narrowness of the 
carapace, which has the antero-lateral borders less arched, so that 
the lateral projection of the carapace beyond the exorbital tooth 
is only about 2/3rds of the greatest diameter of the orbit instead 
of at least 5/6ths as in the larger specimens of Series A. These 
differences, however, are rather less strongly marked in the 
larger than in the smaller specimens of this series. The frontal 
lobes are very prominent in one specimen, and in all there is a 
tendency for the margin of the front to be less deflexed than in 
Series A. The exorbital and epibranchial teeth are very pro- 
minent, and, although the carapace as a whole is much less rough 
than in the largest specimens of the first series, the epigastric and 
postorbital crests are sharply defined and the oblique lines on the 
lateral regions are numerous, sharply cut, and. granulated. The 
marginal granules on the under surface of the merus of the 
chelipeds are very prominent and spiniform. It is worth noting 
that even in the larger of the two males the genital appendages 
are very short, less than half the length of the abdomen, and 
apparently immature. 

This form or variety seems to me to lead in the direction of 
Hydrothelphusa. 


Series C.—The largest female in this series is only 16°25 mm. 
in length, but 1s apparently adult ; the largest male is a trifle 
larger. The carapace in all is rather convex, especially antero- 
posteriorly, and the front is bent downwar ds. The carapace, on 
the whole, is rather wide, the ratio of breadth to length being, in 
one specimen, as high as 1°37. The surface is rather smooth, 


DECAPOD CRUSTACEA FROM MADAGASCAR. 919 


although not more so than in some specimens of Series A. The 
epigastric and postorbital crests are somewhat indefinite; in some 
specimens they are nearly in line, but in others they are separated 
by a distance estimated at 2/5ths of that between the epigastric 
and the frontal margin. ‘The exorbital tooth is not prominent 
and the epibranchial i is small. 

The specimens of this series approach the P. humbloti and 
P. grandidiert of Miss Rathbun’s monograph, especially perhaps 
the latter, although they do not agree exactly with the description 
of either. They differ so much among themselves in all the 
characters that might be used to distinguish them from the 
specimens of Series A, that I cannot regard them as specifically 
distinct ; from Series B they are more easily defined. 


Series D.—In the two specimens of this series the carapace 1s 
very smooth, the rugosities on the anterior part of the branchial 
region being hardly perceptible. The general antero- posterior 
convexity of the carapace is well marked, the various regions are 
also convex, and the grooves between them cor respondingly 
distinct. The most characteristic feature is the inflation of the 
anterior branchial regions, which, in both, but especially in the 
female, is much more marked than in any of the specimens men- 
tioned above. The antero-lateral marginal line is not prominent 
and its granulations are nearly obsolete. In the female the 
distance between epigastric and postorbital crests is nearly half 
that between the former and the front, while in the male the 
proportion is not more than two-fifths. 

These specimens, which I cannot believe to be specifically 
distinct from the smoother specimens of Series A, approach 
P. goudoti in general aspect. and are separated from it chiefly by 
the character of the epigastric crests, which do not project nearly — 
so far forwards, are much less oblique, and far more clearly dis- 
joined from the postorbital crests than they are in that species. 


Series K.—This series consists of four large, dark-coloured, 
male specimens which differ considerably from all those discussed 
above. The carapace as a whole is moderately convex, with the 
regions somewhat inflated and the grooves between shen well 
marked. The surface is fairly smooth, with oblique striz on the 
anterior part and more distinct raised lines on the posterior part 
of the branchial region. The crests are well defined and sharp, 
the epigastric well in front of the postorbital. The epibranchial 
tooth is large and the exorbital moderately prominent. 

According to Miss Rathbun’s analytical key to the species of 
the subgenus Potamon these specimens would be referred to 
P. bombetokense, described by her from a single male specimen. 
From the detailed description and figure of that specimen, how- 
ever, they differ in the rather narrower carapace, in having the 
margin between exorbital and epibranchial teeth straight or 
slightly convex instead of concave, and, apparently, in having the 
inter-regional grooves more strongly marked. 


920 DR. W. T. CALMAN ON 


Poramon (Poramon) Goupori H. Milne-Edwards. 
Potamon (Potamon) goudoti H. M.-K., Rathbun, Nouv. Arch. 
Mus. Paris, (4) vi. 1904, p. 305, pl. xiii. fig. 10. 


Measurements of Potamon goudoti. 


‘ | | | 
Locality, &e. Sex. Length. | Breadth. eee | ae ieee 
‘Wedeeasene (no locality), ty ; 
Methuen, No 55.| ¢ 30°25 41 265 12°75 1°35 
Betsileo, B.M. 82.6............| 6 | 31% 42°5 28 13°5 1°35 
r z, .| 3 | 29 39 25°5 115 1:34 
Antananarivo, B.M. 82.13 ...| 3g | 29 39°5 26 12 1:36 
% 4 3g | 28 38°5 25725) Uo n | ueesiam| 
| 55 Hae 27°25 36°25 24°25 11 | 1°33 | 
| ff , (dry)| ¢ | 29°75 | 41 26°5 1275 1°37 | 
Rs ss ~ | Oh 2 39°25 25°75 12P3 WK 
| i ‘ 5 | & | 286 37-5 PES | aul || kei 
| 


The nine specimens agree in general with the published de- 
scriptions and figures of this species, and present no noteworthy 
differences among themselves. 


PoraMon (PoTAMON) METHUENI, sp. n. (Plate XCI.) 


Measurements of Potamon methueni. 


Locality, &c. Sox.| Length. Breadth. Reo | outa Breese 
Teac hone nee epee ie EE eee 
Methuen, No. 34. | 3 33 46:25 | 26°75 12°5 | 1:4 
9 20 36.| ¢ | 30°75 43°75 25 12 | 1°42 
3 i 39.| g | 2825 | 39 24 11 | 1°38 


Description.—Carapace: length about five-sevenths of its 
breadth, very convex, especially from before backwards, gastric 
region sometimes below level of inflated anterior branchial regions ; 
surface finely punctate, smooth. Oblique portions of cervical 
groove shallow and obscure, if produced forwards they would cut 
the line of the postorbital crest rather within the outer angle 
of the orbit; H-shaped depression well marked; posterior part 
of mesogastric region wider than urogastric, the latter fairly well 
defined posteriorly. On the anterior branchial region is a faintly 
marked oblique ridge nearly parallel to the cervical groove. 


DECAPOD CRUSTACEA FROM MADAGASCAR, 921 


HKpigastric crests not very prominent, more or less rugose, 
hardly oblique, situated entirely behind a line joining epi- 
branchial teeth ; they are continued, without interruption, into 
the protogastric portions of postorbital crest, which are very low, 
rounded, and ill-defined, except in so far as they are limited 
by the transverse depression behind each orbit; in general 
direction the postorbital crest is transverse, and it 1s not trace- 
able external to its intersection by the cervical groove. Groove 
between the epigastric lobes bifurcating widely behind, the limbs 
angularly bent so as to become parallel. Front strongly deflexed, 
but with anterior margin yisible from above; divided into two 
low rounded lobes, most distinct from above, less so from in 
front; with a smooth raised marginal line. Upper margin of 
orbit with a prominent smooth border, inclined forwards, with 
only a faint trace of a lobe in the middle, towards the exorbital 
angle which is not at all prominent or dentiform. Lower margin 
of orbit nearly straight as seen from below, and smooth or 
obscurely crenulated; without any trace of a notch or sinus 
below the exorbital angle. Antero-lateral margin strongly arched, 
with a distinct marginal line, which is smooth or, in the smallest 
specimen, obscurely crenulate, and runs on to dorsal surface 
behind. Postero-lateral border concave, with more or less distinct 
lines. Pterygostomial region and, iess conspicuously, subhepatic 
region beset with tubercles. 

Merus of third maxillipeds slightly wider than long; ischial 
groove rather nearer the inner than the outer margin. 

Chelipeds unequal. Lower surface of merus margined with 
tubercles, which may become indistinct ; a small blunt tooth near 
distal end of inner edge; upper edge rugose with a low sub- 
terminal tubercle. Carpus smooth and punctate externally ; 
with two teeth on inner side and a variable number of small 
granules. Surface of hand smooth and rather coarsely punctate ; 
fingers long, slender, and strongly deflexed. 

Sixth somite of male abdomen shorter than its distal width. 

Holotype, wale, 33 mm. in length, in British Museum. Para- 
types, the two other males mentioned above, in Pretoria Museum. 

Remarks.—This species resembles, on the one hand, P. goudoti 
and, on the other, the smooth form of P. madagascariense described 
under Series Dabove. From the former it differs chiefly in the 
wider carapace (the concavity of the postero-lateral borders has 
the effect of increasing the apparent difference in this respect) ; 
in having the epigastric crests nearly transverse, lying behind 
instead of in front of the line joining the epibranchial teeth, and 
quite continuous with the obsolescent postorbital crests ; in having 
the gastric region less elevated and the transverse convexity of the 
carapace therefore less marked; and in having no notch below 
the external orbital angle. Compared with P. madagascariense 
the same differences hold good, even although in that species the 
epigastric crests are less oblique and less advanced than they are 
in P. goudoti. 


999 DR. W. T. CALMAN ON 


HypRorHELPHUSA AGILIS A. Milne-Edwards. 


Hydrothelphusa agilis A. M.-E., Rathbun, Nouv. Arch. Mus. 
Paris, (4) vil. 1905, p. 266, pl. xvii. (Potamonide, xv.) fig. 7 


Measurements of ye aioe aS 


| 
| | 


| Breadth.| | Exorbital_ |Prontal | | Breadth | 


Locality, &c. Sex.|} Length. | 


| width. | width. | ratio. 
| Ambatonharanana, | | | | | | | 
| Methuen, No. 10) Q 2 | 36°56 PMS | PRS 3) alate | 
is % 11] g | 285 | 32% 255 lal Welelco a la | 
: ‘ 12, g | 295 | 335 25°5 1225 | 113 | 
5 a 131 9 | 305 3455 | 26:5 112;5 10a pela oi 
| i a 14) $ | 31-75 | 3625 | 27 | Be | me 
| Ambohidratrimo, | 
Methuen, No. 28) 9 | 27 BS || HPSS ii 115 
| i i 291 g | 315 | 38 27 135 | 12. | 
| : rs 301 2 | 385 455 32°75 15°5 118 | 
| i cs 31| 9 | 275 325 | 23-5 IS) aeLS | 
a xf 32 g | 26 3025 | 2275 | 1025 | 116 | 


The arrangement of the spinules on the lower orbital margin 
is less regular and constant than is implied by Miss Rathbun’s 
description. In other respects the specimens agree well with the 
published accounts of this species. All the female specimens, 
with the exception of No. 30, are clearly immature. 


2. Systematic Affinities of the Madagascar River-Crabs. 


In his monograph of the Indian River-Crabs (Cat. Crust. Indian 
Mus. pt. i. fase. 1. Potamonide, 1910), and in a short but very 
important later paper on the classification of the family as a 
whole (Records Indian Mus. v. pt. iv. 1910, p. 253), Lt.-Col. 
Aleock has given an entirely new aspect to the system of the 
Potamonide. It is necessary, therefore, to enquire how the 
Madagascar species stand with reference to the new divisions of 
the family. Relying mainly, but not exclusively, on the character 
of the mandibular palp, Alcock has divided the River-Crabs of the 
Old World (apart from the aberrant Deckeniine) into the two 
subfamilies of Potamonine and Gecarcinucine. In the former 
(text-fig. 161, A) the terminal segment of the palp is simple, 
although it may be thickened amd plumose at the base, while in 
the latter (text-fig. 161, C) it is “deeply cut into two lobes which 
embrace the incisor process of the mandible.” Alcock refers the 
Madagascar genus Hydrothelphusa to the Potamonine, but he 
pout out that ‘the thickening at the base of the ter minal joint 


DECAPOD CRUSTACEA FROM MADAGASCAR. 923 


of the mandibular palp is more than ordinary prominent.” I 
find that this statement applies not only to Hydrothelphusa, but 
also, and in an even greater degree, to the other Madagascar 
species that I have examined. In Hydrothelphusa the proximal 
thickening forms a sharply marked ridge ; in Potamon madagas- 
cariense (text-fig. 161, B), P. goudoti, and P. methuent this ridge 
overhangs a little as a free lobe. This evidently forms a transi- 
tion to the bifid palp of the Gecarcinucine, although it is sufli- 
ciently far removed from the typical Gecarcinucine condition as 
seen in Parathelphusa tridentata (text-fig. 161, C), for example, to 
allow us to refer all the Madagascar species with confidence 
to the subfamily Potamonine, with which they also agree in 


Text-fig. 161. 


C. 


Mandibular palp (denuded of sete) of three species of Potamonide. The lower 
figures represent the palp as seen from below, the upper figures the terminal 
segment as seen from infront. A. Potamon fluviatile. B. P. madagascariense. 
C. Parathelphusa tridentata. 


the shape of the male abdomen. It is possible that a similar 
transitional form of the palp may yet be found in species of 
Potamonine from other parts of the world, but it is evident from 
Alecock’s remarks that it does not occur in any of the Indian 
species ; and, in the absence of evidence as to the species of the 
African and Malaysian Regions, it may be provisionally regarded 
as suggesting specially close affinity between the species found in 
Madagascar. 

I do not think that much importance attaches to the fact that 
in all the Madagascar species the palp consists of only two 


g24 DR. W. T. GALMAN ON 


segments. Alcock found three segments distinct in all the 
Indian species of Potamon, but in his later paper he states that 
in the Potamonine the palp has either two or three segments. 
From the examination of a number of African species I suspect 
that it will not be possible to distinguish sharply between those 
species in which the suture-line between the first and second 
segments is well marked and those in which it is almost or quite 
obliterated. 

As regards the generic affinities of the species here discussed, 
there is nothing to forbid the conclusion that they are closely 
interrelated. ‘They have, indeed, been placed in two distinct 
genera, Potamon and Hydrothelphusa, and the species of the 
former might not unreasonably be placed in two subgenera, 
Potamon and Geothelphusa, but the groups indicated by these 
names are probably more convenient than natural. Both Alcock 
and Miss Rathbun comment on the close resemblance of Hydro- 
thelphusa to Potamon, and Miss Rathbun specially mentions its 
affinity with P. madagascariense ; this affinity becomes even more 
striking when comparison is made with some specimens of 
Mr. Methuen’s collection which I have described above as repre- 
senting a form of P. madagascariense (Series B). Potamon 
goudots is considered by Miss Rathbun as forming a link between 
the subgenera Potamon and Geothelphusa, and the new species 
described above as P. methueni goes even further in the direction 
of the latter subgenus. There can, however, be little doubt that 
Geothelphusa, even as restricted by Alcock, has still very slender 
claims to be regarded as a natural or monophyletic group. <A 
general softening of the asperities of the carapace, such as we 
frequently see within the limits of undoubted species, has only to 
be carried a little further to result in the obliteration of the 
postorbital and epigastric crests; and there is no difficulty im 
supposing that so short a step may have been taken independently 
by more than one evolving species. ‘This independence is, in 
fact, implied by Alcock’s transference of certain species of ‘ G’eo- 
thelphusa” to genera of his subfamily Gecarcinucine, and I think 
there is some slight evidence of the same thing among those that 
remain in the Potamonine. At all events there is no necessity 
to assume that P. goudoti and P. methueni are more closely 
allied to the species of ‘‘Geothelphusa” found in Africa and Asia 
than they are to P. madagascariense. 

Potamon madagascariense itself is placed by Miss Rathbun in 
the group of which P. edule (fluviatile) is the type. While it 
certainly resembles the species of this group in the characters of 
the carapace, it differs a little, as already indicated, in the form 
of the mandibular palp, and also in some other small characters, 
such as, for instance, the much broader contact of the basal 
antennal segment with the front. What the exact value of these 
differences may be it is at present impossible to say. 


DECAPOD CRUSTACEA FROM MADAGASCAR. 925 


' 3. Geographical Relations of the Madagascar River-Crabs. 


In addition to the species mentioned above, three others have 
been recorded from Madagascar. The African Potamon (Pota- 
monautes) depressum was recorded by Lenz and Richters, but its 
occurrence is doubted by Miss Rathbun. Potamon (Potamon) 
pittarellu, described by Nobili (Boll. Mus. Zool. Torino, xx. 
No. 507, 1905) since the publication of Miss Rathbun’s mono- 
graph, appears to be allied to P. madagascariense, but is, no 
doubt, a distinct species. Finally, Potamon (Parathelphusa) 
antongilense, described by Miss Rathbun, is evidently a remark- 
able and isolated species, concerning which further information 
is much to be desired ; as far as can be judged from the shape of 
the male abdomen, it is not a Parathelphusa in the sense in which 
that genus has been restricted by Alcock. 

Leaving aside these three species, and considering only those 
that are represented in the collections now examined, it is 
evident that the Potamonid fauna of Madagascar bears no close 
relation to that of Peninsular India, which, as Aleock has shown, 
belongs entirely to the subfamily Gecarcinucine. This is im- 
portant, since geologists seem to be agreed that the connection 
of Madagascar with India existed only at a time (not later than 
the early Oligocene) when Peninsular India was separated from 
the continent of Asia. Of the possible affinities with African 
species it is perhaps unsafe to speak until the rich Potamonid 
fauna of Africa shall have been examined as thoroughly as that 
of India has been by Aleock; but it is worth noting that no 
typical Pofamon seems to be known from East Africa; and if, on 
the one hand, P. madagascariense is the most primitive of the 
Madagascar species, and, on the other, if it is really allied to the 
group of P. flweiatile, then it is hard to guess by what route it 
ean have reached the island.. Further than this it does not seem 
possible at present to go. As I have indicated above, our con- 
ceptions of the phylogenetic relationships of the species rest, as 
yet, on a very narrow basis of morphological fact; and, until 
that basis is broadened very considerably, the group of River- 
Crabs, as a whole, appears to me to be a hazardous subject for 
zoogeographical speculation, 


Parr IJ.—Family Pau #MoN1ID# (River- Prawns), 


The Palemonide of Madagascar have already formed the sub- 
ject of an important memoir by Coutiére (Ann. Sci. Nat. Zool. 
(8) xii. 1900, pp. 249-342, 5 pls.), on which I have relied mainly 
for the determination of the four species in Mr. Methuen’s 
collection. These are as follows :— 

Palemon lepidactylus Hilgendorf. 
dolichodactylus Hilgendorf. 
ritseme de Man. 

7 hildebrandti Hilgendorf. 


Proc. Zoou. Soc.—1913, No. LXIT. 62 


9 


9? 


926 DR. W. T. CALMAN ON 


While all the adult males and a good many of the females in 
the collection have been referred. to one or other of these species, 
there remain over a number of specimens which either lack the 
large chelipeds or are clearly immature. It is possible that these 
may include representatives of some other species, but I have not 
succeeded in identifying them, and it does not seem that any 
good purpose would be served by enumerating them here. 

The most interesting species in the collection is that which I 
identify with Bithynis hildebrandti of Hilgendorf, which does not 
appear to have been re-examined since its description in 1897. 
The series of specimens collected by Mr. Methuen, together with 
those already in the Museum Collection, show, I think, that this 
species is not closely related to the Chihan species which is the 
genotype of Bithynis; that the single character which led to its 
being referred to the genus Bithynis is a matter of individual 
verseision 3 and that, therefore, the species should be removed to 
the genus Palemon. 


Pat#Mon (MACROBRACHIUM) LEPIDACTYLUS Hilgendorf. 


Palemonlepidactylus Milgendort, Monatsber. Akad. Wiss. Berlin, 
1878 (1879), p. 838, pl. iv. figs. 14-16. 

Palemon (Maer obrachium) lepidactylus Coutiére, Ann. Sci. Nat. 
Zool. (8) x1i. 1900, p. 272, pls. x., x1. figs. 1-15 a. 

Localities. Ambatonharanana, 7 males (Methuen); Rapides de 
VIfotry, | male (Wethuen): Betsileo, 3 males, B.M. 80.27. 


In the characters of the rostrum and of the larger cheliped all 
the specimens fall within the range of variation determined for 
this species by Coutiére. In no case, however, does the form of 
the smaller cheliped agree with that which he regards as typical. 
‘The fingers of this appendage are in all the specimens straight or 
nearly so, meeting completely or with only a slight gap, and 
having a scanty provision of bristles on their opposed edges. In 
these respects the specimens agree much better with Coutiére’s 
P. hilgendorfi, and suggest that that species should not be 
regarded as distinct from P. lepidactylus. 


- PaL#Mon (PARAPALAZ[MON) DOLICHODACTYLUS Hilgendorf. 


Palemon dolichodactylus Hilgendorf, Monatsber. Akad. Wiss. 
Berlin, 1878 (1879), p. 840, pl. iv. fig. 18. 

Paleemon dolichodactylus Coutiére, Ann. Sci. Nat. Zool. (8) xii. 
1900, p: 283, pl. xi. figs. 18, 19. 

Localities. Andyanolaho and Tongoroby, Onilahy River, 15 
males, 3 females (Methuen). 

The larger males in the collection agree very well with the 
accounts of this well-marked species given by Hilgendorf and by 
Coutiére. With them are associated some smaller specimens in 
which the woolly covering of the larger chela is very slightly 
developed. The rostral formula is ay wibhit4 Suomi thie upper 


2-4 


teeth postorbital. 


DECAPOD CRUSTACEA FROM MADAGASCAR. 927 


PAL&MON (EuPALzZMON) RITSEM& de Man. 


Palemon(Lupalemon) ritseme de Man, Zool. Jahrb., Abth. Syst. 
Ixe SOs po ci4e op. cits x.) 1698 \plyxxcxvan tie. 70! 

Palemon ritseme Coutiére, Ann. Sci. Nat. Zool. (8) xii. 1900, 
p. 314, pl. xii. figs. 32-33 a. 

Localities. Manambato, streams running into Lake Rasovbe, 
3 males, 8 females (Methwen); Ivondro, 6 males (Methuen) ; 
Ambilo, 2 males (Zethuen); Tamatave, 1 male, B.M. 82.6. 


The specimens collected by Mr. Methuen belong, without 
doubt, to the same species as a specimen in the Museum Collec- 
tion from Tamatave, determined by Mr. Miers as P. ide Heller. 
They differ conspicuously from typical specimens of that species, 
however, in the fact that the surface of the carapace and of the 
tail-fan is completely smooth or presents only very minute and 
inconspicuous traces of spinules, while the chelipeds of the second 
pair are very finely scabrous. Since Nobili has described a 
smooth or nearly smooth variety of P. ide from New Guinea, 
and Coutiére records a similar example from Madagascar, this 
character may not be of specific value. The chelipeds of the 
second pair are shorter, in relation to the body-length, than in 
Coutiére’s specimens of P. ide, the longest being only about 
11 of that length, and most of them being actually shorter. 
Although the carpus is, except in three instances, longer than 
the chela, the difference is very much less than is shown by 
Coutiére’s measurements in specimens of P. ide of similar size, 
and the fingers are always much more than half the length of the 
palm. On the other hand, the rostral formula in nearly all cases 
falls within the somewhat narrow limits assigned to the variations 
of this character in P. ide, and the armature of the fingers agrees 
precisely with Coutiére’s description. 

In all the points mentioned in which these specimens differ 
from the descriptions of P. ide, they approach Coutiere’s account 
of the form which he identifies with P. ritseme de Man. Our 
smaller specimens, however, which alone are directly comparable 
with Coutiére’s largest, have the second chelipeds more asym- 
metrical and apparently a good deal stouter, the chela itself, and 
especially the palm, being relatively a little shorter. The larger 
specimens have many of the proportions rather different, the 
greater relative length of the palm in most of them being 
noteworthy. Finally, in the specimens now examined, the pos- 
~ terior pereopods are, on the whole, stouter than in those 
previously described in this species, the ratio of length to breadth 
in the propodus of the fifth pair varying from 25 to 21, while 
de Man gives the ratio as from 35 to 25 and Coutiére as 25, 

On the whole, the balance of characters seems to be in favour 
of regarding our specimens as older individuals of the species to 
which Coutiére’s specimens belonged, which he has recorded under 
the name of P. ritsema@ ; whether they really belong to de Man’s 
species is, perhaps, a little more doubtful; and I am not at all 

62* 


998 DR. W. T. CALMAN ON 


confident that they would be found to differ essentially from the 
smooth variety of P. ide. It may be noted that, in having parts 
of: the surface of the carapace slightly scabrous, some of our 
specimens agree with P. sintangensis de Man, which Coutiére 
regards as a varietal form of P. ritseme. 

The following table gives (in millimetres) the more important 
measurements of all the specimens that have one or both 
chelipeds of the second pair preserved. The “total length” of 
the body and also the ‘‘ total length ” of the chelipeds of the second 
pair are only approximate, on account of the impossibility of 
straightening the specimens without injury. In an ovigerous 
female the eggs measured *5 x -4 mm. 


Measurements of Palemon ritseme. 


; 
| Chelipeds of second pair. 
heen «| Total | Rostral 
| Locality, &e. Sex Nenoth. ase Tot Hi | | 
hansen | Merus. Carpus. | Chela., Palm. | Fingers. 
| | | | | 
| | | 
| Ivondro, | | | | | 
| Wiktihnen, NOOB ce ING |) PaO BON Sas Wes boi ho 15 
| | | 
E Ba CaS 23 20150 ate ABE S38 6rn 2Mcome Malet 
” BB saps || Ue ay 12 93)| 18 PA) er wri |) Os 
| 
} | | 
2+9 | | | 
x a7...) 6 | 108 | == 88 | 14 2750426 paal lel aaxo 
| Manambato, | 948 | | | | 
| Methuen, No. 42 ...| ¢ 98 ere pe Eee 22 e23rom elo om eee) 
ieera | | | | | | 
| lvondro, | Oey | 9 | | ¢ 
| Methuen, No. 99 ...| g | 95 | —Q> | Ve i | 3 | a | ae aoe 
| Manambato, | 348 | | | 
| Methuen, No. 46...| @ | 92.) == | 2 || 10 165 | 165 |) 85 | 8 
247 | | | 
| Tamatave, B.M.82.6 | 3 90 way | 2028 35 | SBS || BO 13° 
Manambato, I pate | | 
Methuen, No. 47 ...| g | 80 3 AGS, 10 hy) Wy 95) 75 
| | 
. 45 O75 2 2 9 155 96 15 8°5 | 65 
| | 


PALEMON HILDEBRANDTI Hilgendorf. (Plate XCII.) 


Bithynis (?) hildebrandti Hilgendort, Sitz.-Ber. Ges. Naturf. 
Freunde Berlin, 18938, p. 244; Coutiére, Ann. Sci. Nat. Zool. (8) 
xii. 1900, pp. 250, 256, 259; Ortmann, Bronn’s Thierreich, 
Crustacea, ii. 1901, p. 1292; Ortmann, Proc. Amer. Phil. Soe. 
Philadelphia, xli. 1902, p. 274. 


DECAPOD CRUSTACEA FROM MADAGASCAR. 929 


The material which I have examined consists of eleven speci- 
mens,from three localities, in the British Museum (Nat. Hist.) Col- 
lection and nineteen specimens from one locality in Mr. Methuen’s 
collection. ‘The Museum specimens (PI. XCII. fig. 1) agree with 
Hilgendorf’s description of this species, except in one or two small 
details to be mentioned below. Mr. Methuen’s specimens were at 
first regarded as representing a probably new species of Palemon, 
since they possess a well-developed hepatic spine on the carapace 
(Pl. XCIL. fig. 6). Closer examination showed, however, that 
they differed in no other respect from the Museum specimens, 
and their specific identity was finally proved by the discovery, 
among Mr. Methuen’s specimens, of one which has no trace of a 
hepatic spine on one side of the carapace, while on the other 
side its place is indicated by a small and inconspicuons tubercle. 

The following description applies to all the specimens examined. 

The sides of the carapace and the abdominal somites are 
rather coarsely punctate. The rostrum is more or less straight, 
with the dorsal edge moderately arched, of varying length but 
always shorter than the antennal scale, and often shorter than 
the antennular peduncle ; of the dorsal teeth, one is postorbital. 

The tip of the telson (Pl. XCII. figs. 4 & 5) is rather broadly 
rounded, sometimes, but not always, with a minute median point ; 
the inner lateral spines are much longer than the outer. Between 
the inner spines is a fringe of sixteen plumose hairs. 

The third maxilliped does not reach the tip of the antennal 
scale. The chelipeds of the first paix extend beyond the scale by 
a little more than the chela. 

The chelipeds of the second pair (Pl. XCII. fig. 2) are similar, 
smooth, with fine scattered setules, and may extend beyond the 
antennal seale by the length of the chela and carpus. The carpus 
is equal to the merus, widened distally ; the chela slightly com- 
pressed (5:6), a little wider than distal end of carpus; palm 
longer than carpus and equal to fingers; fingers with a sharp 
cutting-edge for the greater part of their length, with two small 
teeth near the base of the dactylus, and one, with an indistinct 
second, on the immovable finger (Pl. XCII. fig 3). 

Posterior legs moderately stout, propodus of last pair abouttwelve 
times as long as wide and about four timesas long as the dactylus. 

The females are ovigerous from a length of about 38 mm. 
upwards. The eggs measure approximately 2 x 1-4 mm. 

It will be observed, from the table of measurements given 
below, that the females are much more numerous than the males 
in the collections examined and also that they considerably exceed 
them in size; both conditions appear to be uncommon among the 
species of Palemon, but they are found also in P. moorei of Lake 
Tanganyika. 

Hilgendorf described the mandibular palpas consisting of only 
two segments. In all the specimens which I have dissected the 
usual three segments are distinct. Hilgendorf also attributes 
three teeth to the dactylus of the second chelipeds and states that 


930 DR. W. T. CALMAN ON 


the carpus “ ragt nur ein wenig unter die Antennenschuppe nach 
vorn,” while in our specimen the dactylus bears only two teeth 
and the carpus may extend beyond the tip of the antennal scale. 
In all other respects, however, Hilgendort’s description applies so 
well to our specimens that their specific identity can hardly be in 
doubt. 

The existence of an affinity between this species and the 
Bithynis gaudichaudii of Chile and Peru, asserted only in a 
hesitating manner by Hilgendorf, is fully accepted by Coutiére, 
but denied by Ortmann, who considers that the resemblance does 
not extend beyond a single character (the lack of a hepatic spine) 
which may easily be supposed to have arisen by convergence. 
Ortmann’s view is strongly supported by the variability of this 
character in the present series of specimens. I have examined 
the appendages of both species for other evidence of affinity 
between them, and have failed to find it. The branchial system 
and the mouth-parts of both are very similar to those of several 
species of Palemon with which I have compared them. Only in 
one point do the mouth-parts of B. hildebrandti present anything 
unusual, and that is the reduced size of the epipodite of the 
first maxillipeds, but this constitutes no resemblance to B. gaudi- 
chaudii, in which the epipodite is quite as large as in the species 
of Palemon examined. On the other hand, the differences in 
general aspect between the two species are considerable ; B. hilde- 
brandti has the chelipeds hardly differing in the two sexes, 
slender, symmetrical, smooth, with the carpus equal to the merus, 
and the fingers armed with only a few small teeth near the 
base ; B. gaudichaudii is a very much larger species, with the 
chelipeds very stout, much more strongly developed in the male 
than in the female, very unequal on the two sides of the body, 
beset with spiny tubercles, with the carpus shorter than the 
merus, and the fingers of the larger chela toothed for half their 
length or more. If it be advisable to maintain the genus Bithynis 
(which seems to me doubtful) it must be for the South American 
species alone, and B. hildebrandti must be transferred to the genus 
Palemon. 

The variability of the hepatic spine in this species recalls th 
cases of “mutation” recently described by Bouvier in certain 
Atyide. It resembles these cases in its discontinuity, only one 
individual out of those examined being in any way intermediate 
between the two forms; and it further resembles some of them 
at least in the fact that it is geographically limited, for the 
specimens of the two forms come from different localities. Both 
of these points, however, require to be tested by further collecting. 
Tt differs from Bouvier’s cases in that if concerns only a single 
character, and one which, were it not for its constancy in other 
Palemonide, might be regarded as of trivial importance. 

The large size of the eggs may be taken to indicate an abbre- 
viation or suppression of the larval development, and this is 
likely to be associated, as jt is in some other Palemonide, with 


DECAPOD CRUSTACEA FROM MADAGASCAR. 


931 


an exclusively fresh-water habitat and a restricted area of distri- 
bution ; it may be not without significance therefore, that, of the 
species of Paleemonide in the present collection, P. hildebrandti is 
the only one that does not occur outside the island of Madagascar. 


Measurements of Palemon hildebrandti. 


Locality, &c. 


Ambatonharanana, | 
Methuen pene eee 


” 


UM 


” 


” 


” 


” 


»” 


a” 


” 


9 


4.6 


38 


36 


36 


Total Rostral | Hepatic | 
‘length. formula.) spine. | 


| 


Chelipeds of second pair. 


| Merus. 


Carpus. 


Chela. 


Palm. | Fingers. 


“Tr 
I 
or 


932 


ON DECAPOD CRUSTACEA FROM MADAGASCAR. 


Measurements of Palemon hildebrandti (continued). 


80 


| Fingers. 


| Chelipeds of second pair. 
Air Total Rostral | Hepatic 
| Locality, &. Sex. length. formula.) spine. | l Nl 
| | Merus. | Carpus. | Chela.) Palm. 
| 
foueh. oe i ae nai aaa ag | | 
| | | | | 
| | 14+5 1 GO) 1. Bo) |) WSO | 
| Betsileo, B.M. 82.6.) g | 50 | ~ 3 0 eo | 6:0 | 145 | 
| 
a sp il CRA Sh A Aan, sia [pesos felt 
ee ieee 
| 147 | 657 57 || 13:0 
99 ” sisi 2 46 | “o | 0 | ry 5-7 67 12°2 
| | 14+6 | (60 60 | 12°2 
aera : ea! Rephieeon BoA OO 55 | 12°0 
mi ey alee Page ae! Oo 6:0 60 | 120 
| | | 
| | | | 
: Apes, Puen ies = ear 60 | 60 | 136 
| | | | 
ig | | 147 | 45 | | 45 8-9 | 
” 3 ? | 38 SEG ios | 0 ae 4°5 | 89 | 
| | | 
Madagascar | 
B.M.79.20........... 9 | 52 | on GACT Ohi HOR ONT eS Fe 
| 
Movie fot @ | a = 0 6:0 60 | 14:0 
| | 
| 
East Imerina, | | 
| BM. 92.7.4.6-7...| 9 | 37 | a8 LO, algae ies Hie! ak | 
| | | | 
| : 14+6 | 
% + | 2 Be To LO eepne tee elm moc ats Ikeaanes 


EXPLANATION OF THE PLATES. 
Puate XCI. 


Potamon methueni, sp. n. 


Fig. 1. Male. Holotype. Natural size. 
2 


at ES 3 Hand of larger cheliped. 14. 
Bs gy ue Abdomen. XX 13. 
Aa oe i Third maxilliped. 23. 


Pruate XCII. 
Pelemon hildebrandti. 


Fig. 1. Typical form, ovigerous female. X 23. 
2 Cheliped. x 5. 
Bo a 4 base of fingers. X 10. 
4., as: Telson. XX 10. 
D. f Tip of telson. ™ 1d. 
6. Form with hepatic spine. Carapace. 2k. 


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LE CHRIORCHIS INERMIS. 


ON A NEW REPTILIAN TREMATODE. 933 


53. A New Trematode of the Genus Lechriorchis from the 
Dark Green Snake (Zamenis gemonensis). By Maris 
V. Lesour, M.Sc., Assistant Lecturer and Demonstrator 
in Zoology, Leeds University *. 
[Received September 20, 1913: Read November 25, 1913. | 


(Plate XCIL4) 


INDEX. 
Page 
Ethology: A new species of Trematode in the body- 
Sy I 
cavity of Zamenis gemonensis .......1.ceeee see eee eee 933 
Geographical Zoology: Southern Europe; Zamenis 
grap sy 5 
gemonensis, a new Trematode from the body- 


CAVAD YE. cr eecc es dence ae nein ste ence retee CHER Gece te Meebect 933 
Systematic: Lechriorchis inermis, sp. n., from the 
body-cavity of Zamenis gemonensis  .................. 933-936 


On June 10th, 1913, a specimen of a Dark Green Snake 
(Zamenis gemonensis), from Southern HKurope, died in the Zoolo- 
gical Society’s Gardens, and loose in the body-cavity were a large 
number of Trematodes which were sent to me for identification. 
These prove to be a new species of Lechriorchis Stafford closely 
related to Z. validus Nicoll (1911). The habitat is interesting as 
all the Trematodes of the group which includes Lechriorchis 
whose habitat is known, occur in the air-passages, pharynx, 
or anterior part of the cesophagus of their hosts. This appears 
to be the first time that one of this group has been found 
occupying the body-cavity. 

An interesting feature in the new species is the absence of 
spines on the body. The cuticle is quite smooth, unlike any of 
the other members of the genus. However, in some specimens 
spines can be seen below the surface of the cuticle, but not 
reaching to the outside, which seems to indicate a gradual dis- 
appearance of spines owing to the worm living in the body-cavity 
of its host, and therefore being in no danger of getting swept 
away as is the case in open passages. In all other ways it agrees 
closely with the other members of the genus. 

On account of the absence of spines I propose for this new 
species the name Lechriorchis mermis. 


LECHRIORCHIS INERMIS, sp.n. (PI, XCIII.) 


Length of body 5-10 mm. Oral sucker 0°40-0°60 mm, 
Ventral sucker 0°60-0°92 mm. Average ratio of oral to ventral 
sucker 2:3. Prepharynx small, esophagus the same length as 
pharynx. Intestinal ceca reaching a good way beyond testes, 


* Communicated by the SpcRETARY, 
¥ For explanation of the Plate see p. 936. 


934 MISS M. V. LEBOUR ON 


Testes behind centre of body, right in front of left. Cirrus-sac 
reaching to posterior margin of ventral sucker. Ovary on right 
side behind or slightly overlapping ventral sucker. Yolk-glands 
reaching from just in front of the level of the anterior margin of 
ventral sucker to centre of testes or behind them. Receptaculum 
seminis uterinum present on right side of body behind right 
testis. 


The body (fig. 1) is convex dorsally and nearly flat ventrally, both 
suckers being conspicuous and the cirrus usually protruding from 
the genital pore. The length of mature worms varies from 5 to 
10 mm.,the last-named being in the case of a specially well extended 
specimen examined under pressure. Unfortunately, this large 
specimen was damaged before being fixed. One immature specimen 
occurred measuring 4.4 mm. in length which contained a few 
eges and had a very short uterus. In breadth the worms 
measured from 1:2 mm. to 1:9 mm., the greatest breadth being 
in the region of the ventral sucker. ‘The average length is about 
5:5 mm., breadth 1:4 mm. 

A good deal of interesting variation occurred in the arrangement 
of the organs. The suckers vary but little, the ratio of the oral 
and ventral suckers being almost exactly 2:3. Both are circular in 
outline with circular apertures. 'The aperture of the oral sucker 
measures about 0°20 mm. across; the aperture of the ventral 
sucker is more than half the width of the sucker and very regular 
and conspicuous. The usual position of the ventral sucker: is 
with its centre almost exactly at the anterior third of the body. 
It may, however, be a little further back. In the young specimen 
it is much nearer the centre of the body, a fact to be expected as 
the hind portion of the body always increases to a much greater 
extent than the fore part. 

The prepharynx is exceedingly short but quite distinct; the 
pharynx is broad and muscular, the breadth slightly greater than 
the length (length 0°20 mm.); the esophagus is about the same 
length as the pharynx.. Surrounding the pharynx are large 
glandular cells which are continued for a short distance down the 
ceesophagus. Pharyngeal glands are mentioned by Stafford in 
L. primus (1905). The intestinal ceca vary in thickness in 
different parts and reach usually about 0°60 mm. beyond the 
testes; beginning near the centre of the body, they gradually 
become more lateral and slightly vential in position until they 
reach the testes, when they become quite irregular and may be 
both dorsal to the testes or one ventral and one dorsal (fig. 3); ‘and 
one may be longer than the other, in one case quite twice as much 
extending beyond the left testis as the right. The usual length 
beyond the testes is 0°60 mm. Sometimes the ceca bend inwards 
so as to be completely internal to the testes. The ends are 
swollen and very much broader in the posterior than in the 
anterior part. 

The excretory vesicle is large and opens at the extreme posterior 


A NEW REPTILIAN TREMATODE. 935 


end, where it is surrounded by large deeply staining cells; just 
in front of the opening the vesicle is much folded, giving it 
the appearance of having been “‘ puckered in” before the narrow 
opening. In front of this pleating it widens out as a large tube 
running dorsal to and parallel with the ascending limb of the 
uterus; in front of the testes and behind the ovary it forks, each 
limb reaching far forward beyond the ventral sucker to about 
midway between the ventral sucker and pharynx. 

The testes are oval bodies more or less lobed on their inner 
margins, usually lying obliquely with the right testis in front of 
the left; the usual situation is for the right to be about 0°15 mm. 
in front of the left, but they may be almost on a level, and in 
two cases the left testis was in front of the right. Curiously 
enough, these variations occurred in the small immature worm 
and in the largest specimen. It is just possible that we may 
have to do here with a different species. In a specimen 5:2 mm. 
in length the testes measured, right 0°90 mm. x 0:44 mm., left 
0°84 mmm. x 0°44 mm., the greater axis lying longitudinally. On 
the inner margin there are two or three lobes. The cirrus-sac is 
long and reaches almost to the posterior level of the ventval 
sucker. Sometimes it reaches the extreme posterior end, and in 
two cases (again the immature worm and the largest) it was 
much shorter and only reached a short way behind the anterior 
margin of the ventral sucker; in this respect being much nearer 
L. validus Nicoll. The vesicula seminalis is much convoluted 
and occupies the posterior third of the cirrus-sac, the ductus 
ejaculatorius being surrounded by an enormous mass of prostatic 
cells and ending ina short protrusible cirrus often to be seen 
exserted from the genital pore. The male opening is on the 
right side of the genital pore, which is situated on a level with 
the centre of the pharynx and midway between it and the lateral 
margin of the body-wall. The ovary is almost round, and lies 
dorsally on the right side of the body immediately in front of the 
right testis. It overlaps the ventral sucker for about a third of 
its diameter; its position varies however, and in some cases it 
may be rather more than this and in others not so much. The 
portion of the ovary that overlaps the cirrus- pouch is on its right 
side. The oviduct receivesa large shell-gland, gives off a Laurer’s 
canal, and receives the vitellarian duct, then runs down as the 
uterus in many coils to nearly the posterior end of the body, 
when it doubles back as a much broader tube, runs forward 
gradually straightening and narrowing and passes the ventral 
sucker dorsally. In the course of the descending portion behind 
the right testis is a receptaculum seminis uterinum. When the 
uterus reaches the level of the anterior ends of the excretory 
vesicle its walls form a vagina with a thick cuticular lining, and 
surrounded by a layer of deeply staining large cells which 
accompany it for the whole of its course until it opens on the 
left side of the genital pore. The eggs are very numerous and 
of adeep brown colour when old, the young eggs being much paler. 


936 ON A NEW REPTILIAN TREMATODE. 


The cap is not very conspicuous until the egg is open. The eggs 
(fig. 2) measure ‘033 mm.—'036 mm. x ‘021 mm.—-023 mm. The 
vitellaria are variable in length, but never reach beyond an ex- 
tremely short distance in front of the ventral sucker, 0°02 mm. is 
quite the furthest they ever reach and usually it is not so much. 
Posteriorly they reach to about the middle of the testes, but are 
extremely variable in this region, and the left may reach to 
behind the left testes and the right hardly reach to the anterior 
portion of the right, or both may reach to the level of the middle 
of the testes. The vitellarian follicles are in small groups lying 
laterally to the outside of the intestinal ceca; small ducts from 
them unite in a transverse duct each side lying just behind the 
ovary, and these unite in a small reservoir in the centre of the 
body which gives off the vitellarian duct to the oviduct. 

Lechriorchis validus Nicoll is undoubtedly the most nearly 
related to the present species. JL. inermis differs from it chiefly 
in its much smaller eggs and position of the vitellaria. The 
position of the testes (right in front of left instead of the reverse) 
and of the vesicula seminalis being rather more variable, do not 
seem to be such important specific characters. 


References. 


Nicouu, W. (1911).—On Three New Trematodes from Reptiles. 
Proceedings of the Zoological Society of London, pp. 677— 
681. 

SrarrorD, J. (1905).—Trematodes from Canadian Vertebrates. 
Zoologischer Anzeiger, xxvii. p. 691. 


EXPLANATION OF PLATE XCIII. 
Lechriorchis inernis, sp. 0. 


The following letters apply to all the figures. 


C.S. Cirrus-sac. | R.S.U. Receptaculum seminis 
K.P. Excretory pore. | uterinum. 
E.V. Excretory vesicle. | 8.GL. Shell-gland. 
G.P. Genital pore. T. Testis. 
I. Intestine. | U. Uterus. 
O. Ovary. | V. Vitellaria. 
(ES. Gsophagus. | VG. Vagina. 
O.S. Oral sucker. | V.S. Ventral sucker. 
PH. Pharynx. V.SEM. Vesicula seminalis. 


P.PH. Prepharynx. 


Fig. 1. Ventral view. Length 5°4 mm. 
2. Eggs, length °033 mm. 
3. Transverse section through unpaired portion of excretory vesicle and testes, 
showing intestinal cca irregularly placed. 
4, Transverse section through ventral sucker, 


IBZ gS) MEMS) Je ZIOINY, 


A.H.Searle del.et lth, Huth imp. 


ZBUGMATOLEPAS MOCKLERI gen.et spn. 


Mb Lat 

Ob Uy 

ATA, 
ANS 


IF) ops LOS) NPIL 2G’ 


Huth imp. 


A.H.Searle del. et lith. 


CALANTICA (TITANOLEPAS) TUBERCULATA, Darwin sp. 


ON FOSSIL CIRRIPEDES. 937 
54. Cirripedes from the Cenomanian Chalk Marl of 
Cambridge. By THomas H. Wiruers, F.G.S.* 
[Received September 25, 1913: Read November 25, 1913.] 


(Plates XCIV. & XCV.7) 


INDEX. 
Page 

Structure of Zeugmatolepas and Titanolepas ..................941, 946 
Lines of Evolution in Pollicipedide .......... ace GOT 
Distribution (Geological), Cretaceous of * Europe, Gengiate 

lepas and Calantica (Litanolepas) . Berane raat .. 939, 944 
Zeugmatolepas mocklert, gen. et SP. N.... 2.0... 2ee cee ree cee cee vee eas 938 
Z. crete Steenstrup sp. .... 94.1 
Scalpellum crete Steenstr up sp., ‘referred to Zeugmatolepas . . 942 
Zeugmatolepas compared with Pollicipes ....... . 941 
Scalpellum tuberculatum Darwin, referred to Titanolepas . 943 
Titanolepas subgen. nov. of Calantica, for Scalpellum 

tuberculatum Darwin ...... ISIS aveh oy aceon eee eke 
Calantica, see Titanolepas and Scillelepas BSCE ace d oaauce ons 947 
Scillelepas compared with Titanolepas ..............0.c000 eee. GAT 


Through the industry of the late Mr. F. Mockler, I have been 
enabled to examine a large series of Cirripede remains which he 
obtained from the Cenomanian Chalk Marl in the neighbourhood 
of Cambridge. By far the greater number can be referred to two 
species, which add materially to our knowledge of the phylogeny 
of the pedunculate Cirripedes. 

One, Zeugmatolepas mockleri, gen. et sp. n., 1s represented by 
two nearly complete capitula and portions of nine others, as well 
as a large number of isolated valves. It is interesting from the 
fact that, while it agrees with: the species of Pollicipes in the 
number of valves of the capitulum, it differs not only in the 
more specialized form of scutum, but also in the size and position 
of the upper latera, which are nearly as large as the scuta, and 
have become elevated to occupy the whole of the space between 
the scuta and terga. This form certainly serves more than any 
other to connect the genus Pollicipes to Scalpellum, and, together 
with S. (?) crete Steenstrup sp., from the Upper Senonian of 
Denmark, is now included in the new genus Zeugmatolepas. 

The second species, Scalpellum tuberculatwm Darwin, is repre- 
sented by a large number of separated valves, including several 
not previously known. These enable us not only to reconstruet 
the capitulum, but also to show that the species must be referred 
to the genus Calantica Gray. Representatives of that genus 
are now found living, and the geologically oldest species of its 
subgenus Scillelepas, to which Scalpellum tuberculatum comes 
nearest, occurs in the Miocene of Italy. The recent and 
Tertiary species have the umbones of all the valves apical, so 
that it is surprising to find in the Cretaceous representative, 


* Communicated by Dr. W. T. Canman, F.Z.S, 
+ For explanation of the Plates see p. 948. 


938 MR. T. H. WITHERS ON 


S. tuberculatum, a more specialized form of scutum, the umbo 
being subcentral. On account of its higher evolutionary deve- 
lopment, S. tuberculatwm is placed in anew subgenus of Calantica, 
namely Z7vtanolepas, and it is here suggested that this form may 
represent an offshoot from the ancestral Scillelepas line, and may 
eventually have given rise to the genus Oxynaspis. 

Both species are remarkable in having an advanced form of 
scutum in which the umbo is subcentral, and, together with a 
small undescribed species of which only the carina is known, 
constitute the oldest known examples of this specialized type of 
valve. It is quite evident that Zeugmatolepas mocklert and 
Titanolepas tuberculata are derived from either the genus 
Pollicipes or from a Pollicipes-like ancestor, and both have 
evolved the same type of scutum. These forms seem to show 
that the transition of the umbo in the scutum from an apical 
to a subcentral position, has occurred independently in unrelated 
forms belonging to distinct lines of development, and recent 
work leads me to believe that this is equally true also of the 
carina in the more advanced forms of Scalpelluwm (sensu lato). 
In 7. tuberculata the position of the umbo in the scutum is 
constant; but in Z. mockleri it changes apparently according to 
age, although some individuals seem to be more advanced in this 
respect. It may be, therefore, that in the scutum of Z. mocklert 
the subcentral umbo was quite a recent development. 

The two species must have been comparatively common in the 
Chalk sea, for of Zewgmatolepas mockleri we have remains of at 
least 100 capitula, and of Z%tanolepas tuberculata more than 30. 


Family POLLICIPEDIDS. 
ZEUGMATOLEPAS *, gen. nov. 


Capitulum longer than broad, comprising at least 34 valves, 
composed of scuta, terga, upper latera, and carina, and three or 
more whorls of subtriangular lower latera, with a rostrum, 
subrostrum, and subcarina ; upper latera almost as large as the 
scuta, occupying the whole of the space between the sorta and 
terga, the apices of the upper latera contiguous with those of the 
scuta ; umbo of scuta subcentral, and of the remaining valves 
apical, 

Genotype.—Zeugmatolepas mocklert. 


ZEUGMATOLEPAS MOCKLERI, sp. n. (Plate XCIV. figs. 1-14.) 


Diagnosis.— Valves thin and generally smooth. Carina narrow, 
usually with a strong, median, rounded keel, not divided off into 
parietes and intraparietes, tage margin acutely angular. Scutum 
subtriangular to trapezoidal in shape, with the umbo varying in 
position from just below the apex to about one-third the length 
of the valve from the apex ; there is in most scuta a deep trough 


* Zedyua = a connecting link. 


FOSSIL CIRRIPEDES. 939 


above the umbo extending almost parallel to the upper occludent 
margin ; apical portion usually constricted and acute ; basilateral 
angle shortly and obliquely truncated. Tergum subrhomboidal, 
<i its upper half curved towards the eeu basal angle 
sometimes acute, but more often shortly and obliquely truncated. 

Material.—Two almost complete capitula and nine portions of 
others. All of these appear to be immature, and there is also a 
large series of isolated valves. 

Holotype.—The nearly complete capitulum, I. 15830 (Pl. XCIV. 
sae, ALT). 

Distribution.—Cambridge Greensand: Cambridge; Cenomanian, 
Chalk Marl: near Cambridge, and Burham, Kent ; Cenomanian, 
A. plena-marls : Oxted, Surrey, and Wantage, Berkshire ; [?Ceno- 
manian |, Chalk detritus: Charing, Kent. 

Measurements.—The two nearly complete capitula (Pl. XCIV. 
figs. 11, 13) each measure 4 mm. from the apex of the scutum to 
the base of the lowest whorl of latera, and the uncrushed 
expitulum (Pl. XCIV. fig. 11) measures 4-2 mm. in breadth. 
Both are immature examples. Among the detached valves the 
largest examples of the upper series measure respectively :— 

Carina, I. 15820. Length (valve shghtly broken at apex) 
5-9 mm.; breadth 1:7 mm. 

Scutum, I. 15822. Length (from apex to lowest point of basal 
margin) 6°4 mm.: breadth 4°5 mm. 

Tergum, I. 15833. Length 7°6 mm.; breadth 3:4 mm. 

Upper latus, I. 15834. Length 4:5 mm.; breadth 3-2 mm. 

There are, however, several fragments which show that these 
valves attained somewhat larger dimensions. A comparison of 
these with the proportions of the valves, as shown in the nearly 
complete capitulum, seems to show that the capitulum of the 
species reached a length of at least 15 mm. 

Description of valves.—Valves thin, smooth, or ornamented 
with excessively fine ridges radiating from their umbones. 

Carina (Pl. XCIV. figs. 1, 2) narrow, widening gradually 
downwards from the apex to ‘the basal margin, either slightly 
bowed inwards, straight, or bowed outwards, noe ‘divided off into 
parietes and intraparietes ; basal margin acutely angular; tectum 
slightly to moderately convex transversely ; in some carine there 
is a Strong, rounded, median, longitudinal keel, nearly as wide as 
the portion of the valve on either side, but in some specimens 
it is less strongly marked and in others barely perceptible. Inner 
surface somewhat thickened near the apex and marked with 
oblique growth-lines ; the extent of the surface thus thickened 
varies, and at the most is one-third of the length of the valve; 
the valve therefore projected freely to a variable extent. 

Scutum (Pl. XCIV. figs. 3-6) varying in shape from sub- 
triangular to trapezoidal, moderately convex transversely, the 
surface of the valve sloping more steeply from the umbo to the 
lateral margin; umbo situated just below the apex in young 
specimens, and in mature examples usually occupying a position 


940) MR. T. H. WITHERS ON 


about one-third the extent of the valve from the apex ; above the 
umbo the valve is usually produced abruptly into an acute angle ; 
lateral margin longer than the basal margin, almost straight in 
its lower part and usually sharply bent inwards and upwards 
above a line with the umbo; basal margin indistinctly marked off 
into three almost equal parts, the two outer parts being obliquely 
inclined towards the apex; lower occludent margin slightly 
convex, about the length of the basal margin; upper occludent 
margin about half the length of the lower occludent margin, 
sometimes nearly in line with it, but more often abruptly bent 
from the umbo towards the lateral margin, and in some cases 
almost in line with the umbo and nearly at right angles to the 
lower occludent margin. Almost all the specimens have a strong 
ridge extending from the umbo to a point just above it on the 
lateral margin, and above this ridge the narrow portion of the 
valve is sharply bent downwards and upwards, and forms a. deep 
trough which is bounded by the raised upper occludent margin ; 
three further ridges, not to be seen in some specimens, extend 
from the umbo to the basal margin, one to the basilateral angle, 
a second to one-third of the distance from the rostral angle, and 
a third to midway between the two. On the inner surface there 
is a deep pit for the adductor scutorum; the inner occludent 
mar. gin is much thickened, and above the nae a portion of the 
valve is bent downwards at right angles to the outer surface in 
some specimens, and in others it is sharply bent inwards; when 
the occludent margin is almost in line with the umbo, this 
downwardly and inwardly bent portion of the valve almost 
entirely overhangs the pit for the adductor scutorum, and forms 
a roof to the upper part of the inner surface. 

Tergum (Pl. XCIV. fig. 7) subrhomboidal, slightly convex 
transversely, with a feebly marked curved ridge extending from 
the apex to the basal angle at about one-fifth the width of the 
valve from the carinal margin, and from this ridge the valve 
slopes steeply to the carinal margin ; upper half of valve curled 
towards the scuta, and in most valves away from the opposing 
tergum; apex acute; basal angle sometimes acute, and sometimes 
slightly and obliquely truncated; carinal margin convex when 
continuous, but when formed of two almost equal portions, as is 
usually the case, the upper margin is convex and the lower margin 
almost straight; scutal or lateral margin straight, almost equal 
in length to the concave occludent margin, which has a slightly 
raised border; the shape of the upper carinal and occludent 
margins is influenced by the extent to which the upper half of 
the valve is curved towards the scuta, a character which is 
very variable. Along the occludent margin, especially in those 
examples conspicuously bowed away from the opposing tergum, 
a portion of the valve is bent downwards and sharply inwards to 
form an obtusely angular ledge on the inside of the valve; this 
ledge, as well as that formed on the inner occludent margin of 
the scutum, was no doubt for the protection of the animal’s body, 


FOSSIL CIRRIPEDES. 941 


for these ledges would serve to fill up the hiatus made through 
the upper portions of the scuta and terga being bowed away 
from each other. In some of the terga from the Chalk Marl of 
Cambridge and Burham, the adoccludent portion of the valve is 
not bent downwards and inwards to form a ledge on the inner 
surface, but extends outwards to form a second raised ridge 
almost parallel with the raised occludent margin. The terga from 
the B. plena- marls have this outer ridge finely denticulated on 
its margin. 

Upper latus (Pl. XCIV. fig. 8) subtriangular, slightly curved 
towards the scuta, almost flat : umbo slightly projecting ; at the 
scutal margin a narrow slip of the valve is bent downwards, and 
during the thickening and growth of the valve a slight ledge has 
been formed which has a tendency to project beyond the umbo ; 
tergal margin slightly convex; scutal margin slightly concave, 
and about the same length as the tergal margin: basal margin 
marked off into three almost equal lines, and to the angles thus 
made two feebly marked ridges extend from the umbo. 

Rostrum (Pl. XCLV. fig. 9) diamond-shape, bowed inwards; a 
strong rounded keel extends from the apex, widens gradually to 
the basal angle, and the portion of the valve on each side slopes 
steeply towards the lateral margins ; apical portion acute; basal 
portion rather less acute than the apical portion, with the basal 
angle slightly rounded. The inside of the valve is deeply concave. 

Structure and Affinities—The capitulum of Zeugmatolepas 
mockleri (see restoration, Pl. XCIV. fig. 14) closely resembles 
that of Pollicipes, especially i in the munmiber of valves, but differs 
(1) in the more specialized form of seutum which has a subcentral 
umbo, (2) in the size and position of the upper latera, which are 
elevated to occupy the whole of the interval between the scuta 
and terga, and (3) in the capitulum being longer than broad, and 
therefore of a more erect and Scalpellwm-like shape. In this 
form the umbo of the scutum appears generally to be almost 
apical in young valves, and to become further removed from the 
apex with age, until it reaches about one-third of the distance of 
the valve from the apex. Some individuals, however, have the 
umbo further removed from the apex than others of the same 
size. All the valves are exceedingly variable, much more so than 
in any fossil pedunculate Cirripede with which I am acquainted ; 
these variations are noted under the descriptions of the several 
valves. 

Zeugmatolepas mockleri agrees more closely with S. (?) crete 
Steenstrup* sp. from the Upper Senonian of Denmark than with 
any other species, and, indeed, S. (?) crete may quite well have 
been directly developed from 7. mockleri. Through the kindness 
of Dr. J. P. J. Ravn I have been able to examine examples of 
the known valves (scuta, terga, and carina) of S. (2) crete from 


* Steenstrup, J., Kroyer’s Neturinete Tidsskrift, Bd. i. 1837, p. 359; Bd. i, 
1839, p. 399, pl. v. figs. 1-3; Darwin, C. R., Pal. Soc. Mon Foss. Lepadidee, 1851, 
p. 45, pl. i. figs. Ll a-e. 


Proc. Zoou. Soc.—1913, No, LXITI, 63 


942 MR. T. H. WITHERS ON 


the collection of the Mineralogical Museum of the University of 
Copenhagen, and Dr. Ravn informs me that this species has been 
found only in the Upper Senonian (Zone of Scaphites constrictus). 
Darwin doubtfully referred S. (%) crete to the genus Scalpellum, 
and observed (1851, p. 45): “ Protessor Steenstrup was originally 
inclined to believe that the capitulum [of SS. (?) crete] was formed 
of only five valves; could this be proved, the species would very 
naturally rank with a small recent one from the Island of 
Madeira, which, owing to the upward growth of the scuta, and to 
certain peculiarities in the animal’s body, I have felt myself com- 
pelled to raise to the rank of a genus, under the name Oxynaspis.” 
An examination of the valves from Copenhagen University, and 
of some valves obtained by myself from the matrix of a specimen 
presented to the Geological Department of the British Museum 
by Copenhagen University, shows, at any rate, that we can no 
longer hold Steenstrup’s view that S. (?) crete had only five valves. 
Among the above valves are some small lower lateral plates, 
which, although not so well preserved as those from the English 
Cenomanian Chalk, obviously belong to the same general type as 
those of Zeugmatolepas mockleri. WS. (%) eretce, together with the 
species now described, is included, for the reasons previously 
given, in the new genus Zeugmatolepas. 

The scutum of Z. crete differs from that of Z. mocklert in that 
the umbo is further removed from the apex, being situated almost | 
midway on the occludent margin, and in a greater portion of the 
valve near the basi-lateral angle being sharply upturned almost at 
right angles to the middle part of the basal margin. The lateral 
(or tergal) portion of the valve is not steeply inclined from the 
umbo, and there is no deep trough, as in’ Z. mockleri, running 
parallel with the upper occludent margin. These and other difter- 
ences result, no doubt, from the difference in position of the umbo, 
and the consequent greater development of the tergal or lateral 
portion of the valve. The carina in Z. cret@ is relatively wider 
than in Z. mockleri, the central ridge is much more obscure, and 
in the tergum there is a much wider groove near the occludent 
margin. The whole of the valves in Z. crete are appreciably 
thinner. These differences, however, while deserving of specific 
distinction, appear to result from the further development of 
valves like those of Z. mockleri, and consequently I consider these 
two species to be closely and probably directly related. 


Genus CALANTICA. 


1825. Calantica Gray, ‘Annals of Philosophy’ (n. s.), vol. x. 


jo OM 

1907. Calantica Gray: Pilsbry, Bull. U.S. Nat. Mus. no. 60, 
Oh (oh 

1908. Calantica Gray : Pilsbry, Proc. Acad. Nat. Sci. Philadelphia, 
p. 106. 


Capitulum with two whorls of valves, the upper comprising 
paired scuta, terga, and a carina, the terga occupying the whole 


FOSSIL CIRRIPEDES. 943 


of the space between the scuta and carina; lower whorl com- 
prising three pairs of latera, a rostrum, and a subcarina. Umbo 
of all valves apical. 

The Oriental group, called by Pilsbry (1908) Calantica s. str., 
has the valves of the lower whorl low and wide, small, not 
concealing the bases of the valves of the upper whorl. 


TITANOLEPAS *, subgen. nov. 


Valves of lower whorl large, high, but not incurved, and over- 
lapping the bases of the valves of the upper whorl, Umbo of 
scutum subcentral and of the remaining valves apical. 

Genotype.—Titanolepas tuberculata. 


CaLAnTICA (TITANOLEPAS) TUBERCULATA Darwin sp. (PI. XCY. 
figs. 1-10.) 

1851. Scalpellum tuberculatuwm C. R. Darwin, Pal. Soc. Monogr. 
Foss. Lepadide, p. 43, pl. 1. fig. 10. 

1854. Scalpellum tuberculatum C. R. Darwin, Ray Soe. Monogr. 
Sub-class Cirripedia, Balanidz, Synopsis et Index Syste- 
maticus, p. 634. 

1854. Scalpellum tuberculatum C. R. Darwin: J. Morris, Cat. 
Brit. Foss. 2nd ed. p. 97. : 

1865. Scalpellum tuberculatum C. R, Darwin: J. W. Salter & 
H.. Woodward, Cat. & Chart Foss. Crustacea, p. 27, pl. 1. 
ime. 0 

Lot. Sealpelen tuberculatum COC. R, Darwin: H. Woodward, 
Brit. Mus. Cat. Brit. Foss, Crustacea, p. 143. 

1886. Scalpellum tuberculatum C, R. Darwin: J. Kafka, Sitz.- 
Ber. k. bohm. Gesell. Wiss. Prag (1885), p. 565, pl, i, 
Tae, “Lc 

1887. Geico tuberculatum C, R, Darwin: A, J. Fritsch & 

J. Kafka, Crust. Bohmischen Kreidef, p, 6, fig. 9. 

1888. Scalpellum tuberculatum C. R. Darwin; A, Peron, Bull, 
Soe. Sci, Yonne, vol. xli. (1887) p. 269. 

Diagnosis.—Valves with fine ribs radiating from their apices 
and standing out as blunt spines at each zone of growth, Carina 
strongly convex transversely, with the parietes narrow and 
striated. Scutum with the umbo situated about one-third the 
extent of the valve from the apex, and with two ridges extending 
from the umbo, one to the basi-lateral angle, and the other to near 
the rostral angle, Tergum elongately diamond-shape, with a very 
acute basal angle. Valves of lower whorl with the inner apical 
half flat and strongly marked with growth-lines. 

Remarking on this species, Darwin said (1851, p. 43): “Through 
the kindness of Mr. Harris, J have examined several valves, which 
1 believe to belong to the same species : the specimens were found 
in the chalk detritus [of Charing, Kent], and, therefore, may have 
come from the Upper or Lower Chalk or Chalk-marl; but more 
probably from the Upper Chalk,” 


* Tivavos = White earth, Chalk, 


Ce 
Os 
%* 


944 MR. T. H. WITHERS ON 


T. tuberculata has since been recorded by J. Kafka (1886, 
p- 965) from the Cenomanian of Kamajk, Bohemia, and by 
A. Peron (1888, p. 269) from the Turonian of Yonne, France. 
Kafka mentions a single tergum only, and if his figure is Aeon. 
it differs in certain “ohamneiers from the original tergum from 
Charing. Peron recorded a small and a very large carina, but 
gave neither measurements nor figures ; he drew attention to the 
fact that Darwin had supposed the species to come probably from 
the Upper Chalk, and said that so far as he knew it came from 
the ‘Turonian. 

T’. tuberculata occurs fairly commonly in the Cenomanian Chalk 

Marl near Cambridge, and we have over 150 valves, including 
99 valves of the lower whorl. From the Turonian 7Jerebr apna 
zone at Whyteleafe, Surrey, we have a single scutum, and from 
the Cenomanian Chalk Marl of Burham, Kent, a single rostrum. 

Type.—The collection of the late Mr. Wm. Harris from the 
Chalk detritus of Charing, Kent, was acquired from his daughter 
in 1881, by the British Museum (Natural History). Several 
valves of 7’. tuberculata are represented in the collection, three 
of which—a carina (I. 14657), scutum (I. 14659), and tergum 
(I. 14658)—are mounted together on paper, and with them is a 
label bearing the name ‘“ S. tuberculatum” in Darwin’s hand- 
writing, These are probably the original specimens figured by 
Darwin, but there is no means of proving this since Darwin gave 
no measurements of the figured valves of this species. A further 
label (not in Darwin’s handwriting) bearing the words “ very 
valuable” was with the specimens, but whether this was meant 
to indicate that they are Darwin’s original specimens must be 
left to individual opinion. I consider them to be the types, and 
since Darwin regarded the carina as the typical valve in Scalpellum, 
that valve is here taken as the holotype. 

Measurements.—The valves of this species are exceedingly 
small, and to judge from these, the largest capitulum could not 
have measured much more than 5 or 6 mm. in length. Of the 
upper series of valves the largest examples measure respec- 
tively :— 

Carina, I. 15850. Length 4:1 mm.; breadth 1:2 mm. 

Scutum, I. 15843. Length (from ‘apex to middle of basal 
margin) 2°5 mm.; breadth 2°6 mm. 

Tergum, I. 15851. Length 4:3 mm.; breadth 2°6 mm. 

Distribution. [¢ Cenomanian]| Chalk detritus: Charing, Kent ; 
Cenomanian, Chalk Marl: near Cambridge, and Burham, Kent ; 
Cenomanian, Korytzaner Schichten: Kamajk, Bohemia; Turonian, 
Terebratulina-zone: Whyteleafe, Surrey ; Holaster planus-zone : 
Armeau and Saint-Julien-du-Sault (Yonne), France. 

Description of Valves—Valves plainly ribbed longitudinally ; 
the ribs are narrow, but become more prominent on the edge of 
each zone of growth, where the ribs are thicker and usually stand 
out as short blunt spines. 

Carina (Pl. XCV. fig. 1) very narrow, either slightly bowed 


FOSSIL CIRRIPEDES. 945 


inwards, straight, or bowed outwards; tectum strongly convex 
- transversely, not carinate; basal margin almost rounded to 
bluntly angular; in young specimens, as shown by the periods 
of growth, the basal margin was almost straight; parietes slightly 
hens inwards, narrow, aie less than half ane width of the 
tectum, and marked with almost parallel growth-lines ; inner 
surface of valve deeply concave for the lower two-thirds, the 
upper third of the valve being solid and marked with lines 
of growth, indicating that the valve jrojected freely to that 
extent. 

Scutum (Pl. XCV. fig. 2) trapezoidal, umbo situated on the 
occludent margin about one-third the extent of the valve from 
the apex; almost flat to strongly convex ; four-sided, the lateral 
or tergal margin slightly longer than the basal margin, with 
which “it makes an angle of about 70°; the basal margin ‘much 
longer than the lower part of the occludent margin, with which 
it makes almost a right angle, the upper and lower portions of 
the occludent margin rane in an angle of about 135°; the upper 
occludent margin, said Darwin, “ homologically corr esponds with 
the tergal margin of the other eretaceous species and with the 
upper, nearly straight, portion of the occludent margin in the 
tertiary S. magnum and the recent S. vulgare.” The edge of the 
upper occludent margin is upturned and thickened, forming a 
prominent ridge, with its apex projecting beyond the lateral 
margin; a second prominent ridge extends from the umbo to the 
basi-lateral angle; a third ridge, barely perceptible in some 
specimens, extends from the umbo to the basal margin, near 
to the rostral angle. On the inner surface there is a deep 
hollow for the adductor muscle ; along the upper occludent margin 
the inner surface is slightly raised, which corresponds to the 
outer thickened ridge, the space between the two ridges being 
concave ; the inner ridge extends to the umbo, whence it becomes 
more pronounced and turns inwards, forming a prominent ridge 
above the hollow for the adductor muscle. 

Tergum (Pl. XCV. fig. 3) elongately diamond-shape, slightly 
angular transversely ; the occludent margin is thickened owing 
to the presence of a ridge on the outer and inner edge, corres- 
ponding with those on the upper occludent margin of the scutum ; 
close and parallel to the occludent margin is a ridge which 
becomes prominent near the scutal margin and extends beyond 
it; a very prominent straight ridge extends from the apex, and 
thickens towards the very acute ‘basal angle; scutal and lower 
carinal margins almost equal in length, and longer than the 
occludent and upper carinal margins, which latter are also of 
almost equal length and together make a right angle. 

Valves of the lower whorl (Pl. XCV. figs. 4-8). These are 
ornamented externally like the scutum, tergum, and carina. 
The apical half of their inner surface is almost flat and marked 
with lines of growth, showing that these valves must have pro- 
jected freely and overlapped the bases of the valves of the upper 


946 MR. 'T. H. WITHERS ON 


whorl ; the lower haif is considerably concave. Umbo of valves 
apical. The valves are much less modified than is the case with 
the valves composing the lower whorl in the more typical forms 
of Scalpellum, and consequently their position in the capitulum 
is not so readily determinable. Seventeen of the valves are 
symmetrical, and five asymmetrical. Of the symmetrical valves, 
nine have an extremely concave basal margin with produced 
lateral angles (Pl. XCV. fig. 4) and probably represent rostra ; 
ae others have a much less coneave basal margin (Pl. XCV. 

fig. 8) and may represent sub-carine. ‘The valves just mentioned 
are thought to be rostra and sub-carine, owing to the pronounced 
concavity of the inner part of the valve near the base. The 
three remaining symmetrical valves probably represent median 
lateral valves (Pl. XCV. fig. 6). They differ from the supposed 
rostra and sub-carine in the smaller concavity of the inner basal 
part of the valve, and in the growth periods being formed of 
two downward curves which meet on the median line, gradually 
becoming more obtuse towards the basal margin. Of the asym- 
metrical valves, two are subtriangular in shape with a recurved 
acute apex, and probably represent carinal latera (Pl. XCV. 
fig. 7); the three others, which may be rostral latera (Pl. XCV. 
fig. 5), are about twice as wide as long, and are divided un- 
equally by a fold extending from the apex to a position on the 
basal margin about two-thirds the distance from the rostral 
angle, the ‘basal mar gin on each side being concave 

Cirencinnre and A Fiiiion, —While Darwin had little doubt that 
the scutum and tergumn figured by him belonged to the same 
species, he did not feel so certain with regard to the carina, and 
he pointed out that this valve had a close general resemblance to 
the same valve in Pollicipes rigidus J. de C. Sowerby. A specimen 
of 7. tuberculata in the British Museum (Natural History), 
registered I. 14656, formerly in the Harris Collection, affords 
valuable evidence on this point, for it consists of the carina and 
the paired terga in position (PI. XCOV. fig. 9), and proves that 
the carina really does belong to the same species. Darwin 
stated also that the apex of the carina did not project freely in 
this species, but an examination of the carine from Charing, 
including the type, and of the numerous carine from the Chalk 
Marl of Cambridge, shows that the upper part of the inside of 
the valve is much thickened to one-third of its length from the 
apex (Pl. XCV. fig. 1c), and this portion is marked with 
growth-lines, showing that it must have projected freely to that 
extent. Proof of this is afforded by the specimen I. 14656 from 
Charing (Pl. XCV. fig. 9), in which the carina and terga are 
in position, for the carina there projects freely. 

The carina, scutum, and tergum only of 7. twberculata were 
known to Darwin, but the specimens from Cambridge include 
several other valves which enable us to add much to our know- 
ledge of the capitulum. The ornamentation of these valves is so 
characteristic that no doubt can be entertained as to their 
belonging to this species, Although there areas many as twenty- 


FOSSIL CIRRIPEDES. 947 


two valves *, other than scuta, terga, and carine, to judge by 
their structure, none appear to have formed upper latera, or at 
all events to fit between the scuta and terga, as do the upper 
latera in most forms of Scalpellum. Moreover, the form of the 
scuta and terga and the structure of their inner surface shows 
that the tergal margin of the scutum and the scutal margin of 
the tergum were in close proximity to each other. In this con- 
nection Darwin said (1851, p. 44): “I have no doubt that the 
ridge along the upper occludent margin of the scuta, and 
that on the occludent margin of the terga, together with 
their projecting points, are related to each other, owing to 
the close contact of these valves.” It follows, therefore, that if 
no upper latera were present, the upper whorl of valves in 
T. tuberculata compr ised only five valves, namely, a carina, a pair 
of terga, and a pair of scuta. The valves of the lower whorl, as 
mentioned in their description, must have overlapped the bases 
of the valves of the upper whorl, and if we are correct in desig- 
nating these valves as rostrum, rostral latus, median latus, 
carvinal latus, and sub-carina, the lower whorl would consist of 
at least eight valves, making thirteen valves for the whole 
capitulum (see restoration, Pl. XCV. fig. 10). 

In the number and arrangement of the valves of the capitulum 
Titanolepas tuberculata approaches the forms of Scalpellum 
grouped by H. A. Pilsbry 7 under the genus Calantica Gray, and 
comes nearest to those oaieeledl by him in the section Scile- 
lepas Seguenza, of which the type is S. carinata Phillipi sp., 
from the Pliocene of Messina, Sicily. The capitula of the species 
grouped under Calantica closely resemble that of Pollicipes, but 
differ in that there is only a single basal whorl of valves, com- 
prising three pairs of latera, a rostrum, and a sub-carina. The 
species included by Pilsbry under the section Scillelepas are 
chiefly characterized by the absence of an upper lateral plate in 
the upper whorl, which consequently consists only of five valves, 
and by the bases of the valves of the upper whorl being over- 
lapped by those of the lower whorl; the umbo in all the valves 
is apical. While 7. tuberculata agrees with these species in 
having five valves only in the upper whorl, and in the valves of 
the lower whorl overlapping those of the upper whorl, it differs 
from all the species included under the genus Calantica and the 
section Scillelepas by the umbo of the scutum being in a sub- 
central position. 

We have in 7. tuberculata, therefore, a species agreeing in 
all essential characters with the forms of Calantica (Scillcelepas), 
except that it has a more specialized form of scutum. The 

* Two further valves have since been noticed among the Cirripede material from 
the Chalk detritus of Charing, Kent, in the British Museum (Natural History). 
These are identical with the valves from Cambridge, which I consider to be rostral, 
and are registered I. 14663, I. 14664. A further rostrum has been found in the 
Chalk Marl of Burham, Kent. 

+ H. A. Pilsbry, “On the Classification of Scapelliform Barnacles,” Proc. Acad. 
Nat. Sci. Philadelphia, 1908, pp. 106-107; “The Barnacles _(Civripedia) contained 


in the Collections of the U.S. National Museum,” Bull. U.S. Nat. Mus. No. 60, 
1907, pp. 8-9. 


948 ON FOSSIL CIRRIPEDES, 


fossil species that can be definitely referred to Calantica (Scille- 
lepas) are found in Tertiary rocks, viz. Scillelepas parone 
from the Miocene of the neighbourhood of Turin, Italy, and 
S. carinata and S. ornata from the Pliocene of Messina, Sicily, 
so that it is an important advance to show that a representative 
of the genus existed so far back in time as the Cenomanian. It 
is certainly remarkable to find that 7’. twberculata had a more 
highly specialized seutum than the Tertiary and recent species, 
and to mark this advance in the evolution of the group, I place 
it in a new sub-genus, 7itanolepas. This course is the more 
desirable since 7. twberculata may represent a side line of develop- 
ment from the ancestral Scillelepas line, which, by the sup- 
pression of the lower whorl of valves, and by the umbo of the 
carina becoming sub-central, may eventually have given rise to 
the genus Oxynaspis. 

For help in connection with this paper, I wish to express my 
indebtedness to Dr. F. A. Bather, Dr. W. T. Calman, Mr. C. P. 
Chatwin, and Dr. H. A. Pilsbry. 


EXPLANATION OF THE PLATES. 


All the specimens here figured are in the collection of the Geological 
Departinent of the British Museum. 


Prats XCIV. 
Zeugmatolepas mockleri Withers, gen. et sp. n. 
Fig. 1, Carina. Cenomanian, Chalk Marl: near Cambridge. I. 15820. a, outer 
view; b, side view; c, inner view. 
2. Carina, I. 15821. a, outer view; 6, side view ; c, inner view. 
3, 4, 5, 6. Scuta (various forms), I. 15822-I. 15825. ) 
7. Tergum, I. 15826. (a, outer view; 6, inner 
8. Upper latus, I. 15827. r view. 
9. Rostrum, I. 15828. y 
10. Several valves in position, including the carina, I. 15829. 
11. Holotype. Nearly complete capitulum showing the left side, I. 15830. 
12. Several valves in position, I]. 15831. 
13. Nearly complete capitulum showing the right side, I. 15832. 
14. Restoration of capitulum based on the above material from the Chalk Marl 
of Cambridge. ¢., carina; 6., scutum; ¢., tergum ; u.l., upper latus ; 
r., rostrum; /.7., lower latera; s.c., sub-carina. 


Figs. 1, 2, 4, 9-18, 10 diameters; figs. 3, 5-8, 14, X 5 diameters. 


——— 


Prate XCV. 
Calantica (Titanolepas) tuberculata Darwin sp. 


Ki 


oR 
e 


. Carina. Cenomanian, Chalk Marl: near Cambridge. I. 15842. a, outer 
view ; 6, side view; c, inner view; d, transverse section hear apex. 

. Scutum, I. 15843 (figure reversed). 

. Tergum, I. 15844. 

. Rostrum, I. 15845. | 

. Rostral latus, I. 15846. r a, outer view ; 6, inner view. 
Median latus, I. 15847. | 

. Carinal latus, T. 15848 (figure rever rsed). 

. Sub-carina, I. 15849. 

. Carina and pair of terga in BART [? Cenomanian| Chalk detritus ; 
Charing, Kent. J. 14656. 

Restoration of capitulum, based on disconnected valves from the Chalk Marl 
of Cambridge. c., carina; s., scutum; ¢., tergum; r., rostrum ; 7./., rostral 
Jatus; m.7., median latus; e./., carinal latus; s.c., sub-carina. 


Figures 1-10 X 10 diameters. 


sae Gene re 


= 
© 


ed pet 


cf; 


oe 


Ins Zo So ISIS, Il, XCVI, 


BATRACHIDERPETON LINEATUM. 


ae 


Noe ee 


Ins B. SB, UNS, IPM, SCV, 


BATRACHIDERPETON LINEATUM. 


ON A COAL-MEASURE STEGOCEPHALIAN. 949 


55. Batrachiderpeton lineatum Hancock & Atthey, a Coal- 
Measure Stegocephalian. By D. M.S. Watson, M.Sc., 
Lecturer on Vertebrate Palzeontology in University 


College, London *. 
[Received May 20, 1913: Read October 28, 1913.] 
(Plates XCVI. & XCVII.,f and Text-figures 162-167.) 


INDEX. Page 
Miorpliol 0 oye tanta tra his. negroes juice | OO. 
Systemiatl ee caer askaeecesd wesc eee ee eee OOO 


Amongst the wonderful series of vertebrate fossils in the 
Hancock Museum, Neweastle-on-Tyne, derived from the Low 
Main Seam of Newsham Colliery, a small stegocephalian Batrachi- 
derpeton is represented by excellent material, which is described 
as completely as possible in this paper. 


SxuLu.—WMaterial. 


I. A nearly complete skull with the ornament and palate well 

preserved. The type specimen. (Atthey collection.) 

II. A complete skull showing with great clearness all the 
sutures on the top of the head. (Atthey collection.) 

III. A much disarticulated skull of importance in ascertaining 
the structure of the palate. (Atthey collection.) 

IV. Half of a badly-preserved skull, showing well the maxillary 
and palatine teeth. (Dinning collection.) 


Shape of the Skull. 


All the specimens are crushed flat, so that the shape of the 
skull has to be settled by indirect means. It is triangular in 
plan, the postero-lateral corners being much produced into long 
‘cornua.” Specimen II. shows that there were almost vertical 
cheeks, as on the right side this part was turned under and 
crushed on to the lower surface of the roof of the skull, whilst 
the left side is pressed outwards. This crushing has only very 
slightly distorted the palate, and the distance between the two 
quadrates is almost identical with that between the spot where 
the cheek is turned under on the right side and the corresponding 
point on the left side. 

There is a pineal foramen of medium size almost exactly half- 
way between the point of the nose and the back of the head in 
the middle line. The orbits are laterally placed and lie entirely 
in front of the middle of the skull. The small anterior nares are 
also laterally placed. 

The structure of the cranial roof will be best understood from 


* Communicated by Prof. J. P. Hin, D.Sc., F.R.S., F.Z.S. 
+ For explanation of the Plates see p. 962. 


950 MR. D. M. S. WATSON ON A 


text-figs. 162 & 164. All the sutures drawn with solid lines in 
these figures can be clearly seen in specimen IT. 
The interesting features are the great development of the 


Text-fig. 162. 


Batrachiderpeton lineatum. Dorsal aspect of the restored skull and clavicular 
apparatus. X13. Skull mainly founded on specimen II., from which all the 
sutures are taken, and the pectoral girdle restored from the bones of specimen III., 
which is a smaller individual, enlarged to correspond in size with the skull. 


Reference letters :—Cu., Clavicle; Cm., Cleithrum; Fr., Frontal; I.Cu., Inter- 
clavicle; Lac., Lachrymal; Mx., Maxilla; P.Mx., Premaxilla; P.O., Post- 
orbital; P.Par., Postparietal; Par., Parietal; Po.FR., Postfrontal; PR. FR., 
Prefrontal ; Sq., Squamosal; Tas., Tabulare. 


COAL-MEASURE STEGOCEPHALIAN, 951 


horns, formed entirely by the squamosals and tabulares, and 
the correspondingly large size of the post-parietals. 
The extremely small size of the nasals is an interesting feature. 
The sutures separating the lachrymals from the premaxille are 
not certain, but there is a separation and displacement of the 
bones at the dotted line which has probably taken place along 
the suture. 


Text-fig. 1635. 


Batrachiderpeton lineatum. , Ventral aspect of the restored skull and clavicular 
apparatus. X13. Skull mainly from specimen I., sutures from specimen III., 
clavicles, etc., from specimen III. 


Reference letters as before with :—Quv., Quadrate; Pat., Palatine; PaR.Oc., Pare 
occipital process; Par.Sp., Parasphenoid; Pr.V., Prevomer; Pr., Pterygoid. 


952 MR. D. M. 8. WATSON ON A 


The palate is of great interest; in the restoration the general 
features are drawn from specimens I., I1.,and 1V., and the sutures 
are introduced from specimen ITI., where the pterygoids, palatines, 
and prevomers are disarticulated. 

Basis cranii.—The basis cranii is much crushed in all the 
specimens and was apparently very thin and delicate. 

The two large exoccipital condyles are clearly shown in I. 
and Il. They are rounded from front to back and to a less 
extent from side to side. The bone between them is extremely 
thin, but no doubt is partly composed of basioccipital. The basi- 
sphenoid in I. is slightly better preserved ; it bears two very small 
basipterygoid processes only very obscurely indicated. In front 
of this region it is produced into a narrow rostrum undoubtedly 
parasphenoid. 

Extending outwards and backwards from the sides of the basis 
cranii are two long, very delicate processes which are shown, 
much crushed and broken, in specimens I., II., and III. In 
specimen I. on the right side, a small foramen is rather obscurely 
indicated on the process about a third of its length from the 
middle line, this opening can only be the fenestra ovale. The 
process is probably mainly opisthotic, with the prootic forming 
some of its front border. Immediately above the exoccipital 
condyle in the same skull is a foramen for cranial nerves ix. 
and x. 

The outer end of the opisthotic articulates as usual with a facet 
on the under surface of the tabulare. 

The quadrates are well shown in specimens J. and It. They 
are small bones with a well-developed trochlear condyle, the outer 
border is articulated with the quadrato-jugal, there being 
probably no foramen between the two bones. 

The upper end of the quadrate is supported by a downwardly 
directed flange from the squamosal, well seen in specimens I. 
and III. The inner border of the quadrate is continued by 
a short thin pterygoid process (specimen I.). 

The pterygoid isa large bone which articulates by a small facet 
with the basipterygoid process of the basisphenoid (specimen I.). 
Behind this it is continued as a deep but thin plate, the posterior 
ramus, which articulates distally with the quadrate and whose 
upper border has a suture with a continuation of the descending 
flange of the squamosal mentioned above. The anterior part of 
the bone forms a broad flat plate covered with a shagreen of small 
sharp teeth. The anterior border of the plate is in contact with 
the prevomer and the lateral border with the palatine. 

The palatine bears a single row of nine medium-sized swollen 
teeth, and its outer border articulates with the jugal and maxilla. 
There is apparently no transpalatine. 

The prevomer is a small bone with a median symphysis with 
its fellow; it forms a good deal of the inner border of the posterior 
nares and bears two teeth placed transversely ; its exact relations 
to the palatine are not certain. 


COAL-MEASURE STEGOCEPHALIAN. 953 


The premaxilla appears to bear three teeth, but the suture 
separating it from the maxilla with six teeth is not quite certain. 


Text-fig. 164. 


Batrachiderpeton lineatum. Weft lateral aspect of the restored skull and clavicular 
apparatus. Skull from specimen IT., clavicle, etc., from specimen III. 1. 


Reference letters as in former figures with :—Jv., Jugal; Qu.Ju., Quadrato-jugal. 


Lower Jaw.—Waterial. 


Specimen III. The perfect left ramus and disarticulated parts 
of the right ramus of the same individual. Although the 
posterior part of the right ramus, composed of the angular, 
surangular, and prearticular, is not well preserved, this specimen 
is of great importance because it gives the most satisfactory 
evidence of the front of the mandible, the dentary and splenial 
being slightly separated and completely free from matrix on 
the outer and largely so on the inner side. 

Specimen V. <A perfect right ramus showing well the suture 
between the prearticular and the angular. (Atthey collection.) 

Specimen VI. A perfect right ramus. (Barkas collection.) 

Specimen VII. An isolated dentary. (Atthey collection.) 

The various specimens are crushed in different directions and 
suggest that the jaw had a considerable cavity and was perhaps 
nearly circular in cross section. 

There is a distinct though short postarticular process. 

The articular is well ossified though small; it bears an articular 
facet adapted to the trochlea quadrati. 

The suture between the surangular and angular is not clearly 
shown in any specimen. The angular is a very large bone 
extending nearly up to the symphysis; it forms the lower part 
of the jaw, and its inner border articulates by a suture (spe- 
cimen V.) with the prearticular, from which it is separated by 
a smali oval internal mandibular vacuity. 

‘The prearticular 1s a large bone covering the inner side of the 

articular and extending well forward. 

The dentary is a fairly large bone overlapping the surangular 
and angular behind; it bears eleven or twelve swollen teeth and 
has a symphysis with its fellow; its lower border has a suture 


954 MR. D. M. S. WATSON ON A 


with the splenial. The splenial has a symphysis with its fellow 
and is entirely a bone of the outer side of the jaw, as is 
conclusively shown in specimen ITI. 


Text-fig. 165. 


Batrachiderpeton lineatum. Right ramus of the lower jaw, outer aspect. X 2. 
Outline and sutures from specimen ITT. 


Reference letters:—Ane., Angular; Den., Dentary; Sp., Splenial; Sur.Ane., 
Surangular. 


The complete jaws show extremely well a great patch of small 
conical teeth lying inside those of the dentary. Specimen III. 
shows conclusively that these do not belong to the dentary, the 
splenial, the angular, or the prearticular, so that it is quite 
certain that there is another element, which can only be the 
coronoid. 


Text-fig. 166. 


Batrachiderpeton lineatum. Left ramus of the lower jaw, inner aspect. X 2. 
Drawn from specimens III., V., & VI. 


Reference letters as on text-fig. 165 with :—Cr., Coronoid; Pr.Arv., Prearticular. 


Between the coronoid, prearticular, and angular there is a small 
vacuity. 


Pectoral girdle. 


Specimen ITI. retains both clavicles and a cleithrum. 

The clavicle is composed of a flat ventral plate with an orna- 
ment of strong radiating ridges, from the upper surface of which 
within the lateral border an almost cylindrical process rises 
abruptly, this slopes slightly backwards and outwards and gives 
attachment to the cleithrum. The dorsal surface of the ventral 
plate of the clavicle shows clearly the impression of the part of 


COAL-MEASURE STEGOCEPHALIAN. 955 


the interclavicle which overlaid it, and enables the width of the 
latter bone to be accurately determined. 


Text-fig. 167. 


Ceraterpeton reticulatum H. & A. Restored dorsal surface of skull and 
clavicular arch. X 4. 


The cleithrum is a T-shaped bone, flat, and ornamented by 
fine grooves, the upright of the T is affixed to the vertical process 
of the clavicle. 


Discussion of the Skull. 


One of the most interesting features of the skull is the reduc- 
tion of the number of bones in the temporal region. The 
primitive amphibian has in the temporal region three bones, the 
squamosal, supratemporal, and intertemporal, all of which occur 
in the primitive reptile Seymouria. It is important to discover 
which of these bones is really present in Batrachiderpeton. 

In the primitive amphibians “ Lowomma” and Pteroplaa all 
three bones are present in a typical manner and also the 
squamosal; that is, the outer bone, which articulates with 
the quadrato-jugal, is well characterized and differentiated frem 
the other two by the fact that its posterior border is bent down 
in front of the otic notch and articulates with the upper edge of 
the quadrate ramus of the pterygoid, it also passes down behind 
the quadrate itself, overlapping a good deal of its posterior 
surface. Neither the intertemporal nor supratemporal has any 
connection whatever with the pterygoid or quadrate. 

The conditions in these primitive amphibia are,those which 


956 MR. D. M. S. WATSON ON A 


always occur in the large Stegocephalia, modified of course by 
changes in the shape of the skull. I have seen them in :— 
“« Loxomma,” Pteroplax, Batrachosuchus, Cyclotosaurus, Plagio- 
sternum, Trematosaurus, Micropholis, Capitosaurus, Aphaneramma, 
Bothriceps, Archegosaurus. 

In Batrachiderpeton the squamosal is united to the pterygoid, 
and covers the back of the quadrate, not by the turning down 
of its hinder edge but by a special descending flange, a difference 
depending entirely on the development of the ‘‘ horn” which has 
carried the otic notch, which, in all amphibia and reptiles where 
it is present, lies between the tabulare and the squamosal, far 
out and backwards. The great sheet of squamosal which les 
behind the quadrate is in fact a new development, the original 
posterior border being now represented by the flange. 

There is hence no doubt that the single temporal bone re- 
maining in Batrachiderpeton is the outer of the original three. 
That this is really the homologue of the mammalian squamosal, 
I hope to be able to show in a subsequent paper. The other 
bones of the top of the skull do not call for special notice. ; 

The palate of Batrachiderpeton is of very great interest. I 
pointed out recently that the type of palate found in ““Loxomma,” 
with a large single basioccipital condyle, basipterygoid processes 
on the basisphenoid, and large pterygoids, which leave only a very 
small interpterygoid vacuity divided by a narrow parasphenoid, is 
the primitive type of amphibian palate, because it agrees very 
closely with that of the Crossopterygian fish, and because no 
palate with large parasphenoid and interpterygoid vacuities is 
known to occur in the Coal-Measures or Lower Carboniferous. 
From known types it is possible to pick a morphological although 
non-phylogenetic series showing exactly how the change from the 
primitive palate to the typical palate of Capitosaurus has taken 
place. Hryops, which, as shown by v. Huene’s recent extremely 
interesting paper, has a quite large basioccipital and basipterygoid 
processes of the basisphenoid, and in which the parasphenoid and 
interpterygoid vacuities are only of moderate size, is exactly 
intermediate between the primitive type and that found in the 
Triassic forms, where the basioceipital and basisphenoid are very 
much reduced and the pterygoids are supported by sutural 
union with the edges of the parasphenoid. The palate of Batra- 
chiderpeton agrees with the primitive type, in having large 
pterygoids meeting in the middle line and articulating with 
basipterygoid processes of the basisphenoid, It differs, however, 
in that it has already completely replaced the basioccipital con- 
dyle by a pair of exoccipital condyles, just as have the later 
large Stegocephalia. 


Iscussion of the Lower Jaw. 


Except that it is not known whether an epicoronoid was present 
or not, the lower jaw of Batrachiderpeton is extremely like that 


COAL-MEASURE STEGOCEPHALIAN. 957 


of “Loxomma” in its structure, particularly in the fact that the 
splenial is entirely a bone of the outer side of the jaw, which 
is proved in the most conclusive manner by specimen III. 

In later Stegocephalia the splenial (as shown, for example, 
in Lryops, Anaschisma, Bothriceps, and “ Labyrinthodon ” lepto- 
gnathus), although it still has a good exposure on the outer 
surface, has also a large flange running up on the inner side of 
the jaw to the coror.oid 

In Reptilia the internal part of the splenial is always larger 
than the external, and in later reptiles this latter part disappears 
altogether. 

It thus seems certain that the trend of evolution of the splenial 
has been directed to gradual growth of an internal flange at the 
expense of the external part of the bone which alone occurs in 
the primitive amphibia. 

In correlation with this development is a reduction of the 
coronoid and a complete loss of the epicoronoid. 

As the angular is to all appearance a serial homologue of the 
splenial, it is probable that it may have tended in early Amphibia 
to pursue a similar course of change, in which case we may 
suppose that in primitive forms the suture between the angular 
and the prearticular probably ran along the lower border of the 
jaw just as does the suture (when there is one) between the 
coronoid and the splenial. 

This type of jaw, which we have arrived at hypothetically, is 
actually realized in Crossopterygian fishes, Megalichthys being the 
best type. 

The structure of the jaw in Paleozoic Crossopterygians was first 
accurately made out by Traquair in Rhizodus and Rhizodopsis. 

I give here an account of the jaw of Megalichthys and have also 
examined those of Holoptychius and Osteolepis, which agree in 
general structure. 

In Megalichthys the dentary is a large bone narrowing from 
front to back, where it ends in a point. It bears throughout its 
border a single row of small pointed teeth, within which at the 
anterior end are one or two large “laniary” teeth. Its lower 
border overlaps three bones, the splenial, angular, and sur- 
angular. 

The splenial has an articulation with its fellow; it lies entirely 
on the outer side of the jaw and its lower border is entirely free. 
Its oblique posterior edge overlaps the angular. 

The angular is a bone similar to the splenial, except that it 
has of course no symphysis and its lower border articulates with 
the coronoid. a 

The surangular is overlapped by the angular, which it greatly 
resembles ; it covers the outer side of the well-ossified articular. 

* The combined prearticular and coronoid is a very large bone 
running forward from the inner side of the articular nearly to 


* A specimen of Glyptolepis paucidens in the Manchester Museum seems to 
show a clear suture between the short prearticular and the coronoid. 


Proc. Zoou., Soc.—i913, No. LXIV. 64 


958 - MR. D. M. 8. WATSON ON A 


the front of the jaw ; in front it ends freely in a point, its lower 
border articulates with the angular and surangular, and its 
upper border with the epicoronoids; the inner surface of the 
bone is covered with a shagreen of small denticles. 

The epicoronoids are two small bones lying between the upper 
edge of the coronoid and’ the dentary. The posterior is a short 
bone forming the front of the suprameckelian vacuity, and 
bearing two Tar ge teeth and the pits for their replacing teeth. It 
is separated from the anterior epicoronoid by a large round 
foramen. ‘The anterior bone bears one tooth and its replacing 
tooth-socket. 

The jaw of Megalichthys differs slightly from that of Ahizodus, 
Rhizodopsis, etc., in there being only three bones besides the 
dentary on the outerside of the jaw, instead of four, and only two 
epicoronoids instead of three. 

Comparison of this Jaw with that of “ Zovomma” will, I think, 
justify the nomenclature used here, even the direction of overlap 
of the bones being the same in the two types. 


Systematic position of Batrachiderpeton. 


The classification of the smaller stegocephalian Amphibia, so 
abundant in the Coal-Measures and Permian Rocks of Europe 
and North America, is in such confusion, to which some 
recent work has added, that it is at present only possible to 
proceed by reference to individual specimens which have been 
well described. The difficulty of identifying species and the 
rashness with which genera often founded on very imperfect 
materials have been extended to include other types often of 
very distinct structure, combined with the great technical 
difficulties of working on small and often badly-preserved 
animals, make any wide divisions at present of very doubtful 
value. 

Batrachiderpeton in the forward position of its orbits and the 
ereat ‘“cornua” at once recalls Ceraterpeton, originally described 
by Huxley from Kilkenny. The type-specimen of this latter 
genus is badly preserved, so that the sutures of the cranial roof 
are not clearly shown, although it seems probable that the 
apparent sutures are real, in which case there are many resem- 
blances between the two types. 

To the type-species Ceraterpeton galvani two other specimens 
have been assigned—Ward’s specimen from North Staffordshire 
described by C. W. Andrews, and a specimen from Kilkenny 
figured by A. Smith Woodward. These two specimens agree 
extremely closely in size and in the shape of the skull, but they 
differ from the type in being only about one-half of the size and 
in having a narrower skull. In the Hancock Museum there are 
three specimens closely similar from the Low Main seam of 
Newsham, one of which is the type of Hancock and Atthey’s 


COAL-MEASURE STEGOCEPHALIAN. 959 


Urocordylus reticulatus. These three specimens agree very closely 
in measurements and in all characters which can be observed ; 
they are undoubtedly cospecific. They also agree exactly with 
Andrews’ and Smith Woodwaxd’s specimens in the proportions of 
the skull and body, and in size, but differ from Andrews’ specimen 
very markedly in the character of their cranial ornament, which 
is almost entirely composed of ridges instead of pits. They must 
hence be regarded as specifically distinct. 

In text-fig. 167 (p. 955) I have given a drawing of the top of 
the head with such sutures as are clearly visible in one of the 
Neweastle specimens preserved in counterpart; this should be 
compared with Dr, Andrews’ figure. It is certain that the ptery- 
goids met in the middle line, as in Batrachiderpeton, although 
the detailed structure of the palate cannot be made out. 

One important feature, shown clearly by the type-specimen of 
‘“« Urocordylus” reticulatus, is the T-shaped cleithrum, also shown 
in Dr. Woodward’s figure, where it is interpreted as scapula. 
Such a shaped bone is known in no other animal except Batrachi- 
derpeton, and seems to show definitely that ‘* Urocordylus”’ 
reticulatus and Datrachiderpeton are closely related. ‘The very 
remarkable clavicle is also identical in the two types. 

Whether Urocordylus reticulatus H. & A. really belongs to the 
genus Ceraterpeton is uncertain, but it is probable that it does ; 
in any case, by calling it Ceraterpeton reticulatwm in full, no 
trouble will arise ; it seems probable that it is not Urocordylus. 

One of the most marked features of Ceraterpeton is the ex- 
pansion and corrugation of the distal ends of the neural and 
hemal spines; the meaning of this condition, which occurs 
throughout the column, is obscure. It also occurs in Ceraterpeton 
galvant, Urocordylus wandesfordii, Scincosaurus crassus, Ptyonius 
estocephalus, and other types. As there is no definite evidence 
that these types cannot be related, 1¢ is natural to assume that 
they are; but it must be remembered that any such relation 
can only be slight, for if Fritsch’s restoration of the skull of 
Scincosaurus crassus is at all correct, it differs very greatly from 
C. reticulatum and Batrachiderpeton. 

Another type which appears to be related is Diceratosaurus 
punctolineatus Cope, recently deseribed by Jekel. The palate 
of this type is extremely like that of Batruchiderpeion, differing 
in minor features of the dentition, in the larger interpterygoid 
vacuity, and in the presence of a transverse bone. 

If we suppose, as is quite probable, that some of the sutures 
of the cranial roof were invisible, it is probable that the skull 
structure was very like that of Ceraterpeton reticulatum. Ou the 
other hand, this type has not the T-shaped cleithrum and peculiar 
neural arches of the latter form. 

Assuming, as seems justifiable, that Batrachiderpeton is allied 
to Ceraterpeton reticulatum, it is of interest to see the differences 
between the two types. Ceraterpeton reticulatwm is obviously the 

64* 


960 MR. D. M. 8. WATSON ON A 


less modified, and the following list shows the direction of change 
in passing to Batrachiderpeton :— 


1. Increase in the relative size of the “ cornua”’ involving 
larger squamosals, postparietals, and tabulares. 

2. More anterior position of the orbits. 

3. Great decrease in size of the nasals. 

4, Broadening of the clavicles and interclavicle. 


The most interesting comparison, however, is with Diplo- 
caulus, a type which Jzkel has already brought into relation 
to Ceraterpeton. 

Comparison of text-fig. 162 (p. 950), the dorsal surface of the 
skull of Batrachiderpeton, with the figures of the skull of 
Diplocaulus given by Williston & Case, shows a very striking 
resemblance, particularly in the way in which the horn is 
developed, by enlargement of the squamosal (prosquamosal of 
Williston & Case), tabulares, and postparietals (supra-occipital 
plate of Williston & Case). 

If the bone regarded by Williston as a nasal is compared with 
the undoubted lachrymal of Batrachiderpeton no doubt of its 
identity can arise, it being evident that the nasals, extremely 
small in Batrachiderpeton, are quite lost in Diplocaulus. 

It also seems extremely probable that the bone called 
squamosal by Williston & Case is really the postorbital dragged 
out from the border of the orbit by the development of the 
horn. 

The only other important difference shown on the top of the 
skull lies in the fact that in the earlier type the frontal is 
excluded from the orbit, whereas in the American form it forms 
much of the border of that opening. ‘This depends on the 
different position of the orbits, which in Batrachiderpeton are 
laterally directed, whilst in Dzplocaulus they look directly 
upwards. It is in every way probable that this difference 
depends entirely on the habits of the animals in question. 

The skull of Batrachiderpeton shows no grooves for lateral line 
organs, and it seems likely that the animal was not very 
aquatic ; and as its skull was probably of only moderate size, it 
is possible that it pursued an ordinary life. 

Case, apparently justly, regards Diplocaulus as living in the 
mud at the bottom of ponds, and its head is so enormous 
that it must have been carried flat on the ground. Such a habit 
of life is inconsistent with laterally placed eyes, which would 
therefore necessarily migrate to the dorsal surface as in all flat- 
bodied bottom-dwelling fishes, e. g., Lophius, Raia, Nolea. 

The clavicular apparatus of Diplocaulus is very similar to that 
of Batrachiderpeton. The clavicles in both types consist of a flat 
ventral plate, from the upper surface of which a cylindrical 
process rises dorsally within the lateral edge. 

The vertebral column of Diplocaulus is now well known. The 


COAL-MEASURE STEGOCEPHALIAN, 96] 


presacral vertebra, 16 in number, have an elongate hour glass- 
shaped “centrum ” and elongated neural arches with low elong gate 
spines. There are two processes for the rib, the upper Grmied by 
the arch, the lower by the “centrum.” There are hypapophysial 
in addition to the ordinary zygapophysial articulations. The 
caudal vertebree have no rib articulations and expanded hemal 
arches fused into the “ centrum.” 

The vertebral column of Ceraterpeton is not well known, but 
the Newcastle specimen of C. reticulatwm seems to show the 
following characters:—'There are about 18 presacral vertebre 
with hour-glass-shaped centra, long neural arches and spines 
almost eontainly with hypayophysial i in addition to the ordinary 
articulations, and a single process neaily in the centre of the 
vertebra for the rib; this may be partly carried by the arch. 
The later caudals differ in having no rib articulation and in 
having large expanded hemapophysial spines fused onto the 
centrum. 

The vertebral columns of the two types are, in fact, very 
similar, differing apparently only in the fact that in Diplocaulus 
the head of the cba is double, whereas in Ceraterpeton reticulatwm 
it 1s single. 

The shortness of the neural and hemal spines of Diplocaulus 
may be due merely to the flatness of that animal. 

The lower jaw of Diplocaulus as described by v. Huene agrees 
very well with that of Batrachiderpeton, the inner row of teeth 
at the front being probably borne on the real coronoid, and 

. Huene’s complementare being an epicoronoid. 

In fact, the only important difference between the two types 
lies in the palate. Dzplocaulus differs in its palate from Batrachi- 
derpeton exactly as does Hryops from Loxomma. 

The dentitions in the two types are identical in general lines. 
The differences are that in Batrachiderpeton there are large 
pterygoids meeting in the middle lne and supported by the 
basisphenoid, in Diplocaulus there are large interpterygoid 
vacuities and the pterygoids are supported by the parasphenoid. 

I have already endeavoured to show that in the large Stego- 
cephaha this change actually takes place, a view which if correct 
removes all ditticulty of regarding Batrachiderpeton and Diplo- 
caulus as very closely allied. 

Finally, it may be noticed that the characters in which 
Diplocaulus differs from Batrachiderpeton are exactly those in 
which the latter differs from Ceraterpeton :— 


1. Increase in relative size of the cornua, involving larger 
squamosals, postparietals, and tabulares. 
More anterior position of the orbits (and their dorsal 
position). 
3. Complete loss of the nasals. 
4. Broadening of the clavicles and interclavicles. 


962 ON A COAL-MEASURE STEGOCEPHALIAN. 


I wish to express my thanks to the Committee of the 
Northumberland and Durham Natural History Society and to 
K. L. Gill, Esq., Curator of the Hancock Museum, for the 
kindness with which they have allowed me to work on the- 
wonderful material in their Museum. 


BIBLIOGRAPHY. 


AnpreEws, C. W.—Note on a Specimen of Keraterpeton galvani 
Huxley, from Staffordshire. Geol. Mag. new ser. Dee. iv. 
vol. 11. pp. 81-84. 1895. 

Bror, EF. v.—Hin Beitrag zur Kenntniss von Diplocaulus 
Cope. Centr. f. Min. Geol. u. Pal. 1902, pp. 536-541. 
Casn, KE. C.—Revision of the Amphibia of the Permian of 
North America. Carnegie Institution, Washington, pub. 

no. 146, 1911. 

J @KEL, O.— Ueber Ceraterpeton, Diceratosaurus u. Diplocaulus. 
Neues Jahrb. f. Min. Geol. u. Pal. 1903, Heft 1, pp. 109- 
134. 

Huxtry, 'T. H.— Description of the Vertebrate Remains from the 
Jarrow Colliery. Proc. Roy. Irish Acad. vol. xxiv. p. 353. 
1867. 

Traquair, R. H.—On the Structure of the lower jaw in Rhizo- 
dopsis and Rhizodus. Ann. & Mag. Nat. Hist. ser. 4, 
vol. xix. pp. 299-306. 1877. 

Watson, D. M. 8.—-The larger Coal-Measure Amphibia. 
Manch. Mem. vol. lvl. pt. 1, pp. 1-13, pl. & figs. 1912. 

Wuuurston, 8. W.—The Skull and Extremities of Diplocaulus. 
Trans. Kansas Acad. Sci. 1909, pp. 123, 131. 

Woopwarp, A. S.—On a New Specimen of the Stegocephalian 
Ceraterpeton galvani. Geol. Mag. new ser. Dee. iv. vol. iv. 
1 AB, LOA 

Hancock, A., and Arrnry, T.—Description of a Labyrinthodont 
Amphibian, a New Generic Form, obtained in the Coal 
Shale at Newsham, near Newcastle-upon-Tyne. Ann. & 


Mag. Nat. Hist. ser. 4, vol. vi. pp. 56-65, pl. 1. 1870. 


EXPLANATION OF THE PLATES. 


Prats XCVI. 


Fig. 1. Skull of Batrachiderpeton lineatum H. & A. Specimen I. (type-specimen), 
14. Dorsal aspect. 
2. Same specimen as fig. 1. Palatal aspect. 
3. Rami of the lower jaw of Batrachiderpeton lineatum, X 2. 


3 


A. Specimen VI. Right ramus, outer aspect. 
B. Specimen III. Left ramus, inner aspect, showing very clearly the patch 
of teeth on the coronoid. 


Prare XCVII. 
Fig. 4. Skull of Batrachiderpeton lineatum H. & A. Specimen II., X 1. Dorsal 


aspect. 
5, Same specimen as fig. 4. Palatal view 


West, Newman 


1-4. PEACHIA QUINQUECAPITATA. 
5-7. BICIDIUM AQUOREZ 8.B.PARASITICUM. 


ON NEW ACTINIANS FROM BRITISH COLUMBIA. 963 


56. On two New Actinians from the Coast of British 
Columbia. By J. PLayrarr McMourrtcu, C.M.ZS. 


[Received August 1, 1913: Read November 11, 1913.] 
(Plate XCVITI.*) 


INDEX. 
Systematic :— Page 
FZCACHIOAGUUNO MECH ILC, ISP ep Mey ersceee steelers eects seieeeect = 963 
Bi cidenmeguoncces Spa Um ceseeeentaee eee een OO 


The two forms described below were obtained during a visit to 
the Marine Station maintained by the Canadian Government at 
Departure Bay, Vancouver Island. They are of especial interest 
from the probability that they represent stages of a single species, 
and belong to a group that has not yet been described as occurring 
on the west coast of North America. Since the specific identity of 
the two forms is only a probability, it has seemed well to regard 
them for the present as distinct, and even, for reasons given 
below, to assign them provisionally to different genera. 


PEACHIA QUINQUECAPITATA, sp. n. (PI. XCVITI. figs. 1-4.) 


This form (fig. 1) was dredged by Dr. C. McLean Fraser 
in Nanoose Bay, Vancouver Island, in 15-20 fathoms. Jn the 
majority of the individuals the base is depressed in the centre 
and smaller than the column. It is thin, and in all cases 
shows clearly the lines of insertion of the mesenteries as well- 
marked invections, so that it can have possessed but very slight 
adhesive powers, if any. In one individual the attachments of 
the mesenteries did not quite reach the centre of the base, but 
ended abruptly a short distance from it, leaving a circular central 

~area which was exceedingly thin, but nevertheless not perforated. 
In other examples this condition did not occur, the insertions 
of the mesenteries extending quite to the centre, so that there 
were no indications of a terminal pore, such as has been described 
by Gosse (1860) in P. hastata. 

The column (fig. 1) is almost cylindrical, though usually 
contracting somewhat towards either extremity and, in some 
examples, is grooved by twelve well-marked invections, although 
in others these are quite indistinct. Examined under a lens the 
surface is seen to be studded with minute elevations, which are 
all the more noticeable by being of a paler colour than the 
general surface. No foreign particles were adhering to these 
elevations in any of the examples studied, and, as I had no 
opportunity for examining living specimens, I cannot say whether 
they had the power of adhering to surfaces with which they 
came into contact, as seems to be the case with the similar 


* Wor explanation of the Plate see p. 972. 


964 PROF. J. P. McMURRICH ON 


structures of P. hastata (Haddon and Dixon, 1885) and P. tri- 
capitata (Andres, 1883). 

There was no differentiation of the upper part of the column to 
form a capitulum, nor was there any distinct margin. 

The tentacles are twelve in number, arranged in a single cycle. 
They are of moderate length and stoutness, obtuse or tapering 
slightly at the tips, but not capitate, and frequently with longi- 
tudinal grooves. 

The lips are elevated considerably above the level of the 
disk, and show only a single siphonoglyph (fig. 2), which is 
completely separated from the rest of the stomatodeeum by the 
fusion of its lips throughout the greater part of their extent. 
At least this was the case in three individuals that I examined 
anatomically, and it would seem, therefore, to be of general 
occurrence. Sedgwick (1884) has described the same peculiarity 
as occasional in P. hastata, but it is not mentioned by other 
authors who have studied the anatomy of that species (Haddon, 
1885, 1889; Faurot, 1890, 1895). 

The general surface of the stomatodeum is longitudinally 
grooved along the lines corresponding to the insertions of the 
mesenteries, and each lip is raised in the intervals between the 
grooves into five rounded elevations, which correspond with the 
intervals between the perfect mesenteries, with the exceptions of 
those between the directives. One of these latter is quite small 
and the other is represented by the margins of the siphonoglyph, 
which, although not elevated to any great extent beyond the rest 
of the lips, nevertheless bear fine tuberculiform or digitiform 
processes (figs. 2, 3) which correspond with the conchular lobes 
of other forms. Two of these processes form a pair lying one 
on either side of the more dorsal portion of the siphonoglyph 
opening, and the three others are situated more ventrally, the 
median one being in the sagittal plane of the body, opposite 
the bottom of the siphonoglyph. In three individuals examined, 
of different sizes, I found no variation in the form or arrange- 
ment of these conchular processes, and would therefore conclude 
that this is their final adult arrangement. 

Size.—The height of the column, measured in individuals pre- 
served in formalin, was in the larger forms about 2°0 em., the 
diameter at about the middle being about 0-9 cm., and at the 
limbus 0°6 cm. The length of the tentacles was 0-4 cm. 

Colour.—Examples preserved in formalin retain a considerable 
amount of the original coloration, and from these it is possible to 
state that the general colour of the column is a more or less 
intense brownish red. In some cases it is distributed over the 
entire extent of the column and in others it is limited to the 
distal portion, the proximal or basal part being almost colourless. 
A closer examination shows that the pigment is not uniformly 
distributed throughout the area in which it occurs, but presents 
the appearance of a brownish red ground-colour upon which 
are scattered numerous minute dots of a paler shade, these 


NEW ACTINIANS FROM BRITISH COLUMTIIA. 965 


representing the small elevations already described as scattered 
over the entire surface of the column. The tentacles seem to have 

been of a paler colour than the upper part of the column and 
marked by four or five rings of a more or less intense brownish 
red (fig. 2). The conchular processes seem to have been of the 
deeper shade, with paler apices. 

Structure.—The column mesoglea is much thicker than the 
ectoderm and is of a finely fibrillar structure, cells being scattered 
among the fibrils. There are no ectodermal muscles, but 
associated with the presence of ectodermal gland-cells there is a 
well-defined layer of nerve-fibres. The adhesive organs appear 
in sections as elevated areas of the ectoderm, composed of slender 
cells considerably longer than those of the adjacent ectoderm ; 
the areas contain no gland-cells and, consequently, appear much 
paler than the general ectoderm in stained preparations. In 
structure, therefore, the organs differ from typical verruce only 
in being elevations of the surface instead of depressions. The 
endodermal musculature is supported on short simple processes, 
and towards the distal portion of the column it becomes weaker, 
the cells being arranged in an almost simple layer, and there is 
no indication of a sphincter. 

The musculature of the tentacles and disk is very moderately 
developed, and there are no muscle-fibres on either the ectodermal 
or endodermal surface of the stomatodeum. The structure of 
the siphonoglyph is quite characteristic. Its ectoderm is much 
higher than that of the rest of the stomatodeum and destitute of 
gland-cells. The surface bears very numerous strong cilia and is 
provided with a distinct cuticular layer, beneath which the nuclei 
of the cells are arranged in very many layers, the basal portion of 
the epithelium containing but few, and appearing as 1f composed 
of a very fine reticulum, which is traversed by a band of nerve- 
fibres. The endoderm is also much higher than it is over the 
rest of the stomatodeeum and its basal portion is also reticular in 
structure, though somewhat coarser than the ectoderm. The 
mesogloea also differs from that of the general stomatodzeum, its 
ground-substance being almost homogeneous and assuming a 
clear blue tint with hematoxylin-erythrosin, while elsewhere the 
colour is decidedly violet. The fusion of the lips, mentioned 
above, is merely an epithelial union; indeed, it appears to be 
due to adhesion of the cuticular layers of the two adjacent 
surfaces. 

The mesenteries are arranged in ten pairs, six of which 
are perfect, the remaining four pairs occupying the lateral 
and sulco-lateral interspaces. Of the perfect mesenteries, two 
pairs are directives, and all have well-developed diffuse muscle- 
pennons (fig. 4) consisting of strong though but slightly 
branched lamellae. A well-marked parieto-basilar muscle is also 
present forming usually a slight fold. The reproductive cells 
could not be distinguished in the individuals examined, but 
the endoderm of the mesenteries in the region proximal to the 


966 PROF. J. P. McMURRICH ON 


mesenterial filament is greatly thickened and very granular, as 
if from the presence of ingested food-material. ‘The eight im- 
perfect mesenteries are destitute of mesenterial filaments and, 
indeed, represent only the muscular portion of the perfect 
mesenteries. Their muscle-processes ave but few in number and 
they do not possess any distinct pavieto-basilar muscle. There 
is no difference in the development of the various perfect 
mesenteries such as Faurot has described for P. hastata; all are 
provided with mesenterial filaments and are alike in all 
particulars. 

The first description of a form that may definitely be assigned 
to the genus Peachia is that furnished by Reid (1848) of a species 
washed ashore in the Bay of St. Andrews and named Actinia 
cylindrica. ‘This name had, however, already been employed by 
Renier (1804) for Certanthus membranaceus, and has therefore 
given place to the term Peachia hastata, proposed by Gosse (1855) 
for a form with which Reid’s species is evidently identical. In 
the following year Koren and Danielssen described under the 
name Siphonactinia boeckit a form that is certainly congeneric 
with the Peachia of Gosse, and that author in 1860 described two 
additional members of the genus, P. uwndata and P. triphylla. 
No further additions to the list of species were made until 1879, 
when Hutton described as P. carnea a form cast up on the beach 
at Dunedin, New Zealand; a little later Andies (1883) described 
as Siphonactinia tricapitata a form from Naples that he had 
originally (1881) considered identical with Gosse’s P. triphylla, 
and in 1893 I described P. korent from off the coast of the 
Argentine Republic. There seems to be no room for doubt as to 
the distinctness of P. guinqguecapitata from P. hastata: the form 
of the conchula, the equality of all the perfect mesenteries, and 
the feebler development of the longitudinal musculature of the 
imperfect ones furnish sufficient bases for their separation, in- 
dependently of their coloration and geographical distribution. 
P. koreni, with only eight tentacles and a simple conchula, may 
also be regarded as distinct, and although the description of 
P. carnea is very incomplete, its geographical distribution is 
prima facie evidence of its distinctness also. 

With regard to the remaining four species, all of which are 
Kuropean, the evidence is not so clear, sce no anatomical data 
concerning them are available. P. wndata, from the Channel 
Islands, is believed by Haddon (1889) and G. Y. & A, F. Dixon 
(1891) to be merely a young example of P. hastata, its conchula 
resembling that of immature examples of the latter species. 
P. triphylla, also from the Channel Islands and the Firth of 
Clyde (Robertson, 1869, 1875), on the other hand, has a conchula 
composed of three lamellar processes, and in this resembles 
P. boeckii; the latter, however, bearing the lamellae upon the 
summit of a tubular prolongation of the siphonoglyph, whereas 
in Gosse’s figure of P. triphylla they are represented as sessile. 
The difenencs may, however, be due toa difference in contraction, 


NEW ACTINIANS FROM BRITISH COLUMBIA. 967 


and it is not improbable that the two forms are identical. It 
must be noted, however, that Faurot (1895) identifies P. boeckir 
with P. hastata, and until anatomical data are furnished it will 
be well to regard it as a distinet form. Finally, in the Medi- 
terranean P, iricapitata the conchula is composed of tubercles ; 
and while the correctness of its original identification by Andres 
(1881) with P. triphylla must still be regarded as a possibility, 
it seems advisable for the present to regard it as a good 
species. 

A provisional arrangement of the known species of Peachia 
may then be stated thus :— 


Tentacles 12, 


Conchular lobes 6-20, irregular ...............0..0........ PB. hastata. 
Couchular lobes 3, lamellar, 
sessile ......... 06.04 P. triphylla. 


borne on tubular prolongation of siphonoglyph. P. boeckii. 
Conchular lobes tuberculiform, 


three in number sodoosdeaoabdsBeosocsonensce | Leo HPUCRY DIAM RES 
five im mumaber 2...) sccseseeesececeenseicre esses ee Quinguecapitata. 
? Poets dsb Aun ote Soetoqupenidn cool Me ra AOU TORO 
MentaclestSii Mi tasts sa yserche copies nba weuietse nian stb ere MONEIUTs 


BIcIDIUM EZQUOREA, sp.n. (Pl. XCVIII. figs. 5-7.) 


This form is not uncommon as a parasite upon the bell of the 
Leptomedusa “quorea forekalii, which is of very common occur- 
rence in the waters of British Columbia. The column (fig. 5) 
has a rounded base, is somewhat conical or urn-shaped and 
wrinkled transversely by contraction in examples preserved in 
formalin. Proximally it usually shows some faint longitudinal 
grooves which mark the lines of insertion of mesenteries, but no 
signs of verruce or other adhesive organs could be seen. ‘The 
margin, which is indistinct, is occupied by a single series of twelve 
short and obtuse tentacles. The mouth (fig. 6) is usually 
widely expanded, so as to completely hide the disk, and shows a 
single siphonoglyph ; no conchula was present. 

Size.— Length about 7 mm., greatest diameter 4 mm. 

Colowr.—There was no trace of colour in any of the examples 
seen. 

Structure—T'be column mesoglea is thinner than either the 
ectoderm or endoderm and has a homogeneous or finely fibrillar 
structure, cells being scattered throughout the ground-substance. 
The ectoderm is of uniform structure throughout its entire 
extent, no verrucal areas being distinguishable. There are no 
ectodermal muscle-fibres, and those of the endodermal surface are 
arranged ina simple layer, there being no mesogleeal lamellee for 
their support and no indications of a sphincter muscle. The 
musculature of the tentacles is also very feeble, the ectodermal 
fibres being arranged in a single uviform layer, while the endo- 
dermal ones are hardly distinguishable. 

The single siphonoglyph is prolonged somewhat below the level 


968 PROF, J. P. McMURRICH ON 


of the rest of the stomatodeum and forms a well-marked deep 
groove, whose endoderm is much higher than that of the rest 
of the stomatodeum, although it does not present a reticular 
formation such as occurs in Peachia quinquecapitata. ‘There are 
six pairs of mesenteries, all of which are perfect and all possess 
mesenterial filaments; no indications of additional mesenteries 
of the second cycle were to be found. ‘Two of the six pairs 
were directives. ‘The longitudinal musculature (fig. 7) formed 
a low pennon extending throughout the entire muscular area 
of the mesentery, the supporting lamelle being palisade-like 
in their arrangement, and is higher than the endoderm that 
covered them. No parieto-basilar muscle was present in my 
preparations, which did not, however, include the most proximal 
portion of the column. 

A number of immature Actinians, a list of which has been 
given by Haddon (1887), have been described as parasites in 

various species of Medusz, and all present structural features 

sufficiently similar to suggest that they are generically identical 
with one another and with the form described above. They have 
been assigned in part to the genus Halcampa and in part to 
Peachia, but none of these has yet been actually shown to trans- 
form into the one or the other of these genera, and until the 
adult condition is definitely known it seems well to retain 
for them the genus Dicidium established by L. Agassiz (1859), 
recognizing, however, that this is but provisional and that the 
forms assigned to it are larval or at least Immature. 

The evidence furnished by the structure, although not con- 
clusive, does however give some indications of the probable 
position of the adult form and, I believe, points towards the 
genus Peachia, rather than to Halcampa. Differences between 
the two genera that are pertinent to this question are to be found 
in the presence of a conchula in Peachia and its absence in 
Halcampa; in the single deep siphonoglyph of Peachia as 
compared with the two shallow ones of Hulcampa; and in the 
somewhat diffuse form of the muscle pennons in Peachia as 
compared with the compact and circumscribed ones of Halcampa. 
The first of these differences is not apparent in all species of 
Bicidium, the conchula probably being late in its development, 
but the other characteristics are constant in all known forms 
whose anatomy has been recorded. 

Since Haddon (1887) reviewed the various species of Bicidium 
some additions have been made to our knowledge of them, and in 
support of my contention as to their affinities to Peachia rather 
than to Haleampa, a reconsideration of them may not be amiss. 
The absence of a conchula in B. equoree is, it is true, opposed to 
its reference to Peachia, but, as has been pointed out, this 
structure may very well be late in developing, and in the nature 
of the siphonoglyph and the muscle pennons the similarity to 
that form and the difference from Halecampa are pronounced. 


NEW ACTINIANS FROM BRITISH COLUMBIA, 969 


The occurrence of P. quinquecapitata in the same locality is also 
suggestive, but the differences between the two forms are too 
marked to warrant their identification, although it is not im- 
probable that these differences are due to age rather than specific 
differences. 

The structure of Actinia clavus of Quoy and Gaimard (1833), 
recently studied by Pax (1912), shows, I believe, that it must be 
considered congeneric with B. equoree, although Pax refers it to 
the genus Halcampa. As in the British Columbian species, there 
are no indications of a conchula; but again, the form of the 
siphonoglyph and the muscle pennons suggest Peachia rather than 
Halcampa, and until further evidence as to its exact position 
is available, it would seem well that it should be known as 
Bicidium clavus.* 

The evidence as to the affinities of B. parasiticwm is more 
definite. It was first described by L. Agassiz (1859) from Cyanea 
arctica and later by Verrill (1864), who showed that it possessed 
a well-marked trilobed conchula, and, still later (1866), assigned 
it to the genus Peachia. Preparations that I have of this species 
show it to have a striking general similarity to 5. equoree, the 
mesenteries being twelve in number, all perfect and all bearing 
mesenterial filaments, and the siphonoglyph single and deep. 
The muscle pennons also are of the same diffuse type, but the 
lamelle (fig. 8) are not arranged in the manner of a palisade, 
but are decidedly branched in a dendritic manner so that 
they present an appearance of being airanged in groups. It 
is to be noted that Verrill (1874) mentions the capture of two 
very large examples of this species, imbedded in gravel at low- 
water mark at Eastport, Maine; he does not, however, give any 
anatomical data concerning them, and until it is definitely known 
that the form develops the four pairs of secondary mesenteries 
. (zygocnemes), it seems-advisable to allow it to remain in the 
genus Sicidium. 

The form which F. Miller (1860) deseribed very completely as 
Philomedusa vogtii is also undoubtedly a Bicidium. It occurred 
upon the meduse Olindias and Chrysaora and was provided with 
a trilobed conchula, twelve tentacles, twelve mesenteries, all of 
which were perfect and furnished with mesenterial filaments, and 


had asingle siphonoglyph. The form described by Graeffe (1884) 


* The observations of Pax make it quite certain that R. Hertwig (1882) was in 
error in identifying a Halcampa from the Kerguelen Islands with Quoy and 
Gaimard’s species. The possibility of Hertwig’s forms being young examples of 
Studer’s Hdwardsia kerguelensis, which Kwietniewski (1896) has shown to hea 
Halcampid, should not be lost sight of, although the differences in the descriptions 
as they stand are too great to permit of a definite identification. Furthermore, the 
contention of Pax that the H. clawus of Tizard and Murray (1881) and Appelléf 
(1897) is distinct from that described by Hertwig is undoubtedly correct, but since 
the forms so named by Appelléf were the actual types of Danielssen’s Halcampoides 
ahyssorwm, it is difficult to understand the necessity for the new name, seplentri- 
onalis, that Pax bestows upon them. 


970 PROF. J. P. McMURRICH ON 


as Haleampa medusophila, occurring on various meduse (Zina, 
Octorchis, and Mquorea) is probably identical with Miller’s 
species, the absence of a conchula being probably due to its 
greater immaturity. 

In 1860 Wright described as Peachia fultoni * a form that 
he found parasitic on a species of Thawmantias (Phialidium %), 
later (1861) changing its name to Haleampa fultoni. This 1s 
probably the same as the form described by Haddon (1887), and 
erroneously, as has been pointed out by Carlgren (1904), regarded 
by that author as the larva of Haleampa chrysanthellum. In 
1887, McIntosh recorded the occurrence of actinian larvee on 
various Thaumantiad meduse occurring at St. Andrews, identi- 
fying them as the young of P. hastata, an opinion in which 
Haddon (1888) concurred after an examination of their structure, 
still maintaining, however, their distinctness from the examples 
previously described as the young of HH. chrysanthellum. They 
possessed eight tentacles and eight large mesenteries together 
with four smaller deuterocnemic ones. It may be that Haddon 
is correct in regarding the two forms he examined as distinct 
species, but even if so it seems clear that they are to be referred 
to the same genus, and since it is still uncertain that they 
actually do develop into Peachia, it will be preferable to place 
them in Biciduin. 

Finally, Dendy (1888) has described from medusz obtained 
at Port Philip a parasitic actinian with twelve tentacles, within 
which there was ‘“‘an inner circle of 12 cushion-like swellings,” 
which in older individuals become saccular outgrowths. It is 
not possible to ascertain from Dendy’s account, whether or not 
these outgrowths represent a conchula, and the exact affinities of 
the form must remain doubtful, although with a probability that 
it represents a Peachia’. 

The available evidence seems, accordingly, to point strongly in 
favour of these various medusophilous forms being young stages 
in the development of Peachia rather than Halcampa, but a 
direct linking up of the immature examples of Sicidiwm with 
their respective adults is necessary to settle the question. 


* Some confusion exists in the references to the literature of this species. Andres 
(1883) gives the date of its first description as 1859 and as references the Pro- 
ceedings of the Physical Society of Edinburgh, vol. 11. 1859, and the New Edinburgh 
Philosophical Journal, vol. xii. 1860. Haddon (1887) repeats the latter reference 
but substitutes for the former, Proceedings of the Royal Society of Edinburgh, 
vol. ii. 1860. There is no paper in the Proceedings of the Royal Society of 
Edinburgh by Wright referring to this species, and the date of vol. i. of the 
Proceedings of the Royal Physical Society of Edinburgh, which does contain such 
a paper, is 1861 and not 1859. ‘The reference to the Edinburgh New Plilosophical 
Journal (this being its correct title) should read “ New Series, vol. xii. 1860.” 

+ Carlgren (1904) mentions, without descriptions or names, two additional forms 
that should probably be referred to this genus, one occurring in the medusa 
Eutimalphes indicans on the Swedish coast and the other on a large medusa from 
Valparaiso. 


“NEW ACTINIANS FROM BRITISH COLUMBIA. 971 


LITERATURE. 


Aaass1z, L.—On some new Actinoid Polyps of the United States. 
Proc. Boston Soc. Nat. Hist. vi., 1859. 

Anpres, A.—Prodromus neapolitane actiniarum faune ete. 
Mitth. Zool. Stat. Neapel, ii., 1881. 

Anpres, A.—Die Aktinien. arma und Flora des Golfes von 
Neapel, ix., 1883. 

APPELLOF, Die Actiniengattunzen Fenja, Algir und Hal- 
campoides, Dan. Bergens Mus. Aarb. x1., 1881. 

CaRLGREN, O.—Kurze Mittheilungen tiber Anthozoen. Zool, 
Anz. xxvii, 1904. 

Dantetssen, D. G.—« Actinida.” Norwegian North Atlantic 
Exped. xix., 1890. 

Denpy, A.—Note on some Actinian larve parasitic upon a 
medusa from Port Philip. Proc. Roy. Soc. Victoria, 
1888. 

Dixon, G. Y. and A. F.—Report on the Marine Invertebrate 
Fauna near Dublin. Proc. Roy. Trish Acad., ser. 3, i1., 
1891. 

Favrot, L.—Sur la disposition des cloisons mésenteroides chez 
la Peachia hastata. C. R. Acad. Paris, ex., 1890. 

Favror, L—Etudes sur l’anatomie, l’histologie et la développe- 
ment des Actinies. Arch. Zool. exp. et gén., ser. 3, ii1., 
1895. 

Gossn, P. H.—Description of Peachia hastata, ete. Trans. 
Linnean Soe. xxi., 1855. 

Gosss, P. H— Actinologia Britannica. London, 1860. 

Graerre, E.—Uebersicht der Seethierfauna des Golfes von Trieste. 
ILI. Celenteraten. Arb. zool. Inst. Wien, v., 1884. 

Happon, A. C., and Drxon, G. Y.—The structure and habits 
of Peachia hastata (Gosse). Sci. Proc. Roy. Dublin Soc., 
n. S. iv., 1885. 

Happon, A. C.—On the arrangement of the mesenteries in the 
parasitic larva of Halcampa chrysanthellum (Peach). 
Proc. Roy. Dublin Soe. n. s. iv., 1887. 

Happon, A. C.—On larval Actiniz parasitic in Hydromeduse 
at St. Andrews. Ann. Mag. Nat. Hist. ser. 6, 11., 1888. 

Happon, A. C.—A Revision of “She eae Netinie. "Part It, 
Sci. Trans. Roy. Dublin Soe. ser. 2, iv., 1889. 

Hurron, F. W.—Contributions to a (Galenterate Fauna of 
New Zealand. Trans. New Zealand Inst. xii., 1879. 

Koren, J., and Dantetssen, D.C.—Nye Actinier. Fauna htt. 
Norvegie, i1., 1856. 

Kwirtniewski, C. R.—Revision der Actinien welche von 
ierrnee Profs Sbuder. ayn gesammelt worden. Jena. 
Zeitschr. xxx., 1896. 

McIntosu, W. C.—On the commensalistic habits of the larval 
forms of Peachia. Ann. Mag. Nat. Hist. ser. 5, xx., 
1887. 


972 ON NEW ACTINIANS FROM BRITISH COLUMBIA. 


McMourricu, J. P.—Report on the Actiniz collected by the 
U.S. Fish Commission Steamer ‘ Albatross’ during the 
winter of 1887-1888. Proc. U.S. Nat. Mus. xvi., 1893. 

Miter, F.—Ueber Philomedusa Vogtii. Arch. f. Naturg. xxvi., 
1860. 

Pax, F.—Revision des types des Actinies décrites par Quoy et 
Gaimard. Ann. Sci. Nat., Zool. sér. 9, xvi., 1912. 

Quoy et GAImMARD.—Zoologie du Voyage de la corvette ‘Astrolabe.’ 
Paris, 1833. 

Ret, J.—Account of a new species of Actinia (A. cylindrica). 
Aun. Mag. Nat. Hist. ser. 2, 1., 1848. 

Rosertson, D.—Living specimen of a rare Sea Anemone, Peachia 
triphylla. Proc. Nat. Hist. Soc. Glasgow, i., 1869. 
Rosertson, D.—On the Sea-Anemones of the shores of the 

Cumbraes. Proc. Nat. Hist. Soc. Glasgow, i1., 1875. 

Sepewrck, A.—On the Origin of Metameric Segmentation ete. 
Quart. Journ. Micr. Sci. ser. 2, xxiv., 1884. 

Tizarp and Murray.—Exploration of the Faroe Channel during 
the summer of 1880 in H.M. hired ship ‘ Knight Errant.’ 
Proc. Roy. Soc. Edinburgh, x1., 1881. 

Verritt, A. E.—Revision of the Polypi of the Eastern Coast of 
the United States. Mem. Boston Soc. Nat. Hist. 1., 
1864. 

Wraieut, 1’. S.—Ohservations on British Zoophytes. Edinburgh 
New Phil. Journ. n. s. xii., 1860. 

Wricut, T. 8.— Observations on British Zoophytes. On 
Halcampa Fultoni, a parasitic Actinia. Proc. Roy. Phys. 
Soe. Edinburgh, u1., 1861. 


EXPLANATION OF PLATE XCVIII. 


r. 1. Peachia quinquecapitata. Nat. size. 
2. Oral view of P. quinquecapitata expanded. X 2? 
3. Oral view of partially contracted individual of P. quinquecapitata, 
showing arrangement of conchular lobes. 
4, Transverse sections of mesenteries of P. guinquecapitata. 
5. Side view of individual of Bicidiwm equoree. X A. 
6. Oral surface of B. equoree. X 4. 
7. Transverse sections of mesenteries of B. equoree. 
8. Transverse sections of mesenteries of B. parasiticum. 


ON SPONGES IN WATERWORKS. 973 


57. Sponges in Waterworks. By W. N. Parner, Ph.D., 
F.Z.S., Professor of Zoology, University College, 
Cardiff, 


[ Received September 17, 1913 ; Read November 11, 1913.] 


INDEX. Pages 
Occurrence in Cardiff Waterworks and elsewhere ... 973, 974 
TBRHL IYO) GSA dion aoa las Buados abe Reb donors nosoad aad pao uaaEos doaaaens oom KIC Cee) 
Reproduction Neato ano PRA nod cass ete rosacachEcceuannseemey ei ONO 
Methods of treatment adopted .....................255 ae 976 


Some three or four years ago I received through my colleague, 
Dr. Scholberg, some specimens of a branched, yellowish-brown 
organism which had been found growing in some of the pipes 
at the Cardiff Waterworks, and which proved to be very fine 
examples of Spongilla lacustris. Subsequently I learnt that 
‘there had been an extensive growth of this sponge in the pipes 
leading to one of the series of filter-beds, which had caused much 
trouble and difficulty; and that, although the growth had 
apparently disappeared for a time after treatment with sulphate 
of copper, it, as | expected, soon appeared again, and became as 
flourishing as ever. In the summer of 1911 the growth was so 
extensive as to cause anxiety, not only as regards reducing the 
flow in the pipes, but also on account of the unpleasant odour 
which had been noticed in the water, due doubtless to the 
living sponge as well as to its decomposition. In December 
of the same year I was requested by the Waterworks Engineer, 
Mr. ©. H. Priestley, to report on the matter to the Waterworks 
Committee; and now that the treatment I subsequently pro- 
posed has—at any rate for the present—been successful, I have 
the permission of the Committee to publish an account of our 
experiences. [I should lke to express my indebtedness to 
Mr. Priestley and his staff for their cooperation and for giving 
me every facility in prosecuting the enquiry; I have also to 
thank Dr. Kemna, of Antwerp, Dr. Harmer, Mr. Kirkpatrick, 
and Professor Minchin for valuable information given at the 
beginning of the investigation. 

_ Although sponges of various species have been recorded at 
numerous other Waterworks *, they do not appear to have been 
the chief source of trouble, as at Cardiff. 

The interesting subject of the fauna of Waterworks has quite 
recently been so fully dealt with by Dr. Harmer in these 
“ Proceedings” (19138, p. 426), that it is unnecessary for me 


* Hg. at Boston, Mass. (Bowerbank. Proc: Zool. Soc. 1863, & Potts, Acad. Nat. 
Sci. Philadelphia, 1881, 1882, and 1887); Hamburg (Kraepelin, Abh. Naturwiss. 
Ver. Hamburg, 1886); Torquay (Chapman, “ Animal Growths in Water-pipes,” 
Transactions of the Institute of Water Hngineers, 1913, & Kirkpatrick—see Harmer, 
P. Z. 8. 1913, p. 486) ; and Aberdeen (Kirkpatrick, loc. cit. p. 489). 


Proc. Zoou. Soc.— 1913, No. LXV. 65 


O74 PROF. W. N. PARKER ON 


to refer in detail to the history and literature of the subject. 
Dr. Harmer’s account of the difficulties which have recently 
occurred in various English Waterworks owing to the presence 
of Polyzoa, shows the importance of biological investigation in 
dealing with such cases, as was pointed out in this country by 
Professor Hickson in ms presidential address to Section D of the 
British Association at Southport in 1903. 

The Cardiff district is supplied with excellent soft water from 
the Taff Fawr reservoirs in Brecknockshire, midway between 
Merthyr and Brecon. From these higher reservoirs the water 
is conducted by conduits into various other storage-reservoir's 
at a lower level, in the near neighbourhood of Cardiff, and 
from those into filter-beds: the latter, of course, prevent the 
access of organic particles and spicules into the service- pipes 
supplied from them. The sponge had hitherto only been found in 
the pipes leading to certain of the filters on the northern border 
of Cardiff known as the “‘ Heath” filters, from which the greater 
part of the city is supplied. The water to these filters comes 
from two storage-reservoirs, at Llanishen and Lisvane, situated 
about a mile farther north. Careful examination has so far not 
revealed any trace of the sponge in other parts of the system, 
which therefore I need not describe further here*. 

The water from Llanishen reservoir passes through metal 
screens with fine meshes, so as to strain off all but minute solid 
particles, into a valve-shaft from which it is conducted by under- 
ground pipes to a valve-chamber at the “ Heath.” It is then 
again screened before flowing into a 3-ft. pipe, with which lateral 
pipes are connected ending in bell-mouths, one to each of the six 
filter-beds. 

The chief trouble occurred in this pipe, which was thickty lined 
with a luxurious growth of the sponge, consisting of dense 
incrustations giving off numerous finger-like processes which 
branched and branched again, and some of which reached a 
length of 8 inches or more, frequently showing concrescence 7. 
Other growths were found in the valve-chamber and _ screen- 
chamber, in the corners and crevices of which it was not easy to 
get access to them. 

On visiting the “ Heath” on December 12th, 1911, the sponge, 
packed with gemmules, showed no signs of dying down for the 
winter, and it appeared to me that had it been desired to culti- 
vate the organism, probably no better plan could have been 
devised than that furnished by the water-chambers and pipes 


* For further details with regard to the Cardiff Waterworks, see a paper by 
C. H. Priestley, M.Inst.C.K., on the “ Development of the Cardiff Water Supply,” 
read at the Sessional meeting of the Royal Sanitary Institute at Cardiff on April 
12th, 1912; also a description of the Waterworks by the same author issued from the 
Cardiff Waterworks Engineer’s Otfice, 1908. 

+ Cf. figures by Bowerbank, “A Monograph of the British Spongiadz,” Ray 
Society, 1864-1882; and Johnston, “A History of British Sponges and Lithophytes 
(Corallines),” London, Edinburgh, and Dublin, 1882. 


SPONGES IN WATERWORKS. LD) 


(cf. Kraepelin, loc. cit.). It can grow as well in the dark, with- 
out zoochlorelle, as in the light ; and being effectively protected 
from frost and other adverse conditions, it seemed possible that 
jt might continue alive during the whole winter. On the same 
day, we made a cursory examination at Llanishen and Lisvane, 
with no result; but on the following day, the reservoir-keeper, 
after emptying the valve-shaft at Llanishen, found a considerable 
growth at a depth of between 10 and 20 feet. 

Since then, a careful look-out has been kept for the sponge 
throughout the entire system, but there has been no positive 
evidence of its occurrence in other parts, either in the open or in 
the underground pipes. 

The two most important problems which presented themselves 
were—the destruction of the sponge in the infected area, and the 
prevention of re-infection throughout the whole system. As 
regards the latter, the only sound method would be the intro- 
duction of pre-filtration through sand, so as to starve the sponge 
by preventing the microscopic organisms (on which it and other 
‘‘microphagous”” organisms which might at any time appear 
depend for their nutriment) from passing into the pipes and 
reservoirs (cf. Kemna, quoted by Harmer on p. 432), as well as 
to keep out larve and gemmules. This method would take 
a long time to carry out and entail considerable expense ; 
moreover, 1t would not be effective unless it could be adopted at 
the lower reservoirs (where there are difficulties as regards level) 
as well as the higher ones, as there would be a reappearance of 
organisms in them. As the matter was urgent, I therefore felt 
it would be better to confine attention at first to the destruction 
of the organism in the infected area and the prevention of its 
regeneration and of reinfection by the gemmules. 

Treatment with any destroying agent except such as would be 
dangerous to human life would be ineffective if applied to the 
adult sponge, as it would not destroy the myriads of gemmules 
protected by their resistant coats. It was therefore necessary in 
the first instance to make a periodical examination of the sponge 
for a year or more, so as to ascertain (1) whether sexual repro- 
duction occurred, (2) when the gemmules germinated, and 
(8) whether the adult growth perished partially or entirely 
during the course of the winter*. 

An examination of specimens at various times of the year has 
so far revealed no larve or any signs of sexual reproduction ; but 
the enormous production of gemmules throughout the crusts and 
branches is quite sufficient to start new growths in any part to 
which the gemmules could gain access. There can be no doubt 
that a sexual stage is unnecessary for the reproduction of the 
sponge from year to year. 


* Cf. Marshall (SB. Naturf. Gesellsch., Leipzig, 1884, & Journ. Roy. Micros. 
Soc. v. 1885) ; Weltner (Archiv f. Entwicklungsmechanik, Bd. xxxiii. Heft 3 u. 4). 


6d* 


976 ON SPONGES IN WATERWORKS. 


On visiting the “ Heath” and Llanishen on February 3rd, 1912, 
when the filter-beds were covered with ice 13 ins. thick, we "found 
that the adult growth was dying down, but that it was still 
closely packed with gemmules, some of which had begun to 
germinate and start new growths among the old. On March 23rd 
gemmation had proceeded further, new growths being abundant, 
and I suggested that in the course of the spring the pipes and 
chambers should be scraped and treated with strong brine, so 
applied as to reach any small spaces and crevices in which 
gemmules could lodge, and thus to attack the sponge when it was 
least capable of resistance. This was done on May 2nd, and 
with the sponge were removed a large quantity of iron corrosive 
nodules—another trial to the Water Engineer, referred to by 
Harmer on pp. 430, 437, & 438 of his paper; the brine was then 
washed out through a bye-pass. The result was apparently satis- 
factory, butin the course of the summer a few small new growths 
were observed by the Foreman: they were very much less 
abundant than in the previous summer, and no smell had been 
noticed since the treatment. An examination on September 24th 
showed that these comparatively few young incrusting sponges 
had not yet given rise to any outgrowths, and that they con- 
tained no gemmules: they were in all probability due to a few 
gemmules which had not germinated when the treatment was 
carried out. I therefore suggested a second application of brine. 
By November 14th, when this was done, the growths had 
increased in size considerably, but apparently no gemmules had 
been formed. Instead of leaving the pipes empty for only a few 
hours, as on the previous occasion, it was possible this time to 
give them three days before being again used, and since then no 
trace of the sponge has been seen, 

It is interesting to note that no Polyzoa were found, but that 
a number of colourless Hydree like those described by Kraepelin 
at Hamburg were obtained by the reservoir-keeper from the 
bottom of the valve-shaft at Llanishen on May 25th; unfor- 
tunately these were all dead by the time they reached me*. 


* Since the above was written, a few young sponge-growths were again observed 
and at once destroyed. Colonies of Cristatella, with numerous statoblasts, had 
also appeared : fortunately this is not one of the “moss’’-forming Polyzoa. (Oct. 
25th, 1913.) 


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ON CYPRINODONT FISHES. 977 


58. A Revision of the Cyprinodont Fishes of the Subfamily 
Peeciliine. By O. Tate Ree@an, M.A., F.Z.S. 


[Received October 11, 1913 ; Read November 11, 1913. ] 
(Plates XCIX,-CI.* and Text-figures 168-173.) 


InDEX. : 

SYSTEMATIC ;— Page 
SHINO MNS OP CUAL) cadcsorcdaosesceinoar essonosnsppasasnooncccasco” IU) 
(Fammleoisas COO, BOs Ws cp cboder sco.oco cog ecuscorogdedanaounndaanel eke 
Gh, DIG RO/TUCHEG Ly Bis DS seoloaseqs asd enovensnasrseadabadananécce els 
GSMElANOSticEa spe We ete ee nee eee eee OO 
(GU OTIRET In SDD ATEN cree amaetun bbe cacianobueis Hebe on bRE bn Sc aWBA BBAAHA Mme Lets, 
Groleqostictaspay ees hee SS 
(CF OTROKSOULOLR, 9). 0B Ws emp pevonesed 200 gadley BaacAsdctien eoqsoHoDsedadco. set 
(EL, CIOUPATAOSOD, SDs We occ avo va scaagndosccasnagnodoanncsoacoass Otek’) 
Sh, CRMMCKOTUSIS, SDs Wy sovscdose sposen osecgnsoacesepadsonss00n000 CARD) 
Priapichthysy seus NOs cece eee eree acess ee OO 
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IP, GQADTIEDSUS, BOs To.“ enobennnescescososdsuteacedadbeneaancnooncsée . COME 
Brachyrhaphis, SN. MOV. ........5... 0220 ye eects eee OOF 
JEGROTOTA HEH ODOR, FON NON on), onapsa son don se 204 s0Heed ons bedanacogona. WKS 
TECH HOO ROAMALO FRG TKO soo enoweonne obs coc esasqocnaceobondee 1003 
IPO NLOPHD (REN; SUOWooee005 oor oon penoss soe 960 vOdopmorenpecgsannde LOE} 
JRO AUT FONARTSS (Ds V1.0\ 900 abe soawnesnovog Bhan mecrnoopAnBenennsasaer NOUN, 
Mollienisia gracilis, NOM. NOV. ..........2-.2..;20001-. LOIZ 
LMM MNIGMOfASCLALA, IS) Wen ten een see cece ae yl Olle 
PE RORNOLAL SPs Mercer iccan ce eeesueue eee teens sro peace ome eee LOLG, 
1 Ds OLUTION 0G gp aunese sto ocubns cananBaproneecocnmapoayasomdaacoas M6) 
Pr CaUdOfaS etal GSP. Mer. .co, arouse nosso eeee es eoenenensey LONG, 
Oy WAKER, BID We, sonora covondovocbasducavoascepsancay cocoon CML 7/ 


The Peeciliine are viviparous Cyprinodonts in which the anal 
fin of the male is advanced and modified into an intromittent 
organ, They are American, ranging from Carolina to Argentina 
and from Arizona to Ecuador ; many of the species live in the 
sea as well as in fresh water. Most of the species are quite 
small, two or three inches long; the giant of the group, Belonesox 
belizanus, attains a length of eight inches, In some forms the 
males are much smaller than the females, and the adult male of 
Heterandria formosa, 15 to 20 mm, long, is one of the smallest 
fishes known; in several species males are rather scarce, 

In Giinther’s classification (Cat. Fish. vol. vi.) the Cyprinodonts 
were divided into Carnivore, with the intestine short or but 
little convoluted and the bones of the lower jaw firmly united, 
and Limnophage, with the intestine long and convoluted and 
the bones of the lower jaw loosely connected. It is now known 
that both types oceur in two distinct subfamilies, Characodontinza 
and Peeciliine, and there is good evidence that in the latter the 


* For explanation of the Plates see p, 1018, 


978 MR. C. TATE REGAN ON 


limnophagous structure has been evolved three or four times 
independently. 

The intromittent organ is freely movable and is supported 
internally by bony stays, 2 to 5 in number, that project down- 
wards and forwards from the bridges that connect the parapo- 
physes of the posterior precaudal vertebre. It is formed by the 
prolongation of the third, fourth, and fifth anal rays, the first 
and last of which are stout and are more or less expanded trans-' 
versely so as to margin a groove on one side or the other, or they 
may even meet so as to form a tube. The urogenital orifice is 
directly in front of the base of the fin, and may often be covered 
by the pelvic fins, which, doubtless, help to conduct the seminal 
fluid into the groove, or in certain forms where the pelvic fins of 
the male are considerably enlarged they may be closely applied 
to the intromittent organ, converting the grooves into closed 
tubes. The distal segments of the prolonged rays may be 
variously modified into spines, hooks, barbs, ete., which may help 
to retain the organ in position during coition. 

The differences in structure of the intromittent organ are of 
great systematic importance; hitherto they have been almost 
neglected, except by Higenmann (Proc. U.S. Nat. Mus. xxxil. 
1907, p. 425), who has described them in the genera Lebistes, 
Girardinus, Glaridichthys, Phalloceros, and Phalloptychus *. 

Many of the species eat Mosquito larve, and the ‘“ Millions ” 
fish (Lebistes reticulatus) has been introduced into many countries 
in the hope that it will check malarial fever t. On account of 
their small size, pretty appearance, and interesting habits the 
Peeciliine are great favourites with European aquarium-lovers, 
especially in Germany; several of the new species described 
below have been sent to me by my valued correspondents in 
Hamburg, Herr J. P. Arnold and Herr A. Rachow, as newly 
imported aquarium-fishes. In the German aquarium-journals 
and in two booklets by Stansch (Die lebendgebiirende Zahnfarkfen, 
Leipzig, Wenzel, 1910, 1911) have appeared many accounts of the 
habits of different species: from these one gathers that the 
broods follow each other at intervals of only a few weeks, but 
that the number of young varies greatly according to the species ; 
thus a pair of Pecilia pare had broods of 8, 11, and 10 at 
intervals of four weeks, but in P. vivipara 124 were counted in a 
single brood. ‘The new-born young are usually from 5 to 10 mm. 
long and grow rapidly, in many species attaining sexual maturity 
inabout three months. The breeding-habits are of great interest : 
in species where the males are brilliantly ornamented (J/ol- 
lienisia, Xiphophorus, Pecilia brannert) they dart about displaying 
their beauty ; in these and in related forms the females appear 
to encourage their advances, but in others they are quite shy 
and their mates have to exercise cunning to get near them. It 


* Some additional figures are given by Langer in a paper on the morphology of 
these fishes, just published (Morph. Jahrb. xlvii.). 
+ See P.Z.S. 1910, p. 146; 1912, p. 906, 


CYPRINODONT FISHES. 979 


is of some interest to note that in Pecilia and related genera 
the females welcome the attentions of the males and the intro- 
mittent organ is quite short, but as in Gambusia and Phalloceros 
the females try to make the males keep their distance it is 
much longer. 
Synopsis of the Genera *. 
I. Lower edge of caudal peduncle sharp, without a 
median series of scales; bones of lower jaw firmly 
united ; teeth conical or villitorm. 

Body rather deep ; pelvic fins well developed, behind base 
of pectorals; anal tin of male without long appen- 
COlEKe =I} Mea nan Gun ttn Sagon some ud Sema TOES Can Coe eee cae 

Body elongate; pelvic fins absent in female, minute and 
below gill-openings in male; anal fin of male ending 
in a pair of long appendages with spine-like processes 2. Temeurus. 


1. Alfare. 


II. Lower edge of caudal peduncle rounded or obtuse, 
with median series of scales. 
A. Pelvic fins similar in both sexes. 
1. First produced ray of anal fin of male without 
long processes or appendages. 

a. Third and posterior branch of second pro- 
duced ray of anal fin of male each ending 
in a retrorse spine or hook. 

Jsws not produced; teeth conical or villiform; anal fin 
of male with distal part of first produced ray serrated, 
the terminal serrations directed towards its tip; 
third produced ray and posterior branch of second 
each ending in a retrorse hook. Dorsal 6-12; origin 
belindithatiotanalieeeeneee case reece ee eres Bonn Guaaebustian 

Jaws produced ; teeth slender, pointed, depressible; anal 
fin of male with terminal segments of first produced 
ray not forming serrations; third produced ray and 
posterior branch of second each ending in a retrorse 
SPUN CBee eer See ese aac COREE ep ae eloweson: 

6. Third and posterior branch of second pro- 
duced rays of anal fin of male not bearing 
retrorse hooks or spines, 

a, Anal fin of male ending in a more or less 
distinct antrorse hook, the anterior branch 
of the second produced ray curved for- 
ward. 

* Mouth moderate, with distinct lateral 
cleft ; teeth conical or yilliform, 
+ Dorsal 7-12; origin behind that of anal. 

Extremity of anal fin of male supported equally by third 


and anterior branch of second produced rays ......... 5. Priapichthys, 
Extremity of anal fin of male supported equally by first 
and anterior branch of second produced rays ......... 6, Priapella. 


+t Dorsal 11-17; origin in advance of 
that of anal; extremity of male anal 
fin a strong hook formed by the un- 
segmented end of the anterior branch 
of the second produced ray ............... 7, Pseudoxiphophoyus, 
** Mouth small, without distinct lateral 
cleft; hook at end of anal fin of male 
formed by anterior branch of second 
produced ray. 


* The differences in the structure of the male intromittent organ used in this 
synopsis are illustrated in text-tigs. 168, 169 (genera 1 to 4 and 11), 170 (genera 5 to 
10), 171 (genera 10 and 13), 172 (genera 14 to 22), and 178 (genera 23 to 26). 


” 


980 MR. C. TATE REGAN ON 


Teeth somewhat compressed, pointed; anal fin of male 
with first produced ray not serrated, third longer 
than posterior branch of second . 

Teeth broad incisors; anal fin of male with first pro- 
duced ray serrated not far from tip, third shorter 
than posterior branch of second .. bere ee 

*** Mouth small, fanevere teeth TnI 
oar-shaped. Extremity of anal fin of 
male supported equally by first and 
anterior branch of second produced 
rays; first not serrated ..................055 

B. Anal fin of male short, ending in a small 
retrorse hook formed by second and 
third produced rays; teeth conical or 
villiform 

y. Anal fin of male Tone erg ‘dlandie, me 
ending in an antrorse hook. 

Bones of lower jaw firmly united; teeth conical or villi- 
form, fixed .. : 

Bones of lower jaw ‘loosely connected ; ‘teeth more or less 
ease and compressed, mov able : 

. First produced ray of anal fin of male maine 
in an antrorse appendage; mouth small, but 
bones of lower jaw rather firmly joined. 

Appendage forked, each fork antler-like; teeth oar- 
shaped .. a de rsedcr sae edeateri aaah eernbbicia 

Appendage not forked, ‘long, doubly curved, belnteds 
teeth chisel-shaped abit 

3. First produced ray of exell fin a et ean 
a pair of curved horn-like processes not far 
from its end. 

a. Bones of lower jaw firmly united. 

Teeth chisel-shaped ; outer series close-set 

Teeth spear-shaped ; outer series spaced UNS AS 

6. Bones of lower jaw ey connected ; teeth 

ROWAN, sccnonadocvne 


B. Pelvic fins enlarged in the “amallas ‘dhe ceuol | ray 
longest. 

1. Bones of lower jaw firmly united; outer series 

of teeth slender, pointed ..................... 000005 

2. Bones of lower jaw rather firmly united; outer 

series of teeth broad incisors .............0.00.0.. 

3. Bones of lower jaw loosely connected; outer 

series of teeth slender, curved, oar- or spoon- 
shaped ; intromittent organ short. 

a. Extremity of intromittent organ unpro- 
tected; first produced ray with terminal 
hook and strong subterminal serrations; 
anterior branch of second hooked forward ; 
posterior branch of second with distal seg- 
ments ‘produced into serrations beyond 
extremity of third. 


Third produced ray without hook; anterior branch of 


second unsegmented distally ; caudal similar in both 
SEXES) PL OFSAl OIA Re ate re mmeNins Ree eP A a) a epee Ser 
Third produced ray ending in a retrorse hook ; anterior 
branch of second segmented throughout; caudal 
of male with lower rays a into a long 
ee appendage. Dorsal 11-15.. ee F 
. Extremity of intromittent organ Spretened Ty 
a hood of thick skin that can be slipped off 
frontwards, being attached only at its base 
to the first prolonged ray. 
a, First prolonged ray tapering evenly, sharply 
serrated distally. 


8. Heterandria., 


9. Pseudopecilia. 


10. Peciliopsis. 


. 11. Brachyrhaphis. 


12. Leptorhaphis. 


. 13. Phalloptychus. 


. 14. Phalloceros. 


. 15. Cnesterodon. 


... 16. Glaridichthys. 
17. Toxus. 


. 18. Girardinus. 


19. Pamphorichthys. 


20. Pamphoria. 


21. Platypecilus. 


22, Xiphophorus. 


COYPRINODONT FISHES, 981 


Kirst prolonged ray without antrorse spine; last without 

LOUIE! FORCES ocd. ado condor sho adecaoocsougoscqdecnd sea doaneeeos ap JECHEC CG? 
Kirst prolonged ray without antrorse spine; end of last 

bearing a pair of processes directed outwards and 


towards)theybasevor the) sm) sesscueee ce sess sn cedasejaccies ses: 24. Lebistes. 
First prolonged ray bearing a small antrorse spine at or 
near its end; last with processes as in Lebistes ...... 25, Mollienisia. 


8. First prolonged ray becoming abruptly 
slender at some distance from end, not 
sharply serrated, bearing a small antrorse 
spine at or near its end; last prolonged 
‘ray without terminal processes ............ 26. Limia. 


. 1. Atraro Meek, 1912. 


Petalosoma (non Lewis) Regan, Ann. Mag. Nat. Hist. (8) i. 
1908, p. 462. 

Alfaro Meek, Field Mus. Publ. Zool. x., Sept. 1912, p. 72. 

Petalurichthys Regan, Ann. Mag. Nat. Hist. (8) x., Nov. 1912, 
p. 494. 

This genus resembles Gambusia, except for the sharp lower edge 
of the tail. The intromittent organ (text-fig. 169 F, p. 990) is 
simple in structure, and the pelvic fins are enlarged in the male. 


1, ALFARO CULTRATUS. 

Petalosoma cultratwum Regan, Ann. Mag. Nat. Hist. (8) ii. 
1908, p. 462. 

Alfaro acutiventralis Meek, Field Mus. Publ., Zool, x.1912, p.72. 

Depth 33 in the length. Dorsal 7-8; when laid back nearly 
reaching caudal. Anal 9-10, in advance of dorsal. 32 to 35 


scales in a longitudinal series. Total length 45 to 90 mm. 
Costa Rica. 


2. ALFARO AMAZONUM. 

Petalosoma amazonum Regan, Ann. Mag. Nat. Hist. (8) viii. 
1911, p. 659, figs. 

Depth 32 in the length. Dorsal 8-9, when laid back not 
reaching caudal. Anal 10, in advance of dorsal. 33 or 34 scales 
in a longitudinal series. Total length 35 to 51 mm. 

R. Amazon at Obidos. 


2. Tomreurus Eigenm., 1909. 
Ann. Carnegie Mus. vi. p. 53. 


TOMEURUS GRACILIS. 


‘Kigenm. |. c. and Mem. Carnegie Mus. v. 1912, p. 460, pl. Ixv. 
fies) 7—8: 


“British Guiana. 
3. GamBusiA Poey, 1855 *. 
Mem. i. p. 382. 
Gambusia (part.) Giinth. Cat. Fish, vi. p. 333 (1866); Garman, 


* G. picturata Poey (‘Synopsis,’ p. 410, 1868) is a doubtful species of uncertain 
position. 


982 MR. CG. TATE REGAN ON 


Mem. Mus, Comp. Zool. xix. 1895, p. 82; Regan, Biol. Centr.- 
Amer., Pisces, p. 93 (1907). 

Paragambusia Meek, Publ. Columbian Mus., Zool. v. 1904, p. 133. 

This genus, as now restricted, includes species in which the 
male intromittent organ is moderately long, nearly 4 of the length 
of the fish (without caudal), and is formed on a definite plan 
which is subject to considerable modification in the different 
species. The distal segments of the first prolonged ray are pro- 
duced into processes directed more or less towards its apex ; the 
anterior branch of the second prolonged ray is slender distally 
and at some distance from the “end has an antrorse projection 
which may be termed the “ elbow”; the posterior branch of this 
ray ends in a retrorse pointed hook or barb, and the segments 
immediately proximal to the elbow of the anterior branch are 
produced backwards into serre ; the third prolonged ray ends in 
a hook more or less similar to that of the second. 

Of the 17 species here recognized I have seen males of 9, the 

distal part of the anal fin of each of them is figured. Of Gam- 
busia holbrookii (text-fig. 169 A) [ have examined two adult males, 
of G. oligosticta (text-fig. 169 B) four, of G. nicaraguensis (text-fig. 
168 A) two, of G. wrayi (text-fig. 168 B) four, of G. gracilior 
(text-fig. 168 C) three, of G. dominicensis (text-fig. 169 C) two, 
of G. nigropunctata (text-fig. 168 F) one, of G. punctuta (text- 
fig. 168 D) three, and of G. senilis (text-fig. 168. E) one. These 
suffice to prove that the differences shown in the figures are really 
specific; the number, form, and size of the modified segments 
of the first prolonged ray, the number of segments distal to 
he elbow, the form of the hook, and the number and length 
of the serre of the second prolonged ray, and the form of the 
hook of the third, vary but slightly for each species. The most 
noticeable variation is in G. oligosticta, the other examples 
differing from the one figured in the addition of a segment 
to each hook, so that the anterior hook has a short stem and 
the posterior quite a long one. 

The following table gives the number of dorsal rays, of scales 
in a longitudinal series, and the distribution of the species :— 


Dz. be Se | Distribution. 
Ws Cho OOUDROOIPIE 250255 580 200406 8 30 | Virginia to Alabama. 
Ds (Cis FOOPEIOUIS oc0650 000005 sn6060 a 31-32 | Florida to Texas. 
Bb CR GIWES. don ceo s0ecncs00ecs00¢ 6-7 30-31 | Florida to Tampico. 
215 (CR EAOUMS Soadoasasosaccn soon 8-9 30 Chihuahua, Mexico. 
| 5. G. nicaraguensis............ 7-8 28 |S. Mexico to Nicaragua. 
| RE WO ed ccantacadace bag sagees sl iq 32 | Nicaragua. 
| 7. G. caudovittata ............ | 8 31 | R. San Juan, Colombia. 
Waanclymumctate me (cue emlln aLO 31-33 | Cuba. 
| O, @ nigropunctata ......... 9 30 | 9 
| 10. G. melanosticta ............ | BNO AVS) gg 
| ll. G. puncticulata : 9 | =830 hg 
| 12. G. melanopleura............| 11-12 | 30-81 | Jamaica. 
IS “Cro GUAGE Tooc'sed soe x60 oon seu deh 8 | 31-33 5 
\ 1a. Gy Oligosticta pen. ees ee) SH9 29-31 | eS 
WG (Cro GPRACH VOR a5 30408 600500006 8-9 31-33 5¢ 
16. G. dominicensis ............ 9 | - 28-29 | Haiti. 
17. G. caymanensis ............ 9 | 30-31 ) Grand Cayman. 


983 


CYPRINODONT FISHES, 


Text-fig. 168. 


Distal part of intromittent organ o 


C. G. gracilior 


Py 10), Et, 


p- 61 
334 (1866) ; 


Len} 
lor 
Ye 
pwc’ 
| 
Dee 
Ds 
Do eS 
Cate 
as, 
« fo: 
hee, 
FG te 
eres 
= Se 
lea 
Se ae 
@ 
oS) 2s 
a ace 
x AO 
Sl ee . 
Ses 
ee 2 


Gi 


kit 


t holbroo 


Cambusic 


1. GAMBUSIA HOLBROOKII. 
Fleterandria holbrooktt Gir: 


984 MR. ©, TATE REGAN ON 


Garman, Mem. Mus. Comp. Zool. xix. 1895, p. 82, pl. x1. 
figs. 4-13. 

Haplochilus melanops Cope, Proc. Amer. Phil. Soc. xi. 1877, 
p. 457, 

Zygonectes atrilatus Jord. & Brayton, Bull. U.S. Nat. Mus. x11. 
1878, p. 84. 

@. Depth of body 34 to 4 in the length, length of head 4. 
Diameter of eye 3 to 34.in length of head, interorbital width 2. 
30 scales in a longitudinal series, Dorsal 8; origin above 
posterior end of anal, equidistant from base of pectoral and base 
of caudal; longest rays 3 length of head, Anal 10. Pectoral 
8 length of head; pelvics extending to origin of anal. Least 
depth of caudal peduncle 2 length of head. Olivaceous ; usually 
a, dark bar below the eye; a blackish spot above the vent; dorsal 
and caudal fins with small dark spots. 

¢. Origin of dorsal fin equidistant from eye and base of 
caudal. 

Virginia to Alabama. 

Here described from specimens from North Carolina and 
Virginia, 3 females 38-48 mm, in total length and 2 males of 
26 mm. 


2. GAMBUSIA PATRUELIS, 


Heterandria patruelis Baird & Girard, Proc. Acad. Philad. 1854, 
p. 390. 

Gambusia patruelis Girard, U.S. Mex. Bound, Surv., Fish. p. Us 
pl. xxxix. figs, 1-7. 

2. Depth of body 3 in the length, length ef head 4. Diameter 
of eye 34 in length of head, interorbital width 2. 31 or 32 
scales in a longitudinal series. Dorsal 7; origin behind end of 
base of anal, nearer to base of caudal than to base of pectoral ; 
longest rays 2 the length of head. Anal 10; first branched ray 
longest. Pectoral 4 length of head; pelvies extending to vent. 
Least depth of caudal peduncle # length of head, Brownish 
above, yellowish below ; dorsal and caudal fins with small dark 
spots. 

Florida to Texas. 

Here described from two specimens from Pensacola (Jordan), 
AT and 50 mm, in total length, 


3. GAMBUSIA AFFINIS. 


Heterandria afinis Baird & Girard, Proe. Acad. Philad, 1854, 
| B05 
js Gambusia afinis Girard, U.S. Mex. Bound. Surv., Fish. p. 72, 
pl. xxxix. figs. 12-15 (1859); Giinth. Cat. Fish. vi. p. 336 
(1866). 
Gambusia speciosa Girard, Proc, Acad. Philad, 1859, p. 121. 
Gambusia gracihs Girard, |. ¢. 
Gambusia humilis Giinth. t. ¢, p. 334. 


CYPRINODONT FISHES. 985 


Zygonectes brachypterus Cope, Bull. U.S. Nat. Mus. xx. 1880, 
p- 34. 


Zygonectes inurus Jord. & Gilb. Proc. U.S. Nat. Mus. 1882, 
p- 043. 

2. Depth of body 33 to 44 in the length, length of head 
31 to 41, Diameter of eye 32 to 4 in length “of Head) interorbital 
width re to 2. 30 or 31 scales in a longitudinal series. Dorsal 
6-7; origin above posterior part or end of anal, equidistant from 
some part of operculum or base of pectoral and base of caudal ; 
longest rays 4 to $ length of head. Anal 10-11. Pectoral from 
2 to nearly as lone as head; pelvics reaching vent or origin of 
omell Brownish or olivaceous above, yellowish or silvery below ; 
scales dark-edged; usually a dark suborbital bar; dorsal and 
caudal fins with small dark spots. 

Florida to Tampico; Mississippi. 

Numerous females, measuring up to 50 mm. in total length, 
from Florida and Louisiana; one from Tampico. 


4, GAMBUSIA SENILIS. (Text-fig. 168 E.) 


? Heterandria nobilis Baird & Girard, Proc. Acad. Philad. 18538, 
ps 390; 

¢Gambusia nobilis Girard, U.S. Mex. Bound. Surv., Fish. p. 71, 
pl. xxxix. figs. 8-11 (1859). 

Gambusia senilis Girar d, Proc. Acad. Philad. 1859, p. 122 


2. Depth of body 3 to 3} in the length, length of head 32 6 4, 
Diameter of eye 34 in length of head, interorbital width 2. 
30 scales in a longitudinal series. Dorsal 8-9; origin above 
middle or posterior ‘part of anal, equidistant from base of pectoral 
and base of caudal ; longest rays 3 length of head. Anal 9-10. 
Pectoral 2 length of head ; pelvics reaching vent. Least depth 
of caudal ‘peduncle 2 a length of head. Brownish above, yellowish 
below; scales dark- ed ged, especially on sides of abdomen ; a dark 
suborbital bar and a “dusky. lateral band from eye to base of 
caudal. 

3. Origin of dorsal fin equidistant from preeoperculum and 
base of caudal. 

Chihuahua, Mexico, 

Several females, 45 to 50 mm. in total length, and one 
male of 25 mm. 


5. GAMBUSIA NICARAGUENSIS. (‘Text-fig. 168 A.) 


Gambusia nicaraguensis Ginth, Cat. Fish. vi. p. 336 (1866), and 
Trans. Zool, Soc. vi. 1868, p. 483, pl. irxxii. fig. 3; Regan, Biol. 
Centr.-Amer., Pisces, p. 96 (1907). 

Paragambusia nicaraguensis Meek, Publ. Columbian Mus., Zool. 
v. 1904, p. 135. 

2. Depth of body about 3 in the length, length of head 
31 to 4. Diameter of eye 3 to 3% in length of head, inter- 
orbital width 2. 28 scales‘in a longitudinal ° series. Dorsal 7-8 ; 


986 MR. C. TALE REGAN ON 


origin above or a little behind posterior end of anal, twice as far 
from middle of eye as from base of caudal; longest rays # or 2 
length of head. Anal 11; second branched ray longest, edge of 
fin emarginate. Pectoral a little shorter than head; _pelvics 
reaching vent or origin of anal. Least depth of caudal peduncle 
2 length of head. Upper part of body with 2 or 3 series of small 
dark spots along the rows of scales; dorsal and caudal fins 
spotted. 

3. Origin of dorsal fin twice as distant from end of snout as 
from base of caudal. 

Southern Mexico; Nicaragua. 

Several females, 40 to 55 mm. in total length, and two males 
of 30 mm., including the types from Lake Nicaragua, and speci- 
mens from El Hule (Aeek) and Coaxacoaleos (Arnold). 


6. GAMBUSIA DOVII, sp. n. 
2. Depth of body 3? in the length, length of head 3}. Dia- 


meter of eye 31 in length of head, sneer bital width 13. 32 scales 
in a longitudinal series. Darel 7; origin behind ond of anal, 
twice as “ion from end of snout as from base of caudal. Anal 10; 
anterior branched rays longest, but edge not emarginate. Least 
depth of caudal peduncle slightly more than } length of head. 
Olivaceous ; traces of spots on back and on caudal fin. 

Lake Nicaragua. 

A single specimen of 33 mm., collected by Captain Dow. 


7. GAMBUSIA CAUDOVITTATA. 
Regan, Ann. Mag. Nat. Hist. (8) xii. 1913, p. 471. 


Recently described from a single specimen from the Condoto, 
a tributary of the San Juan, W. Colombia. 


8. GAMBUSIA PUNCTATA. (Text-fig. 168 D.) 


Gambusia punctata Poey, Mem. i. p. 384 (1855); Giinth. 
Cat. Fish. vi. p. 334 (1866); Jord. & Everm. Bull. U.S. Nat. 
Mus. xlvii. 1896, p. 680. 

2. Depth of body 3 to 4 in the length, length of head 34 
to 4. Diameter of eye 3 to 31 in length of head, interorbital 
width 2. 31 to 33 scales in a ‘longitudinal series. Dorsal 10; 
origin above posterior end of anal, nearer to base of caudal than 
to head; longest rays 2 to 1 the length of head. Anal 11; last 
simple or first. branched ray longest ; edge of fin straight or 
slightly emarginate. Pectoral # length of head pelvics re aching 
vent. Least “depth of caudal ‘peduncle tt o 8 length of nena 
Upper part of side with a dark spot on each scale, forming 3 to 5 
regular longitudinal series; usually small dark spots on dorsal 
and caudal. 

3. Dorsal origin about equidistant from head and base of 
caudal ; longest rays } length of head. 

Cuba. 

Here described from 16 females, measuring up to 85 mm. in 
total length, and 3 males of 40 to 50 mm. 


CYPRINODONT FISHES, 987 


9. GAMBUSIA NIGROPUNCTATA, sp. n. (Text-fig. 168 F.) 

2. Depth of body 34 to 32 in the length, length of head 
32 to 32, Diameter of eye 3 to 3+ in length of head, interorbital 
width 2 to 21. 30 scales in a longitudinal series. Dorsal 9: 
origin above posterior part of anal, equidistant from base of 
pectoral and base of caudal. Anall0-11. Least depth of caudal 
peduncle less than # length of head. Coloration as in related 
species. 

3. Depth of body 4 in the length, length of head 32. 
Diameter of eye 3 in length of ea mieror pital width mL, 
Dorsal origin equidistant from middle of opercle and base of 
caudal ; longest rays > length of head. Pectoral 2 length of 
head. 

Cuba. 

Here described from two females of 34 and 50 mm. anda male 
of 25 mm. from Fermina, Bemba. 


10. GAMBUSIA MELANOSTICTA, Sp. 0. 


2. Depth of body 3 in the length, length of head 34 to 3!. 
Diameter of eye 3 to 31 in the Tenesh of head, interorbital 
width 2. 29 to 30 scales in a longitudinal series. Dorsal 9-10; 
origin ALOE end of anal, equidistant from base of pectoral ain 
base of caudal ; longest rays } to 2 engin of head. Anal 10-11; 
first. branched ray Tlongest, Pectoral 2 to 2 length of head ; 
pelvics reaching vent. ‘Least depth of cma peduncle 2 a leneth 
of head. Brownish above, golden below; upper par ts with 
seattered small dark spots; dorsal and caudal fins spotted. 

Cuba. 

Three females from Havana, 35 to 42 mm. in tctal length, 
received from Dr. D.S Jordan. 


11. GAMBUSIA PUNCTICULATA. 


Gambusia puncticulata Poey, Mem. 1. pp. 386, 390, pl. xxxi. 
figs. 6, 7 (1855); Gunth. Cat. Fish. vi. p. 334 (1866) ; Garman, 
Mem. Mus. Comp. Zool. xix. 1895, p. 87. 

@. Depth of body 34+ to 34 in the length, length of head 
4 to 41. Diameter of eye 33 in length of head, interorbital 
width 2. Mouth smaller than in any other species of the genus. 
29 or 30 scales in a longitudinal series. Dorsal 9; origin above 
middle or posterior part of anal, equidistant from head and base 
of caudal; fin rounded, the longest rays 3 length of head. 
Anal 11, pointed. Pectoral 2 or # length of head ; pelvics small. 
Least depth of caudal peduncle 3 " length of head. Upper parts 
with scattered small dark spots ; 2 or 3 series of small black spots 
on dorsal and caudal fins. 

3. Origin o dorsal equidistant from eye and base of caudal; 
longest rays 3 length of head. 

Cuba. 

Here described from two females, 35 mm. in total length, pre- 
sented by Capta:n Vipan. 


988 MR. CG. TATE REGAN ON 


12. GAMBUSIA MELANOPLEURA. 


Pecilia melanopleura Gosse, Soj. in Jamaica, p. 84, pl. i. fig. 3 
(1851). 

Haplochilus melanopleurus Giinth. Cat. Fish. vi. p. 317 (1866). 

Gambusia melanopleura Garman, Mem. Mus. Comp. Zool. xix. 


1895, p. 88. 


Depth of body 3 in the length, length of head 33. Diameter 
of eye 3 to 31 in length of head, interorbital width 1}. 30 or 31 
scales in a longitudinal series. Dorsal 11-12; origin above 
posterior part of anal, equidistant from base of pectoral and base 
of caudal; longest rays 4 length of head. Anal 10-11; branched 
rays slightly decreasing from first or second. Pectoral 7 length 
of head; pelvics reaching vent. Least depth of caudal peduncle 
3 or 2 length of head. Traces of spots on dorsal and anal fins 
and in some of a few spots on the back. 

Jamaica. : 

Several females, the largest 47 mm. in total length, types of the 
species. 


13. GAMBUSIA WRAYT, sp. n. (PI. XCIX. figs. 3, 4, and Text- 
fig. 168 B.) 

2. Depth of body 33 to 4 in the length, length of head 
32 to 34. Diameter of eye 3 to 33 in length of head, inter- 
orbital width 2. 31 to 383 scales in a longitudinal series. 
Dorsal 8; origin above posterior part of anal, equidistant from 
base of pectoral and base of caudal; longest rays 3 length of 
head. Anal 10-11; second branched ray longest. Pectoral 
8 length of head; pelvics reaching vent. Caudal rounded or 
subtruneate. Least depth of caudal peduncle 3 to 2 length of 
head. Coloration probably as in G. gracilior. 

3. Dorsal origin equidistant from middle or posterior part of 
operculum and base of caudal; longest ray # length of head. 
Least depth of caudal peduncle # or 2 length of head. 

Jamaica. 

Hight females measuring up to 55 mm. in total length, and 
four adult males of 35 to 40 mm., collected by C. A. Wray. 


14. GamBustIA oLicosticta, sp.n. (Pl. XCIX. figs. 1, 2, and 
Text-fig. 169 B.) 

2. Depth of body 22 to 34 in the length, length of head 
3 to 34. Diameter of eye 3 to 34 in length of head, inter- 
orbital width 2. 29 to 31 scales in a longitudinal series. Dorsal 
8-9; origin above last 2 or 3 rays of anal, equidistant from base 
of caudal and base of pectoral; fin rounded, longest rays 4 length 
of head. Anal 10-11; first branched ray longest, the rest 
regularly decreasing. Pectoral length of head ; pelvics reaching 
vent. Caudal rounded or subtruncate. Least depth of caudal 
peduncle 2 or 2 the length of head. Brownish above, yellowish 
below ; edges of scales darker; usually a dark lateral stripe and 
a few scattered spots; dorsal and caudal with small spots. 


CYPRINODONT FISHES. 989 


$. Dorsal origin equidistant from eye or postorbital part of 
head and base of caudal; longest rays 2 the length of head. 

Jamaica. 

Six females, measuring up to 47 mm. in total length, and seven 
males, the largest 33 mm. Some of these were collected by 
C. A. Wray, others by the Rev. J. Seed Roberts. 


15. GAMBUSIA GRACILIOR, sp. n. (PI. XCIX. figs. 5, 6, and 
Text-fig. 168 C.) 

2. Depth of body equal to or a little more than length of 
head, which is 3} (young) to 4 in the length of the fish. 
Diameter of eye 3 in length of head, interorbital width 2. 
31 to 33 scales in a longitudinal series. Dorsal 8-9; origin 
above end of anal, a little nearer to base of caudal than to base 
of pectoral ; fin rounded, longest rays } length of head. Anal 
LO=11; first or second br: snched rays longest, the rest regularly 
decreasing. Pectoral as long as head without snout; pelvics 
reaching vent. Caudal rounded or subtruneate. Caudal ‘peduncle 
slender, its least depth + the length of head. Brownish above, 
yellowish below; usually a narrow dark lateral stripe, some- 
times a few scattered dark spots; dorsal and caudal with small 
spots. 

3S. Dorsal further forward and more elevated ; origin about 
equidistant from middle of opercle and base of caudal; longest 
vays about 2 the length of head. Least depth of caudal peduncle 
3 the length of head. 

_ Jamaica. — 

Hight females, measuring up to 50 mm. in total length, and 
five males, the largest 37 mm., collected by C. Wray. These were 
mixed up with the examples of G. oligosticta, and could be picked 
out by their more slender form and smaller head, characters 
found to be associated with differences in the structure of the 
intromittent organ. ‘ 


16. GAMBUSIA DOMINICENSIS, sp. n. (PI. XCIX. fig. 7, and 
Text-fig. 169 C.) 

2. Depth of body 34 in the length, length of head 3%, 
Diameter of eye 34 in length of head, interorbital width 2. 
28 or 29 scales in a longitudinal series. Dorsal 9; origin above 
posterior end of anal, nearer base of caudal than base of pectoral ; 
longest rays 3 length of head. Anal 10-11; first branched ray 
longest. Pectoral # length of head; pelvics reaching origin of 
anal. Least depth of caudal peduncle 2 length of head. 
Brownish, scales dark-edged; a faint lateral stripe; a few 
scattered dark spots on upper parts; abdomen golden; dorsaland 
caudal fins with series of small dark spots. 

¢. Dorsal origin equidistant from head and base of caudal. 

Haiti. 

A female of 52 mm., and two males, each 25 mm. in total 
length, presented by Herr A. Rachow. 


Proc, Zoou. Soc.—1913, No. LX VI. 66 


990 MR. C. TATE REGAN ON 


17. GAMBUSIA CAYMANENSIS, Sp. n. 
Depth of body 3} to 4 in the length, length of head 3 to 31, 
Diameter of eye 3 to 34 in length of head, interorbital width 


Text-fig. 169. 


ee 
oS 


oA 


} 


oo 
== Spy 
male 


Distal part of intromittent organ of A. Gambusia holbrookii. B. G. oligosticta. 
C. G. dominicensis. D. Brachyrhaphis rhabdophora. HK. Belonesow beli- 


zanus. FE. Alfaro amazonum. 


CYPRINODONT FISHES. 991 


2 to 24. 30 or 31 scales in a longitudinal series. Dorsal 9 ; 
origin above middle of anal, equidistant from operculum and base 
of caudal ; longest rays nearly $ length of head. Anal 10; first 
branched ray longest. Pectoral # length of head; pelvics reaching 
vent. Least depth of caudal peduncle 4 length of head. Oliva- 
ceous ; scales dark-edged; a dark bar below eye; dorsal fin with 
two series of dark spots; caudal faintly spotted. 

Grand Cayman. 

Two females, 22 and 26 mm. in total length. 


4. Brtonzesox Kner, 1860. 


Sitzungsb. Akad. Wien, xl. p. 419. 


Related to Gambusia, but the jaws produced, with broad bands 
of slender, pointed, depressible teeth. Anal fin of male very 
similar to that of Gambusia, differing in that the terminal seg- 
ments of the first produced ray are not serrated, while rather 
small retrorse spines at the ends of the third and the posterior 
branch of the second produced rays represent the hooks of 
Gambusia. 

A single species. 


BELONESOX BELIZANUS. (Text-fig. 169 E.) 


Belonesox belizanus Kner, Sitzungsb. Akad. Wien, xl. 1860, 
p- 419, fig.; Giinth. Cat. Fish. vi. p. 333 (1866); Meek, Publ. 
Columbian Mus., Zool. v. 1£04, p. 135; Regan, Biol. Centr.-Amer., 
Pisces, p. 93 (1907). 

Depth of body 4 to 6 in the length, length of head nearly 3. 
Snout 4 the length of head. 55 to 65 scales in a longitudinal 
series. Dorsal 9-10. Anal 10-12, in advance of dorsal. 

Atlantic Slope of Vera Cruz, British Honduras, and Guatemala. 

The largest species of the group, the males attaining 100 mm., 
the females 200 mm. in total length. Here described from ten 
specimens from Perez, Belize, and Lake Peten. 


5). PRIAPICHTHYS, gen. nov. 


Differs from Gambusia in that the intromittent organ is longer, 
when laid back nearly reaching the caudal fin, and is quite 
differently formed ; the terminal part is hooked forward and the 
prolonged rays taper distally and have no specially modified 
segments, except the usual serrz of the posterior branch of the 
second ; the first prolonged ray does not extend to the apex, and 
the hook is supported by the anterior branch of the second and 
the third (text-fig. 170 B). 

It is here assumed that five species from Central America and 
Colombia are congeneric with P. annectens, the only species of 
which I have examined males; in all but P. parismina males are 
known and agree with those of P. annectens in their long intro- 
mittent organ. 

66* 


992 MR. C. TATE REGAN ON 


Synopsis of the Species. 


T. Origin of dorsal fin in, or a little in advance of, middle of 
entire length (including caudal fin), above anterior part or 
middle of anal. 


A. D. 10-12; no black spot on anal fin ...................:...... anmnectens. 

B. D. 8-9; a blackish spot on anal fin. 
Length of head 33 to 32 in length of fish (without caudal) ......... episcopi. 
Length of head 43 to 4% in length of fish (without caudal) ......... nigroventralis. 


II. Origin of dorsal fin much nearer to end of caudal fin than 
to tip of snout, above posterior part or posterior end of 


anal. D. 7-8. 
A blackish spot on anal fin .. ue nagonogovevasccaoonca  fXGMPOSTOONR. 
7 or 8 dark vertical bars on posterior part. of body... spnosnaancosse — UPC AOCOEPRo 
8 or 9 dark bars on anterior # of body ee eee eR eI CLUE IULOCULCTUS7 Ss 


1. PRIAPICHTHYS ANNECTENS. (Text-fig. 170 B.) 


Gambusia annectens Regan, Ann. Mag. N. H. (7) a, IOOG, 
p. 259, and Biol. Centr.- Amer., I2nCSS, [9s Oi, jk omy, IGE, Ds 6 
(1907). 

Costa Rica. 


2. PRIAPICHTHYS EPISCOPI. 

Gambusia episcopi Steind. Sitzungsb. Akad. Wien, Ixxvul. 1878, 
p. 387, pl. 11, figs. 3, 4. 

Panama. 

3. PRIAPICHTHYS NIGROVENTRALIS. 


Gambusia nigroventralis Kigenm. Indiana Univ. Studies, 1912, 
No. 8, p. 26. 


Rio San Juan, Colombia. 
4, PRIAPICHTHYS PARISMINA. 


Gambusia parismina Meek, Publ. Field. Mus., Zool. x. 1912, 
Oa Mlk: 
Costa Rica. 


5. PRIAPICHTHYS TRIDENTIGER. 


Gambusia tridentiger Garman, Mem. Mus. Comp. Zool. xix. 
1895, p. 89. 


Panama. 


. PRIAPICHTHYS TURRUBARENSIS. 
phage turrubarensis Meek, Publ. Field Mus., Zool. x. 1912, 
joa Ibs 
Costa Rica. 


Perhaps a synonym of P. tridentiger. 
6. PRIAPELLA, gen. nov. 


This genus shows relationship to Priapichthys in the structure 
of the long intromittent organ as well as in other characters. 


CYPRINODONT FISHES. 993 


PRIAPELLA BONITA. (Text-fig. 170 E.) 


Gambusia bonita Meek, Publ. Columbian Mus., Zool. v. 1904, 
p. 132, fig. 39; Regan, Biol. Centr.-Amer., Pisces, p. 95 
(1907). 

Rio Papaloapam in Mexico. 


7. PskupoxreHoPHorws Bleek., 1863. 


Atl. Ichth. iii. p. 140; Giinth. Cat. Fish. vi. p. 332 (1866). 


Differs from Priapichthys in the longer dorsal fin originating 
in advance of the anal (9) and in the structure of the intro- 
mittent organ (text-fig. 170 C), with the anterior branch of the 
second prolonged ray ending in an unsegmented antrorse hook, 
at the base of which the first and third prolonged rays terminate. 

There are three species from Mexico and Central America, but 
males of one only (P. bimaculatus) have been described. 


1. PSEUDOXIPHOPHORUS TERRABENSIS. 


Gambusia terrabensis Regan, Ann. Mag. N. H. (7) xix. 1907, 
p- 260; Biol. Centr.-Amer., Pisces, p. 97, pl. xii. fig. 7 (1907). 

Dorsal 12-14; origin nearly equidistant from tip of snout and 
base of caudal. Anal 9-10; origin nearly below middle of dorsal. 
No dark spot above pectoral. 

Rio Grande de Terraba, Costa Rica. 


2. PSEUDOXIPHOPHORUS JONESII. 


Mollienisia jones Giinth. Ann. Mag. N. H. (4) xiv. 1874, 
otal 
: Pseudoxiphophorus pauciradiatus Regan, Ann. Mag. N. H. (7) 
xii. 1904, p. 256 and xvi. 1905, p. 362. 

Gambusia jonesti Regan, Biol. Centr.-Amer., Pisces, p. 97, pl. xii. 
fig. 8 (1907). 

Dorsal 11-13; origin equidistant from tip of snout and middle 
or posterior part of caudal. Anal 9-11, below anterior part of 
dorsal. A dark spot above pectoral. 

Mountain lakes and streams of Central Vera Cruz, Mexico. 


3. PSEUDOXIPHOPHORUS BIMACULATUS. (Text-fig. 170 C.) 


Aiphophorus bimaculatus Heck. Sitzungsb. Akad. Wien, i. 1848, 
[> ASO, Jol, me sles, My A, 

Pseudoxiphophorus bimaculatus Ginth, Cat. Fish. vi. p. 332 
(1866) ; Garman, Mem. Mus. Comp. Zool. xix. 1895, p. 81. 

Gambusia bimaculata Regan, Biol. Centr.-Amer., Pisces, p. 98, 
pl. xiv. fig. 4. 

Dorsal 13-17 ; origin nearly equidistant from tip of snout and 


994 MR. C. TATE REGAN ON 


base of caudal. Anal 9-11, below anterior part of dorsal. A 
dark spot above pectoral. 


Southern Mexico, British Honduras, and Guatemala. 


Text-fig. 170. 


Distal part of intromittent organ of A. Heterandria formosa. B. Priapichthys 
annectens. C. Pseudoxiphophorus bimaculatus. D. Poeciliopsis presidionis. 
KE. Priapella bonita. ¥. Pseudopeeilia feste. 


CYPRINODONT FISHES. 995 


8. Hereranpria, Agass., 1853. 


Amer. Journ. Sci. xvi. p. 135. 

Girardinus (part.) Giinth. Cat. Fish. vi. p. 351 (1866). 

Heterandria (part.) Garman, Mem. Mus. Comp. Zool. xix. 1895, 
[Ds BOs 

Differs from Gambusia and Priapichthys in the small transverse 
mouth and compressed pointed teeth, the enlarged outer teeth 
forming a close-set series. The intromittent organ is elongate, 
as in Priapichthys, and its structure indicates a close relationship 
to that genus. It differs in that the third prolonged ray is 
scarcely longer than the posterior branch of the second, so that 
the hook is supported only by the anterior branch of the second. 

Two species from the United States and Mexico. 


1. HereranpriA Formosa. (Text-fig. 170 A.) 


Heterandria formosa Agass. Amer, Journ, Sci. xix. 1855, p. 136 ; 
Garman, Mem. Mus. Comp. Zool. xix. 1895, p. 91, pls. iv. fig. 7, 
viii. fig. 8, and xi. figs. 1-3 ; Jord. & Everm. Bull. U.S. Nat. Mus. 
xlvii. 1896, p. 687. 

Girardinus formosus Giinth. Cat. Fish. vi. p. 354 (1866). 

Depth of body 32 to 4 in the length, length of head 33 
to 4. Diameter of eye 3 to 32 in length of head, interorbital 
width 2. 28 to 30 scales in a longitudinal series. Dorsal 7-8 ; 
origin above middle of anal, scarcely nearer to end of caudal than 
to tip of snout. Anall0. A dark lateral band crossed by several 
vertical bars ;,a blackish spot on basal part of dorsal, another on 
anal. 

South Carolina to Florida. 

Hight specimens, the largest female 30 imm., the males 15 to 
20 mm. in total length. 


2. HETERANDRIA FASCIATA. 


Gambusia fasciata Meek, Publ. Columbian Mus., Zool. v. 1904, 
p. 129, fig. 37; Regan, Biol. Centr.-Amer., Pisces, p. 95 (1907). 

Very similar to the preceding in form and coloration, except 
for the absence of the lateral band, differing especially in the 
more posterior dorsal fin, its origin above end of anal and much 
nearer to end of caudal than to tip of snout. Dorsal 8. Anal 
9-10. 32 scales in a longitudinal series. 

Southern Mexico. 

Total length 55 mm. 

Six specimens from Oaxaca and Tequesixtlan (Gadow). 

The male of this species has not yet been described. 


9. PSEUDOPGCILIA, gen. nov. 
Closely related to Heterandria, but with the teeth broad 
truncated incisors and the intromittent organ somewhat different 
in structure. 


996 MR. C. TATE REGAN ON 


PSEUDOPGCILIA FESTH. (Text-fig. 170 F.) 

Pecilia feste Bouleng. Boll. Mus. Zool. Torino, xii. 1898, 
No. 329, p. 13. 

Santa Elena, Western Ecuador. 


10. PasciLiopsis, gen. nov, 


This genus has the mouth and dentition of Pecilia, but in the 
males the pelvic fins are neither enlarged nor modified, and the 
intromittent organ is long and slender. The first prolonged ray 
and the anterior branch of the second extend to the end of the 
fin and are somewhat curved forward distally; the posterior 
branch of the second is short and the third still shorter and 
slender distally, so that the serrations of the second project 
beyond it (text-figs. 170 D and 171 B). 


Mexico, Central America, and Colombia. 


Synopsis of the Species. 


1. Dorsal origin behind that of anal. 


Sides of body with a series of vertical bars ......... 1. presidionis. 
A lateral series of spots, pave more than 8 in number and 


smaller than the eye 2. lutzi. 
A lateral series of 4 to 8 spots, , each about as lar: oe | as the eye 3. pleurospilus. 
No bars or spots Se Metteaaetaesanees rc puede 4. retropinna. 

2. Dorsal origin in advance of that of anal. 

Diameter of eye 3 to 34 in length of head, which is 3} to 32 in 

the length of fish .......... .... 5. isthmensis. 
Diameter of eye 2% to en in length of head, ‘which is 4 to 4h in 

the length of fish . Las diaratinelst bisGemiaus Cannel et aa tenes eieenesen), OTM DUCUCeIAUs 


1. Pa@cILiopsis PRESIDIONIS. (Text-fig. 170 D.) 

Pecilia presidionis Jord. & Culver, Proc. Calif. Aead. (2) v. 
1895, p. 413, pl. xxix. 

Girardinus presidionis Regan, Biol. Centr.-Amer., Pisces, p. 99 
(1907). 

As I have stated in the ‘ Biologia’ this form and the next seem 
to differ from P. pleurospilus only in coloration, and should perhaps 
be regarded as subspecies of that species. 

Rio Presidio in Sinaloa. 

Heterandria colombianus Kigenm. (Indiana Univ. Studies, 1912 
No. 8, p. 27), from brackish water at the mouth of the R. Dagua, 
Colombia, seems to be extremely similar to P. presidionis. 


2. Pa@cILIoPsIs LUTZI. 


Heterandria lutzi Meek, Publ. Colombian Mus., Zool. v. 1904, 
p. 148, fig. 47. 

Girardinus lutzi Regan, Biol. Centr.-Amer., Pisces, p. 99 
(1907). 


Oaxaca, Mexico; R. Motagua in Guatemala. 


CYPRINODONT FISHES. 997 


3. PdCILIOPsIs PLEUROSPILUS. 

Girardinus pleurospilus Giinth. Cat. Fish. vi. p. 355 (1866), 
and Trans. Zool. Soc. vi. 1868, p. 486, pl. Ixxxvil. fig. 1; Regan, 
Biol. Centr.-Amer., Pisces, p. 100 (1907). 


Lakes Duenas and Nacasil in Guatemala. 


4, PacILIOPsIS RETROPINNA. 

Pecilia retropinna Regan, Ann. Mag. Nat. Hist. (8) 11. 1908, 
p- 458. 

Costa Rica. 


. P@CILIOPSIS ISTHMENSIS, sp.n. (PI. C. figs.3, 4; and Text- 
fig. 171 B.) 

@. Depth of body 23 to 3 in the length, length of head 35 to 33. 
Diameter of eye 3 to 34 in length of head, interorbital width 
14 to 2. 26 to 28 scales in a longitudinal series. Dorsal 9-10; 
origin equidistant from anterior edge of eye and base of caudal; 
longest ray # length of head. Anal 10; origin below fourth or 
fifth ray of dorsal: ; first branched ray longest, 3? length of head or 
more. Pectoral a little shorter than head ; pelvics reaching anal. 
Least depth of caudal peduncle 2 or ? length of head. Olivaceous ; 
scales dark-edged ; a blackish spot aber ‘the vent; fins dusky, the 
dorsal with blackish basal band and dark edge. 

3. Dorsal origin equidistant from tip of snout and base of 
caudal, or nearer former; intromittent organ, when laid back, 
nearly or quite reaching caudal fin; indistinct cross-bars on body. 

Colon, Panama, 

8 females, 35 to 60 mm. in total length, and 4 males of 35 to 
42 mm., presented by Herr A. Rachow. 


6. PascILIOPsIs PITTIERT. 

Pecilia pittierts Meek, Field Mus. Publ., Zool. x. 1912, p. 71. 

Closely related to the preceding, but described as with a smaller 
head and lax ger eye. 


La Junta, Costa Rica. 
Total length 43 to 65 mm., males to 53 mm. 


11. BrRacHyRHAPHIS, gen. nov. 


Differs from Gambusia in the shorter intromittent organ ending 
ina small retrorse hook formed by the second and third pro- 
duced rays. 


BRACHYRHAPHIS RHABDOPHORA, (Text-fig. 169 D.) 


Gambusia rhabdophora Regan, Ann. Mag. Nat. Hist. (8) ii. 
1908, p. 457. 

Costa Rica, 

Specimens recently received show that in this species the origin 
of the anal fin may be behind below or in advance of that of the 
dorsal. 


998 MR. C. TATE REGAN ON 


Gambusia wmbratilis Meek (Publ. Field Mus., Zool. x. 1912, 
p- 70) seems to differ from B. rhabdophora only in the larger 
eye. 

12. LeprorHaPHIS, gen. nov. 


Differs from Gambusia only in the structure of the intromittent 
organ, which is longer and more slender than in that genus and 
slightly curved backwards distally. The first produced ray is not 
serrated, gradually tapers distally, and extends nearly to the tip 
of the fin; the second is slightly longer and has the posterior 
branch almost as long as the anterior and with the usual serra- 
tions, which project externally ; the third ray also seems to reach 
nearly to the tip of the fin and appears to meet the first and form 
a complete tube. 


LEPTORHAPHIS INFANS. 

® Xiphophorus gracilis Heck. Sitzungsb. Akad. Wien, i. 1848, 
p. 300, pl. ix. figs. 3, 4. 

Gambusia infans Woolman, Bull. U.S. Fish. Comm. 1894, 


Text-fig. 171. 


Distal part of intromittent organ of A. Phalloptychus januarius and 
B. Peciliopsis isthmensis. 'The rays have been separated. 


CYPRINODONT FISHES. 999 


p- 62, pl. 11. fig. 3; Meek, Publ. Columbian Mus., Zool. v. 1904, 
p- 131, fig. 38; Regan, Biol. Centr.-Amer., Pisces, p. 96 (1907). 
Gambusia gracilis Meek, t. c. p. 130. 
Rio Lerma and Rio Balsas in Mexico; ? Orizaba. 


13. PHatitorprycHus Higenmann. 
Proc. U.S. Nat. Mus. xxxii. 1907, p. 430. 


Mouth and dentition of Pecilia. Intromittent organ long 
and slender; third produced ray crossing second on right side 
near base, meeting first and forming a closed tube; second 
twisted distally so that the serrations are directed laterally or 
even forwards. 


PHALLOPTYCHUS JANUARIUS. (PI. C. figs. 7, 8, and text- 
terse L/S) 

Girardinus januarius Hensel, Arch. f. Nat. xxxiv. 1868,’ 
p. 360, and xxxv. 1869, p. 89. 

Girardinus theringw Bouleng. Ann. & Mag. Nat. Hist. (6) 
iv. 1889, p. 266. 

Girardinus zonatus Schreiner, Arch. Mus. Rio Janeiro, xii. 
LOSS ps We 

S.E. Brazil; La Plata. 

This species may at once be recognized by its coloration, 
having several narrow blackish vertical stripes on each side of 
the body. Dorsal 9; origin above ora little behind that of anal. 


14. PHALLOcCEROS Higenm. 
Proc. U.S. Nat. Mus. xxxii. 1907, p. 431. 


Mouth small, but bones of lower jaw rather firmly joined ; 
teeth oar-shaped, slender, curved, as in Pecilia. Intromittent 
organ long (about # length of fish to base of caudal) ; last segment 
of first produced ray modified into an antrorse appendage which 
bifurcates, each fork antler-like in form ; anterior branch of 
second ending just beyond the first in an antrorse process. 


PHALLOCEROS CAUDOMACULATUS. (PI. C. figs. 5, 6, and text- 
fig. 172 C.) 

Girardinus caudimaculatus Hensel, Arch. f. Nat. xxxiv. 1868, 
p. 362, and xxxv. 1869, p. 89. 

Glaridodon janwarius Garman, Mem. Mus. Comp. Zool. xix. 
1895, p. 42. 

S.E. Brazil; La Plata. 

The coloration readily distinguishes this species from Phallo- 
ptychus januarius. Plain olivaceous, usually with a vertical 
blackish spot on side below dorsal fin ; sometimes a lateral series 
of dusky bars. Dorsal 7-8; origin above middle or posterior 
part of anal. 


1000 MR. C. TATE REGAN ON 


15. CNESTERODON Garman. 


Mem. Mus. Comp. Zool. xix. 1895, p. 43. 

Gulapinnus Langer, Morph. Jahrb. xlvii. 1913, p. 207. 

Mouth small, but rami of lower jaw rather firmly joined and 
teeth chisel-shaped, broader than in Pecilia. Intromittent 
organ long (nearly 2 length of fish to base of caudal); last 
segment of first produced ray an appendage which is not forked, 
but is very long, peculiarly curved, and pointed distally; second 
ending in a process which is not or scarcely antrorse ; third 
terminating in a hook, as in Gambusia. 


CNESTERODON DECEMMACULATUS. (Text-fig. 172 B.) 


Pacilia decemmaculata Jenyns, Zool. ‘ Beagle,’ Fish. p. 115, 
pl. xxii. fig. 1 (1842). 

Pecilia gracilis Cuv. & Val. Hist. Nat. Poiss. xviii. 1846, 
p- 133. 

Cnesterodon decemmaculatus Garman, Mem. Mus. Comp. Zool. 
xix. 1895, p. 44. 

Cnesterodon carnegiet Haseman, Ann. Carnegie Mus. vii. 1911, 
p. 385, pl. Ixxxiil. 

La Plata; Rio Grande do Sul. 

A small species, easily recognized by the lateral series of dark, 
rounded or vertically expanded spots. Dorsal 8-9; origin above 
or a little behind that of anal. 


16. GLARIDICHTHYS Garman, 1896. 


Glaridodon (non Seeley, Proc. R. Soc. xliv. 1888, p. 135); 
Garman, Mem. Mus. Comp. Zool. xix. 1895, p. 40. 


This genus is well distinguished by the dentition and by the 
structure of the intromittent organ. It includes a single species 
from Cuba and perhaps one from Chihuahua, but males of the 
latter have not been described. 


1. GLARIDICHTHYS UNINOTATUS. 


Girardinus uninotatus Poey, Mem. ii. pp. 309, 383 (1861); 
Giinth. Cat. Fish. vi. p. 351 (1866). 

Glaridodon uninotatus Garman, Mem. Mus. Comp, Zool. xix. 
1895, p. 41. 

Glaridichthys faleatus Higenm. Bul]. U.S. Fish. Comm. xxii. 
p. 224, fig. (1904). 

Glaridichthys torralbasi Kigenm. t.c. p. 225, fig. 

2. Depth of body 3} to 44 in length, length of head 4 to 5. 
Hye large, more than 4 of length of head. 28 to 31 scales in a 
longitudinal series. Dorsal 9-11; origin nearly equidistant from 
head and base of caudal. Anal 10; origin well in advance of 
that of dorsal. Olivaceous ; edges of scales usually darker ; often 


1001 


CYPRINODONT FISHES. 


Text-fig. 172. 


intromittent organ of A. Girardinus metallicus. B. Cnesterodon 


Distal part of 


caudomaculatus. D. Platypecilus macu- 


latus. KE. Xiphophoru 


1002 MR. C. TATE REGAN ON 


a lateral series of short vertical bars; usually a blackish spot 
above origin of anal fin. 

3. Dorsal origin nearly equidistant from eye and base of 
caudal. Bars on side well marked; lateral spot indistinct; a 
blackish streak behind first produced ray of anal. 

Here deseribed from several specimens up to 75 mm. in total 
length from Cuba, including a male of 47 mm. extremely similar 
to the type of G. torralbasi, and co-types of G. faleatus. The last 
is a pale variety without the lateral spot, but structurally similar 
to G. uninotatus; the slight production of the vertical fins is 
found in several specimens with the typical coloration. 


2. GLARIDICHTHYS LATIDENS. 

Glaridodon latidens Garm. Mem. Mus. Comp. Zool. xix. 1895, 
p. 42. 

Chihuahua ; Mexico. 


17. Toxus Higenm., 1904. 
Bull. U.S. Fish. Comm. xxii. p. 226. 


This genus differs from the preceding only in the dentition. 


TOXUS RIDDLEI. 


Higenm. |. c¢. figs. 


Cuba. 
18. GrrarpDrINus Poey, 1855. 


Mem. i. pp. 383, 390; Garman, Mem. Mus. Comp. Zool. xix. 
1895, p. 45. 

Mouth and dentition as in Pecilia, but intromittent organ 
formed exactly as in the two preceding genera. 

A single species from Cuba. 


GIRARDINUS METALLICUS. (Text-fig. 172 A.) 

Poey, Mem. i. pp. 387, 391, pl. xxx1, figs. 8-11 (1855); Giinth. 
Cat. Fish. vi. p. 351 (1866); Garm. Mem. Mus. Comp. Zool. xix. 
1895, p. 46. 

Girardinus denticulatus Garm, t.¢. p. 47. 

Girardinus creolus Garm. |. ¢. 

Girardinus garmani Higenm. Bull. U.S. Fish. Comm. xxii. 
p. 226, fig. (1904). 

Heterandria cubensis Kigenm. t. c. p. 227, fig. 

Depth of body 3 to 4 in length, length of head 4 to 41. 
Diameter of eye 23 to 3 in length of head. 27 to 31 scales in a 
longitudinal series. Dorsal 9-10; origin nearly equidistant 
from base of pectoral and base of caudal (a little nearer head 
than caudal in fg). Anal 10-12; origin well in advance of 
dorsal. Olivaceous; edges of scales darker; usually a lateral 


CYPRINODONT FISHES. 1003 


series of dark bars; dorsal with anterior edge dark and usually 
a blackish spot or band posteriorly near base. 

Cuba. 

Several specimens up to 70 mm. in total length. 


19, PAMPHORICHTHYS, gen. nov. 


Mouth small; bones of lower jaw firmly united; a series of 
firmly set, slender, pointed teeth and a band of much smaller 
teeth within. Intestine long, coiled. Males with anal and 

elvic fins below pectorals, the former a slender pointed organ 
about + the length of the fish (without caudal), the latter also 
modified, the outer rays produced, the inner about 4 as long and 
forming a projection near their ends. 

This genus is based on Garman’s Heterandria minor, which 
seems to differ from Heterandria in the more slender teeth, 
longer intestine, and shorter intromittent organ, but especially in 
the modification of the pelvic fins in the male. 


PAMPHORICHTHYS MINOR. 


Heterandria minor Garman, Mem. Mus. Comp. Zool. xix. 
1895, p. 92. 

Amazon. 

20. PAMPHORIA, gen. nov. 

Mouth small; bones of lower jaw rather firmly joined; a series 
of broad incisors and a band of minute tricuspid teeth within, 
Intestine long.- Males with anal fin much advanced, forming an 
intromittent organ that is longer than the head, and with the 
second ray of the pelvics produced. 


PAMPHORIA SCALPRIDENS. 
Cnesterodon scalpridens Garman, Mem. Mus. Comp. Zool. xix. 
1895, p. 45, pl. v. fig. 12. pl. viii. fig. 17. 
Amazon. 
21. Piatypacitus Giinth., 1866. 


Cat. Fish. vi. p. 350. 

Mouth small, transverse ; bones of lower jaw loosely connected ; 
an outer series of slender curved teeth, somewhat expanded and 
compressed towards their apices, spoon-shaped or oar-shaped. 
Males with the pelvic fins enlarged, nearly as long as the intro- 
mittent organ, which is unprotected by a hood, but has the 
distal end remarkably modified (text-fig. 172 D). 

Two species from Mexico, with the intromittent organ precisely 
similar in structure. 


1, PLATYP@CILUS COUCHIANUS. 

Linvia couchiana Girard, Proc. Acad. Philad. 1859, p. 116. 

Pecilia couchiana Regan, Biol. Centr.-Amer., Pisces, p. 104, 
pleavarticoIN( 190m): 


1004 MR. C. TATE REGAN ON 


Depth of body about 2? in the length, length of head about 4. 
23 to 27 scales in a longitudinal series. Dorsal 9-11; origin 
equidistant from snout and anterior part of caudal. Anal 7-8, 
opposite posterior 4} of dorsal. Blackish or brownish above, 
yellowish below, the two colours sharply separated. 

Rio San Juan at Monterey. 


2, PLATYPa@cILUS MACcULATUS. (Text-fig. 172 D.) 

Platypecilus maculatus Giinth. Cat. Fish. vi. p. 350 (1866). 

Pecilia maculata Regan, Biol. Centr.-Amer., Pisces, p. 105 
(1907). 

Closely related to P. couchianus. Depth of body 2 to 22 in 
the length, length of head 34 to 32. Anal 8-9. Olivaceous; a 
dark spot or crescent or a pair of spots at base of caudal. 

Atlantic Coast streams of Mexico and Guatemala. 


22. XipHopHorus Heckel, 1848. 

Sitzungsb. Akad. Wien, 1. p. 291. 

Closely related to the preceding, as is shown by the great 
similarity in the structure of the anal fins of the males (text-fig. 
172K). The species occur in rivers of the Atlantic Slope from 
Tamaulipas to Guatemala. After examination of a large series 
of specimens I conclude that only 3 species are well established. 


1. XIPHOPHORUS MONTEZUM2. 


Jord. & Snyder, Bull. U.S. Fish. Comm. 1900, p. 131, fig. 11 ; 
Regan, Biol. Centr.-Amer., Pisces, p. 107 (1907). 

Dorsal 11-13. Anal 6-8. Scales 27 to 29. Depth about 3 
in the length, length of head about 4. Yellowish; scales of 
upper part of body dark-edged ; usually a dark lateral stripe. 

Rio Panuco, Mexico. 


2. XIPHOPHORUS HELLERI. (Text-fig. 172 EK.) 


Heck. Sitzungsb. Akad. Wien, i. 1848, p. 291, pl. viil.; Giinth. 
Cat. Fish. vi. p. 349 (1866); Regan, Biol. Centi.-Amer., Pisces, 
p. 107 (1907). 

Xiphophorus guentheri Jord. & Everm. Bull. U.S. Nat. Mus. 
xlvii. 1896, p. 702. 

Xiphophorus jalape Meek, Publ. Columbian Mus., Zool. v. 
1903, p. 136, pl. x1. 

NXiphophorus strigatus Regan, t.c. pl. xiv. fig. 7. 

Dorsal 11-14. Anal 8-10. Scales 26 to 30. Depth 3 to 4 
in the length, length of head 4 to 42. Sides of body blue, with 
red stripes between the series of scales. 

Southern Mexico ; Guatemala. 

Mexican examples have nearly always a strong stripe from eye 
to base of caudal (strigatis), the males sometimes with another 
backwards from axil of pectoral (helleri, jalape); in specimens 


CYPRINODONT FISHES, 1005 


from Guatemala there is, as a rule, no strong lateral stripe 
(guentheri): 

Xiphophorus rachovii Regan (Ann. Mag. Nat. Hist. (8) viii. 
1911, p. 373), from Puerto Barrios, Guatemala, has a pair of 
alinge spots at the base of the caudal foe other examples received 
later from Puerto Barrios lack these spots, but are not very 
different in other respects. Whether this is another variety of 
X. helleri or a hybrid with Platypecilus maculatus, as some 
aquarium-writers in Germany believe, still remains to be settled. 


3. XIPHOPHORUS BREVIS. 

Regan, Ann. Mag. Nat. Hist. (7) xix. 1907, p. 65, and Biol. 
Centr.-Amer., Pisces, p. 108, pl. xiv. figs. 8, 9. 

Dorsal 13-15. Anal 9-10. Scales 27. Depth 24 to 22 in 
length, length of head 32 to 32. No strong lateral stripe. 

British Honduras. 


23. Pacri1A Schneider, 1801. 


Bloch’s Syst. Ichth. p. 452. 
Pecilia (part.) Giinth. Cat. Fish. vi. p. 339 (1866); Garman, 
Mem. Mus. Comp. Zool. xix. 1895, p. 52. 


Mouth small, transverse; teeth slender, curved, expanded 
towards the tip, spoon- or oar-shaped, forming a series with a 
band of minute teeth within; bones of lower jaw loosely con- 
nected. Pelvic fins in male enlarged, the second ray prolonged. 
Intromittent organ short, its tip protected anteriorly by a cuta- 
neous hood; first prolonged ray without terminal spine, with 
several segments not far from the end forming acute serrations ; 
last ray simple, without appendages (text-fig. 173 A-C). 

As now restricted this genus only includes four (or five) 
species from South America. 


Synopsis of the Species. 


I. Dorsal 7-9; origin nearly abovelthatotanalu ners esse eee 1. vivipara. 


II. Dorsal 6-7. 
Origin of dorsal above posterior end of anal; usually a dark spot 
above pectoral, equidistant from head and dorsal fin ...... 2. pare. 
Origin of dorsal above middle or posterior part of anal; a dark 
spot on each scale and dark stripes between the series of 


SGAIES Gin SIGS Or |WOGhY soosdonsasnonc0 souBoodon aouiddesduadéode Hoe unboRReDD 3. picta. 
Origin of dorsal above middle of eee a blackish spot or bar 
at base of caudal fin .. noddonee candog cet onan cdacoooanécasennasanaoas — Gly MRED 


1. Pa@criia vivipaRA. (Text-fig. 173 C.) 

Pecilia vivipara Schneid. Bloch’s Syst. Ichth. p. 452, pl. Ixxxvi. 
fig. 2 (1801) ; Gunth. Cat. Fish. vi. p. 344 (1866); Garman, Mem. 
Mus. Comp. Zool. xix. 1895, p. 53. 

Pecilia schneideri Val. in Humboldt, Obs. Zool. ii. p. 159 
(1828); Cuv. & Val. Hist. Nat. Poiss. xviii. p. 135 (1846). 

Proc. Zoou. Soc.—1913, No. LX VII. 67 


1006 Mk. C. TATE REGAN ON 


Pecilia surinamensis Val. t. c. p. 158, pl. li. fig. 1; Cuv. & Val. 
t.c. p. 120. 

Pecilia unimaculata Val. t.c. p. 158, pl. li. figs. 2,5,6; Cuv. & 
Val. t.c. p. 128; Giinth. t.c. p. 346. 

2. Depth of body 22 to 4 in the length, length of head 34 to 44. 
Diameter of eye 3 to 4 in length of head, mterorbital width 
12 to 2. 25 to 27 scales in a longitudinal series. Dorsal 7-9 ; 
origin nearly above that of anal and equidistant from some part 
of operculum and base of caudal. Anal 8-10. Pectoral a little 
shorter than head; pelvics reaching vent. Least depth of caudal 
peduncle ? to # length of head. Olivaceous ; edges of scales usually 
darker ; body often with dark cross-bars ; often a blackish spot, 
which may be pale-edged, on upper part of side 2 or 3 scales in 
front of dorsal fin; often a pair of blackish spots at base of caudal 
above and below, extending along margins of fin ; fins sometimes 
immaculate, but dorsal and caudal sometimes with series of small 
spots, or dorsal with one or two blackish bands, 

3. Dorsal a little further forward and higher than in 9. 

Venezuela and Leeward Islands to the La Plata. 

Numerous examples, the largest female 80 mm. and the 
largest male 60 mm. 


2. Pacinia PARE. (Text-fig. 173 B.) 

Pecilia vivipara pare Kigenm. Ann. N.Y. Acad. vii. 1894, 
p- 629. 

? Pacilia amazonica Garman, Mem. Mus. Comp. Zool. xix. 
1895, p. 64. 

Acanthophacelus bifurcus Kigenm. Mem. Carnegie Mus. v. 
1912, p. 459, pl. Ixv. figs. 4-6. 

2. Depth of body 34 in the length, length of head 3}. 
Diameter of eye 34 in length of head, interorbital width 2. 28 
scales in a longitudinal series. Dorsal 6 ; origin above posterior 
end of anal, nearly equidistant from head and base of caudal ; 
middle rays longest, 4 length of head. Anal 8-9. Pectoral ? 
length of head; pelvics extending to origin of anal. Least 
depth of caudal peduncle 3 length of head. Olivaceous ; usually 
a pale lateral stripe; a vertically expanded dark spot with pale 
margin on upper part of side, nearly equidistant from head and 
dorsal fin ; fins immaculate. 

3. Dorsal origin nearer to head than to base of caudal ; 
longest rays ? length of head. A dark spot above pectoral (as in 
2), a larger one just above anal fin, a third on caudal peduncle ; 
dorsal and caudal sometimes spotted, latter with dark upper edge. 

Amazon; Guiana. 

Four specimens, two females of 28 mm. and two males of 22 
-and 24 mm. in total length, received from Prof. Eigenmann ; 
these are co-types of the species from Para and of A. bifureus 
from British Guiana. The intromittent organ is figured 
(text-fig. 173 B). 


Garman’s description of P. amazonica applies to P. pare, 


eee 


CYPRINODONT FISHES. 1007 


except that the dorsal origin is said to be a little in advance of 
that of anal. 


3. Pacruia PrcTaA, sp. n. (PI. C. figs. 1, 2, and Text-fig. 173 A.) 


Acanthophacelus melanzonus (part.) Kigenm. Ann. Carnegie 
Mus. vi. 1909, p. 51; Mem. Carnegie Mus. v. 1912, p. 457, 
pl. Ixiv. fig. 6. 

2. Depth of body 53 to 4 in the length, length of head about 4. 
Diameter of eye 33 to 4 in length of head, interorbital width 14. 
26 to 29 scales ina longitudinal series. Dorsal 6-7, origin above 
middle or posterior part of anal and equidistant from base of 
pectoral and base of caudal; fin small, rounded. Anal 9-10, 
pointed. Pectoral # length of head; pelvics reaching vent or 
origin of anal. Least depth of caudal peduncle about # length of 
head. Olivaceous; sides of body with a dark brown spot on each 
scale and dark brown longitudinal stripes between the series of 
scales. 

3. Depth 3 to 34 in the length. Dorsal origin nearly equi- 
distant from eye and base of caudal; fin elevated. Least depth 
of caudal peduncle 3 to ? length of head. Coloration sometimes 
as in female, but usually there are a few large dark spots on 
posterior part of body; dorsal fin usually with blackish spots ; 
often an ocellus on upper part of base of caudal fin. 

Demerara. 

Numerous examples, the females measuring up to 45 mm., 
the males to 30 mm. in total length, presented in 1872 by F. G. 
Beckford, Esq. 

A. melanzonus.is based on a male of Lebistes reticulatus (type) 
and several females of P. picta. 

In this species the intromittent organ differs somewhat from that 
of P. vivipara, but is formed on the same plan (text-fig. 173 A, C). 


4. P@cILIA BRANNERI. 


Peeilia brannert Eigenm. Ann, N.Y. Acad. vii. 1894, p. 629. 
Pecilia heteristia Regan, Ann. Mag. Nat. Hist. (8) iii. 1909, 
p. 235. 


2. Depth of body 33 in the length, length of head 4. Dia- 
meter of eye 33 in length of head, interorbital width nearly 2. 
27 or 28 scales in a longitudinal series. Dorsal 6-7; origin 
above middle of anal and nearly equidistant from base of pectoral 
and base of caudal. Anal 8, pointed. Pectoral ? length of head ; 
pelvics extending to origin of anal. Least depth of caudal 
peduncle 2 length of head. Olivaceous ; edges of scales darker ; 
some blackish vertical streaks on the side and a blackish spot or 
vertical bar at base of caudal fin. 

3. Dorsal origin equidistant from middle of operculum and 
base of caudal; two last rays produced into long filaments. 
Caudal spot larger than in Q ; sometimes a blackish stripe near 
upper edge of caudal fin. 

67* 


1008 MR. C. TATE REGAN ON 


Para. 

Here described from the types of P. heteristia, ¢ and 9, each 
35 mm. in total length. 

The intromittent organ is very like that of P. vivipara, but the 
anterior branch of the second ray has fewer segments, as in 
J OUCH 


24, Lepistes Filippi, 1861. 


Arch. Zool, Anat. Fisiol. i. p. 69. 

Acanthophacelus Kigenmann, Proc. U.S. Nat. Mus. xxxii. 1907, 
p- 426, fig. 1. 

Differs from Pecilia only in that the third prolonged anal 
ray of the male has its last segments modified into a pair of 
projections which are directed obliquely outwards and towards 
the base of the fin. 

A single species from South America. 


LEBISTES RETICULATUS. (Text-fig. 173 D.) 


Pecilia reticulata Peters, Monatsb. Akad. Berlin, 1859, p. 412; 
Garman, Mem. Mus. Comp. Zool. xix. 1895, p. 458. 

Lebistes pecilioides Filippi, Arch. Zool. Anat. Fisiol. 1. 1861, 
p- 69, pl. iv. f. 6; Giinth. Cat. Fish. vi. p. 356 (1866). 

Girardinus reticulatus Gunth. t. c. p. 352. 

Girardinus guppy Ginth. t.c. p. 353; Regan, Proc. Zool. Soc. 
1906, p. 390, pl. xxi. figs. 1, la. 

Acanthophacelus reticulatus Kigenm. Proc. U.S. Nat. Mus. 
xxxil. 1907, p. 426, f. 1; Mem. Carnegie Mus. v. 1912, p: 458, 
pl. Ixv. figs. 1-3. 

Acanthophacelus melanzonus (part.) Eigenm. Mem. Carnegie 
Mus. v. 1912, p. 457, pl. Ixiv. fig. 5. 

2. Depth of body 3 to 4 in the length, length of head 3 to 44. 
Diameter of eye 3 to 33 in length of head, interorbital width 14 
to 2. 26 to 28 scales in a longitudinal series. Dorsal 7-8, 
rounded or obtuse; origin equidistant from eye or operculum and 
base of caudal. Anal 9-10, pointed; origin below or a little in 
advance of that of dorsal. Pectoral $ length of head ; pelvics ex- 
tending to origin of anal. Least depth of caudal peduncle 2 2to ? 
length of head, Olivaceous; edges of scales darker, forming a 
agin: fins immaculate, exeEpe sometimes a small done spot on 
middle of caudal, 

3. Dorsal more elevated than in 9, when laid back nearly 
reaching base of caudal fin; pelvics nearly as long as the short 
intromittent organ. @oloration very variable ; often a blackish 
spot above or behind base of pectoral, another above anal fin or 
on middle of side, a third at base of caudal peduncle; often also 
several bluish-silvery spots, which may unite to form a lateral 
band, margined above and below by dark longitudinal stripes ; 
dorsal and caudal sometimes spotted, often dark-edged. 

Venezuela and St. Lucia to Santos. 


1009 


CYPRINODONT FISHES. | 


Text-fig. 173. 


Distal part of intromittent organ of :—A. Pecilia picta. B. P. pare. C. P. vivi- 


EF. Mollienisia 


KE. Limia versicolor. 


s reticulatus. 


S 


D. Lebist 


sphenops. ‘The pre 


pare. 


D. 


receding figures note that these ave reversed, the first 
y being on the right, the third on the left. 


uce-like hood is indieated in fig. 


D 


iI 


Tn comparing with p 


nn 


prolenged 1 


1010 MR. C. TATE REGAN ON 


Numerous examples, including co-types of the species received 
from the Berlin Museum and the types of Girardinus guppyi ; 
females to 5d mm., males to 25 mm., in total length. 


25. Moutrenis1a Le Sueur, 1821. 


Journ. Acad. Philad. ii. p. 3; Giinth. Cat. Fish. vi. p. 347 
(1866); Garman, Mem. Mus. Comp. Zool. xix. 1895, p. 49. 


This genus differs from Pacilia in having a small antrorse 
spine at the end of the first prolonged ray of the male anal fin, 
as in Limia,and at the end of the third a paired process, probably 
the modified terminal segment, directed obliquely outwards and 
towards the base of the fin, as in Lebistes (text-fig. 173 F), 

Some species hitherto referred to Pecilia (M. sphenops, 
M. gracilis) have an intromittent organ exactly similar to that of 
Molhienisia latipinna and M. petenensis, and as the little-known 
M. formosa is intermediate between the two groups in the size of 
the dorsal fin, all may be placed in JMollienisia. Some other 
species, with the dorsal fin further back than the anal, are pro- 
visionally included ; all that is known of the males is Garman’s 
statement that in P. cwneata the anal fin of the male is shorter 
than the head, indicating that the species is neither a Peciliopsis 
nor a Girardinus, although it may be a Limia. 


Synopsis of the Species. 


I. Dorsal 11-16; in adult males origin nearer end of snout than 
base of caudal, base longer than distanee from caudal. 
A. Caudal rounded or subtruncate (@ ) or lower angle slightly 
produced (4); scales 28 to 80. Dorsal 15-16 ............. 1. petenensis. 
B. Caudal rounded; scales 26 to 28. 


Dorsal 14-16; in females origin equidistant from end of snout 
and base of caudal (young) or nearer snout (adult); base equal 
to (young) or more than (adult) distance from caudal . 

Dorsal 11-13; in females origin a little nearer base of caudal than 
end of snout, even in the adult ; base equal to (adult) or less 
than (young) distance from cauiclalee Male ore ers ee Cee 3. formosa. 


II. Dorsal 10-11; origin nearly equidistant from end of snout 

and base of caudal fin; base shorter than distance from 

caudal; anal origin below fourth ray of dorsal; scales 28 

POND O)) Peper erate case seins sip ye ONE cave etiolee neaosmee Serteetee ue nae Meee CUCU LOSE 
III. Dorsal 7-11; origin nearly equidistant from end of snout 

and middle or posterior end of caudal fin; base shorter than 

distance from caudal. 


to 


. latipinna. 


A. Origin of anal below or a little behind that of dorsal ...... 5. sphenops. 

B. Origin of anal in advance of that of dorsal. 
Dorsal 8-10. Anal 8-10. Scales 29-33 . 6. elongata. 
Dorsal 7-9. Anal 9-11. poe: 27-380.. 7. oceidentalis. 
Dersal 8. Anal 9. Scales 2 8. spilwrus. 


1. MoLuLIEeNIsiA PETENENSIS. 


Giinth. Cat. Fish. vi. p. 348 (1866), and Trans. Zool. Soc. vi. 
1868, p. 485, pl. cart. figs. 1-3; Regan, Biol. Centr.-Amer., 
Pisces, p. 106 (1907). 


CYPRINODOND FISHES. LOLi 


2. Depth of body 23 to 3 in the length, length of head 4. 
Diameter of eye 4 in length of head, interorbital width 13. 28 
to 30 scales in a longitudinal series. Dorsal 15-16; base about 
as long as its distance from en of SOUS, longer than its distance 
from caudal fin; longest rays 3 or 2 2 length of head. Anal 9-10; 
origin below middle of dorsal. Pector al as long as head ; pelvics 
reaching origin of anal. Caudal rounded or subtruncate. Least 
depth of caudal peduncle nearly equal to length of head. A 
small spot on each scale, most conspicuous posteriorly; dorsal 
and caudal with series of small spots. 

3. Base of dorsal 14 to 13 its distance from end of snout and 
about twice its distance from caudal; longest rays longer than 
head. Anal below anterior part of dorsal; produced rays 2 length 
of head ; second pelvic ray about as long. Lower angle of caudal 
slightly produced. Spots on lower part of dorsal forming undu- 
lating stripes; a series of large spots, one on each interradial 
membrane, along middle of fin. Lower margin of caudal blackish. 

Lake Peten in Guatemala. 

Three females, 100 to 110 wm., and three males, 100 to 130 mm. 
in total length, types of the species. 


. MOLLreNniIsiA LATIPINNA. 


t Sueur, Journ. Acad. Philad. 11. 1821, p. 3, pl. ii.; Cuv. & 
Val. Hist. Nat. Poiss. Sanu, [os Bos jole WAT 3 Giimth. Cat. Fish. vi. 
p. 348 (1866) ; Garman, Mem. Mus. Comp. Zool. xix. 1895, p. 50, 
jolts 30g § dork, ay Everm. Bull. U.S, Nat. Mus. xIvii, 1896, p. 699. 

Pecilia multilineata Le Sueur, t. ¢. p. 4. 

Pacilia lineolata Girard, U.S. Mex. Bound. Sury., Fish. pac Or 
pl. xxxv. figs. 9-11 (1859). 

Lima pecilioides Girard, t.c. p. 70, pl. xxxviil. figs. 8-11 ; 
Proc. Acad. Philad. 1859, p. 115. 

Limia matamorensis Girard, Proc. Acad. Philad. 1859, p. 116. 

2. Depth of body 23 to 3 in the length, length of head 33 to 4. 
Diameter of eye 34 to 4 in length of head, interorbital width 14 
to 2. 26 to 28 scales in a longitudinal series. Dorsal 14-16; in 
adults base somewhat shorter than its distance - ‘om end of snout, 
longer than its distance from caudal; rays 3 length of head. 
Anal 9-10; origin below middle of dorsal. Pectoral a. little 
shorter than head; pelvics reaching vent or origin of anal. 
Caudal rounded. Least depth of caudal peduncle about ? length 
of head. Spots on scales forming longitudinal sieinen: dorsil 
and basal part of caudal with series of small spots. 

3. Dorsal base, in adult, about 14 its distance from end of 
snout or twice its distance from caudal; rays longer tonne head. 
Anal below anterior part of dorsal ; produced rays 7 length of 
head, a little longer than second pelvic ray ; amd rounded. 
Spots on lower part of dorsal forming longitudinal stripes ; a 
series of large spots, one on each interradial membrane, along 
middle of fin; caudal spotted superiorly; with blackish lower 
edge. 


1012 MR. C. TATE REGAN ON 


Southern United States. 
Five females, 40 to 68 mm., and two males, 65 and 78 mm., 
from Pensacola and New Orleans. 


3. MOLLIENISIA FORMOSA. 


Limia formosa Girard, Proc. Acad. Philad. 1859, p. 115. 

Mollienisia formosa, Giinth. Cat. Fish. vi. p. 349 (1866). 

2. Depth of body 23 to 3 in the length, length of head 31 to 4. 
Diameter of eye 3 to 32 1 in length of head, interorbital width 12 
to 2. 26 to 28 scales in a longitudinal series. Dorsal 11- 14; 
origin a little nearer to base of caudal than to end of snout ; base, 
in adult, nearly equal to its dhisusinse from caudal, but less than its 
distance from head; longest rays } to 2 2 length of head. Anal 
9-10; origin below fourth ray of dorsal. Pectoral a little shorter 
than head; pelvics reaching vent or or igin of anal. Caudal 
rounded. Heast depth of camel peduncle 2 to ? length of head. 
A spot on each scale ; dorsal usually spotted. 

3. Dorsal origin nearer to end of snout than to base of caudal ; 
base as long as its distance from eye; longest rays nearly ? 2 length 
of head. 

Atlantic slope of Mexico and Central America. 

Several small specimens (¢ and @ ) up to 45 mm. from Tampico 
(Jordan). Also three large females, 65 to 80 mm. in total length, 
from Colon (Rachow) ; hese are structur ally similar to the bohene: 
but have the coloration obscured by blackish mottling. 


4. MOLLIENISIA GRACILIS, nom. nov. 

Pecilia petenensis (non Moll tenisia petenensis Ginth.) Ginth. 
Cat. Fish. vi. p. 342, and Trans. Zool. Soc. vi. 1868 »P- 484, pl. Ixxxv. 
figs. 3, 4. 

2. Depth of body 4 to 43 in the length, length of head 42 
to 42. Diameter of eye 3} to 3? in length of head, inter orbital 
width 2. 28 to 30 scales ina loneieudinall series. Dorsal 10- Lil 
origin equidistant from preeorbital and base of caudal; anterior 
branched rays 2 length of head. Anal 8-10, acutely. pointed ; 
origin below fourth ray of dorsal. Pectoral as long as head. 
Least depth of caudal peduncle ? length of head. Olivaceous; 

scales dark-edged ; dorsal and cnndlel speckled. 

3. Depth 33 to 3% in length ; depth of caudal peduncle equal 
to length of head. Dorsal origin equidistant from end of snout 
and base of caudal; posterior rays longest, as long as or longer 
than head. 

Lake Peten in Guatemala. 

Two females, 120 and 150 mm. and three males, 95 to 120 mm. 
in total length, types of the species. 


5. MOLLrenisia SPHENOPS. (Text-fig. 173 F.) 

Pecilia sphenops Cuv. & Val. Hist. Nat. Poiss. xviii. p. 130, 
pl. 526 (1846); Regan, Biol. Centr.-Amer., Pisces, p. 102, pl. xiii. 
(1907) (with synonymy). 


CYPRINODONY FISHES. 1013 


Girardinus Suse sesoaNs Steind. Dealeselau, Akad. Wien, xli. 1880, 

p- 87, pl. vi. figs. 4, 5 
Pecilia sulvatoris Regan, Ann. Mag. Nat. Hist. (7) xix. 1907, 
p- 65; Biol. Centr.-Amer., Pisces, p. 104, pl. xiv. figs. 2, 3 (1907). 

Pecilia amates Miller, Bull. Amer. Mus. xxii. 1907, p. 108. 

? Platypecilus tropicus Meek, Publ. Columbian Mus., Zool. vii. 
1907, p. 146. 

Pecilia tenwis Meek, t. c. p. 147. 

Pecilia spilonota Regan, Ann. Mag. Nat. Hist. (8) 1. 1908, 
p. 460. 

2. Depth of body 24 to 4 in the length, length of head 33 
to 43. Diameter of eye 3 to 4 in length of head, interorbital 
width 14 to 2. 25 to 30 scales in a longitudinal series. Dorsal 
8-11; origin equidistant from eye or operculum and base of 
caudal, Anal 8-10; origin below or a little behind that of 
dorsal. Pectoral as long as or shorter than head. Least depth 
of caudal peduncle equal to or less than length of head. Dark 
greenish to pale olivaceous; edges of scales sometimes darker ; 
often a dark spot on each scale of side of body ; sometimes narrow 
vertical bars on sides; dorsal and caudal sometimes immaculate, 
often with series of small spots, or with a dark basal spot which 
may extend over most or all of the fin. 

3. Dorsal further forward and more elevated ; spots on sides 
rarely present, dark cross-bars often distinct. 

Coasts and rivers from Sinaloa to Panama and from Tamau- 
‘lipas to Colombia, Venezuela, and the Leeward Islands. 

Numerous examples to 120 mm. in total length, from all parts 
of the range of the species. 


6. MOLLIENISIA ELONGATA. 

Pecilia elongata Ginth. Cat. Fish. vi. p. 342, and Trans. Zool. 
Soe. vi. 1869, p. 484, pl. Ixxxv. fig. 2; Regan, Biol. Centr.-Amer., 
Pisces, p. 102 (1907). 

? Pecilia cuneata Garman, Mem. Mus. Comp. Zool. xix. 1895, 

-p. 62. 


Panama. 


7. MOLLIENISIA OCCIDENTALIS. 


Heterandria occidentalis Baird & Girard, Proc. Acad. Philad. 
1853, p. 390. 

Pecilia occidentalis Garman, Mem. Mus. Comp. Zool. xix. 1895, 
p. 71; Regan, Biol. Centr.-Amer., Pisces, p. 102 (1907). , 

Pacific Coast Rivers from Arizona to Jalisco. 


8. MoLLIENISIA SPILURUS. 


Pecilia spilurws Gunth. Cat. Fish. vi. p. 345 (1866); Regan, 
Biol. Centr.-Amer., Pisces, p. 101 (1907). — 


Mexico o1 ential America. 


1014 MR. C. TATE REGAN ON 


26. Limra Poey, 1855. 


Mem. Cuba, 1. p. 383. 

Acropecilia Hilgendorf, Sitzungsb. Ges. Nat. Freund. 1889, 
p. 52. 

This genus is well marked off from Pacilia by the structure of 
the intromittent organ (text-fig. 173 E). At some distance from 
its end the first produced ray becomes abr uptly slender ; the distal 
part is not serrated, and near its extremity it bears an antrorse 
spine; the second and third produced rays are nearly as in 
Pecilia. As in Pecilia, the intromittent organ is short, usually 
shorter than the head, and its extremity is protected anteriorly 
by a cutaneous hood; in fully adult males the second pelvic ray 
is more or less prolonged. 

Of eight species seven are from Cuba, Haiti, and Jamaica, and 
one from Venezuela. I have examined adult males in all but 
L. ornata. 


Synopsis of the Species. 


TI. Origin of dorsal fin about equidistant from middle of eye 
and base of caudal; origin of anal below fifth ray of 
OUOS SFT fies es eee omarpliatlar Ucn tn ugdan vad nan gameeosarobaccowues dl uouuuandae 
TI. Origin of dorsal nearer to te of Peandedl re to eye. 
A. Origin of anal below anterior part of dorsal. 


Dorsal 8-9; length of head 33 to 4 in length of fish ............... 2. dominicensis. 
Dorsal 10; length of head 32 to 4 in length of fish ............... 3. nigrofasciata. 
Dorsal 10-11; length of head 3 to 3} in length of fish ............ 4. arnoldi. 


B. Origin of anal below or very slightly in advance of that 
of dorsal. 
1. Length of head 2 length of fish (in adults of 50 mm.) ; 
a lateral series of dark bars in both sexes. 
Tuterorbital width less than } length of head 5. ornata. 


Interorbital width 5 length of head, OLSMLONe wees eeee .. 6. caudofasciata. 


2. Length of head less than 4 length of fish Gna a specimen 
of 43 mm.). ¢ with 3 or 4 blackish cross-bars, g 
with a blackish lateral stripe anteriorly............... 7. heterandria. 


C. Origin of anal always distinctly in Ra vaaee’ of that of 
dorsal ; head 4 length of fish (in adults of 50-60 mm.). 8. versicolor. 


1. LIMIA VITTATA. 


Pecilia vittata Guichenot in Ramon de la Sagra, Hist. Nat. 
Cuba, Poiss. p. 146, pl. v. fig. 1 (1850); Giinth. Cat. Fish. vi. 
p. 339 (1866); Garman, Mem. Mus. Comp. Zool. xix. 1895, 
De Diss 
Limia cubensis Poey, Mem. Cuba, 1. p. 388, pl. xxxi. figs. 12-13 
(1855). 

Limia vittata Poey, t. c. p. 389, pl. xxxi. figs. 14, 15. 

? Platypecilus perugie Evermann & Clark, Proc. U.S. Nat. 
Mus) xxx. 1906) ps 8515 fig: 

2. Depth of body 24 to 3 in the Jength, length of head 33 
to 48. Diameter of eye 3 to 34 in length of head, interorbitel 
maiden 14 to 2.. 26 to 28 scales in a longitudinal series. Dorsal 


CYPRINODONT FISHES, 1015 


9-11; origin nearly equidistant from middle of eye and base of 
caudal ; branched rays subequal, or the middle ones the longer, 
3 or 3 2 length of head. Anal 10; origin below fifth ray of dorsal. 
Pectoral = length of head or more ; pelvies reaching vent. Least 
depth of caudal peduncle * to + length of Theme) Olivaceous ; 
edges of scales darker ; often a dark lateral band or 1 to 3 series 
of dark spots along the series of scales on middle of side ; ; dorsal 
and caudal usually with small dark spots. 

36. Dorsal origin equidistant from snout and base of caudal ; 
posterior rays longest, longer than head in the adult. Intro- 
mittent organ a little shorter than head; second pelvic ray 
produced, inner rays ending ina knob. Least depth of caudal 
peduncle nearly equal to length of head. Usually dark cross-bars 
on body; spots on dorsal and caudal fins larger and blacker than 
in females. 

Cuba; ?San Domingo. 

Several specimens from Cuba, the males 45 to 70, the females 
40 to 100 mm., in total length. ; 


2. LIMIA DOMINICENSIS. 

Pecilia dominicensis Cuv. & Val. Hist. Nat. Poiss. xviii. p. 131, 
pl. 526. fig. 1 (1846). : 

Pecilia melanogaster Ginth. Cat. Fish. vi. p. 345 (1866). 

Menigpoeen 8 Ge ees Evermann & Clark, Proc. U.S. Nat. 


Depth of Body 2 Nes ie 3 a the length, length of head 32 to 4. 
Diameter of eye 3 to 33 in length of head, interorbital width 13. 
26 or 27 scales in a longitudinal series. Dorsal 8-93; origin 
equidistant from occiput or operculum and base of caudal, or tip 
of snout and end of caudal, first or second branched ray longest, 
3 length of head. Anal 10; origin below second or third ray of 
dorsal; anterior branched rays longest. Pectoral ? length of 
head; pelvics reaching vent. Least depth of caudal peduncle 
2 to #? length of head. Olivaceous; sometimes traces of dark 
bars on upper part of sides and of a dark spot at base of caudal ; 
a blackish spot on basal part of posterior 3 of dorsal fin; ripe 
females with abdomen golden in front, blackish behind, the two 
colours separated by a ventral line at insertion of pelvic fins. 

San Domingo; ? Jamaica. 

Seven specimens; two of the types of the species, ¢ and 2, 
38 and 42 mm. in total length, and five females of 50 to 60 mm., 
types of P. melanogaster. 


3. LIMIA NIGROFASCIATA, sp.n. (PI. CI. figs. 1, 2.) 


2. Depth of body 23 to 3 in the length, length of head 33 
to 4. Diameter of eye 34 to 33 in length of head, interorbital 
width 14. 27 or 28 scales in a longitudinal series, Dorsal 10 ; 
origin equidistant from cuidate of operculum and base of caudal ; 
fin rounded, the longest rays 4 length of head. Anal 9, obtuse; 


1016 MR. C. TATE REGAN ON 


origin below anterior + of dorsal. Pectoral 3 length of head ; 
pelvics not or barely reaching vent. Least depth of caudal 
peduncle = or ? length of head. Body with 7 to 9 blackish 
vertical bars ; a dark spot near base of posterior part of dorsal. 

3. Depth 27 to 24 in the length. Dorsal origin equidistant 
from eye or preoperculum and base of caudal; longest rays 
3 length of head, or in adult nearly as long as bead, 

Miragoane, Haiti. 

Two females, 43 mm. in total length, an immature male of 
41 mm., and an adult male of 48 mm., presented by Messrs. J. 
Paul Arnold and A, Rachow. 


4, LimraA ARNOLDI, sp.n, (PI. CI. fig. 5.) 


2. Depth of body 3 to 3% in the length, length of head 
3 to 33. Diameter of eye 3 to 34 in the length of head, inter- 
orbital width 2. 26 or 27 scales in a longitudinal series. Dorsal 
10-11; origin equidistant from postorbital part of head and base 
of caudal ; fin rounded, the longest rays 3 length of head. Anal 
9-10, rounded or obtusely pointed ; origin below anterior part of 
dorsal. Pectoral 7? length of head; pelvics extending to vent. 
Least depth of caudal peduncle 3 to # length of head. Body with 
8 to 12 narrow dark vertical bars; dorsal sometimes spotted, 
often with a small dark spot posteriorly near its base. 

3. Depth of body 3 in the length. Dorsal origin equidistant 
from eye and base of caudal; longest rays 2 length of head. 
Intromittent organ } length of fish (to base of caudal); second 
pelvic ray stout, a little produced beyond third and fourth, which 
ave truncated distally. Least depth of caudal peduncle ? length 
of head. No dark vertical bars, but several series of vertically 
expanded spots. 

Miragoane, Haiti. 

Ten females, 26 to 42 mm. in total length, and three males of 


28 to 38 mm., presented by Herr J. Paul Arnold. 


5. Limia orNatA, sp.n. (Pl. CI. fig. 7.) 


Depth of body 33 to 4 in the length, length of head 3 to 34. 
Diameter of eye 34 to 4 in length of head, interorbital width 
21 to 23. 28 scales in a longitudinal series. Dorsal 8-10; 
origin equidistant from some part of operculum and base of 
caudal; fin rounded, the longest rays 4 length of head. Anal 10; 
origin below that of dorsal. Pectoral 2 or 3? length of head ; 
pelvics reaching vent or origin of anal. Least depth of caudal 
peduncle 2 the length of head. Head, body, and usually vertical 
fins with blackish spots more or less developed ; body also with a 
series of dark bars or large vertically expanded spots. 

Haiti. 

Five females, 31 to 55 mm. in total length, presented by Herr 
J. Paul Arnold. 


CYPRINODONT FISHES. 1017 


6. Limia CAUDOFASCIATA, sp. n. (PI. CI. fig. 6.) 


@. Depth of body 3 to 34 in the length, length of head 
3 to 34. Diameter of eye 3} to 33 in length of head, inter- 
orbital width 14 to 2. 26 scales in a longitudinal series. Dorsal 
8-10; origin equidistant from some part of opsxomtou and base 
of caudal; anterior branched rays longest, 4 length of head. 
Anal 10; origin below or a little in advance of ‘Tak of dorsal. 
Pectoral ? length of head; pelvics reaching vent. Least depth 
of caudal peduncle 3 to # length of head. Olivaceous or 
brownish ; edges of scales darker: a series of dark vertical bars 
on posterior part of body ; a dark spot near base of posterior part 
of dorsal; abdomen yellowish, or when tumid golden anteriorly 
and blackish posteriorly, as in ZL. dominicensis. 

3. Dorsal origin about equidistant from eye and base of 
caudal; longest rays 3 to 2 (adult) length of head. Intromittent 
organ less than } length of fish; second pelvic ray produced in 
adult, nearly as long as prolonged anal rays. Least depth of 
caudal peduncle 2 length of head. 

Jamaica. 

Several females, 28 to 42 mm. in total length, and six males of 
25 to 48 mm., collected by C. A. Wray. In males of 25 to 
30 mm., the anal fin has not attained the adult structure and the 
second pelvic ray 1s scarcely prolonged; males of 35 to 48 mm. 
have the anal fully formed, but only the largest has the pelvies 
strongly produced. 


7. LIMIA HETERANDRIA, sp.n. (PI. CI. figs. 3, 4.) 


2. Depth of body 33 in the length, length of head 32 to 44. 
Diameter of eye 3 to 31 in length ‘of head, interorbital width 2. 
26 scales in a longitudinal series. Dorsal 8; origin equidistant 
from some part of operculum and base of candle fin rounded, 
longest rays a little more than 4 length of head. Anal 9, 
pointed; origin below that of dorsal. Pectoral # length of head ; 


a 


pelvics reaching vent. Least depth of caudal peduncle 4 or # 
length of head. Olivaceous; a dark longitudinal stripe from 
above pectoral to level of origin of anal; a dark spot above vent ; 
a black spot near base of posterior part of dorsal. 

3. Dorsal origin egemchisceuat from middle of eye and base of 
caudal ; longest rays 2 length of head. Intromittent organ about 

2 length of fish (to ‘base “oi caudal) ; second ray produced, 3 3 as 
inne as prolonged anal rays. Stripe on side of body and spot 
above vent absent : body with 3 or 4 narrow dark vertical bars. 

La Guayra, Venezuela. 

Two females, 43 and 27 mm.,and a male of 25 mm. (adult), 
presented by Herr J. Paul Arnoid. 


8. Limia verstcotor. (Text-fig. 173 E.) 
Pecilia donunicensis (non Cuv. & Val.) Giinth. Cat. Fish. vi, 


1018 ON CYPRINODONT FISHES. 


p- 346 (1866); Garman, Mem. Mus. Comp. Zool. xix. 1895, 
oe Dee 

Girardinus versicolor Giinth. t. ¢. p. 352. 

Acropacilia tr idens Hilgendorf, Sitzungsb. Naturf. -Freunde, 
1889, p. 52. 

2. Depth of body 3 to 4 in the length, length of head 
34 to 4. Diameter of eye 3 to 34 in leneth of head ; inter- 
orbital width 12 to 2. 26 to 28 scales in a longitudinal series. 
Dorsal 7-9 ; origin equidistant from some part of. operculum and 
base of eam ‘middle rays longest, about 2 the length of head. 
Anal 8-10; origin a little in advance of that of dor sel Pectoral 
a little shorter ‘than head ; pelvics about reaching vent. Least 
depth of caudal peduncle 2 402 = the length of Inn Olivaceous, 
edges of scales darker; an indistinet Jateral band or a series of 
short vertical bars ; sometimes irregular dark spots on the back ; 
a blackish spot on base of posterior part of dorsal; caudal 
sometimes with vertical bands. 

3. In specimens of 30 mm. dorsal and pelvic fins as in 9, but 
in those of 40 to 45 mm. dorsal more elevated, the longest rays 
? length of head, second pelvic ray somewhat produced send i inner 
alate rays expanded and truncated at ends. 

San Domingo. 

Ten females, 35 to 60 mm., including the types of the species, 
and four “anoles, 30-45 mm. 


EXPLANATION OF THE PLATES. 


PratE XCIX. 


2. Gambusia oligosticta, 9 & g. 
4, Be wrayi,2? & SB. 

, 6. a gracilior, 2 & g. 
a es dominicensis, 9. 


Pram C: 


2. Pecilia picta, 2 & S. 

, 4. Paciliopsis isthmensis, 2 & 3. 
6. Phalloceros caudomaculatus, 2 & 3. 
8. Phalloptychus januarius, 9 & 3. 


PratTE CI. 


. Limia nigrofasciata, 2 & g. 
»  heterandria, 2 & g. 
» arnoldi, 2. 

» caudofasciata, 6. 

»  ornata, 


STH OVE bo 


P. ZS) (er ie Gil 


d.Green del. et lith. 
1LBUFO HYPOMELAS. 2. AGALYCHNIS CALCARIFER. 


tee arts) . USNS) IE, CHIL. 


d.Green del.et. Chromo. 


AGALYCHNIS SPURRELLEI. 


IAS HOS) ell) (CIN. 


J.Green del.et Chromo. 


1. DENDROBATES AUROTZANIA 
2-4.D.TINCTORIUS,var.COCTAI, 5-6.D.TINCTORIUS,vs.CHOCOENSIS. 


P45: iss Pisey. 


J.Green del.et lith. 


CINOSTERNUM SPURRELLI. 


. 


sre 


learn oe ieols) JP i. (Wl 


J.Green del.et ith 


CINOSTERNUM SPURRELLI. 


PP. Zeon LSP ies CMe 


J.Green del.et lith. 


LANOLIS BREVICEPS. 2. ANADIA VITTATA. 


Ms Sa LORS. Pil. CVA. 


————— 


d.Green del.et lith. 


1. HOMALOCRANIUM CORALLIVENTRE. 2.ELAPS MICROPS 
&. LEPTOGNATHUS SPURREEEI 


1913- Proc.Zool.Soc.London 1913:1019-38 ,figs.174-178 
b1s.102-108 


ON COLOMBIAN BATRACHIANS AND REPTILES. 1019 


59. On a Collection of Batrachians and Reptiles made by 
Dr. H. G. F. Spurrell, F.Z.S8., in the Choco, Colombia. 
By G. A. BoutencEr, F.R.S., F.Z.S.* 


{Received October, 10th, 1913: Read November 11th, 1913.] 


(Plates CIT.—CVIII.7 and Text-figures 174-178.) 


INDEX. 

5 Page 

Geographical Zoology: Choco, Colombia, Batrachians and 
Reptiles... sates er Autenaetncereeo eres LOLS. 
Development : bere of late omnes. Sicoatene Leeseranen LO2ZS 
Ethology: Nesting-habits of Agalychnis spurr aIB.. LOA: 
Variation: Dendrobates CPaerAl ie aen RL RUSS 1026 
New species: Cecilia wmtermedta@ 2.2 0..c.028 senses LOZO 
BF) I CUMTOGE Osecntitactiina bauer steed baaeeecesoos deo I MUr-Ut 
Bufo hypomelas .. Jehiccsis watan ene neon steer O22 
Agalychnis poor cE see Saanpnduce ORE! 
Dendrobates paraensis (Cam Brazil) Jaisueaiee 1028 
op GUROUGHOED con cornoecussenconcocencccs IKOFE) 
Cinosternum spurrellt ...........1............-...-- 1030 
AYRES OPRESUCGIB.:0 snocca noo cdaconcanocecconieccseoeces OBI 
Anadia wittata ........... Becmecnnuoebaon cucie 40ssxs, 
Homalocranium cor aUnenivel Weeeennuntn: LOSS: 
JONCGIS MACPODE bacco socdooqovocass0 cen vpaeae svonenove NOS 
Leptognathus spucamnclll.. ese LOSS 


Dr. Spurrell, to whom the Zoological Society is indebted for so 
many interesting additions to its collection of Reptiles, has now 
transferred his activity as a collector from West Africa to South 
America. The series of beautifully preserved Batrachians and 
Reptiles brought together by him during the first few months 
of his stay in the Choco, Colombia, and presented by him to the 
British Museum, is one of great interest and shows how much 
remains to be done in the exploration of that part of South 
America. Unless otherwise stated, the specimens are from 
Pena Lisa, Condoto, altitude 300 feet, Within the last few 
years, series of Batrachians and Reptiles brought together by 
Mr. M.G. Palmer in the same district had been acquired by the 
Trustees of the British Museum and furnished the types of 
several new species described by met. They showed a close 

agreement with the fauna of N.W. Keuador, for a knowledge of 
which we are indebted chiefly to Mx. Rosenberg §—an agreement 


8 
which is further exemplified by Dr. Spurrell’s collection. 


Published by permission of the Trustees of the British Museum. 

For explanation of the Plates see p. 1038. 

Ann. & Mag. N. H. (8) ii. 1908, p. 515, and vii. 1911, p. 19. 

Boulenger, P. Z.S. 1898, p. 107, and Ann. & Mag. N. H. (7) ix. 1902, p, 51. 


wntt+ % 


1020 MR. G. A. BOULENGER ON 


BATRACHIA., 
APODA. 


1. CxcILIA ISTHMICA Cope. 
Known from the Isthmus of Darien and Western Heuador. 


2. CMCILIA INTERMEDIA, sp.n. (Text-fig. 174.) 

Intermediate between C. tentaculata L. and C. pachynema Gthr. 
Snout as in the former, more strongly projecting than in the 
latter. Outer mandibular teeth larger than in the former and 
smaller than in the latter. Snout rounded, very prominent, as 
long as the distance between the eyes; tentacle on the lower 
surface of the snout, nearer the edge of the mouth than the 


Text-fig. 174. 


B 


Cecilia intermedia. 
A. Side view of head. B. Lower view of head and anterior extremity of body. 
C. Lower view of posterior extremity of body. 

nostril. Body cylindrical, its diameter 43 to 56 times in the 
total length. 200 to 250 circular folds, mostly widely interrupted 
on the back and on the belly ; from the posterior fourth of the 
body short intermediate folds appear and gradually extend across 
the back; the last. 60 to 80 folds close together and complete on 
the back, the 15 to 30 hindermost completely encircling the body. 
Uniform. blackish ; head of young yellowish or pale brown. 

I have examined six specimens of this species, which was 


COLOMBIAN BATRACHIANS AND REPTILES. 1021 


first discovered in N.W. Ecuador by one of Mr. Rosenberg’s 

collectors :-— 
1. 8. Javier, N.W. Ecuador.—Total length 950 mm., dia- 
meter of body 17. 

2. Paramba, N.W. Ecuador, 3500 ft.—Total length 820 mm., 
diameter of body 19. 

3. Pambelar, N.W. Ecuador.—Total length 690 mm., dia- 
meter of body 12. 

4, Pefia Lisa, Condoto, Choco (had been swallowed by an 
Elaps rosenbergti 940mm. long).—Total length 690mm. ; 
diameter of body 13 *. 

Same locality (taken from the stomach of a Streptophorus 
atratus swallowed by an Elaps corallinus).—VTotal length 
280 mm.; diameter of body 6. 

6. Same locality.—Total length 270 mm.; diameter of 

body 5. 
I avail myself of this opportunity to define a further species, 
from a collection made‘in the Choco by Mr. M. G. Palmer, recently 
acquired by the British Museum :— 


ot 


Cecilia palmeri, sp. n. (Text-fig. 175.)—Head and dentition as 
Text-fig. 175. 


Cecilia palmeri. 
Lettering as in text-fig. 174. 


* The copulatory organ of this specimen is extruded; it measures 10 mm. 
in length and terminates in a four-lobed “ glans.” 


Proc. Zoot, Soc.—1913, No. LX VIII. 68 


1022 MR. G; A. BOULENGER ON 


in C. pachynema. Diameter of body 58 times in total length? 
200 circular folds, mostly interrupted on the back and belly ; 
from the posterior fourth of the body short interrupted folds 
appear and gradually extend across the back; the last 95 folds 
close together and complete on the back, the 16 hindermost com- 
pletely encircling the body. Uniform blackish. Total length 
700 mm. ; diameter of body 2s 

A single specimen from Novita, Rio San Juan, 200 ft. 


3. CHCILIA NIGRICANS Bler. 

Described in 1902 from a single specimen from the Rio Lita, 
N.W. Ecuador. 

The specimen in Dr. Spurrell’s collection measures 395 mm., 
diameter of body 7. 


ECAUDATA. 
4, Buro HYPOMELAS, sp.n. (PI. CII. fig. 1.) 


Crown without bony ridges; snout truncate, moderately pro- 
minent, with angular canthus and vertical loreal region ; inter- 
orbital space as br oad as the upper eyelid ; tympanum moderately 
distinct, half diameter of eye. Fingers moderate, first not 

extending as far .as second ; toes short, barely ihe webbed ; 

subar Hoole tubercles feebly pr ominent, single ; ; two small meee 
tarsal tubercles; no tarsal fold. Tlarso-metatarsal articulation 
reaching the tip of the snout. Skin smooth ; parotoid gland 
narrow, shorter than the head. Black, elegantly marked with 
grey lines above, vermicular or ring-shaped ; a whitish streak on 
each side in the lumbar region, and another on the arm ; lower 
parts with small greyish-white spots. 

From snout to vent 20 mm. ‘ 

This small toad, represented by a single specimen from the 
upper waters of the Condo, altitude 1200 feet, differs from all its 
American congeners in the shortness of the inner finger. 


Buro Hamatiticus Cope. 
Buro mARiNuS L. 


Buro rtypHontus L. 


Dr. Spurrell, in his notes, remarks: ‘Said to be the toad from 
which Indians extract arrow-poison.” See further on, p. 1028. 


8. Hyna MAxima Laur. 


‘“¢ Apparently strictly nocturnal. Pupil of eye at night large 
and round, in daylight diamond-shaped. One caught at night on 
top of partition between two rooms in bungrlow was almost 
uniform chocolate-brown with a dark median streak from snout 
to middle of back. By daylight it turned to a hght yellowish 
brown, with very pronounced markings of warm darker brown on 


COLOMBIAN BATRACHIANS AND REPTILES. 1023 


the body and limbs, greyer to sepia-brown on the head; a patch 
of lighter yellow across loins. Iris golden brown; lower eyelid 
veiled with gold.” 

It is characteristic of this species to possess a rudiment of 
pollex appearing as a small tubercle in females and a larger 
tubercle in males. In the male sent by Dr. Spurrell, this pollex 
projects externally as a long sharp bony spine at right angles to 
the metacarpal of the inner finger and piercing the skin ; whether 
this is due to the desiccation of the specimen, or occasionally 
happens during life as in the case of the pollex of Rana holst or 
of the ends of the ribs in the Pleurodele Newt, I cannot say. 


Text-fig. 176. 


Hyla maxima. Inner fingers of male. 


9. Hyna BpAaupint Daud. 


When taken out of a dark hole these frogs. are brown above, 
usually with a metallic-green streak along the upper lip. After 
exposure to light the colour changes to light yellowish grey, with 
small green spots on the back and bars on the hind limbs. One 
of the specimens, a female, is remarkable for the longer hind 
limbs, the tibio-tarsal articulation reaching beyond the tip of 
the snout. 

These frogs were pairing on the 3lst May. 

Larve, ab different stages of development, agree in essential 
characters with those of H. arborea : position of eyes, spiraculum 
and vent, shape of tail, structure of the mouth and lips. Three 
long series of teeth in each lip, the third of the upper lip inter- 
rupted, the others continuous; beak broadly edged with black. 
Total length 60 mm.; body 21; width of body 13; tail 39; 
depth of tail 16. 


10. AGALYCHNIS CALCARIFER Blgr. (Pi. CII. fig. 2.) 


Described in 1902 from a single specimen from the Rio 
Durango, N.W. Ecuador, 350 feet. 

. The ‘only example obtained by Dr. Spurrell is described in his 
noes as brilliant sage-green above, flanks and upper surface of 
thighs (with the exception of a narrow green streak) rich orange- 
yellow with black bars; hands and feet and lower parts orange- 

68* 


1024 MR. G. A. BOULENGER ON 


yellow; iris grey, bordered with orange-yellow ; lower eyelid 
transparent, edged with turquoise-blue. 


11, AGALYCHNIS SPURRELLI, sp.n. (PI. CIII.) 


Tongue oval, feebly emarginate behind. Vomerine teeth in 
two strong, transverse or slightly oblique series on a level with 
the front edge of the rather large choane. Head moderately 
depressed, as long as broad or slightly broader than long; snout 
rounded, not projecting beyond the lower jaw; canthus rostralis 
feeble ; loreal region concave and very oblique; interorbital space 
broader than the upper eyelid ; tympanum close to the eye and 
two-thirds to three-fourths its diameter. Fingers much flattened 
and nearly entirely webbed, the disks nearly as large as the eye 
or about two-thirds its size; toes rather short, much flattened, 
entirely webbed (three-fourths webbed in the young), the disks a 
little smaller than those of the fingers; subarticular tubercles 
very prominent. The tibio-tarsal articulation reaches between 
the eye and the tip of the snout. Skin smooth, granular on the 
belly and on the basal half of the lower surface of the thigh; a 
regular series of granules extends along the whole length of the 
lower surface of the thigh ; a narrow dermal fold along the outer 
edge of the forearm, the inner edge of the tarsus, and across the 
heel ; a few scattered (white) flat warts on the back, Green above, 
the dorsal warts, two to nine in number and irregularly disposed, 
white, edged with violet-black ; belly yellowish white; iris ruby- 
red ; lower eyelid with an open meshwork of golden lines. Male 
without vocal sac, with a patch of black nuptial asperities on the 
upper surface of the inner finger. 

From snout to vent 95 mm. 

Very closely allied to the Central American A. moreleti 
A. Dum., this handsome frog differs in the more extensively 
webbed digits, the presence of white warts on the back, and the 
absence of vocal sac in the male. 

Four specimens are in the collection. The first was reported 
to Dr. Spurrell to have been found at the top of a high tree that 
had been felled, on March 30th. On April 2nd Dr. Spurrell took 
a pair in embrace on a leaf overhanging a pool of water two feet 
below ; from the sketch accompanying his notes, the amplexus is 
similar to that of Hylu arborea. The pair were in the act of 
breeding, and the eggs, as they were extruded, were being fixed 
to the upper surface of the terminal third of the leaf, in double 
rows following more or less regularly the venation, the very 
prominent ribs affording a support for their attachment. On the 
leaf sent with the specimens, the breeding operations of which 
were suddenly interrupted, 59 eggs are attached, and 7 more 
adhere to the right foot of the female, showing that she uses her 
feet for the purpose of fixing the eggs. 

According to a sketch by Dr. Spurrell, which is here re- 
produced (text-fig. 177), another leaf on the same stalk was 
already entirely beset with eggs (laid by the same female 2) 


COLOMBIAN BATRACHIANS AND REPTILES. 1025 


and folded over to form a nest as in Phyllomedusa hypo- 
chondrialis*. The eggs resemble those of Alytes obstetricans 
in size (diameter 3 to 4 mm.) and in the tough gelatinous 
capsule, but differ in the upper pole being brown and in not 
being strung together. 


Text-fig. 177. 


Agalychnis spurrelli spawning. 


The leaf, which I have not been able to get identified, probably 
belongs to the family Anonacez. 


12. LEPTODACTYLUS PENTADACTYLUS Laur. 
13. Hy opEs pALMATUs Bler. 
14, HyLopks consPprciLuAtus Gthr. 


15. GLOSSOSTOMA ATERRIMUM Gthr. 


Originally described from Costa Rica, this Engystomatid has 
since been found on the Rio Durango, N.W. Ecuador, 


* Budgett, Q. J. Micr. Sci. xlii. (1899) p. 318, pl. xxviii. 


1026 MR. G. A. BOULENGER ON 


The specimen in Dr. Spurrell’s collection measures 55 nim, from 
snout to vent. It was shining jet-black in life, with the belly 
grey. The thick skin of the ventral region forms a strong fold 
on each side, which extends across the pubic region. 


16. Rana PALMIPES Spix. 


A single female specimen, measuring 95 mm, from snout 
to vent. 

Tympanum three-fourths the diameter of the eye. First 
finger extending well beyond second. Skin of back finely 
granulate; upper surface of tibia with longitudinal ridges 
formed by confluent granules. Tibio-tarsal articulation reaching 
half-way between the eye and the tip of the snout. Back pale 
olive-grey (appears to have been green in life), without spots ; 
belly uniform white. 

IT have examined smaller specimens (40 to 70 mm.) from the 
same locality (Condoto) from Mr. M. G. Palmer’s collection. In 
the smallest the tympanum measures barely two-thirds the 
diameter of the eye, the upper parts are blackish brown and 
the lower parts profusely spotted with black. 


17. PHYLLOBATES PRATTI Blegr. 


A single specimen, 15 mm. long from snout to vent. 

This species was described in 1899 (Ann. & Mag. N. H. [7] in. 
p. 274, pl. x1. fig. 3) from two specimens obtained by Mr. A. E. 
Pratt at Santa Ines, north of Medellin, at an altitude of 3800 feet. 
Other specimens have since been found in the Choco, at Novita, 
altitude 150 feet, by Mr.M.G. Palmer. In Dr. Spurrell’s specimen, 
the white line in front of the thigh is produced forward to the 
upper eyelid, dividing obliquely the black lateral band. 


18. DENDROBATES TINCTORIUS Schneid. (PI. CIV. figs. 2-6.) 

The extraordinary colour variations to which this species is 
subjected are only paralleled by those of the African frogs of the 
genus Rappia, and afford the same ditticulties to the systematist. 
Some of these variations are local, others are merely individual ; 
great is the perplexity in having to decide which deserve to be 
recognised by a special varietal name. In the collection made by 
Dr. Spurrell the specimens fall under two principal categories which 
1 will designate as vars. coctei and chocoensis respectively. It is 
not an easy matter to fix the typical form of D. tinctorius; the 
following remarks will not be superfluous, té justify the course 
followed by me. 

Schneider (Hist. Amph.1i. p. 175, 1799) is the author of the 
name tinctorius, but the frog itself was unknown to him. He 
refers us to Lacepéde (Quady, ONnalos joy DO, jal xxoabe,. 17s), 
who regards the “‘ Raine a tapirer” as a variety of Hyla rubra 
Laurenti, and describes it as reddish with two irr egular yellowish 
white longitudinal bands; the figure is hopeless for the purpose 


COLOMBIAN BATRACHIANS AND REPTILES. 1027 


‘of identification. But as the name ‘“ Raine a tapirer”’ is based 
on an observation recorded by Buffon (Hist. Nat. Oiseaux, vi. 
p. 235, 1779) to the effect that the frog is used by the savages of 
Guiana and the Amazon to dye parrots, and is small, azure blue 
with longitudinal golden yellow bands, and is to be found pre- 
served in the ‘‘Cabinet du Roi,” the best thing to do is to go 
straight to the description of the first author after Lacepede who 
had access to the specimens alluded to by Buffon, and this author 
is Daudin (Hist. Rain. etc., 1802). The larger of the three speci- 
mens from Guianain the Paris Museum, presumably those alluded 
to by Buffon and Lacepéde, is figured in a perfectly recognisable 
manner on Daudin’s pl. vii. fig. 1, and corresponds very nearly 
with one from Cayenne in the British Museum, which I therefore 
regard as the typical form of Dendrobates tinctorius. Upper parts 
blackish brown with symmetrical white (yellow) markings forming 
two bands on the back, meeting on the coccygeal region and ex- 
tending, above the canthus rostralis, to the end of the snout after 
expanding on the upper eyelid; a transverse bar connects these 
bands across the middle of the back, and another branch extends 
to the shoulder; white markings on the flanks.; belly paler brown, 
with black spots. This is the var. daudini Steindachner, Verh. 
zool.-bot. Ges. Wien, xiv. 1864, p. 262. 


Var. coctei.__Hylaplésie de Cocteau, Dum. & Bibr. Erp. Gén. 
pl. xe. fig. 1; Dendrobates tinctorius, var. B, Dum. & Bibr. op. cit. 
vill. p. 654 (1841); D. tinctorius, var. cocteaui Steind. Verh. 
z001.-bot. Ges. Wien, XIV. 1864, p. 260. 


“ D’un brun marron ou dune teinte lie de vin, .... une tache 
dun blane jaunatre sur Je museau, une autre beaucoup plus grande 
et de forme ovalaire sur chaque flane, .... un large bracelet de 
la méme couleur autour de chaque bras et, de chaque jambe. 
Assez souvent les taches des flanes se confondent sur la région 
abdominale.” Dum. & Bibr. Locality not stated. 

Six specimens, all from the Choco, in the British Museum, are 
referable to this variety :— 

Tado, Rio San Juan, 230 ft. (M. G. Palmer). Dark reddish 
brown above, with black spots, black beneath. An oval yellow 
spot on the top of the head, another occupying the whole flank, 
widely separated from its fellow on the other side; other large 
spots are present, one on the forearm, one on the thigh, one on 
the tibia, and one on the throat. 

b. Same locality, same collector. Similar to the preceding, but 
the spots on the head, throat, and femur absent; the large spot 
on the flank extends to. the shoulder, and those on the forearm 
and tibia completely encircle the limbs. 

c. Upper waters of Condoto River, 1200 ft. (Spurrell). Dark 
brown above, with bright vermilion- red, sharply defined spots, 
black beneath. The br ight spots are as in a, except that the one 
on the top of the head is situated further back; those on the 
throat and forearm. are absent, but others are present on the left 


1028 MR. G. A. BOULENGER ON 


side of the snout and on the left foot ; the forearm is completely 
encircled by the yellow spot. 

d. Same locality asc. Differs from ec in the lateral spot being 
produced as far as the eye on the left side, where it is confluent 
with the oval spot situated on the upper surface of the head as 
in @; the spot on the tibia surrounds the limb; a yellow spot on 
the right foot instead of the left. 

e, f. Same locality as cand d, and very similar in the markings, 
which, however, were noted by Dr. Spurrell as of a vivid 
yellow instead of red. e has the lateral spots very large and 
narrowly separated from each other on the belly, that on the 
left side extending to the temple; a spot on the right foot; fore- 
arm encircled by the red spot. jf has the spot on the right foot 
instead of the left. 

A further Colombian specimen, from Noanama, 8. Juan 
(M. G. Palmer), is not unlike the preceding, but the spots are 
smaller and less regular and of a vivid magenta-red. A figure is 
given to show the disposition of the markings on the black body. 


Var. chocoensis. — Phyllobates chocoensis Posada Arango, 
Memoire sur le poison de Rainette des sauvages du Choco (Paris, 
1869); Deyrolle, Rev. et Mag. de Zool. 1872, p. 465, pl. xxvii. 

The frog noticed by Posada Arango as furnishing the Choco 
Indians with a deadly poison for their arrows is described as 
bright yellow on the head and back and bluish black on the limbs 
and lower parts. A specimen answering to this description is 
preserved in the British Museum, from Juntas, Rio San Juan, 
Choco, 320 ft. (M. G. Palmer). 

Nine specimens, obtained by Dr. Spurrell at Petia Lisa, 
Condoto, 300 feet, may be referred to this form, and show 
considerable variation in coloration. A snake which started 
eating one of these frogs was observed to drop it hurriedly. 
‘Ground colour of upper parts a rich orange-yellow, belly dark 
brown ; markings on back and limbs varying from dark reddish 
orange to chocolate-brown, not sharply defined nor regularly 
symmetrical ; much variation in colour and arrangement of 
markings; ground colour of upper parts sometimes lemon-yellow 
or brownish yellow.” ‘The specimens which most nearly approach 
Posada Arango’s description have the bead and back uniform 
yellow or with only one dark spot, on the middle of the back, 
just behind the head; the sides are blackish brown, and the upper 
surface of the limbs yellow. 


The numerous varieties of Dendrobates tinctorius are much in 
want of revision. Among those already described there is one 
which is unquestionably entitled to specific rank and for which I 
wish to propose the name Dendrobates paraensis. Disks of fingers 
and toes much larger than in D. tinctorius; a small but very 
distinct tubercle on the inner side of the tarsus, nearer the meta- 
tarsal tubercles than the tibio-tarsal articulation. In some 


COLOMBIAN BATRACHIANS AND REPTILES, 1029 


specimens the upper parts, with the exception of fore limbs and 
feet, are uniform greyish or yellowish white (yellow in life ?), the 
rest of the animal deep black; in others the black predominates, 
the white being restricted to the upper surface of the head and to 


Text-fig. 178. 


n B 
Dendrobates tinctorius (A) and D. paraensis (B). 


Upper view of hand. 


more or less confluent blotches on the back ; further specimens 
are intermediate between the two extremes. From snout to vent 
38 mm. Hight specimens, from Para, are preserved in the 
British Museum, three of which are mentioned in the Catalogue 
of Batrachians, p. 143 (var. B, h, 7, £). 


19, DENDROBATES AUROTANIA, sp. n. (PI. CIV. fig..1.) 


Snout truncate, barely as long as the eye; loreal region 
vertical; interorbital space as broad as the upper eyelid; 
tympanum not very distinct, three-fifths the diameter of the eye. 
Fingers slender, first extending beyond second, with rather small 
disks; toes slender, disks as large as those of the fingers; sub- 
articular tubercles very feeble; two scarcely prominent meta- 
tarsal tubercles and a very feeble fold on the inner side of the 
tarsus terminating club-shaped half-way between the metatarsal 
tubercles and the tibio-tarsal articulation. The tarso-metatarsal 
articulation reaches far beyond the tip of the snout, the tibio- 
tarsal falling between the eye and the end of the snout; tibia 
half as long as head and body. Skin smooth, of dorsal region 
very porous. ‘“ Jet-black, with a metallic orange-yellow streak ” 
from the end of the snout along the canthus rostralis and the 
upper eyelid to the loin; ‘‘a few fine gold freckles on thighs and 
arms.” 

From snout to vent 30 mm. 

A single specimen from Pefia Lisa, Condoto, 500 feet. 

This frog approaches very closely D. trivittatus Spix, differing 
only in the shorter snout and the absence of tubercles on the 
back. 


1030 MR. G. A. BOULENGER ON 


REPTILIA,. 
CHELONIA. 
1. CINOSTERNUM SPURRELLI, sp.n. (Pls. CV. & CVI.) 


Jaws strong; beak strongly hooked. Carapace smooth and 
without keels, flattened on the vertebral region, profile descending 
very abruptly behind; deep grooves between the shields; 
posterior border serrated. Plastron smaller than the opening of 
the shell, not emarginate posteriorly; lobes well movable, front 
one longer than the fixed portion and slightly shorter than the 
hind one; gular shield not half the length of the front lobe; 
suture between the pectoral shields much shorter than that be- 
tween the humerals; axillary and inguinal shields elongate and in 
contact with each other. A patch of small horny keeled tubercles 
on the back of the leg, opposed to one on the thigh. Tail ending 
in a claw-like scute. Carapace blackish brown, plastron yellow 
with blackish blotches; head, neck, and limbs blackish above, 
whitish below; a broad oblique yellowish band on each side of 
the head, from behind the eye to the neck, passing above the 

. tympanum; jaws yellowish horn-colour. 

This very distinct species is represented by a single male 
specimen, the shell of which measures 115 mm. The curious 
shape of the first vertebral shield, depicted on Pl. CV., is probably 
an individual peculiarity and has therefore not been alluded to in 
the description. 

The position of C. spwrrelli in the system is in group II. A of 
my Synopsis (Cat. Chelon. p. 38), which group, up to the present, 
was unrepresented south of Mexico. 


2. Nicorza wAsura Bler. 


Described in 1902 from specimens obtained at Bulun and on 
the Rio Durango, N.W. Keuador. 

The specimen in the present collection is quite young. Shell- 
length 70 mm. ‘The digits are fully webbed. 


EMYDOSAURLIA. 


3. CAIMAN SCLEROPS Schn. 


LACERTILIA. 
4. ANOLIS FASCIATUS Bley. 


Only known from Guayaquil and. N.W. Eeuador (A. elegans 
Blgr.) ‘*Green above, with darker green markings, yellow on 
belly; crown of head and neck with faint orange markings. 
Gular appendage white with six broad orange bands.” 


5. ANOLIS MACULIVENTRIS Bler. 


First described from N.W. Ecuador. 
“ Gular appendage crimson.” 


COLOMBIAN BATRACHIANS AND REPTILES. 1031 


6, ANOLIS PRINCEPS Bler. 


This handsome lizard, one of the largest of the genus, was 
originally described from specimens from N.W. Heuador. It has 
since been found in Choco, at Condoto and T'adé, by Mr. M..G. 
Palmer. 

“Yellowish grass-green, brighter about head and neck, with 
yellow blotches round eyes; diagonal markings brownish. black ; 
gular appendage white.” 


7. ANOLIS BREVICEPS, sp. n. (Pl. CVII. fig. 1.) 


Head once and two-thirds as long as broad, slightly shorter 
than the tibia; snout obtuse; forehead and interorbital region 
concave; frontal ridges short and divergent ; upper head-scales 
very small, keeled; scales of the supraorbital semicircles enlarged, 
separated by four or five series of scales; enlarged, strongly 
keeled supraocular scales ; occipital as large as or larger than the 
ear-opening, separated from the supraorbital by three or four 
series of scales; canthus rostralis sharp, canthal scales three or 
four ; loreal rows seven or eight; seven upper labials to below 
the centre of the eye; ear-opening moderate, vertically oval. 
Gular appendage very small in males, barely indicated in females ; 
gular scales keeled. Body not compressed ; no dorso-nuchal fold. 
Dorsal seales very small, keeled, gradually merging into the 
minute granules of the sides; ventrals larger, roundish-hexagonal, 
keeled. Limbs slender; the adpressed hind limb reaches the 
tip of the snout, or between the eye and the tip of the snout ; 
digits feebly dilated; 13 to 15 lamelle under phalanges IT and 
IIT of the fourth toe. Tail cylindrical, once and a half to once 
and two-thirds as long as head and body, without vertebral series 
of enlarged scales. Male with enlarged postanal scales. Grey- 
brown to dark brown above, with a ventral series of blackish 
dots or small V-shaped markings; a dark brown, black-edged 
cross-band between the eyes and another band from the end of 
the snout, through the eye, to the side of the neck, very broad 
and sharply defined on the temple; a more or less distinct 
whitish streak may be present below this band. 


Sh anes 2 

Hbovellilemethiee anne nece: 108 124 mm. 
ASS clapwetn tars lte ons lee datasets Ne 13 
Wardithwot headless e Sein ) 

'EBOdy) iia. sete eeieiae seeps BySO 30 
Thome lin) soa.5ccedosbeccees 21 21 
dla Glielimmllo eye eee. ses 40, 40 
TPilotans abo. setae: assis ede 18} 133 
Sans REAR MNEs AIR 66 76 


Three specimens. 

Near the Central American A. godmant Blgr. Distinguished 
by the shorter head, the smaller gular appendage, and the shorter 
tail. = 


1032 MR. G. A. BOULENGER ON 


8. ANOLIS NOTOPHOLIS Blgr. 
The gular appendage is red in both sexes. 


9. ANOLIS MACROLEPIS Blgr. 


10. Potycurus currurosus Berthold. 


Two specimens, male and female. 

Male green above and on the belly, cream-colour on the sides ; 
a vertebral series of cordiform pale reddish-brown black-edged 
spots pointing forwards, connected by a bright orange vertebral 
band. Female reddish brown above and beneath, with angular 
dark brown bands across the back; a few dull green blotches on 
the head, and on the sacral and anal regions. 


11. CoRYTHOPHANES CRISTATUS Gray. 


This remarkable lizard was believed to be restricted to Central 
America (specimens from British Honduras, Guatemala, Nicaragua, 
and Costa Rica are preserved in the British Museum), but a 
specimen from the Rio Tamana, Choco, was in the collection 
made by Mr. Palmer, and two are in Dr. Spurrell’s collection. 
“Shows rapid changes of colour when alive.” 


12. BASsILIscUS AMERICANUS Laur. 


Its habitat extends from Panama and Costa Rica to Venezuela 
and Southern Colombia. 


13. Bastniscus GALERITUS A. Dum. 


Blackish above, dotted with yellow ; upper surface of head and 
a band on each side of the belly reddish brown; throat white ; 
belly and lower surface of limbs greenish white, speckled with 
black. 


14. ENYALIOIDES HETEROLEPIS Bocourt. 


Originally described from Veragua. Numerous examples from 
Colombia and N.W. Ecuador are preserved in the British 
Museum. 


15, DipLocLossus MonorRoPIs Wiegm. 


The largest specimens are very strikingly coloured. Head 
lemon-yellow with greenish tinge; back chequered brownish 
black and lemon-green; throat, belly, and sides of body ver- 
milion-red ; tail lemon-yellow above, with dark bars, vermilion- 
red beneath. 


16. AMEIVA BRIDGESII Cope. 


Young black above, with seven whitish longitudinal lines ; 
belly bluish grey ; tail blue. 


COLOMBIAN BATRACHIANS AND REPTILES. 1033 


17. Ametva Festiva Licht. 


A single, half-grown specimen. Blackish above, with a pale 
green vertebral band originating on the end of the snout and 
prolonged on the basal half of the tail; two interrupted white 
lines along each side of the body. 

A specimen from Medellin, Andes of Colombia (Coll. A. KE. 
Pratt), is in the British Museum. 


18. AnwapiA viTTATA, sp.n. (Pl. CVII. fig. 2.) 


Head depressed, with long, acutely pointed snout ; body much 
elongated. Nostril in the suture between two nasals. Fronto- 
nasal pentagonal, a little longer than broad, longer than the 
prefrontals, which form a long suture; frontal octagonal, 
narrow behind, in contact with three supraoculars; two large 
parietals on each side, both in contact with the interparietal, 
which is not twice as long as broad and followed by three sub- 
equal occipitals; four supraoculars, first smallest, second and 
third equal; four or five supraciliaries; a loreal, a freno-orbital, 
and a row of infraorbitals; temples with rather small shields ; 
seven upper labials, first narrowly in contact with the fronto- 
nasal; six lower labials; chin-shields, one anterior and four 
pairs, three pairs forming a median suture; gular scales sub- 
equal, squarish, juxtaposed ; collar-scales nine, feebly enlarged. 
Scales on body forming regular whorls, the dorsals quadrangular- 
subhexagonal, as long as broad, the laterals a little narrower, the 
ventrals quadrangular, those of the two median rows larger; 30 
scales round the middle of the body, 58 from occiput to sacrum 
inclusively. Preanal plates in three rows, posterior median pair 
largest. Two przanal and one femoral pore on each side. Limbs 
and tail as in A. ocellata. Pale brownish white above, with three 
dark brown longitudinal stripes, the vertebral edged with black 
and breaking up into three black lines behind the nape; lower 
parts white; tail with three serrated or zigzag brown bands. 


Total length (tail injured) .................. 112 mm. 

JELCE Vel GM eenie te rote none Sun Ae ed ent ee Bee 10 

Wel Glitlnrotmne adnate ercrtrcksS eacey isola ons 5 

From end of snout to fore limb ............ 15 
MMR ae Sean VECINGL craiancaiet hes cis wilt a 40 

Ronen imilbpessccc ee eee ate es see 10 

Tina lelinrmllneerieete set sere iate ince cece ees aces i, 


A single specimen. 
This species is closely allied to A. ocellata Gray. 


19. ALopocLossus copit Blgr. 


Originally described from Hastern Ecuador, this species has 
since been found in Western Hcuador, but had not been reported 
from Colombia. 


1034 MR. G. A. BOULENGER ON 


20. AMPHISBENA FULIGINOSA L. 


Received alive by the Zoological Society. 


OPHIDIA. 
21. TRACHYBOA BOULENGERI Peracca. 


A single young specimen of this species, recently described 
by Peracca, Ann. Mus. Zool. Univ. Nap. iu. 1910, No. 12, was 
referred by me to 7. gularis Peters, P. Z.S. 1898, p. 109, 
pointing out the characters in which it differed from Peters’s 
description and Jan’s figure. The spine-like raised scales on 
the snout and above the eyes, which are so striking in larger 
specimens, are merely indicated in this specimen. 

The British Museum possesses two adult specimens from N.W. 
Keuador, and two from the Choco are now added by Dr. Spurrell. 
1 append particulars of these four specimens :— 


1. 2, St. Javier, N.W. Keuador. Total length 430 mm.; 
tail 40. Se. 27; V. 139; ©. 24. Sixth labial entering 
the eye on the right side. 

2. 3, with claw-like anal spurs. Same locality. Total length 
205 mime wall SOs Sc 30) View sor Co 20.0 atiiuhalaloiall 
entering the eye. 

3. &¢, with claw-like anal spurs. Petia Lisa, Choco. Total 
length 230 mm.; tail 27. Sce.30; V.131; C.23. Labials 
all excluded from the eye. 

4. 2, same locality. Total length 270 mm.; tail 25. Se. 33; 
V. 137; C. 20.- Labials all excluded from the eye. 


_ The type specimen described by Peracca, the habitat of which 
is unknown, has only 23 scales round the thickest part of the 
body. The number of scales in 7’rachyboa is probably as variable 
as it is in Atheris. 


22. STREPTOPHORUS ATRATUS Hallow. 

A specimen taken from the stomach of an Hlaps rosenbergii, 
after having swallowed a Cecilia intermedia. 

23. DRYMOBIUS DENDROPHIS Schleg. 

24, PHRYNONAX FASCIATUS Peters. 

25. HmRPETODRYAS CARINATUS L. 


26. LEPTOPHIS BILINEATUS Gthr. 


One of the three specimens is bright green above, as in the 
type, the two others are of a metallic bronze on the back and 
sides, the head dark green, the sides of the neck green, the lower 
parts pale green. 


The habitat of this species extends from the Isthmus of 
Tehuantepec to N.W. Ecuador. 


COLOMBIAN BATRACHIANS AND REPTILES. 1035 
27. LEPTOPHIS OCCIDENTALIS Gthr. 
28. LioPHIS ALBIVENTRIS Jan. 


29. ATRACTUS MULTICINCTUS Jan. 


In its coloration in the fresh condition (occipital blotch bright 
red) this harmless snake is not without resemblance to Llaps 
mipartitus. 


30. ATRACTUS IRIDESCENS Peracca. : 


This species was described (Boll. Mus. Torin. xi. 1896, No. 252) 


from a single specimen, origin unknown. Three specimens in 
Dr. Spurrell’ s collection may, I think, be referred to it. 


G QD, Bs mnwi,e wall BG, We 144s Ces Bre 
Oo, QQ imi, 2 wan BA, Wo s¥bs O, Beh 
Wore a0 2 mia tavlelle Ven See 20: 


The iridescence from which the species derives its name is 
strongly marked in these specimens. Dark brown above, with two 
dorsal series of transverse blackish spots, which may alternate or 
form cross-bars, and may be bordered by yellow dots or short 
streaks; in specimen @ the belly is white spotted with black in 
front, uniform black behind; in specimen 6 it is white with a 
median series of black spots and a series of smaller spots ‘on 
each side. 


31. PETALOGNATHUS NEBULATA IL. 


32. HIMANTODES CENCHOA L. 
33. LePTopIRA ALBOFUSCA Lacep. 


34. OXYRHOPUS CL@LIA Daud. 


A specimen two metres long. 


35. HoMALOCRANIUM MELANOCEPHALUM IL. 


36. HOMALOCRANIUM CORALLIVENTRE, sp. n. (Pl. CVIILI. fig. 1.) 


Kye about half as long as the snout. Rostral a little broader 
than deep, the portion visible from above about half as long as its 
distance from the frontal; internasals about two thirds the length 
of the prefrontals; frontal hexagonal, obtuse-angled in front, 
acute-angled behind, nearly once and a half as long as broad, 
longer than its distance from the end of the snout, shorter than 
the parietals ; nostril between two nasals, the posterior of which 
is in contact with the preocular; two postoculars; temporals 
1+ 1; seven upper labials, third and fourth entering the eye, 
seventh largest; four lower labials in contact with the anterior 
chin-shields, which are a little shorter or a little longer than the 


1036 MR. G. A. BOULENGER ON 


posterior and in contact with the symphysial. Scales in 15 rows. 
Ventrals 135-141; anal divided; subcaudals 56-59. Dark brown 
above, scales edged with black; a large yellowish spot on the 
upper lip behind the eye, and a yellowish occipital bar encroaching 
on the posterior part of the parietal shields; gular region white, 
rest of lower surfaces bright coral-red. 

Total length 275 mm. 

Two specimens, male (V. 135; C. 59) and female (V. 141; 
C. 56). 

Very closely allied to H. alticola Blgr., from Santa Rita, north 
of Medellin, Colombia (9000 feet), but eye larger and frontal a 
little longer in proportion to its width. 


37. STENORHINA DEGENHARDTI Berthold. 
38. ELAPS CORALLINUS L. 


39. ELAPS ROSENBERGII Blegr. 


J have examined several specimens from various localities in 
N.W. Ecuador, and also one obtained by Mr. M. G. Palmer at 
Tado, Choco. 


40. Evaps mrcrops, sp. n. (Pl. CVIII. fig. 2.) 


Kye extremely minute, barely one-fourth as long as its distance 
from the mouth. Rostral broader than deep; frontal a little 
broader than the supraocular, once and one-third as long as 
broad, as Jong as the prefrontals, half as long as the parietals, 
which are longer than their distance from the internasals; one 
pre- and two postoculars ; temporals 1-+-1; seven upper labials, 
third much longer than fourth, third and fourth entering the eye, 
the latter very narrowly; four lower labials in contact with the 
anterior chin-shields, which are as long as the posterior. Scales 
in 15 rows. Ventrals 205; anal divided; subeaudals 20. Tail 
ending obtusely. Black above, with 46 narrow white, black- 
spotted cross-bars; these widen on the belly, which is barred 
black and white; an orange band across the head, from between 
the eyes to near the end of the parietal shields; tail orange-red, 
with a black ring and a black terminal spot surrounding a small 
white spot. 

Total length 650 mm. ; tail 25. 

A single female specimen. 

This new Hlaps resembles strikingly #. mipartitus D. & B. in 
the coloration, but is well distinguished by the much smaller eye 
and by the proportions of the head-shields. 


41, LeproGNATHUS SPURRELLI, sp.n. (PI. CVIII. fig. 3.) 


Body very slender, strongly compressed. Hye large. Rostral 
small, about once and a half as broad as deep, not visible from 
above ; internasals one-third the length of the prefrontals ; 


COLOMBIAN BATRACHTANS AND REPTILES. 1037 


frontal as long as broad, nearly as long as its distance from the 
end of the snout, much shorter than the parietals; nasal divided ; 
loreal once and a half as long as deep, bordering the eye; no 
preocular ; prefrontal entering the eye; two postoculars, lower 
very narrow ; temporals 3 + 3; eight upper labials, fourth and 
fifth entering the eye; four pairs of chin-shields, the anterior 
very small and in contact with the symphysial, the second larger 
and a little longer than broad. Scales in 15 rows, vertebrals 
strongly enlarged, nearly as long as broad. Ventrals 208; anal 
entire; subcaudals 132. Reddish brown above, with very large 
blackish-brown blotches separated on the back by narrow inter- 
spaces of the ground colour, these interspaces gradually widening 
down the sides, where the blackish blotches are edged by small 
white spots: these blotches form nearly complete rings on the 
anterior part of the body, whilst further back they are gradually 
more widely interrupted by the yellowish-white colour of the 
belly ; lower surface of tail brown, with a few small whitish 
spots; head uniform reddish brown above; lips white, with 
small black spots. ‘‘ Iris golden.” 

Total length 680 mm, ; tail 235. 

A single male specimen. 

Closely allied to Z. annulata Gthr., from Costa Rica, and 
LL. temporalis Werner, from Ecuador. ; 


42. LAcHuEsIS mutus L. 


This is the othrops acrochordus of Garcia*. Grows to 
24 metres. 


43. LacuEsis atrox L. 


44, LACHESIS MONTICELLII Peracca. 


This Pit-Viper, easily distinguished from ZL. lanceolatus by 
the shape of the rostral shield, which is much deeper than broad, 
was described along with Zrachyboa boulengeri from a specimen 
of unknown origin. It is interesting to find the two snakes 
associated in this collection. 

Two female specimens :— 


a= 9A0hmmes tailelGOr seu29)-) Vie 200. 86. 
bs SOO amties 4) MOO; 4, 0 a IAS eile 


Lachesis punctatus Garcia, l.c. p. 31, pl. —, perhaps represents 
the same species. 
45. LACHESIS BRACHYSTOMA Cope. 


The range of this species is now known to extend from the 
Isthmus of Tehuantepec to N.W. Ecuador. 
Described and figured by Garcia as Thanatophis sutus. 


* Los Ofidios venenosos del Cauca (Cali, 1896), p. 23, pl. —. 
Proc. Zoot. Soc.—1913, No. LXIX. 69 


1038 


ON COLOMBIAN BATRACHIANS AND REPTILES. 


EXPLANATION OF THE PLATES. 


Prater CII. 


Fig. 1. Bufo hypomelas, p. 1022, enlarged, and outline natural size. 


Fig. 


2. Agalychnis calcarifer, p. 1023. 

2a. A % Side view. 

2b. Be es Open mouth. 
Puate CITI. 


Agalychnis sgurrelli, p. 1024. Female and eggs. 


Prater CIV. 
. 1. Dendrobates aurotenia, p. 1029. 
2-4, 3 tinctorius, var. coctei, p. 1027. 
5-6. 5 5 var. chocoensis, p. 1028. 


Prats CV. 


Cinosternum spurrelli, p. 1030. Upper view. 


Prate CVI. 


Cinosternwm spurrelli, p. 1030. Lower view. 


Prate CVILI. 


. 1. Anolis breviceps, p. 1031. 


Wes “op % Side view of head. 

Dae) as 5 Upper view of head, enlarged. 

2. Anadia vittata, p. 1033. 

Hols” 5 a Upper view of head, enlarged. 

PA al ey - Side view of head, Pa 

Poh | cn Hf Ventral and anal region, enlarged. 
Bit cs 5 Lower view of head, 53 


Pruate CVIII. 


1. Homatlocranium coralliventre, p. 1035. 

la. is BS Upper view of head, enlarged. 
1b. Side view of head, 43 

2. Hlaps micr ops, Dp. 1036. : 

BGs gp . Upper view of head, enlarged. 

2b. Side view of head, Ss 

3. Leptognathus spurrelli, p. 1036. 

3a. y, Bs Upper view of head, enlarged. 

3b. a a Side view of head, 3 

3c. is 3 Lower view of head, a} 


ON THE PERONEAL MUSCLES IN BIRDS. 1039 


60. The Peroneal Muscles in Birds. By P. CHALMERS 
Mircpety, M.A.,D.Sce., LL.D., F.R.S.,F.Z.8., Secretary 
to the Society. 

[Received October 28, 1913: Read November 25, 1913.] 


(Text-figures 179-190.) 


INDEX. 
STRUCTURE. Page 
Myotoey. 

Peroneal Muscles in Chauna ........- ‘ haere nate LOAD: 

Systematic Description of Peroneal Muscles in Wire caeheiaesets 1043 
JNTIOLOGY. 

Distribution of Varieties of Peroneal Muscles in the 

System .. M4 Sac qune eat ea dae Foti adnate. LOGS 

Adaptation andl Deseon) Sho tunssebnareacmensbncstasesctdcesaaeten LOMO) 
SysTEMATIC. 

RAN AUO Na Cress 5 OAR eee LAE ee EN LOS OSMO72 

(Obyayenrihel (eal: Cannaaceratecnsoce cdc osmGaaodand sted ecoemeadtodsauace ya UO a0} 


In the dissection of an example of Baleniceps rex, which formed 
the basis of a recent contribution to the Society’s Proceedings 
(supra, pp. 644-703), I was interested to notice that one of the 
very few characters in muscular anatomy which the Shoe-bill 
shared with Herons, to the exclusion of Storks, was the presence 
of the leg muscle known as the Peroneus brevis or profundus. On 
looking up the literature relating to the peroneal muscles in birds, 
I found that even Dr. Gadow’s careful descr iption of these muscles 
(Gadow in Bronn’s ‘ Thier-Reich,’ Aves, pp. 180-182) was based 
on a relatively small number of individuals and types, and that 
his work, together with the insignificant contributions of earlier 
and later writers, supplied only a vague and dubious picture of 
the anatomical facts in Aves. I wished to make a survey of 
these muscles as nearly as possible complete. Accordingly I have 
dissected them in the birds that have died in the Society’s Collec- 
tion for some months, have made use of the rich spirit material 
stored in the Prosectorium (most of it dating from the time of 
Garrod and Forbes), and have been allowed to examine some 
specimens in the stores of the British Museum (Natural History) 
and of the Museum of the Royal College of Surgeons, and 
Dr. Gadow has also obliged me with material from Cambridge. 
My observations therefore relate to several hundred birds, dis- 
tributed, as will be seen later, fairly well over the whole Avian 
system. Theoretically, no doubt, it is necessary to work out the 
complete anatomical structure of an animal if one would under- 
stand any portion of it, but in practice this is impossible, and in 
my opinion the examination of a single structure or even portion 
of a structure through the whole series by a single observer can- 
not fail to be interesting and instructive. The method, moreover, 


69* 


1040 DR. P. CHALMERS MITCHELL ON THE 


has the practical advantage that the anatomist gains, at least 
temporarily, a useful dexterity in exposing the structures on 
which he is engaged and is able to cover much ground in a 
relatively short space of time. 

After giving an account of the muscles in a generalized type, I 
shali proceed to a systematic description of the conditions presented, 
following the classification given by Dr. Gadow in the article 
Birp, in the Eleventh Edition of the Encyclopedia Britannica, 
and finally shall discuss some of the problems suggested by the 
degree of coincidence between the varying conditions of these 
muscles and a classification which is based on other anatomical 
facts. 


The Peroneal Muscles in CHAUNA CHAV ARIA (text-fig. 179). 


In some respects the Screamers are generalized birds with 
affinities pointing in many directions, and it happens that the 
condition of the peroneal muscles in other birds can be easily 
described as specializations of the Screamer condition chiefly by 
loss of parts. : 

Peroneus superficialis seu longus.—TVhis is the most superficial 
mass of muscle seen when the skin is removed on the outer, upper 
aspect of the leg from the knee downwards. Its very large fleshy 
mass arises from the anterior and lateral crista of the tibia, from 
the head of the fibula and partly from the fascia covering the 
knee. Laterally, where it is bounded by the external head of the 
gastrocnemius on the one side, and the tibialis anticus on the 
other, its fasciz are closely attached to the fasciz of these muscles. 
The muscular mass is roughly triangular, and from the broad 
base at its origin from the upper end of the tibia and fibula, 
it converges to a flat tendon which is apparent about three 
quarters way down the leg and rather more on the outer than the 
anterior aspect. There is a deep origin formed by a comb of 
fibres projecting inwardly as a keel along nearly the whole length 
of the triangular portion of the muscle and arising partly from 
the whole length of the fibula and partly from the tibial surface 
immediately under the fibula. When the tendon has nearly 
reached the tarsal joint, it gives off a very broad and strong anchor 
(text-fig. 179, Anch.) to the cartilaginous sustentaculum which is 
pierced by some of the flexor tendons and joined superficially by 
the gastrocnemius tendon. It is then continued outwards and 
downwards across the ankle-joint, immediately under the skin, 
but running through a synovial membrane which has to be cut 
open to expose it fully, and ultimately, about a quarter of the 
distance down the tarso-metatarsal shaft, it fuses with the flexor 
tendon of the third digit (flexor primus seu perforatus IIT) (text- 
fig. 179, P.L. IIL), It is supplied by a branch of the ischiadic 
nerve. 

The function of this muscle is to straighten the tarsus meta- 
tarsus on the tibio-tarsus and then to flex the third toe. When 


PERONEAL MUSCLES IN BIRDS. 1041 


Text-fig. 179. 


Peroneal muscles in Chauna chavaria. 
Right leg, external aspect. Muscle striped: tendon dotted. 


P.LON. Peroneus longus. ANCH. “Anchor”: attachment of P. longus tendon 
to Sustentaculum of Flexor tendons. P.L. III. Slip of P. longus tendon passing’ 
across the ankle-joint to fuse with III, tendon of Flexor perforatus of the third 
toe. 

P.BRE,. Peroneus brevis. P.B.1. Tendon of insertion of P. brevis passing across the 
tarsal jot to insertion to proximal end of the tarsus-metatarsus. P.B.2. Slip 
of Peroneus brevis tendon, not usually present in birds. 

Tar. Lig. Ligament crossing the tarsal joint, deep of P.L.III and superficial to 
JEIB5 Ils 

Gastr. External head of gastrocnemius muscle. 

Flex. Flexor tendons. 


1042 DR. P. CHALMERS MITCHELL ON -THE 


the tendon has been dissected out of its synovial groove, it 
appears too long for its purpose, but if its action be tested before 
this dissection has been made, it is clear that the straightening of 
the foot is carried out by the action on the sustentaculum and 
that subsequently the third toe is bent by the special slip. 

The arrangement is odd and very striking, and it is extremely 
difficult to believe that it has been independently developed in the 
large number of different groups of birds in which it occurs. In 
my opinion the disposition of the muscle and its tendons shown 
in Chauna can be explained best if it be regarded as the spe- 
cialized derivative of a condition in which the perforated flexor 
muscles and their tendons were continuous with each other and 
with the peroneus ; as the foot and its digits became specialized, 
the fasciz of insertion became strengthened along the lines 
leading to each digit, and finally by the disappearance of the 
connecting fascize the continuous sheet became broken up into 
separate tendons ; as the tibio-tarsus became longer the peroneal 
slip to the tendon of digit III assumed its odd course. All the 
stages in the possible differentiation of continuous fascie into 
highly specialized tendons still exist in the case of the alar 
tendons of birds. The differentiation of the foot, however, must 
have taken place before the groups of birds became separated, 
and, as I shall show later, the various conditions of the peroneus 
in the different groups of birds can all be explained as differ- 
entiations of the condition in Chauna, in particular by the loss 
of some part. 

Peroneus profundus seu brevis (Tibialis posticus of R. W. Shu- 
feldt, “‘ Myology of the Raven,” 1870, p. 228).—This is a much 
simpler and more slender muscle arising fleshy from the fibula 
and the adjacent surface of the tibia on their anterior aspects, 
beginning about the level of the insertion of the biceps tendon 
and extending a little beyond the distal extremity of the fibula, 
where it passes into a stout rounded tendon (text-fig. 179, P- BRE). 
The greater portion of the muscle is concealed by the edges of the 
tibialis anticus and of the superficial peroneal which meet above 
it. The tendon as it approaches the end of the tibio-tarsus is 
very close to the bone and is deep of the peroneus longus tendons ; 
it flattens out, runs through a synovial channel, crosses the tarsal 
joint overlain by a ligament (text-fig, 179, Tar. lig.), which also 
crosses the joint, and is inserted to the proximal edge of the tibio- 
tarsus on its posterior, outer aspect (text-fig. 179, PAB ail) same lol 
Chawna it also has a second attachment to the antero-median end 
of the tarsal shaft (text-fig. 179, P.B. 2). 

This muscle flexes the tarsus-metatarsus on the tibio-tarsus, and 
at the same time slightly rotates the foot, depressing the axial, or 
great toe side, and raising the abaxial, or fourth toe side. The 
latter action varies with the exact position of the insertion and 
is often very feeble, as, indeed, is the action of the muscle as a 
whole, 


PERONEAL MUSCLES IN BIRDS. 1043 


SysTEMATIC DESCRIPTION. 


I have found that the following points are the most notable :— 

Peroneus longus.—Origin : “superficial,” 7. e. forming a fairly 
broad external sheet at the proximal end of the tibio-tarsal shaft 
and arising chiefly from the head of the tibia and fibula and from 
the fascize over the knee-joint; “fascial,” from the fascie of the 
tibialis anticus on the one side and from that of the external 
gastrocnemius and the underlying flexors on the other ; ‘‘ deep,” 
from the side of the fibular shaft and the adjacent area of the 
tibia on the opposite side from the origin of the peroneus brevis. 
The “ fascial ” origin is the least important morphologically and is 
correlated with the strength and width of the muscle. ‘ Anchor,” 
to the cartilaginous sustentaculum through which the flexor 
tendons run at the tarsal joint. ‘Slip to III,” the superficial 
tendon which runs from the peroneus over the tarsal joint to join 
the perforated flexor tendon of the third toe. 

Peroneus profundus.—Origin. Presence or absence of a bony 
or fibrous bridge at the lower end of the tibio-tarsus. Tendon of 
insertion. 

Relative size of the two muscles. 


RATITAL, 


Struthiones. Struthio.-—-The peroneus longus is enormous. 
Superficial origin extends over to the femoro-fibular ligament and 
the patella. Fascial origin present. Deep origin represented by 
a separate head from the lower end of the fibula. Anchor is 
broad and flat but short. Slip to III strong and rounded. 
Peroneus brevis absent, but there is a short stout ligament which 
may represent either the lower portion of the muscle, or, in my 
opinion, more probably the ligament which crosses the tarsal 
joint in Chauna. 

Rhee. LKhea americana (two examples).—P. longus as in 
Struthio, except that the deep origin extends for the ereater part 
of the length of the fibula. P. Tire evis absent, but tar soil hgament 
present, 

Casuaril. Casuarius uniappendiculatus.—P. longus enor- 
mous ; superficial and fascial origins very extensive, deep origin as 
in Rhea. Anchor short, very wide and partly in slips. Slip to 
III stout. P. brevis absent but there is a stout tarsal ligament. 

Dromeus nove-hollandie.—As in Casuarius, but P. longus not 
quite so large. 

Apteryges. Apterym (@ species) (three examples).—P. longus 
very large; superficial and fascial origins good, but deep origin 
very slight. Anchor broad and short ; slip to III a rounded 
tendon. P. brevis very much reduced but present as a small flat 
tendon arising from a few muscular fibres and running in the 
normal position under the slip to IIT. This is notably different 
from the tarsal ligament in the other “struthious birds” and is 
quite certainly a rudiment of the Peroneus brevis. 


1044 DR. P. CHALMERS MITCHELL ON THE 


In the Ratites the P. longus is the dominant muscle and does 
not differ exceptin minor details from the type in Chawna which, 
for convenience, I may call the “normal” arrangement. The 
Peroneus brevis is at least functionally absent, but the Apteryges 
stand apart from the others in possessing an undoubted vestige 
of it. 


CARINAT A. 
CoLYMBOMORPH. 
COLYMBIFORMES. 


Colymbi. I have not examined a Diver. 

Podicipedes. Podiceps cristata and P. minor.—In both these 
Grebes the P. longus was badly developed, with delicate super- 
ficial and deep origins passing into a long rounded tendon about 
half-way between the knee and the tarsal joint. In the Crested 
Grebe this tendon passed straight to the sustentaculum, so that 
its insertion was represented only by a narrow anchor, there 
being no slip to Il]. In the Little Grebe the anchor was similar 
but I was able to trace a minute but normal slip to II]. On the 
other hand, in the Crested Grebe there was a minute and certainly 
functionless tendon representing the P. brevis which I did not 
find in the Little Grebe. | 


SPHENISCIFORMES. 


Sphenisci. Spheniscus demersus.—P. longus rather weak 
but all origins present; anchor and slip to III both slender and 
short. P. brevis was also slender but arose fleshy from the 
greater length of the shaft of the fibula and tibia and was 
inserted by the usual slender flat tendon running in a synovial 
groove under the slip to III. Gadow mentions that it is present 
but weak in Aplenodytes. Beddard’s statement that 1t does not 
occur in Penguins is erroneous. ‘ 


PROCELLARITFORMES. 


Tubinares.—Daption capensis. Procellaria (?species). Ocea- 
nodroma (¢ species).—In the Cape Pigeon and the two Petrels the 
P. longus was rather small with only the superficial origin, which 
quickly passed into a very long, slender tendon ending in a 
delicate anchor and a slender slip to III. The P. brevis had a 
short but strong origin from high up the fibula ; this passed into 
a strong and very long tendon with the usual flat insertion. The 
P. brevis is rather stronger than the P. longus. The examples 
of Petrels which I examined were part of the material of W. A. 
Forbes and had been preserved in spirit for at least thirty years. 
I was interested to note that they still retained a very strong 
musky odour, so clinging that although I used several kinds of 
soap and petrol I could not get rid of it for several days. 


PERONEAL MUSCLES IN BIRDS. 1045 


Tt is clear that the Colymbomorphe present no coherent 
picture, the Petrels being sharply marked off from the others 
by the relative dominance of the P. brevis. 


PELARGOMORPH®. 
CICONIIFORMES. 


Steganopodes. 


Phaéthontide. Phaéthon.—P. longus normal but rather small, 
with superficial and deep origin, broad anchor and slip to ILL. 
P. brevis normal, with long muscular origin and short tendon with 
usual flattened insertion. 

Sulide. Sula bassana.—P. longus normal and large but with 
only superficial and slight fascial origin. Broad short anchor and 
ship to III. P. brevis absent. 

Phalacrocoracide. Phalacrocorax (?species).—P. longus normal 
and large; origin, anchor and slip to Ill as in Sula. P. brevis 
strong but very short, arising only from the lower end of the 
tibial shaft, distal to the fibula, and with normal insertion by 
flattened tendon. 

Fregatide. Mregata.—Peroneus longus large and strong but 
origin practically only deep from the whole length of fibula and 
adjacent surface of tibia on side turned from the brevis. Broad 
anchor and good slip to III. P. brevis minute but quite distinct 
and normal, with long slender belly and tendon passing under a 
fibrous bridge and then flattening out to pass to usual insertion. 

Pelecanide. Pelecanws.—I have not had a recent opportunity 
of examining a Pelican, but from my old notes I find that the 
P. longus is large and normal with the usual anchor and slip to 
III, and that the P. brevis is present but minute. 

Ardez (from these I exclude Scopus and Baleniceps). 
Ardea egretta, A. herodias, A. herodias lessoni, A.purpurea. Ardetta 
minuta. Nycticorax gardeni, NV. violaceus (2 examples). Botaurus 
stellaris, Cancroma zelodoni.—In all these the P. longus is fairly 
large (text-fig. 180), but has only the superficial origin with shght 
fascial origin. The anchor is rather long, broadens out, and at 
least in one case (A. herodias lesson) was intwoslips. The P. brevis 
is well developed usually from about three-quarters the length of 
the shaft but chiefly from the fibula. The long tendon passes 
through a fibrous bridge and then flattens out to the normal inser- 
tion. I think there must be some mistake in Gadow’s statement 
that the P. brevis is absent in Ardea. I find that its presence is 
one of the differences between Herons and Storks. In Ardetia 
minuta and Cancroma it is particularly strong. 

Scopide. Scopus umbretta.—The P. longus is large with 
superficial and slight fascial origin, broad anchor and good slip 
to III. The P. brevis is either actually absent or represented by 
a very small ligament. 

Balenicipitide. Baleniceps rexw.—P. longus a strong broad 


1046 DR. P. CHALMERS MITCHELL ON THE 
muscle with only superficial and fascial origins. Anchor broad 
and good slip to III. The P. brevis is strong but short, arising 


only from the tibia distal to the end of the fibula; its flattened 
tendon passes over to the usual insertion. 


Text-fig. 180. 


(he r 
/ Ni) 


Peroneal muscles in Nycticoraa gardeni. 
Description and lettering as in Text-figure 179 (p. 1041). 
Br. Bridge traversed by the tendon of the P. brevis. 


Ciconie, Ciconia nigra. Dissura episcopus. Leptoptilus crumen- 
iferus (2 examples). Mycteria americana. Pseudotantulus ibis 
and Z'antalus loculator.—In all these Storks the conditions were 
practically identical. The P. longus was large and strong with 
superficial and fascial origins and no deep origin, very broad 
anchor and good slip to III. The P. brevis was absent, but 
running across the tarsal joint parallel with the slip to III, and 
therefore approximately at 1ight angles with the normal position 
of the P. brevis tendon, was a stout elastic ligament which kept 
the tarsal joint flexed, so that after death the joint could not be 
straightened without force until this had been cut. At first I 
thought that this ligament might represent the brevis, but its 
direction is different, and it does not resemble the indubitable 
vestiges of the brevis which occur in some other birds. 


PERONEAL MUSCLES IN BIRDS. 1047 


Tbididee. Carphabis spinicollis. Platalea leucorodia. Plegadis 
Jalcinellus.—In these the P. longus is normal with superficial and 
fascial origins, only a few fibres of deep origin, good broad anchor 
and slip to III. The P. brevis is present but very weak, arising 
by a few thin fibres from the greater part of the length of the 
fibula and passing into a slender but long tendon with the usual 
flat insertion. 

Pheenicopteri. Phanicopterus antiquorum.—In the Flamingo 
the P. longus is strong with only the superficial and fascial 
origins, with good wide anchor and slip to I1I. The P. brevis is 
absent. 

The general picture presented by the Ciconiiformes is of a 
large well formed longus muscle chiefly with superficial origin, 
and a brevis muscle almost invariably slender and showing a 
strong tendency to disappear, but none the less with the presence 

absence following the minor divisions of the assemblage, 
and not being sporadic in the genera. 


ANSERIFORMES. 


Palamedee. Chauna chavaria.  Palamedea cornuta.—Il 
have already described the conditions in these (supra, p. 1040). 

Anseres. Cygnus olor, C. wigricollis, dx galericulata 
and 4a sponsa. Cereopsis nove-hollandie. Chloéphaga inornata. 
Dendrocygna fulva. Querquedula castanea. Metopiana peposaca. 
“demia nigra. Mergus serrator.—In the Swans, Geese and 
Ducks the P. longus is large with a very broad superficial origin 
wrapped round the front of the knee, partly covering and partly 
with fascial origin from the tibialis anticus and a lesser extension 
towards the gastrocnemius. The muscle narrows rapidly (text- 
fig. 181) to a rounded tendon from which a very broad anchor, 
frequently i in separate slips, is given off, the main tendon forming 
the slip to III. The P. brevis is always present and usually 
large, with a central tendon and pinnate slips of fibres from the 
eveater part of the fibula and adjacent surface of the tibia. It 
frequently dips under a fibrous bridge and then flattens out and 
has an unusually wide insertion to the abaxial side of the end of 
the tibio-tarsus. There is usually one, and not infrequently a 
second separate tendon running across the tarsal joint. ‘The 
variations within the group are small, all the normal parts being 
present. The deep origin of the longus is usually very shght, but 
as an exception it is well- marked in Dendrocygia. The anchor 
is very broad and short, and usually in separate slips, but in the 
ducks generally and in Dendrocygna it is rather more fan-shaped. 
In @demia the brevis is very short, arising only from the tibia 
below the fibula and from a small portion of the fibula; its 
tendon, moreover, instead of crossing under the slip to II] runs 
parallel with it, and is inserted to the axial edge of the tarsus- 
metatarsus. 

In the Anseriformes the peroneals are both well developed, 


1048 DR. P, CHALMERS MITCHELL ON THE 
the longus, however, being relatively the larger, but the brevis is 
actively functional and in addition to its flexion of the tarsal 


joint has a notable rotating action. 


Text-fig. 181. 


/ 


| 


Mi 
Th] fy, 


ANCH: 748. 


Peroneal muscles in Ax sponsa. 
Description and lettering asin Text-figure 179. 


The Per. longus has had the proximal portion removed to show the P. brevis. 


FALCONIFORMES. 


Catharte. Cathartes wwra(2 examples).—The P. longusis a 
strong muscle, with good but rather narrow superficial origin, with 
fascial origin chiefly on the side of the tibialis anticus, and with 
extensive deep origin. The anchor is broad and short and the 
slip to IIT is very strong. The P. brevis is very slender, arising 
only from the distal two-thirds of the fibula, then passing into a 
rounded tendon much weaker than the slip to III under which it 
runs, and then flattens out to the normal insertion. The domin- 
ance of the longus over the brevis recalls the condition in the 
Ciconiiformes and differs from that present in Accipitrine birds 
(excluding Serpentarius). 


PERONEAL MUSCLES IN BIRDS. 1049 


Accipitres. Serpentariidee.—I have had no recent opportunity 
of examining a Secretary bird, but I infer from Dr. Gadow’s 
statement (‘‘ Aves” in Thier-Reich) that both peroneals are present, 
and that as P. longus is very large the condition is generalized and 
thus ike what exists among the Ciconiiformes. 

Vulturide and Falconide. Gypactus barbatus. Tinnanculus 
alaudarius. Falco peregrinus (2 examples). Aquila verreauat. 
Haliaétus leucogaster. Ictinia mississipiensis. Circus gouldi.—In 
all these birds both peroneals are present, but the P. brevis is the 
more powerful and the P. longus tends to be reduced only to its 
deep origin. In the Laimmergeier there is the greatest re- 
semblance with the normal arrangement. The P. longus has all: 


Text-fig. 182. 


P LON. 


SS 


i 


ANCH>-</ 


Pas A 


Peroneal muscles of Falco peregrinus. 
Description and lettering as in Text-figure 179. 


Fib. Anterior edge of fibula. 


three origins, but the superficial origin is rather narrow and 
partly reduced. In all the others the superficial and fascial 
origins have practically disappeared, so that the muscle, as shown 
in the case of the Falcon (text-fig. 182), almost exactly balances 
the P. brevis, arising from the other side of the fibula and the 
adjacent area of the tibia along the greater part of the length of 


1050 DR. P. CHALMERS MITCHELL ON THE 


the shaft. The anchor is always present and is usually rather 
long and slightly fan-shaped. The slip to III is always present 
and normal. The P. brevis is very strong; it arises in the 
Liimmergeier only from the lower end of the fibula and the tibia 
distal of this; in J/etinia its origin begins higher up; in the 
others, as shown in text-fig. 182, it arises from nearly the 
whole length of the fibula below the biceps insertion, from the 
adjacent surface of the tibia and from the tibia distad of the end 
of the fibula. Its very strong tendon passes under a fibrous 
bridge which may be calcified in old birds, then flattens out and 
passes in the usual synovial sheath under the slip to III to its 
normal insertion. 

It is plain that in these birds-of-prey the superficial origin of 
the P. longus has been partly or completely lost and that the 
P. brevis has greatly increased in relative importance, until it 
may surpass the P. longus. 

Pandionide, Pandion haliaétus—In the Osprey I found no 
trace of the P. longus, in which Iam confirmed by Dr. Gadow. 
The P. brevis was short and not very strong, arising from the 
usual origin limited to the lower half of the shaft and passing 
into a weak tendon, which after traversing a bony bridge, 
flattened out and had the normal insertion of this muscle. As 
ornithologists, arguing from other features, have differed as to 
placing the Osprey with the Hagles and Vultures, or with the 
Owls, it is of some interest to note that in the Owls also only the 
P. brevis is present. 

Apart from Pandion, it is clear that the Falconiformes 
generally show a gradual reduction or specialization of the 
P. longus, always retaining, however, the deep origin, the anchor 
and a strong normal slip to ILI, and a gradual increase in size of 
the P. brevis. 

Gadow unites the Ciconiiformes, Anseriformes, and Falconi- 
formes into the ‘‘ Legion” Pelargomorphe. Again excepting 
Pandion, it appears that throughout the Legion, the P. longus 
remains a large and important muscle always with a good anchor 
and good slip to III, but that the P. brevis offers every condition 
from that of perfect equality with or even superiority to the 
longus in the specialized Falcons, to complete absence. 


ALECTOROMORPH &. 


TINAMIFORMES. 


Crypturi. Rhynchotus rufescens (4 examples). Nothura 
maculosa. Calodromas elegans (3 examples).—In all the Tinamus 
J examined, the P. longus was well developed, with very large super- 
ficial and fascial origins and long deep origin. ‘The anchor was 
strong and broad and the slip to III strong. J noticed that in a 
Rufous Tinamu the P. longus first straightened the tarsal joint and 
then flexed sharply the second as well as the third digit. On dis- 
secting out the tendons, I found that the tendon of the perforated 


PERONEAL MUSCLES IN BIRDS. 1051 


flexor muscle of the third digit, after being joined by the tendon 
from the peroneus, gave off not only the slip to the perforated and 
perforating flexor of digit ITI which occurs in birds belonging to 
many different groups, but also a slip to the perforated flexor 
tendon of digit II, an arrangement which I have not noted or 
found recorded in any other bird. 

The P. brevis was present in all the Tinamus I examined; in 
Rhynchotus it was very feeble and slender, arising from the greater 
part of the length of the fibula and passing into a thin tendon 
which flattened out, and passing under the slip to III had the 
normal insertion. In Wothura its origin was from the lower 
part of the fibula, and in Calodromas it was extremely minute 
and reduced to a few fibres and a tendon just visible, but 
running the usual course. The definite presence of the P. brevis 
in Tinamus is in contrast with its equally definite absence in 
most of the Struthious birds, but resembles the condition in 
Apteryx. 


GALLIFORMES. 


Mesitidee.—I have had no opportunity of examining Jesites, 
and Milne-Kdwards does not refer to the peroneal muscles. It 
would be interesting to know their disposition, as in Hurypyga 
the tendon is characteristically long, and very different from the 
arrangement in Galliform birds. 

Turnices. Turnix dussumieri.-—The P. longus is present and 
rather strong, with the three origins, a good anchor and slip to 
If. The P. brevis is slender but arises from nearly three-quarters 
of the shaft, from the fibula and tibia, and passes into a short 
tendon which flattens out to the normal insertion. 


Galli. 


Megapodiide. Talegalla lathami.—The P. longus is very strong 
with large superficial and good fascial origins but no deep origin. 
Anchor strong but narrow, and good slip to III. The P. brevis 
is also good, from the posterior three-quarters of the shaft from 
fibula and tibia. Tendon is very short, passes under a fibrous 
bridge and then flattens out to pass to usual insertion. 

Cracide. Ortalis ruficauda. Mituatuberosa. Pauais galeata. 
Penelope purpurascens.—In all these the P. longus is enormous 
but chiefly from superficial and fascial origin, the deep origin being 
just present. The anchor is very broad, short and strong, and the 
ship to III is good. In some the tendon was ossified down to 
the anchor, The P. brevis was strong, arising from the distal 
three-quarters of the shaft, from the fibula, adjacent surface of 
tibia and tibia distal to the end of the fibula. The tendon then 
passed through a long fibrous bridge, partly calcified in Ortalis, 
and flattened out to usual insertion. 

Gallide. Guttera cristata, G. edowardi, and G. pucherani. Mele- 
agris gallopavo. Argus giganteus, Gallus gallus (2 examples), 


1052 DR. P. CHALMERS MITCHELL ON THE 


Calophasis ellioti. Crossoptilon mantchuricum (2examples). Lol- 
lulus roulroul. Ortyx virginianus (2 examples).  Lophortyx 
douglasi. Coturnix coromandelica. Francolinus infuscatus.—In 
all these Galline birds the P. longus was enormous, with very 
strong superficial, well-marked Suen and rather slight deep 
origins. The anchor was stout and in most cases rather broad and 
short, but in the Francolin was narrow and rounded. The slip to 
IIT was invariably strong. The tendon of the longus was partly 
ossified down to the anchor in the older birds. The P. brevis 
was present in all, but in most cases rather slender, and its origin 
was from the distal three- quarters of the shaft including the 
fibula, adjacent surface of tibia and tibia distal to the fibula, 
The rather short tendon (long in one of the Quails) usually 
passed through a fibrous bridge and then flattened out to 
normal insertion. In Pucheran’s Guinea-fowl, however, the 
tendon did not pass across the tarsal joint, but ended on a knob 
on the distal end of the tibio-tarsus in about the position 
where it occasionally passes through a fibrous or calcified bridge. 
The P. brevis was relatively weak in all these birds, and the 
condition in the Guinea-fowl is at least complete functional 
degeneration, but I have found no case of complete absence, 
although [ examined Penelope and Gallus gallus, in which Dr. 
Gadow found it absent. 

Opisthocomi. Opisthocomus cristatus (2 examples).—The P. 
longus is of moderate size with only a rather narrow superficial 
origin. Good anchor and slip to III present. The P. brevis is 
relatively rather strong, arising from the distal three-quarters of 
the shaft in the normal fashion and with a short tendon flattening 
out to the usual insertion. 

In the Galliformes the P. longus remains the dominant muscle 
and is always well developed, with, however, seldom much deep 
origin. The P. brev is 1s present nae obviously less important. 


GRUIFORMES. 


Rallide. Mulica leucoptera. Gallinula chloropus and G. phani- 
cura. Tribonyx mortiert. Hydrornia allent. Porphyrio porphyrio 
(2 examples). Ocydromus australis. Porzana carolina. Aramides 
chiricote and A. ypecaha. Hypotenidia philippensis (2 examples). 
Rallus longirostris, R. maculatus, and R. celebensis. 

In all these Rails (text-fig. 183) the P. longus is a strong 
muscle usually with an extensive superficial and good fascial 
origin and rather little deep origin. The muscular fibres converge 
to a powerful tendon, frequently ossified, about the middle of the 
length of the shaft. The anchor is very strong and tends to be 
broken into separate slips; the slip to IIT is always present, with 
the usual course. The P. brevis is alsoalways present and rather 
strong, arising from the upper half of the fibula and the adjacent 
surface of the tibia. It then passes into a stout tendon, some- 
times ossified, and having traversed a fibrous or calcified bridge 
flattens out to the usual insertion. The text-figure, drawn from 


PERONEAL MUSCLES IN BIRDS. 1053 


a Green-backed Porphyrio, gives the arrangement usual in the 
Family. 

Gruide. Gruine. Grus australasiana. Anthropoides virgo. 
Balearica chrysopelargus.—In the Cranes the P. longus is rather 
large with extensive superficial, fascial and deep origins, very long 
tendon sometimes ossified, very wide anchor and good slip to IIT. 
The P. brevis is present and arises from nearly the whole length 
of the fibula but is very weak. The tendon, sometimes ossified, 
has the usual insertion by a flattened extremity which passes 
under both the slip to III and a very long tarsal ligament. 


Text-fig. 183. 


Peroneal muscles of Porphyrio porphyrio. 
Description and lettering as in Text-figure 179 (p. 1041). 
Br. Fibrous bridge. 


Aramine. Aramus scolopaceus.—The P. longus is large with 
superficial and fascial origins but without deep origin. It has a 
stout anchor and good slip to III. The P. brevis is either 
altogether absent or represented by a very small ligament. 

Psophiine. Psophia crepitans, P. leucoptera, and P. obscura.— 
The P. longus is large with chiefly superficial and fascial origins. 
The anchor is broad and the slip to III is present and stout, but 
peculiar in so far as after giving off a stout anchor to the per- 
forated flexor of III in the region where it usually fuses with that, 
it passes on and joins much more distally the tendon of the 
perforated and perforating flexor of the same digit. The P. brevis 


Proc. Zoou. Soc.—1913, No. LXX. 70 


1054 DR. P. CHALMERS MITCHELL ON THE 


is present, arising from nearly three-quarters of length of the 
shaft and with usual insertion. 

Dicholophide. Cariama cristata (3 examples).—The P. longus 
is large with superficial and fascial and slight deep origins and a 
very long tendon which gives off a very broad anchor and is con- 
tinued as aslip to III. The P. brevis is represented by a minute 
muscular head arising from the fibula and tibia opposite the 
biceps ligament, and a very long slender tendon close to the tibia 
which after crossin g¢ the tarsal joint flattens out to the usual 
insertion. 

Otidide. Otis tarda (2 examples).—The P. longus is very 
strong with superficial, fascial, and separate deep origins. The 
tendon is not longand after giving off a broad anchor is continued 
as the slip to III. The P. brevis was absent. 

Rhinochetide. Rhinochetus jubatus.—The P. longus was very 
large with all three origins, a stout anchor and good slip to IIT. 
The P. brevis was also ‘well developed with an extensive origin 
along the length of the fibula and adjacent surface of the tibia 
and had the usual flattened insertion ; 1t appeared to me to be 
larger in the Kagu than in any of the other Gruiform birds. 

Kurypygide.  Hurypyga helias—The P. longus was_ well 
developed with all three origins, a broad anchor and good slip to 
III. The P. brevis was ale relatively strong, arising from the 
upper half of the fibula and adjacent tibia and passing ‘into a long 
tendon which flattened out as it approached the normal insertion. 

Heliornithide. Heliornis fulica seu Podoa surinamensis (2 
examples). Podica senegalensis.—The P. longus was large in all 
these but with only superficial and slight fascial origins, the 
deep origin being practically absent. The anchor was very broad 
and strong, but the usual slip to II, although certainly present, 
was delicate and could hardly have been functional. The P. 
brevis was large and strong, with a rounded origin from the 
proximal portion of the fibula, then a very long tendon partly 
ossified which passed under a partly calcified bridge to flatten out 
to the usual insertion. 

Of the Gruiform birds the Rails and the Kagu seem to 
present the most generalized condition of the peroneals since 
both are present “and actively functional, the longus being 
the dominant muscle. In the Cranes, Seriemas and “Basinzils. 
the brevis shows signs of disappearing. The Sun-bitterns and the 
Finfoots stand apart from the others, the former recalling the 
condition most frequently found in the Limicole, and the latter, 
with the tendency for the longus to be reduced, being quite 
peculiar in the group. 


CHARADRIIFORMES. 


Limicole. 


Charadriide. ZLimosa lapponica. Actitis hypoleucus. Gallinago 
celestis. Machetes pugnax. Scolopax rusticula (2 examples). 


PERONEAL MUSCLES IN BIRDS. 1055 


Himantopus nigricollis.  Recurvirostra avocetta.  Hamatopus 
ostralegus. Vanellus vanellus (3 examples). Charadrius pluvialis. 
Rhynchea capensis. 

In all these the P. longus is rather a small muscle with a broad 
superficial, slight fascial, and very little deep origin from the edge 
of the fibula. It narrows rapidly to a long tendon which gives 
off a broad ancbor and ends in the usual slip to III. The 
P. brevis is always reduced, arising only from the proximal part 
of the fibula and passing into a lopg slender tendon which 
passes down the leg parallel with the tendon of the longus, dips 
under a fibrous bridge at the distal end of the tibio-tarsus, and 
then passing under the slip to III flattens out to the usual in- 
sertion. In the Avocet, twoexamples of the Lapwing, and in the 
Oystercatcher, the brevis was reduced toa slender ligament which 
arose from the distal end of the tibio-tarsal shaft in about the 
position where it would normally pass through a fibrous bridge, 
and passed across the tarsal joint to be lost in the fascize under 
the shp to IIT. 

Chionide. Chionis alba.—The P. longus is present with all 
three origins, the deep being very feeble. The muscle passes into 
a very long tendon which has the usual broad anchor and slip to 
Ill. The P. brevis is also present but extremely feeble, a few 
fibres from the fibula passing into a delicate tendon with the usual 
insertion. 

Glareolide. Glareola pratincola (2 examples).—Precisely as in 
Chionis, the tendon of the feeble P. brevis being still longer and 
more slender. i 

Thinocorythide. ZThinocorys ? species.—The P. longus is as in 
the Glareolide and Chionide, but the P. brevis is stronger, with 
an extensive origin from the fibula passing into a long tendon 
with usual insertion. 

(Edicnemide. Wdicnemus scolopax.—The P. longus is large 
and strong, with all three insertions passing into a rather shorter 
tendon with broad anchor and strong slip to III. The P. brevis 
is represented at most by a vestigial tendon. 

Parride. Hydrophasianus chirurgus. Phyllopezus africanus. 
Jacana jacana. Asarcia variabilis.—In all these long-toed, long- 
legged Jacanas the P. longus is rather large with superficial, 
fascial, and good deep origins. The broad band of muscle then 
passes into a very long tendon which gives off a wide but short 
anchor and is continued as the slip to III. The P. brevis arises 
from the upper third of the fibula and adjacent tibia and gives 
rise to a very long tendon which flattens out to the usual 
insertion. 


Lari. 

Laride. Larus argentatus, L. ridibundus.—The P. longus has 
only the superficial origin, and the muscle converges to a very 
long tendon which gives off a broad anchor and then forms the 


usual slip to IIT. The P. brevis arises only from a small 
Oe 


1056 DR. P. CHALMERS MITCHELL ON THE 


proximal portion of the fibula and then forms a very long tendon 
which runs down the leg parallel with the tendon of the longus, 
dips under the slip to III, and has the usual flattened insertion. 

Alcide. Alcea torda.—The P. longus is fairly large, with super- 
ficial and fascial origins, then a long tendon which gives off a 
broad anchor and ends as the normal slip to IIT. The P. brevis 
is also rather strong but much inferior to the longus. It has an 
extensive origin from the fibula and then passes into the usual 
tendon with normal insertion. 


Pterocles. Pterocles alchata (2 examples).—The P. longus 
is broad and rather strong, with all three origins and broad 
anchor and slip to IIi. The P. brevis is absent, so that in this 
respect the Sand-Grouse agree with the general tendency of the 
plover-like birds and differ from the game-birds. 


Columbe. Starnenas cyanocephala.  Leucosarcia picata 
(2 examples). Phlogwnas cruentata and P. luzonica. Geophaps 
plumifera. Phaps chalcoptera and P. elegans. Columbula picur. 
Geopelia cuneata and G. tranquilla. Zenaidura carolinensis. 
Carpophaga rufigula. Columba livia (many examples, wild and 
domestic). Osmotreron bicincta (2 examples). 

The Doves and Pigeons present an interesting series of modifi- 
cations. A state of affairs very closely resembling what occurs 
in Limicolous birds is not infrequent. In Leucosarcia, for 
instance, the P. longus is very strong, with good superficial and 
fascial origins and a considerable deep origin from the proximal 
portion of the fibula. The muscular mass narrows to a very long 
tendon, which eventually gives off a broad anchor and runs on 
to form a good slip to Ill. The P. brevis similarly arises chiefly 
from the pr roximal portion of the fibula and the adjacent surface 
of the tibia, passes into a long tendon which after traversing a 
fibrous bridge flattens out to the usual insertion. The P. longus 
1s definitely the dominant muscle, but the brevis is well formed 
and functional. In Columba the conditions are similar, but the 
extent of the fibular or deep origin of the longus varies, and the 
length of the tendon varies inversely with it. In Carpophaga 
the fibular origin of the longus is still more important, and in 
many of the smaller Pigeons and Doves, and in Osmotreron, the 
deep origin is the more important, the superficial and fascial 
origins getting weaker; the tendon is shorter, but the whole 
muscle is relatively feebler. So also in the smaller Doves and 
Pigeons, the P. brevis tends to degenerate, arising only from the 
lower end of the fibula and the tibia distad of aie, so that the 
tendon is very short and very different from the typical Limicoline 
condition. On the other hand, in Osmotreron, the P. longus has 
become smaller and reduced to its deep origin with a relatively 
short tendon, but the P. brevis has increased in size and is 
actually stronger than the longus. 

In Charadriiform birds generally the P. longus is the dominant 
muscle and the P. brevis tends to become weaker or to disappear. 


PERONEAL MUSCLES IN BIRDS, 1057 


In the case of both muscles, the normal arrangement is for the 
muscular bellies to be limited to the proximal portion of the leg 
and to give rise to very long tendons. The Pigeons and Doves 
show thie type of the Chavadviiform group, but also indications of 
moving away from it. 


In the Alectoromorph group generally, the P. longus remains 
as the dominant muscle and, except in the aberrant Finfoots, is 
always large and powerful. ‘The P. brevis varies, sometimes 
being very feeble or absent, but in some of the Columba, as: an 
exception, shows signs of outstripping the P. longus. ‘The great 
lengthening of the tendons of both muscles in some of the 
Ralliform and most of the Charadriiform birds may easily be 
regarded as in correlation with the long legs of these birds. It 
is interesting to note that the Columbze show signs of a former 
Jong-legged Charadriiform condition. 


CoRACIOMORPH &. 
CUCULIFORMES. 


Cuculi. 


Cuculidee. Cuculus canorus (2 examples). Guira piririqgua 
(2 examples). Hierococcyx varius.—In the Guira cuckoo the 
P. longus is a large muscle with good superficial, fascial, and 
deep origins. Its tendon gives off a long narrow anchor and 
then passes on toform the shp to Jil. In Hierococcyx the longus 
is relatively smaller and the anchor is broader. In Cuculus the 
deep origin is much the most important, and is chiefly from the 
distal end of the shaft. In all the P. brevis is well developed, 
with a strong origin from the tibia below the fibula and a varying 
extension up the fibula. ‘The tendon of insertion runs the usual 
course under the slip to III, and flattens out to an insertion 
placed so that the muscle not only flexes the foot but has a con- 
siderable power of rotation. 

Musophagide. Corythaix persa (2 examples). Twracus cory- 
thaix.—The P. longus is a large muscle with extensive superficial 
and fascial origins and a small deep origin extending down the 
fibula. The anchor is very broad, and the slip to IIT well marked. 
The P. brevis is a long rounded muscle arising from about the 
distal three-quarters of the length of the shaft, from the fibula 
and tibia below the fibula. The tendon passes through a fibrous 
bridge and then flattens out to the usual insertion. 

In the Cuculi the P. longus remains the dominant muscle, but 
there are signs of its being reduced to a deep origin with corre- 
sponding increase of the importance of the P. brevis. The group 
shows, in fact, a transition from the condition which I regard 
as more generalized towards the particular type of specialization 
which becomes more and more marked in other Coraciomorphines. 


1058 DR. P. CHALMERS MITCHELL ON THE 


Psittaci. Stringops habroptilus. Melopsittacus undulatus. 
Nymphicus uveensis.  Nanodes (Lathamus) discolor. Cyano- 
rhamphus alpinus.  Platycercus eximius. Aprosmactus cyano- 
pygius.  Paleornis fasciata. LHclectus pectoralis (2 examples), 
E. roratus. Poeocephalus meyeri. Caica melanocephala. Pachynus 
brachyurus. Chrysotis (¢ species). Myopsittacus monachus. 
Conurus jendayi. Calopsittacus novee-hollandie. Cacatua sul- 
phurea. Trichoglossus novee-hollandice.  Loriws domicella. 

Parrots carry further the specialization of the generalized type 
which is already indicated in the Cuculi. In every case both 
muscles are present, but there are different degrees to which the 
longus is reduced and the brevis increased. In Stringops (text- 
fig. 184) the P. longus is a large muscle with good superficial and 
fascial origins and a deep origin extending nearly the whole 
length of the shaft. The muscular belly narrows to a short 
tendon which is inserted to the capsule of the flexor tendons 
representing what I term the anchor in this memoir. A few 


Text-fig. 184. 


Peroneal muscles of Stringops habroptilus. 
Description and lettering as in Text-figure 179. 


Br. Fibrous bridges. 


delicate strands can be made out passing in the direction of the 
usual slip to III, but that structure is certainly absent func- 
tionally, even if there be a trace of it morphologically. I have 
found a small superficial origin of this muscle in several other 
Parrots ; it was relatively large in Calopsittacus, and present in 
Cyanorhamphus and Caica. Usually, however, it has been lost, 
and the normal condition in Parrots is for the P. longus to be 
reduced to a deep origin. ‘This may be large as in £electus, 
long but slender as in Platycercus, or it may be so small as in 


PERONEAL MUSCLES IN BIRDS. 1059 


Chrysotis and so closely attached to the fasciz of the P. brevis 
that Dr. Gadow has described it as fused with that muscle. 
I found a minute but distinct slip to III in Calopsittacus nove- 
hollandie, and faint traces of fibres in the direction usually 
taken by that slip in Caica, Myopsittacus, and one or two 
others. The P. brevis in Stringops (text-fig. 184) is a stout 
rounded muscle arising from high up the shaft and passing into 
a strong rounded tendon, which after traversing two fibrous 
bridges flattens out to the usual insertion. In most cases, 
however, 1t is much stronger than the P. longus, arising from 
the whole length of the fibula below the biceps insertion, from 
the adjacent area of the tibia and from the tibia distad of the 


Text-fig. 185. 


Peroneal muscles of Platycercus eximius. 
Description and lettering as in Text-figure 179. s 


Br. Fibrous bridge. 


fibula. Its strong tendon usually passes under a fibrous bridge 
and has the normal flat imsertion to the proximal end of the 
tarsus-metatarsus shaft. Platycercus (text-fig. 185) shows the 
most usual condition of this muscle in Parrots. In Trichoglossus 
and some others the P. brevis is even larger, arising from the 
head of the fibula proximad of the biceps insertion. 

Parrots thus show conditions of the peroneals which link them 
with the arrangements found in the groups I have already 
discussed, but as a group are moving away from the normal type 
in the Coraciomorphine direction. 

The Cuculiformes retain marked indications of their former 


1060 DR. P. CHALMERS MITCHELL ON THE 


possession of the generalized condition of the peroneals, but have 
moved away from that condition, the Parrots further than the 
Cuckoos and Plantain-eaters. 


CoRACIIFORMES. 
Coracie: 


Coraciidee. Coracias garrula, C. indica (2 examples). Hurystomus 
orientalis.—The P. longus is rather weak, but retains a small 
superficial, fascial, and slight deep origins. The anchor is very 
short, and the slip to III is extremely slender but present, at 
least occasionally. The P. brevis is stronger than the longus ; 
it arises from about the distal three-quarters of the shaft, from 
the fibula and tibia, passes under a fibrous bridge and is inserted 
by the normal flat tendon. 

Momotide. Momotus lessoni and M. subrufescens.  Aspatha 
gularis.—In these the P. longus is well developed with all three 
origins. It has a short anchor and is continued as a good slip 
to II]. The P. brevis is large and its tendon is just stronger 
than that of the longus. It arises from the proximal half of 
the fibula with the adjacent surface of the tibia and gives rise 
toa rather long, rounded tendon which flattens out at the normal 
insertion. 

Alcedinide. Alcedo ispida (2 examples), A. asiatica and A. 
bengalensis. Ceryle aleyon, C. americana, C. inda, and C. maxima. 
Ceyx -rufidorsa. Cittura cyanotis and C. sanghirensis. Dacelo 
gigantea (3 examples). Halcyon pileata and H. rufa. Sauwropatis 
chloris, S. sancta, S. sordida, and S. vagans.—As I have already 
pointed out in an account of the Anatomy of the Kingfishers 
(‘ Ibis, 1901, p. 97), the P. longus is present in Kingfishers but 
is plainly degenerating, possibly in association with the degene- 
ration of the fibula. It is best-marked in Dacelo; certainly I 
cannot confirm Beddard’s statement (‘Structure and Classification 
of Birds,’ p. 199) that it is absent in that bird, and he makes no 
mention of its presence or absence in other Kingfishers. It has 
a superficial origin reduced to a narrow tendon from the external 
corner of the tibial crest, joined by a few fibres from the tibia 
aleng the region of the fibula representing the normal deep 
origin. It is ‘inserted to the capsule of the flexor muscles, this 
being the usual anchor, but there is no trace of a slip to ITI. 
This is the most common condition, but in a few, notably 
Ceryle, Halcyon, and Ceyx, the muscle is reduced to a simple 
tendon with only a few muscular fibres, the insertion being the 
anchor, The P. brevis is always present and strong, arising 
from the area of the tibia usually covered by the lower end of 
the fibula and passing into a stout tendon which flattens out to 
the usual insertion. 

Meropide. Merops apiaster, M. philippensis—The P. longus 
is present but is very much reduced, having only a narrow super- 
ficial and a few fibres of deep origin. A very short anchor is 
present, and the shp to II although present is very slender. 


PERONEAL MUSCLES IN BIRDS. 1061 


The P. brevis is much stronger, with a good muscular origin from 
the distal three-quarters of the shaft; its tendon passes under 
a fibrous bridge and flattens out to the usual insertion. 


Upupidee. 

Upupine. Upupa epops (2 examples).—The P. longus is absent. 
The P. brevis is large, from the distal three-quarters of the shaft, 
including both fibula and tibia. The tendon flattens out to the 
normal insertion. 

Bucerotinze. Bucorvus (? species).  Rhytidoceros undulatus. 
Lophoceros erythrorhynchus.—I\n the Hornbills the P. longus is 
absent, although I find in my notes that there is a degenerate 
set of tendinous fibres which might possibly represent a vestige 
of it. The P. brevis is very strong, arising from the lower 
half of the shaft, from the fibula and its fibrous continuation, 
and from the adjacent surface of the tibia. The strong tendon 
flattens out to form the usual insertion. In Secorvus the 
P. brevis was equally strong but rather shorter. 


Striges. Athene noctua (3 examples).  Bubo lacteus (2 ex- 
amples), B. maculosus, B. maximus (2 examples). Asio otus. 


Text-fig. 186. 


Peroneal muscles of Strix flammea. 


Right leg, outer view. Muscle striped; tendon dotted. 


P. BR. Peroneus brevis. P. B.1. Tendon of P. brevis. 
Gastr. External head of Gastrocnemius. 

Flex. Flexor muscle of foot. 

Tib. Ant. Tibialis anticus. 

Sl. Sling of tibialis anticus. 


1062 DR. P. CHALMERS MITCHELL ON THE 


Strie flammea. Speotyto cunicularia.—There is no trace of the 
P. longus in any of the Owls. The P. brevis is enormous, but 
usually arises only from the tibia below the fibula. In Strix 
Jlammea (text-fig. 186) it is relatively longer, arising from all 
the fibula distad of the insertion of the biceps, from the adjacent 
tibial surface, and from the tibia distad of the end of the fibula. 
In Athene it is equally long. The stout tendon flattens out after 
passing through a fibrous bridge (absent in Strix) and is inserted 
in the normal fashion to the proximal end of the tarsus-metatarsus. 
Its chief action is to rotate the foot on the tibial shaft so as to 
depress the great toe side and elevate the fourth digit side. It 
comes into action after the enormous tibialis anticus has flexed 
the foot. 

Caprimulgi. 

Steatornithide. Steatornis caripensis.—Vhe P. longus is absent. 
The P. brevis is large and strong, with origin from high up the 
shaft from both fibula and tibia and fairly long tendon which 
flattens out to usual insertion. 

Podargide. Podargus cuviert. Nyctidromus albicollis. df go- 
theles nove-hollandie.—In Podargus the P. longus has a broad 
superficial origin with no fascial or deep origin. It gives off a 
broad anchor and is continued as a slender slip to III. The 
P. brevis is stronger than the longus. It arises from fibula and 
tibia just below the insertion of the biceps, and its rounded 
muscular belly passes into a stout tendon which flattens out to 
the usual insertion. In Wyctidromus the P. iongus is practically 
the same as in Podargus, but the P. brevis is absent. In #go- 
theles nove-hollandie the P. longus is absent, and the P. brevis 
is well developed, as in Steatornis. 

Caprimulgide. Caprimulgus europeus.—The P. longus is large 
with superficial, fascial, and deep origins. It has a broad anchor 
and a good slip to II]. The P. brevis is absent. 

I have followed the usual arrangement of the genera of Capri- 
mulgi that I have examined, and certainly I do not propose to 
rearrange them simply on the evidence of the peroneal muscles. 
It is plain, however, that so far as these muscles are concerned, 
Podargus has remained in the primitive condition, with both 
muscles present and normal; Steatornis and #gotheles have 
specialized in the same direction as the Owls, by losing the 
P. longus; Caprimulgus and Nyctidromus present a condition 
very aberrant amongst Coraciiform birds, and by retaining the 
P. longus and discarding the P. brevis recall the condition which 
frequently occurs in the great assemblages which I have already 
passed in review. 


Cy pselli. 
Cypselide. Cypselus apus.—The P. longus is absent. The 
P. brevis is large and strong, from the proximal end of the shaft 


including the head of the fibula and adjoining area of the tibia 
down three-quarters of the shaft, then a very strong tendon 


PERONEAL MUSCLES IN BIRDS. 1063 


passing over the bridge of the tibialis anticus to usual flat 
insertion. 

Trochilidee. Amazilia felicie.—The P. longus was absent and 
the P. brevis exactly as in the Swift. 

Colu. Colius capensis.—The P. longus was present in a ves- 
tigial condition, with only superficial origin and a*slender tendon 
ending in the anchor, with no slip to III. The P. brevis was 
very large and strong, arising from nearly the whole length of 
the shaft, involving both fibula and tibia. Its strong but rather 
short tendon flattened out to the usual insertion. 

Trogones. Zrogon puella and 7’. atricollis—The P. longus 
was present but not so large as the P. brevis; its origin is chiefly 
deep, and it has a broad anchor and fair slip to III. The 
P. brevis is very large and strong, arising from the greater part 
of the length of the shaft and ending in a short but stout tendon 
which flattens out to the usual insertion. 

Pie. 

Galbulide. Galbula albirostris, G. rufiventris. Urogalba para- 
disea.—My material for examining these was not good, con- 
sisting of partly dissected specimens which had been in spirit for 
many years. ‘The P. longus was certainly very small, but there 
was a distinct tendon running to form an anchor, and in one 
case a trace of the slip to Ifl. The P. brevis was larger than 
the longus and had a stout tendon flattening out to the usual 
insertion. I should be glad, however, to examine fresh specimens. 

Capitonide. Megalema virens.  Cyanops flavifrons. — The 
P. longus was a large muscle with a wide superficial, and short 
fascial and deep origins. The triangular muscular belly rapidly 
narrowed to a stout tendon, which, however, was not so strong as 
the tendon of the brevis, gave off a long and rather narrow 
anchor and was continued as a good slip to III. The P. brevis 
arose from the fibula beginning just below the biceps tendon, 
and from a considerable part of the tibia, and passed into a 
strong tendon which flattened out to the usual insertion. (Text- 
fig. 187.) 

Rhamphastidee. Aulacorhamphus prasinus.  Pteroglossus in- 
scriptus. Lhamphastos discolor (2 examples). Selenidera macu- 
lirostris (2 examples).—-In all these Toucans (text-fig. 188) the 
P. longus was of fair size but with chiefly fascial and long deep 
origins. The short tendon gave off a narrow anchor and was 
continued as a slender slip to III. The P. brevis was a stronger 
muscle, its stout tendon forming the axis of a muscular mass 
arising from the tibia and fibula for three-quarters the length of 
the shaft. The tendon then passed through a strong fibrous 
bridge and had the normal flat insertion. 

Picidee. Dendrocopus major. Brachypternus aurantiacus. Co- 
laptes mexicanoides. Gecinus vittatus. Hypoxanthus rivolit.—In 
the Woodpeckers I found the P. longus always present but rather 
weak and with chiefly superficial origin. Its tendon was very 
thin and flat and formed the usual anchor, and gave off a thin 


1064 DR. P. CHALMERS MITCHELL ON THE 


Text-fig. 187. 


psy / 
iti Ih Hy 
Vth Vy Hiff) 
mn by 


Peroneal muscles of Megalema virens. 
Right leg, outer view. Muscle striped; tendon dotted. 


P. LON. Peroneus longus. ANCH. Anchor to Sustentaculum. PL. III. Slip 
to Flexor of III toe. P. BRE. Peroneus brevis. 


Text-fig, 188. 


Peroneal muscles of Rhamphastos discolor. 
Description and lettering as in Text-figure 186 (p. 1061). 
P. LON. Peroneus longus. ANCH. Anchor. P.L. III. Slip to III. 


PERONEAL MUSCLES IN BIRDS. 1065 


strand which in a fresh example of Dendrocopus formed the usual 
slip to IIT, but which in the other Woodpeckers, of which I had 
only very old spirit specimens, I could not trace completely to 
their insertion. The P. brevis was strong, arising from the distal 
three-quarters of the shaft, and ending in a short tendon which 
flattened out to the usual insertion. 


The Coraciiform birds form an interesting and difficult series, 
but it is clear that the general tendency in the group is for the 
P. brevis to increase and for the P. longus to be reduced. The 
series, however, is not very coherent. A certain number have 
remained almost in the generalized condition with the P. longus 
possibly slightly reduced in its origin and equal to or weaker 
than the P. brevis, but showing the anchor and slip to ITI, and 
with the P. brevis a strong but not excessively strong muscle. 
These central types are the Coracize, Momotide, Meropide, 
Podargus amongst the Caprimulgi, and the Trogones, Capitonide, 
and Rhamphastidee. In others the P. longus is still further reduced 
and has lost the slip to III, whilst the P. brevis has relatively 
still further increased. These are the Alcedinide (Dr. Gadow, 
however, states that Pelargopsis has proceeded still further to 
the complete loss of the P. longus), the Coli, the Galbulide 
(probably), and most of the Picide. In yet another set the 
specialization has proceeded to the complete loss of the P. longus, 
and the P. brevis is always very strong. These are the Buce- 
rotide, Upupide, Striges, and, amongst the Caprimulgi, Steatornis 
and AMgotheles, the Cypselide and the Trochilide. Finally, in 
Nyctidromus and Caprimulgus, although the P. longus is partly 
reduced, the P. brevis is absent. 


PASSERIFORMES. 


Passeres Anisomyode. 
Subclamatores. Hurylemus ochromelas. Cymbirhynchus 
macrorhynchus. 
Clamatores. Picolaptes affinis. Chasmorhynchus nudicollis. 


Tyrannus melancholicus.  Myiarchus tyrannulus.  Pitangus 
sulfuratus. Pitta strepitans. 


Passeres Diacromyode. 

Suboscines. Menura superba. 

Oscines :— 

Corvide, Cracticus destructor. Creadion carunculatus. Cyano- 
corax luxuosus. 

Paradiseidee. Parotia lawesii. Mluradus melanocephalus. 

Sturnide.  Hnodes erythrophrys. Mimo dumonti. <Acrido- 
theres fuscus. 

Icteridx. Jcterus jamaicai. ‘Ostinops decumanus. 

Ploceide. Hstrelda phenicotis. 


Tanagride. Tanagra sayaca. Rhamphocelus brasilius. 
Cerebide. Coreba cyanea. 


Meliphagide.  Hntomyza cyanotus. Acanthorhynchus (2 
species). 


1066 DR. P. CHALMERS MITCHELL ON THE 


Nectariniide. <Avrachnechthra zeylonica. 

Troglodytide. Campylorhynchus wnicolor. 

Hirundinide. Hirundo rustica. 

Laniide. Lanius excubitor. 

Artamidee. Artamus lewcogaster. 

Dicruride. Dicrurus (? species). 

Oriolide. Oriolus galbula. 

Parride. Liothrix luteus. 

Turdide. Zurdus tristis. Merula tamaulipensis.  Geocichla 
citrina. Mimus orpheus. 


The Passeriform birds that I examined were well distri- 
buted over the divisions into which systematists have attempted 
to divide this group, and presented a fair sample of the manifold 
types of habit and size that occur in the group, a sample that I 
take to be fairly representative, as I found extremely little 
divergence. The type which recurs throughout the group with a 
uniformity that is almost tiresome, is well shown in the figure of 
Parotia lawesit (text-fig. 189). Both peroneals are present, well 


Text-fig. 189. 


Peroneal muscles of Pavotia lawesii. 


Description and lettering as in Text-fig. 188 (p. 1064). 


developed and functional. The longus has a broad superficial, 
good fascial, and fairly long deep origins; the muscular mass is 
an elongated triangle which ends in a short tendon which almost 
at once forks, the shorter and stouter fork forming a long narrow 
anchor, the other forming the “slip” to III, This ‘‘V ”-shaped 
forking is very different in appearance from the broad and short 
anchor which is the more common type in other groups, but it 
occurs also in other birds and can be regarded only as typically 
not characteristically passerine. The P. ‘brevis arises from the 


PERONEAL MUSCLES IN BIRDS. 1067 


fibula below the insertion of the biceps tendon and from the 
adjacent surface of the tibia; the fusiform belly gives rise to a 
short tendon which frequently, but not invariably, traverses a 
fibrous bridge and then flattens out to the normal insertion to 
the proximal end of the tarsus-metatarsus. 

The minor differences which are to be found, but too irre- 
gularly to be correlated with size, habit, or systematic position, 
relate to the relative sizes of the two muscles. Sometimes the 
P. longus is rather feebler with little deep origin and a relatively 
narrow superficial origin; the P. brevis, on the other hand, 
becoming very thick and strong. The muscular bellies of both 
muscles may be shorter and their tendons longer. 

In only two of the Passeriform birds examined did I find 
notable differences. In Menwra superba, the inclusion of which 
amongst the Passeriformes I doubt on other grounds, the anchor 
of the P. longus was much shorter and rather wider than usually 
occurs In the group, and the P. brevis was relatively considerably 
weaker. In Hirundo rustica the P. longus had the usual broad 
superficial origin but very little deep or fascial origin, and the 
slip to IIIT was absent. The P. brevis was of the usual Passerine 
type and relatively weaker than in the Swifts. 


The Coraciomorphine birds present many different phases of 
the degeneration of the longus and of the increasing importance 
of the brevis, and in this respect are in marked contrast with the 
other Legions into which Dr. Gadow has arranged the Orders of 


birds. 
SUMMARY AND CONCLUSIONS. 


The Peroneus longus muscle shows every gradation from 
elaborate structure and apparently important function to com- 
plete absence, and the facts fall into a coherent picture if we 
suppose that we have to deal with loss of parts originally present. 
The presence and absence of origin from the fasciz of the neigh- 
houring muscles are the most irregular and may well be interpreted 
as in direct relation to function. If the muscle be large and 
important, it overgrows its bounds and comes in organic contact 
with the adjacent fasciz; if it is smaller and less important it 
remains isolated. When the muscle as a whole appears to be in 
process of degeneration, this may proceed in two ways. Occasion- 
ally, but rarely, the deep origin begins to disappear first, and the 
superficial origin gradually narrows until it becomes only a thin 
superficial tendon. More often the process starts with the 
degeneration of the superficial origin, and this is often accom- 
panied by strengthening and lengthening of the deep origin, 
until finally nothing but the deep origin is left. This next 
gradually shortens until the whole muscle disappears. Of the 
tendons of insertion, the slip to ILI goes first and the anchor 
persists. There are many cases in which the slip to III has been 
lost, the anchor remaining as the only insertion, but I have found 
no instance in which the anchor has disappeared leaving only the 


1068 DR. P, CHALMERS MITCHELL ON THE 


slip to III. The pull on the anchor by the contraction of the 
muscle not only straightens the tarsal joint, but steadies the 
sustentaculum and therefore assists the action of all the flexors 
of the toes; the slip to III acts merely as an accessory to the 
proper flexor of the third toe. 

I find the Peroneus brevis very difficult to understand. Its 
rotating action is often extremely slight, and in its action in 
flexing the tarsal joint it appears to do little more than assist 
the much more powerful tibialis anticus, the mechanical arrange- 
ment of which is more favourable. So far from it being sur- 
prising to find that it has degenerated or completely disappeared 
in so many groups, it is remarkable that it should have been 
preserved at all. In the cases in which it has become stronger 
and has surpassed the longus, it seems to me that its power of 
rotation is greater, and in a number of cases, particularly in 
Passerines, its contraction appears to have the effect of partly 
flexing the toes and the great toe, as if its action would assist in 
perching. I am afraid, however, that an equally elaborate 
investigation of all the muscles of the lower leg would be necessary 
before the varying conditions of the peroneals could be inter- 
preted in terms of function. 

On the information at my disposal I find it extremely difficult 
to associate the conditions of the peroneal muscles with differences 
in habit that point directly to functional adaptation. On the 
other hand, with a few, a very few, exceptions to which I shall 
refer later, there is a close conformity between the condition of 
the peroneals and what appear to be the most securely founded 
systematic divisions. Birds seem to have this or that type of 
peroneal muscle, not because they are arboreal or terrestrial, 
swimmers or waders, scratchers, predatory or vegetarian, but 
because it is the type occurring in this or that systematic division. 
Whatever be their habit they seem to make shift with the type 
of peroneal which occurs in their group. All the four Legions 
into which Dr. Gadow groups the Orders of Carinate birds, and 
most of the orders themselves, show certain members with what 
I have assumed in this paper to be the generalized, possibly the 
more ancestral condition of the peroneals, and all show signs of 
moving in a definite direction away from this generalized con- 
dition. In the Ratites the longus is dominant but the group has 
moved away from the central type by extreme reduction or total 
loss of the brevis. Of the Colymbomorphe the Penguins show 
the central type, the Grebes have a weak longus with the slip 
to III occasionally absent, but the brevis is always rudimentary 
or absent. In the Petrels the longus has always at least a super- 
ficial origin, the anchor and slip to IIT, but the brevis has rela- 
tively increased in size. In the Pelargomorphe (except Pandion), 
the longus remains the dominant muscle and always has both the 
anchor and the slip to III, and the brevis, except in the Falconi- 
formes, tends to disappear. Some Steganopods, Herons and 
Tbises, Screamers, and Ducks and Geese remain in the central 
position. Other Steganopods and Storks and Flamingos have 


PERONEAL MUSCLES IN BIRDS. 1069 


lost the brevis altogether. The Falconiformes show an interesting 
series. Serpentarius isin quite the central position; in Catharte 
the superficial origin of the longus is slightly reduced, but the 
muscle remains the dominant of the pair. In the Falconide 
(omitting Pandion) the superficial origin and the longus generally 
tend to be reduced, but the anchor and slip to III always persist 
and are strong and functional; the brevis increases greatly and 
may equal or surpass the longus. In so far the Falconiformes 
present a parallel with the Owls, but are to be distinguished from 
that group by the retention of the longus with its. anchor and 
slip. In the Alectoromorph Legion the great majority remain in 
the central condition. The longus is invariably present, the 
superficial origin is almost invariably the stronger, the anchor is 
always present and the slip to ITI absent only in the Heliorni- 
thide. The brevis is almost invariably the weaker muscle and 
shows a strong tendency to disappear, e. g. among the Tinamus 
it may be excessively feeble, it 1s feeble or degenerate in some 
Galliform birds, absent or reduced in some Gruide and some 
Charadriide. The Columbe are specially interesting; the longus 
is always present with anchor and slip to III, but there seems to 
be a tendency for it to be reduced and for the brevis to increase, 
especially in the smaller and most Passerine-like Pigeons, 

In the Coraciomorphine Legion the characteristic tendency is 
for the reduction of the longus at the expense of the brevis. The 
Orders, however, show marked differences in the extent to which 
this process has oecurred. Of the Cuculiformes, the Cuckoos and 
Plantain-eaters remain almost in the central condition, and, 
although the brevis is always well developed, it is surpassed by 
the longus. The Parrots, like the Pigeons, show within the 
group all stages from an almost central condition where the longus 
is complete and surpasses the brevis, through stages in which the 
longus is still well developed, although it has lost the slip to III, 
to the final stage in which there is anne no trace of the loner 
remaining. The Coraciiformes have moved furthest from the 
central condition. A very few, the Motmots, some of the King- 
fishers, Podargus among the Caprimulgi, and Trogons and Barbets 
show almost the central condition, but even amongst them the 
longus is usually very little superior to the brevis, although it 
retains its parts. In most Coraciiformes the longus is at least 
feeble, has usually lost its slip to III and is often entirely absent. 
The Passeriformes, on the other hand, are remarkably constant 
and very near the central condition, although the brevis may 
equal or nearly surpass the longus, and in one case (H irundo) 
the slip to III has been lost. In the diagram reproduced as 
text-fig. 190, I have endeavoured to show in a diagrammatic 
fashion the general trend of the modification of the peroneale in 
the groups of birds. The central space represents the central or 
primitive condition, the area to the left the degeneration of the 
brevis, that to the right the degeneration of the longus. The 
placing of the named enclosures represents roughly the position 


Proc, Zoou, Soc.—1913, No. LX XI. ie 


1070 DR. P. CHALMERS MITCHELL ON THE 


of the different groups with regard to the central avian condition. 
It is to be noted that nearly every group seems to have made 
experiments in both directions, but as we read from above down- 
wards in the diagram, we pass from movement towards the left to 
movement towards the right. And it may also be noticed that, 
in a general way, what are usually regarded as the groups ex- 
hibiting the lower modifications of avian structure are towards 
the top of the diagram, those representing the higher types 
towards the bottom. If I had felt justified in expanding this 
diagram, by placing the names of the minor groups in their 
proper orientation with regard to the peroneals, it would have 
been found in the same fashion, that inside each group, on the 
whole, those which are wenerally regarded as the higher types 
were towards the right hand side of the spaces, those representing 
the lower types towards the left hand. In other words, I think 
JI may say that the higher types of avian modification are associated 
with a tendency for the degeneration of the Peroneus longus and 
an increase of the Peroneus brevis. 

Adaptation, direct or associated, may be the fundamental 
explanation of the facts that I have tried to set out, but it is 
difficult to follow, and it is easy to see that kinship appears to be 
a more important factor. The Eagles and Vultures have many 
adaptive resemblances with the Owls, but the former contain 
members directly linking them with the primitive condition and 
have never moved far from it, and the latter show the extreme 
modification of the Coraciiform group. Swifts and Swallows have 
many points in common, but the former, in the condition of the 
peroneals, are extreme Coraciiform birds, the latter very slightly 
modified from the true Passerine condition. So also Humming- 
birds are extreme Coracuformes in this respect, and Sun-birds 
are true Passerines. The family tradition appears even in many 
of the minor changes; all the Storks have lost the brevis and 
the Herons have retained it. 

There remains to say a word as to the few birds which do not 
lie comfortably, so far as the peroneals are concerned, in the 
positions usually assigned them. In a memoir dealing with a 
similar large series of facts in bird anatomy (“On the Intestinal 
Tract of Birds,” Trans. Linn. Soc., Zool. viii. p. 173), I pointed out 
what seems a logical necessity (frequently, i overlooked 
by those who use anatomical characters for systematic purposes), 
that if we have reason to believe a particular character to have 
been ancestral, we cannot assume that animals now without it 
are more nearly allied than those that have retained it. There 
is no @ priort ground for assuming that it may not have been lost 
twice or several times independently. It follows therefore that 
the loss of the longus muscle, or of any important part of it, or 
of the brevis muscle, is no valid clue to systematic position. 

Tt is equally clear that the common retention of the ancestral 
condition is no ground for placing the descendants of a particular 
ancestral stock together, if relative affinity and not convenience 
is to be the basis of classification, 


PERONEAL MUSCLES IN BIRDS, 1071 


A glance at the table (text-fig. 190) will show why anatomists 


on the hunt for convenient systematic characters would take 


Text-fig. 190. 


<—— Peroneus longus greater ; ———> 
than P. brevis, with super- 
P. brevis ficial and deep origins, P. longus 
disappearing anchor and slip to Ill. disappearing 


P. brevis functional. 


Colymbomorphee 


Pelargomorphe 


Alectoromorpha2 


Cuculiformes 


Coraciiformes 


Passeriformes 


Diagram of Peroneal museles in Birds. 


The central space represents the condition regarded as primitive in this memoir, 
the P. longus being fully developed, with superficial and deep origins, anchor 
and slip to III, and stronger than the P. brevis, which, however, is present and 
functional. The various enclosures represent the groups of birds indicated by 
the lettering. Coincidence with the central space indicates that the primitive 
condition of the peroneals is present; displacement towards the left indicates 
reduction of the P. brevis, displacement to the right reduction of the P. longus. 


le 


1072 ON THE PERONEAL MUSCLES IN BIRDS. 


little interest in the peroneals. Some of the Colymbomorphs, 
most of the Pelargomorphs, most of the Alectoromorphs, most 
of the Cuculiformes, a few Coraciiformes, and most of the 
Passeriformes would have to be placed together. We know from 
other reasons that the association would be absurd, but it is not 
more illogical than to place Ornithorhynchus and H chidna in the 
same Oren because they have retained a reptilian legacy that 
was probably once the common property of all mammals. The 
Ratites, some of the Colymbomorphs, some of the Pelargomorphs, 
and a few of the Alectoromorphs would have to be placed to- 
gether because of the absence of the brevis. Some of the 
Colymbomor phs, and a few Pelargomorphs, a few Alectoromorphs, 
a few Cuculiformes, most of the Coraciiformes, would similarly 
have to be placed together because of their loss of the longus. A 
rather tedious piece of work may be of some use, if only it serve 
to direct attention to the confusion that must arise if characters 
be not carefully analysed before they are employed for systematic 
purposes, and to the information which apparently unruly facts 
may afford, if they are plotted out over a sufficiently compre- 
hensive field. 

On the other hand, although the common retention and common 
loss of ancestral character are no proof of affinity, I myself, like 
most anatomists who have taken an interest in trying to correlate 
their facts, have been impressed by the fashion in which allied 
animals march along the path ef modification in the same 
direction. I have found this to be the case in Pigeons (Journ. 
Linn. Soe., Zool. xxvii. p. 210), in Kingfishers (‘ Ibis,’ 1901, p. 97), 
in Gruiform birds (P. Z.8. 1901, p. 629), and in Limicoline birds 
(P. Z.8. 1905, p. 155), and the general results which I have been 
stating in this communication point in the same direction. It 
appears to me therefore that the existence of notable unconformity 
at least suggests that the position assigned in the System calls 
for enquiry. 

The first notable instance is Pandion, which in the matter of 
the peroneals differs from all the Eagles and Vultures and con- 
forms with the Owls. I cannot find that this point in its 
structure has been recorded previously, but many systematists 
have seen in Pandion a link between the nocturnal and diurnal 
birds of prey, and others have actually placed it amongst the 
Owls. The question calls for re-examination. 

T am less impressed with the cases of Vyctidromus and Capri- 
mulyus. So far as the longus is concerned they agree closely with 
Podargus, but although the latter, like all other Coraciomorphine 
birds, has a well-developed brevis, the two former have no brevis. 
IT have already pointed out that at the best the action of the 
brevis is difficult to understand, and appears to be frequently 
superfluous. I am therefore of the opinion that its loss in these 
isolated cases, however curious, is insignificant. 


1HqdAUE VaaLlLadoONy We 


: e eon 
cepeemer tet _ Ee 


“SID lel “SIG “S Z al 


ON BRYDE’S WHALE. 1073 


61. On the External Characters and Biology of Bryde’s 
Whale (Salenoptera brydet*), a new Rorqual from 
the Coast of South Africa. By SOrsan OLsEn, 
Zoological Laboratory, Christiania University f. 

[Received June 27, 1913: Read November 25, 1913. ] 
(Plates CIX.—-CXIII.4) 


INDEX. Page 
Balanoptera brydet 0.0 ieee eeetee reese es LOTS 
Distributions seater cnt ean ene ene OOD, 


Introductory remarks. 


Before the Norwegian whaling trade started in South Africa in 
1909, opportunities for studying whales from that part of the 
world seldom occurred, but when at rare intervals one of the 
bigger whales was stranded on the 8. African coast, if it was not 
at once eaten by sharks it was, as a rule, quickly destr oyed by the 
warm climate and heavy sea breaking on the beach. 

The country is very extensive, the communication along the 
sea-shore bad, and journeys difficult and expensive. If, there- 
fore, one of the few scientists of the country received a report of 
a stranded whale, it was as a rule impossible for him to reach 
the spot before it was too late. So when we take all these 
difficulties into consideration it is not surprising that the bigger 
whales from this part of the world have remained unknown until 
quite recently. 

Mr. W. L. Sclater has only the following to say about the 

enus Balenoptera in the standard work ‘ Fauna of South Africa’ 
(1901) :—* Fin-whales are not uncommon off the coast of 8. Africa, 
but so far as I know, no specimen has ever been secured for a 
museum, or been examined by a competent authority.” After 
this he gives the characteristics of the four northern fin-whales, 
in order to facilitate comparison with the Cape species of the 
same genus. 

The South African whaling industry was founded by Consul 
Johan Bryde, from Sandefjord, Norway, who in 1909 erected the 
first factory in Durban and another in the following year in 
Saldanha Bay on the west coast. 

When the Norwegian whalers returned home after having 
finished their first season in Saldanha Bay, they mentioned a 
whale which had been found outside the latter place, and which 
was very similar to the ‘“ seihval ” (Balenoptera borealis). 

As the ‘“seihval” (Rudolphi’s whale) had been only known 


%* (I have consulted the issue of the Norwegian newspaper ‘Tidens Tegn’ dated 
November 12th, 1912, in which this name was first published by the author. 
Technically the species dates from the description in that Journal, but no details 
are given there which are not fully explained here.—Ep1ror. | 

ar “Communicated by the SECRETARY. 

+ For explanation of the Plates see pp. 1089, 1090. 


1074 MR. GRJAN OLSEN ON 


until then as inhabiting the eastern parts of the North Atlantic, 
it was very desirable from a scientific point of view to get further 
information about the species from Saldanha Bay, and also about 
other South African whales, by studying them on the spot. 
Mr. Johan Bryde, who has often displayed his interest in and 
generosity towards science, then offered me the necessary pecuniary 
assistance, and thus enabled me to undertake a journey to South 
Africa to study the above questions and some other scientific 
matters there. During this expedition, which lasted nearly a 
year, I visited both the east and west coasts and was able to 
make many interesting observations. 

The ‘‘seihval” from Saldanha Bay proved to be very different 
from Rudolphi’s whale (B. borealis), and is described in this paper 
as a new species, named after Johan Bryde, to whose generosity 
IT am so much indebted. I have also to thank Capt, Andr. 
Ingebrigtsen, Capt. P. J. T. Larsen, and Mr. Lars Iversen for 
their kind help. 


BALENOPTERA BRYDEI Olsen. 
‘Tidens Tegn,’ November 12, 1912. (A Norwegian newspaper.) 


Synopsis. 


The average total length is 13 and the maximum nearly 
15 metres. The shape is very elongated; the greatest height 
13-14 per cent. of the total length. The distance from the tip of 
the snout to the angle of the mouth is about 20 per cent. of the 
total length. The flippers are slender and pointed, 8-10°6 per 
cent. of the total length. The dorsal fin is small; its vertical 
height 2-2-4 per cent. of the total length. The distance from 
the tip of the snout to the anterior margin of the dorsal fin is 
69-70 per cent. of the total length, occasionally shorter; the 
minimum is 65 per cent. The distance from notch of flukes to 
vent is about 27°2 per cent. of the total length; the number of 
ventral furrows 42-54; they extend backwards to the umbilicus, 
about 1°2 metres behind the tip of the flippers. The number of 
baleen-plates (without the anterior rudimentary baleen) 250-280 ; 
their greatest length nearly 0°50 metre. The bristles of the 
baleen are very thick, long and stiff, not curling ; their colour is 
erey. ‘The anterior baleen is asa rule more or less white; the 
other greyish-black. The upper side of the flipper is bluish- 
black, the under side grey. The colour of the body is bluish- 
black above, with oblong light-coloured spots; the throat and 
an area up to about 0°65 metre below the flippers on each side 
are dark bluish-grey; the other parts of the under side are 
white, more or less yellowish, often with a grey band across the 
belly in front of the umbilicus. 


General Characteristics and Measurements. 
Balenoptera brydei is a comparatively small species, with an 
average total length of about 13 metres, or occasionally a little 


EGE 


1913 


ESF 


bs 


12 


BAL AGNOP TE RAY BRY DEI: 


Es Zo & ISIS, Pl, CX. 


BALAINOP TERA BRYDEM 


il 


. 


9, lla. BALATNOPTERA BOREALIS. 


Ls S, IGS, Pl, Cit, 


IO, Wis, 13. IIR WIDISI. 


aah Ae ly mae Rk 


14: 


BALATINOPTERA BRY] 


I 4. 3. Ils, Pil 


DEI. 


» Cox, 


BRYDE’S WHALE. 1075 


more. The longest specimen measured by me, and one of the 
largest captured “by the Norwegian whalers in South Africa, was 
a female from Durban measuring 14°95 metres. The female 
seems to attain a slightly larger size than the male. 

During my stay at the whaling stations in Durban and at 
Saldanha’ Bay I was able to examine twelve specimens in ail ; 
and the following table gives the measurements and some other 
characteristics of three which showed comparatively great 
differences. 


Measurements of Balenoptera brydei (in metres). 


a WE II Sho Geakllss ye ley ane 


per | per | | per | 
| | cent cent.) cent. 
| Total length .. wse-+e+2-| 12°35 | 100 |] 13:07 | 100 | 14:95 | 100) 
Tip of snout to anterior end of eye ene ae | 2741 |19°5 |) 2°40 )183)| 29 19-4 
| s os angle of mouth .... ..| 2°50 202) 2°48 |18°9)) 2-9 |19°4 
nS i anterior end of dorsal fin .| 8°55 |69°2) 85 (650) 105 '70:0 
a S blowholes ..........:.. .| 185 |15:0 || 2°05 |15°6|, 2°36 |15°6 
flippers (anterior margin) . 4:0 |32°3 || 40 |30°6]) 49 |32°7 
| From notch of flukes to vent.. 3°38 |27°3 | 3°56 |272| 45 (301 
a5 3 s hinder side of penis | | 
OW WRUNG, co5cc0 ce: 4°43 1350 | 4°61 35°24 48. |32°1)} 
Length of the flippers (from axilla) ......... 10 | 80] 1:39 |10:6|; 1:23 | 8:3} 
Greatest breadth of the flippers............... | 0:33 | 2°6|| 086 | 2°7||- 0-40 | 2:6} 
Breadth of the flippers at base ...............) ... || 028 | 21); 0°35 | 2:3 
Vertical height of the dorsal fin ............ 0:30 | 2°4|) 0°30 | 2:°2)| 0:31 | 2:0 
Miami eben ofeyer grat watercourse re vase GAO | cco |] OL Woo. |) CLO 
Length of the eye-opening .....................} 0°75 | 
' Length of the blowholes.. I oes Hh coo, ff OPED ip nce fh, | OS | 
| Distance between the anterior ends of the| 
blowholes 0:02 | 0:04 
Distance between the posterio ior ends of the 
Dow Oleserssee sees cacceteens sateen ide soe. I) OPER cos! I OPUS 
VETER N OH (WOE AYROS Joo cco sacaHo cou sspacaces006b0 OD | oon |) Ole |) oon ii} Owes 
Length of the penis or vulva.................. WEY ooo, |i LOS | | 018 
rameter Of pemis|abeoase mes eer eee 0°20 
Number of baleen-plates m each jaw (the | 
rudimentary baleen not included) ...... jca. 260) ... jca. 250) ... |jca. 280) 
Greatest length of the baleen-plates (with- 
MUUG LORTSHAIOS). Gao sovodoseandssasocas soo asated ene O38a |e 0°32 | ... || 0°49 
Greatest breadth of the longest baleen (at II 
| thes base)ertiee see tere ee manawa se: 018}... | O14 {| 0:23 
Greatest thickness of the longest baleen ...| 0°05 | ... a ca.0°03 
| Length of the anterior (rudimentary) | 
LOEW IEEIN | cee aanoy Sebban senate SoodapocuEt a ccaGaene OMS | Vee | Onk2 
Length of the longest bristles (on the 
Loneestabaleen)) meee aetna eae 0;20 |... I OWl4 |... (02 
Number of furrows between the flippers...) 48 | ... 54 , 54 
Height of the body at the flippers............ ca. 1°8 |14°7 
Distance between the flippers on the ventral 
RIGID -tedeaoeoctiag oan sen azeeceRan een aeee eae | mes son || 2RO 
lt if 


The shape of B. brydei, as of the other species of the genus 
Balenoptera, varies somewhat, but as a rule it is remarkably 
elongated (PI. CIX.), often even more so than in the Fin-whale 


1076 MR. ORJAN OLSEN ON 


(B. physalus). The greatest height of the body (at the anterior 
end of the flippers) is usually about 14 per cent, of the total 
length, occasionally less, and only in a single case (a male of 
Saldanha Bay. 12.9.12) is it as much as 147 per cent. The 
greatest breadth is almost the same as the height, or a httle less. 
In five specimens measured by me in Saldanha Bay the pro- 
portion of the greatest breadth (at the flippers) compared with 
the total length was as follows :—12°8, 12:9, 14, 14:2 per cent., 
which gives an average breadth of 13-4 per cent. Ina single 
EOI OF very thin male—the breadth was only 11°8 per 
cent. The body is$therefore of a far less powerful build than in 
B. borealis and B. acutorostrata, and resembles more that of the 
Fin-whale (B. physalus). The hinder part of the body (from the 
genital opening to the tail) is comparatively less compressed than 
in B. borealis, but decreases gradually and slowly towards the 
rear up to the root of the tail-flukes. \'The caudal part forms a 
thin ridge above and below, which is at its sharpest along the 
dorsal line (Pl. CXI. fig. 7). In order to facilitate the com- 
parison between B brydei and the other species of the genus 
Balenoptera, 1 have added a table showing the most remarkable 
differences. J have not considered it necessary to mention the 
Blue whale (4. musculus), as its size, dorsal fin, colour, and 
many other characteristics make any confusion with B. brydei 

impossible. 


TABLE 


showing the most notable differences between Balenoptera 
brydei and the other species of the same genus (except 
the Blue whale). 


B. acuto- B. B. B. 
rostrata. borealis. brydei. physalus. 
Total length (maximum) ............. 10°3m, 163m. 15m. 24m. 


Per cent. of total length. 


From tip of snout to blowhole ...... 13°5 lbp 15°3 tise 

ys . % angle of mouth 13-17 178 19°4-20:2 ea. 20 

5 5 3 MOY DVRS sos cos oo oy 29°8 30°6--32°7 33 

i dorsal fin ...... La 61-68 65-70 69-73 

Height of body at the flippers ...... 20 18 14 sth 
Height of the dorsal fin ......... 4-5'6 3°3-4:7 2-2°4 oe 
Length of the flippers (from axilla) ibe 87-93 8-10'6 §°9-11:3 
Greatest breadth of the flippers com- aslto35  aslto 

pared to their length ............... a or 3°6 2°2 oes 
Number of ventral furrows ......... ar iss 42-54 60-100 
Number of baleen-plates oH ithout oy x oF 5 

the rudimentary baleen)........ he AS) ABZBD | AHO 200 
Greatest length of baleen in | adult “ ? ; ~ ; 

specimens (without bristles) ...... ‘ DeaDiee 6 Oe ee Dee ia SRO a 


Distance from notch of 
flukes to anus. 
Ventral furrows............ 


Colours, PO eae Re 


Bristles of baleen ......... 
Food and habits 


BRYDE'S WHALE. 


B. borealis. 


31:6 per cent. (of the total 
length). 


Extend to the middle of 


the belly one foot behind 
tip of flippers. 

The throat always white: 
the underside behind anus 
bluish black, like the 


back. 


Unusually fine, short, curl- 
ing and wool-like, white. 

Always small crustaceans, 
chiefly Calanide. Never 
takes fish, 

Blows as a rule only once 
or twice during each visit 
to the surface. 


Migratory. 


1077 


B. brydei. 


27°2 per cent. (of the total 
length). 

Extend to the umbilicus, 
3°5 feet behind tip of 
flippers. 

The throat dark bluish- 
grey: the underside 
behind anus zhite, 
sometimes yellowish or 
whitish grey. 

Very thick, long and stiff, 
not curling, colour grey. 

Fish. When it occasion- 
ally takes crustaceans, 
they are Euphausiide, 
like the Blue whale. 

Blows 5-6 times during 
each visit to the surface 
(ike the Fin- and Blue 
whales). 

Stationary. 


The distance from the tip of the snout to the angle of the 
mouth is, as we have seen, 19°4—20°2 per cent. of the total 
length, about the sameas on the fin-whale, whereas in Rudolphi’s 
whale it is only about 17°8 per cent. (according to R. Collett). 
The tips of the snout and of the lower jaw are also far more 
acute than in B. borealis, more resembling those of B. physalus. 

The flippers are very slender and pointed, generally com- 
paratively short, but varying in length from 8-10°6 per cent. 
of the total length, and they are thus sometimes remarkably 
longer than has ever been observed in B. borealis. ‘Their greatest 
breadth (at the middle) compared to the length is as 1 to 2°2. 
They differ in this respect very much from those of &. borealis, 
where the proportions are as 1 to 3°5 or 3°6, and they are more 
like the flippers of B. physalus. There is often a cut of about 
an inch in breadth and depth in their posterior margin, near the 
axilla. P 

The dorsal fin is of very moderate size, and its vertical height 
only about 2°2—2°4 per cent. of the totallength. Itisthus smaller 
than is usual in the fin-whale, and very different from the large 
dorsal of B. borealis and B. acutorostrata. Its length from the an- 
terior to the posterior end is only slightly greater than its vertical 
height. The front margin is convex, with a slightly curved apex, 
and the hinder margin is deeply concave and often has a small 
cut in it near the base. The point of the fin is very thin and 
sharp. ‘The dorsal fin is placed noticeably further back than is 
the case in Rudolphi’s whale—often where it would be on the 
fin-whale. In twelve adult specimens examined by me the 
distance from the tip of the snout to the anterior end of the 
dorsal fin was in most cases 69-70 per cent. of the total length, 
and in one instance only 63 per cent. In B. borealis the dorsal 


1078 MR. ORJAN OLSEN ON 


is usually placed anterior to the beginning of the last third of the 
body (according to R. Collett). 

The tail-flukes ave broad and comparatively somewhat shorter 
than in B. borealis. The length of each fluke is from 10-11 per 
cent. of the total length, or about the same as in the fin-whale. In 
amale specimen, of which the total length was 13:07 metres, their 
thickness at the base was about 0°20 metre and 0-12-0'14 metre 
halfway between the base and the tip. 

On each side of the tail, above and below, there is a thin ridge 
which gradually becomes lower towards the rear, and disappears 
between the flukes, about O°-10 metre from the end of the 
body. 

The eyes are comparatively larger than in the fin-whale, and 
placed just above the angle of the mouth, in the same place as in 
the fin-whale, but further back than in Bb. borealis. 

The distance from the tip of the snout to the blowholes is about 
15°3 per cent. of the total length. In adult specimens the length 
of the blowholes is from 0°32-0-40 metre. They are placed 
on a broad ridge on the head in two long furrows, which converge 
very much towards the front. Between them there is a shorter 
central furrow. 

The ventral furrows run along the whole length of the lower 
jaw and extend to the umbilicus, about 1:2 metres behind the 
tip of the flippers (as in the fin- and blue whales). In B. borealis 
they end about 0°30 metre beyond the tip of the flippers 
(R. Collett). Above the long furrows there are 5-8 shorter ones 
with a length of 0°30—-0°60 metre, and these terminate in the 
axilla. In front of the flippers there are also a few short furrows 
which extend forward into the angle of the mouth. On the belly, 
in the area below the angle of the mouth, some shorter furrows 
are as a rule wedged in amongst the others. Just in front of the 
umbilicus, where the long furrows terminate, a number of quite 
short furrows are also to be seen. The number of long furrows 
varies from 42-54. 

The anus is noticeably further to the rear than in B. borealis, 
and the distance from the anus to the notch of the flukes is as a 
rule about 27:2 per cent. of the total length. 

The penis is rather big and with the vulva is placed ina generally 
dark-coloured furrow 1-1°5 metres long, about two-thirds of 
which run anterior to the genital opening, and about one-third 
behind. 

A varying number of whitish-grey oblong spots, about 7-8 cms. 
in length and about 3 ems.in breadth, are distributed over the 
whole body. Along the centre of their long axis there runs 
usually a dark line, from which fine radii of similar lines run in all 
directions. These patches probably show places once attacked by 
parasites. In addition to these whitish spots a number of fresh 
wounds, with a length of as much as 10 cms. and 3-4 cms. deep, 
can be seen in most cases. They are marks caused by parasites— 
generally a Pennella. 


BRYDE’S WHALE. 1079 


The majority of specimens of Bryde’s whale are very thin, and 
their stratum of fat has an average thickness of only 4—5 ems. 
In a single instance—that of an old and probably diseased female 
specimen from Durban—the thickness was only about 3°5 cms. 
Some big specimens, especially females with foetus, may be quite 
fat and their stratum of fat may attain a thickness of as much 
as 7 ems., at any rate on the dorsal side. Specimens as fat as 
this may yield 15 or occasionally nearly 20 barrels of oil, while 
the common yield of this species is only 6-8 barrels. 


Baleen. 


During my stay in 8. Africa I was able to examine the baleen 
of twelve adult specimens, and also a large quantity of whale- 
bone which lay heaped up outside Mr. Bryde’s whaling station in 
Saldanha Bay. 

The baleen of B. brydei is very distinctive of this species, and 
compared with the size of the whale, very small (see P]. CXII. 
fig. 11 B). The longest baleen-plates measured by me, and from 
a female specimen of nearly 15 metres in length, were only 
0-49 metre long (the bristles not included), while baleen from a 
Rudolphis whale of the same size attains a length of about 
0-70 metre. The baleen differs remarkably in shape from that 
of B. borealis; whereas the baleen-plates of 5. borealis are very 
long and slender, those of Bb. brydet are comparatively very broad 
and curve inwards along the inner margin. The comparative 
proportions of the breadth of the largest baleen-plates at their 
base and their length were in four specimens of B. brydei:— 
43, 43-1, 46-9, and 47 per cent., which gives an average of 45 per 
cent. In B. borealis the same proportion is about 27:2 per cent. 
(see Pl. CXIT. fig. 11 4). The baleen of B. brydei seems also to be 
comparatively somewhat thicker and more strongly built than in 
B. borealis. The number of plates in twelve specimens examined 
by me was, as a rule, about 260 in each jaw, though the number 
varied between 250 and 280. But if the rudimentary plates are 
included, the number may be about 350. ‘The number of baleen- 
plates in B. brydei is remarkably small ; for Rudolphi’s whale the 
number (without the rudimentary baleen) is from 320 to 340, 
and in the fin-whale from 360 to 400. 

The rudimentary plates along the tip of the upper Jaw are not 
compressed, and attain a length of 0-10-0°15 metre; they are 
numerous and difficult to count. The first compressed plate 
is seen about 0:25-0:30 metre behind the tip of the snout. 
There is also some rudimentary baleen above the angle of the 
mouth. 

The bristles (Pl. CXII. fig. 10) are longer than in B. borealis 
and thick and stiff, not curling, and on the whole of very strong 
construction—even comparatively stronger than in the fin-whale. 
Their average thickness is about 1 mm.; a little more at the 
distal end of the baleen, and perhaps a little less at the base. 


1080 MR. ORJIAN OLSEN ON 


B. brydei differs very much in this respect from B. borealis, 
and it would hardly be possible with its imperfect straining 
apparatus to keep back such small crustaceans as the Calanide, 
which form the chief food of B. borealis. 

The colour of the baleen in the anterior part of the jaws, 
and about 0°70 metre backwards from the tip of the snout, is 
as a rule more or less white, sometimes perfectly white, but 
more frequently with grey stripes; further back it is greyish 
black, and after death perfectly black. The bristles are grey, 
whitish grey or yellowish in the anterior part of the mouth. 
The colour of the baleen is on the whole rather similar to that of 
the fin-whale, and the whalers told me that in some cases they 
had even seen the white colour asymmetrically placed, as in the 
fin-whale. The baleen might in one jaw be white over a com- 
paratively large area, and in the other jaw fairly uniformly dark- 
coloured. A male caught in Saldanha Bay, March 5, 1913, had 
only dark-coloured baleen. In this respect too B. brydei differs 
considerably from B. borealis, in which the whalebone is only 
rarely white-mottled and as a rule uniformly black with white 
bristles. 

In connection with the description of the whalebone, it is 
worth mentioning that some time after I had given the first 
preliminary report on 5. brydei in a Norwegian newspaper, 
Mr. Bryde wrote to me saying that he could now understand the 
reason for a law-suit in which he had been implicated some time 
before. He had sold a consignment of baleen from his factories 
inS. Africa which his agents there considered to be “ seihval ” 
(B. borealis). The buyers, however, complained and returned 
the whalebone, saying that 1t was not from #&. borealis but from 
another species of whale. The result was a law-suit in which 
the authorities who examined the whalebone gave judgment 
in favour of the buyers, and Mr. Bryde was obliged to pay a 
large fine and take back the whole consignment. 

The Norwegian whalers in South Africa said too that this whale 
(B. brydet) was not the proper “ seihval”; but as it was most 
like the “seihval” in size and colour, they generally called it 
that. 

Hairy covering. 

In all the specimens examined by me (except in the case of 
an old and probably diseased female specimen from Durban, on 
which I could see no hairs at all), I found two rows of hairs on 
the tip of the lower jaw, with twelve hairs in each row, thus: 


Mouth. 


BRYDE’S WHALE. 1081 


The distance between these rows in adult specimens was about 
18 mm., or a little more than between the successive hairs 
in each row. Outside the upper end of these rows were two 
other hairs on each side, placed at the same distance from the 
long rows as between the rows themselves. I noticed no hairs on 
the upper jaw. 

The number of hairs is thus 28 in all. On two foetuses, with 
a length of 3°78 and 0:935 metres, the hairy covering was as well 
developed as in adult specimens. The length of the hairs was 
from 5-10 mm. 


Colowr. 


The upper side of the body—above the flippers—is bluish black, 
and in colour much resembles the fin-whale. After death the 
colour on this part becomes almost entirely black. On the sides 
of the body the colour becomes gradually lighter, and there is no 
sharply detined border between the dark colour of the upper 
side and the white belly. The flippers are of the same colour as 
the back above, or sometimes darker; below they are grey, 
occasionally whitish grey. The under side of the head, and a 
band extending to about 0°70 metre below the flippers and 
backwards to their tip are bluish grey. The centre of the belly 
is a more or less yellowish white. In the dark area below the 
flippers some isolated white spots may sometimes be seen ; similar 
white patches are occasionally distributed along the centre of the 
under side of the lower jaw, and more rarely a small white line 
runs out from the white area on the belly to within one metre or 
less behind the tip of the lower jaw. On the border between the 
dark colour of the throat and the white belly there are often a 
number of bluish grey patches or stripes, and these are more 
thickly sprinkled towards the front, and then gradually form a 
more uniform dark colour in front of the angle of the mouth. 
Between the tip of the flippers and the umbilicus—at the posterior 
end of the long furrows—the white colour of the under side is 
generally interrupted by a bluish grey belt running across the 
stomach. This belt, which has a breadth varying between 0-70 
and 1°5 metres, is often formed by a large number of smaller 
and larger patches or stripes, not of uniform colour; in other 
cases it 1s very little developed. 

Considerable variations occur in the breadth, size, and shape of 
the white area on the belly. 

From the genital opening to the tip of the tail-flukes the 
whole under side is dirty white, sometimes yellowish or whitish 
ervey. This white area begins above the genital opening at the 
middle line of the sides of the body, and in some specimens is 
further back and becomes gradually narrower. 

From the tip of the upper jaw, near the roots of the baleen, 
sometimes a white stripe runs backwards, and the length of 
this is about 0°5 metre and the breadth 5-7 ems. 


1082 MR. GRIAN OLSEN ON 


The dark patehes which are sometimes found on the under side 
are generally situated in the long furrows, or in the furrow at the 
genital opening. 


Occurrence and Capture. 


Balenoptera brydei seems to be common everywhere along the 
south coast of Africa. It is especially so in Saldanha Bay, 
though also in Mossel Bay and near the Cape of Good Hope this 
species is captured in rather large numbers, and is often observed 
from whaling steamers going along the $8. African coast. 

In Darban &. brydet is comparatively rare, but 1s occasionally 
to be seen at all seasons of the year. The total number captured 
from three whaling stations on the Natal Bluff during the season 
1912 was only sixteen, of which one (captured early in 1 November) 
was only about 8 metres long, and not fully developed. In 1911 
six specimens were captured from Myr. Bryde’s station in Durban. 
Of these two were taken on June 5th, one on the 14th, and one 
on the 15th of the same month, one on October 10th, and one on 
November 7th. In 1912 the following were caught from the 
same station:—one on June 4th, one on July 18th, one on 
Aug. 19th, and one on Nov. loth. Some specimens were seen 
but not pursued because, as the whalers told me, they were 
very shy and difficult to catch, and at the same time their value 
was comparatively small. At Mr. Bryde’s whaling station 
near Inhambane, Portuguese EH. Africa (23° 30'S. 1.), B. brydet 
was very seldom seen, and only a single specimen of it was 
captured. Further to the north at Bazaruto Is. (21° 45'8.1.), 
where the floating factory ‘‘ Mangoro” was stationed for two 
months, B. brydet was never seen.  Bryde’s whale therefore 
seems to be a southern species, avoiding the warm waters of the 
tropics, and it is therefore rare on the east coast where the hot 
Mozambique current runs southward. 

Almost all the specimens of 2B. brydet captured in Saldanha 
Bay (33° 8.1.), were found between this bay and Capetown, and 
the whalers told me that this species was much more rarely seen 
north of Saldanha. This fact 1s quite in accordance with the 
experiences of Mr. Green, the manager of Storm Bull’s whaling 
station in Walfisch Bay. He considered the “‘seihval” to be very 
rare along the coast of German S8.W. Africa, and he had not 
observed this species at all in Walfisch Bay. 

I was somewhat surprised therefore when Capt. Andr. Inge- 
brigtsen, who had been stationed with a floating factory at 
Porto Alexandre, Benguela (15° 45'S. 1.) for a couple of years, 
told me that he had observed rather a large number of ‘ seihval ” 
there. The same statement was made by Capt. L. Fredriksen, 
who had seen many specimens of this whale outside Lobito Bay, 
and single ones as far north as 7° 8.1. But when we remember 
that a cold antarctic current runs along the west coast of Africa 


BRYDE’S WHALE. 1083 


far to the north, it is not so surprising that B. brydei may be able 
to live outside the tropical parts of the coast. The conditions of 
existence, however, for this species cannot be very good there, 
because the 3 or 4 specimens captured (during nearly three years) 
outside Pt. Alexandre were all unusually thin and did not yield 
more than 2-3 barrels of oil, or sometimes even less. All the speci- 
mens seen at this place were in very bad condition, with numerous 
wounds from parasites over the whole body, and so thin that the 
whalers did not care to pursue them. They were never seen 
accompanied by young ones outside Pt. Alexandre, and they were 
all lying almost motionless and did not make any efforts to escape 
from the steamer. Unfortunately not a single specimen has been 
examined from this part of the African coast, and very few were 
captured, because the whalers always preferred to take the more 
valuable “humpback” whales, which were to be found in the same 
locality. It has therefore not yet been proved whether the 
‘“‘seihval” from the coast of tropical W. Africa really is Baleno- 
ptera brydet. 

Capt. Andr. Ingebrigtsen also told me that the ‘“seihval ” out- 
side Pt. Alexandre appeared in these waters every year about the 
middle of June and disappeared again about September or 
October. 

Tf this whale follows the same route as the ‘“ humpback” 
generally does—along the coast—we might expect to see migrating 
specimens pass Saldanha Bay, and to find a greater number of 
them there in the spring, October-November, but this is not the 
case. It is probable therefore that they choose another route, 
perhaps further out to sea, where the current is stronger and the 
water cooler. 

It is not impossible that some of the ‘“sei”-whales from 
Portuguese W. Africa may have been Rudolphi’s whale (B. bore- 
alis), a typical specimen of which was captured outside Saldanha 
Bay in November 1912. This is the first S. African specimen of 
B. borealis which has been reported further south than off Cape 
Blanco (20° 45' N. 1.). It was easily recognized as the proper 
““seihval” by the Norwegian whalers; and Capt. P. J. Larsen 
kindly presented me with a quantity of the baleen, which was 
quite distinctive of B. borealis (Pl. XCII. fig. 9). 

As I have mentioned above, Saldanha Bay is the locality where 
the greatest number of 5. brydei are captured. The whaling 
trade was started there in 1910, and only twenty-four specimens 
of the ‘“ seihval” were captured in the first year, and all of 
them in the spring. The first three specimens were caught on 
March 9th and 29th, and on April Ist. Five were captured 
from the 20th to the 30th of April, four from the Ist to the 
10th of May, six from the 10th to the 20th of May, and six from 
the 20th to the 30th of May. After that time the “‘ humpback ” 
was found at this spot in large numbers. 

The following list shows the number of ‘ seihval” captured at 


1084 MR. GRIAN OLSEN ON 


Mr. Bryde’s station in Saldanha Bay during the season of 1911 
(with two whaling steamers), and in the season of 1912 (with three 


whaling steamers). 


April 10-20 
April 20-30 


May 1-10 
May 10-20 
May 20-30 
June 1-10 
June 10-20 
June 20-30 
July 1-10 
July 10-20 
July 20-30 


Sept. 10-20 


Sept. 20-30 
Oct. 1-10 
Oct. 10-20 
Oct. 20-30 
Nov. 1-10 
Nov. 10-20 
Nov. 20-30 
Dec. 1-10 
Dec, 10-20 


— 
Neo) 
— 
= 


CODbHSO0D0000 


CCC eCn CCC CnC) 


Cr ee ara) 


ee ee ww peewee 


Cn 


eee em ee et ene eee 


Ce Cee 


eet eer erp tet e ne 


Ciena eC a 


ea Pa) es | NOM ro KStomdoeunscae | | | a 


CC een a eed 


i ee aa 


CC ee er aC 


— 
No} 
_— 
bo 


bobo | 
NNwornwmnonnwnnton 


| Er egGo Gy Mee sl ENG eS dS | 


The table shows that the majority of 6. brydei are captured in 
the autumn (April and May), and at this time this species also 
seems to be the most numerous outside Saldanha Bay, as well as 
Durban. But we see also that 1t is captured in greater or less 
numbers during the whole season, and it is seen outside Saldanha 
Bay all the year round. 

The difference between the real number of B. brydei in the 
autumn and at other times of the year is in fact not so great as 
the capture seems to indicate, because the whalers always prefer 
to take the more valuable “ humpback” whale when this species 
appears on the spot in May. In 1912 an unusually small number 
of ‘““humpbacks” was seen, and the trade depended on the 
« seihval” to a greater extent than before. 

In 1913 the first specimen was caught on March 5, and from 


BRYDE'S WHALE. 1085 


that date until I left Saldanha Bay a fortnight later, the total 
eatch of 2. brydei consisted of eleven specimens. Almost all of 
these specimens were found unusually far (40-70 miles) from 
the sea-shore, surrounded by large numbers of crustaceans, Eu- 
phausiide. As a rule the whales are to be found quite close 
to the coast, and it is probably the richness of crustaceans 
further out to sea that has tempted them to go out there. It 
is worth mentioning that these crustaceans were Euphausiide, 
which the Blue whale prefers, and not Calanide, the chief food 
of Rudolphi’s whale. 


Food. 


The food of B. brydei consists chiefly of fish, apparently 
-usually a variety of herring which is often found in large 
quantities (many hectolitres) in its stomach. This, for instance, 
was the case with a male examined by me in Saldanha Bay on 
September 12th, 1912. It sometimes takes a species of mackerel 
one foot or more in length, and in Durban more than a hectolitre 
of this fish has been found in its stomach. 

B. brydet is very voracious on the whole—more so than any 
other species of its genus. As an illustration of this, Capt. L. 
Fredriksen told me that he had many times seen it hunting 
among large crowds of small sharks, and that he had found 
sharks of a length of more than two feet in its stomach. <A case 
from Saldanha Bay which was related by Captain Christoffersen, 
and mentioned by many others, was most astonishing: they had 
found there no less than 15 large penguins (Spheniscus demersus) 
and “malagass” (Sula capensis) in its stomach. These birds, the 
moment the whale reached the surface of the water, had probably 
dived down into its open mouth endeavouring to catch fish in 
that abundant hunting-ground, and had thus themselves been 
involuntarily captured by the whale. In similar cases observed 
among B. borealis, the birds have always been spat out of the 
mouth again, and it seems hardly possible that such large birds 
as those mentioned could be swallowed by this species at all. 

L. brydet is not dependent for food on the occurrence of 
crustaceans in the sea and so does not migrate, but is generally 
seen very close to the coast pursuing fish. 

Like the fin-whale, B. brydei cecasionally takes crustaceans and 
then of a larger kind, the Kuphausiide. 


Liology, ete. 


In its biology B. brydei is most like B. acutorostrata and 
BL. physalus, and when the Norwegian whalers started their trade 
in South Africa they were doubtful as to whether they should 
consider this species to be a fin-whale or a “sei”-whale (B. 
borealis). In Durban many of them preferred to call it a 


Proc. Zoou. Soc.—1913, No. LX XII. 72 


1086 MR. ORJAN OLSEN ON 


‘“sildehval” (herring whale), because it hunted herrings. But as 
the colour and size were most like that of the “seihval,” they 
generally called it so, although they knew that it was not the 
proper ‘‘seihval.” 

In Saldanha Bay, where the largest numbers were captured, it 
was often called “bastard” or ‘“seihval bastard,” and considered 
to be a hybrid between the fin-whale and the ‘seihval,” because 
it had the baleen of the first species, but some other characteristics 
of the latter. Besides this ‘“ bastard,” the whalers in Saldanha 
Bay told me about another variety of “ seihval” with exceedingly 
dark-coloured baleen. It was of somewhat smaller size than the 
“bastard,” quicker in its movements, dived deeper down in the 
sea, and was more difficult to catch. It was seen in approxi- 
mately the same numbers all the year round, and in the same 
localities as the “bastard,” but seemed to occur in greater 
numbers northward of Saldanha Bay than the latter, as proved 
by Capt. Christoffersen, who had captured it many times 
outside Paternoster Bay, When the whaling steamers went to 
Capetown for coaling, these two varieties were often seen in 
greater numbers together, and the whalers picked out the 
“bastard” as more valuable and easier to catch. 

Unfortunately, during my stay at Saldanha Bay, not a single 
specimen of this smaller variety was captured, and it is therefore 
difficult to say anything about them, but I consider it probable 
that they are only younger specimens of B. brydei. 

B. brydet is as a rule to be found very close to the coast, 
generally in large numbers, and it is comparatively rarely found 
further out to sea. The whalers in Saldanha Bay often found it 
pursuing fish only a few hundred yards from the beach between 
Robben or Dassen Island and the continent. In Durban it was 
also seen close to the coast (occasionally as far out as 15 miles 
from shore), but never in company with the small ‘ minkehval” 
(probably B. acutorostrata, ov a subspecies closely akin) nor with 
other whales. As mentioned above it has occasionally been found 
60-70 miles from the coast feeding upon Kuphauside. 

It is most like the fin-whale in its manner of swimming, and 
can easily be distinguished from the small “‘minkehval.” When 
coming to the surface to breathe, it may be seen under the water 
for some time before it appears on the surface. During each 
such visit it blows four or five times (like the fin- and blue 
whales, whereas Rudolphi’s whale blows only once or twice) 
before diving down into deep water again. Between these four 
or five blows it does not go too deep down to be seen from the 
ship and followed. It blows far more strongly than does 
B. acutorostrata, and its breath smells worse than that of the 
other whales. In Durban it was observed to remain for an 
unusually long time under water between each visit to the 
surface. It is not seen to take crustaceans in the same peculiar 
way as Rudolphi’s whale (B. borealis). When shot it often swims 


BRYDE’S WHALE. 1087 


round in circles very quickly and gets the line wound round it, 
and is on the whole difhicult to manage. Large females are less 
ditiicult to shoot than the others, as they are not so quick in 
their movements. 

Gravid females do not seem to be followed by their males as is 
the case with the humpback whale. For instance, two females 
captured in Saldanha Bay, March 12, 1913, and with feetuses 
measuring 0°935 and 3°78 metres, were swimming about quite 
alone. Some males captured at this time were also solitary. 

They are gravid at very different times, and females were found 
with foetuses of different sizes both in Saldanha Bay and Durban, 
at the same time as others were followed by young with a length 
of about 6-7 metres. 

It may be mentioned as a curious fact that almost all the 
“sei ”-whales captured in Saldanha Bay from March 5-13, 1913, 
went southwards when the whaling steamers began to hunt 
them. 

The flesh of B. brydet contains less oil than does that of other 
species of its genus, with the exception perhaps of B. borealis. 
When fresh and served like beef-steak it tastes quite good, as I 
had an opportunity of proving when at Saldanha Bay. 


The Fetus. 


About an equal number of each sex of B. brydei seem to be 
eaught. Many of the females were gravid, and, as mentioned 
above, with fetuses of very different sizes :—ée.g., in two gravid 
females examined by me in Saldanha Bay, March 12, “1913 
the fetuses had a length of 0°935 and 3°78 metres (PI. CX LIT. 
figs. 12, 13). The mammee in both these specimens were very 
well developed, and, -when they were dragged out of the water, 
about a half-gallon of mucus ran out of their genital openings, 
as has sometimes been observed shortly after the birth of the 
young in other whales. The largest foetus was full-grown, and 
fell out when the men began to remove the stratum of fat from 
the whale. 

The following table shows some measurements of the two 
foetuses. 

In the larger foetus the penis was placed in a furrow which 
ran backwards from the umbilicus for about 12 cm. The 
baleen had just appeared and was thick and soft, with a length 
of about 3 cm. The bristles had an average length of 6— 
8 mm. and were generally only to be seen at the base of the 
baleen. The anterior and posterior baleen only appeared as 
small ridges. The dorsal fin was bent down to the back on the 
left side, both the apex and the tip being curved to an unusual 
extent, and the hinder margin deeply concave. The tail-flukes 
were strongly bent together towards the median line of the body. 


The colour above was a uniform bluish black, and whitish grey 
(ie 


1088 MR. ORJAN OLSEN ON 


| Male. Female. 
per cent. “per cent. 
| Total length .. a |) 3°78 100 || 0:935 | 100 
| Tip of snout to anterior end of eye .... | 063 1EiGH OMS AS ae) 
e es angle of mouth | 0°70 18°5 | Ong ee | 
ae Fe anterior end of dorsal fin . 4 2°47 65°33 || -0°655 70:0 
a »  blowholes .. seocasuosae| OE mies) | Os} 13:9 
o flippers (anterior margin). 1:2 31:7 || +030 32:0 
From notch of flukes to vent ...... ee esl ee OAs 27°5 | 0°255 O72 | 
vulva or penis ...| 1°35 Sry |, ORS, Peril | 
Leneth of flipper .. sib sgaasdaongagaa scale Oy if a IOES}) I) OOe 66 | 
| Greatest breadth of flipper Per eee GOLD Meson Oe: 32 | 
| Breadth of flipper at its base...... ue teddies wna 27 || 0042 4-4 | 
| Vertical height of dorsal fin ..................; 013° | 34° |! 0:026 iy 
Length of dorsal fin at its base............. le hat ae O38 4:0 
| WRaieaeIEe OP GO uo sudenscassadvegaucocasooieonan) OOH — If cu, | 0:023 | 
HIG EEAN One ChVSAOHANUE?.  cgassncsvarcasouecsseoal| , lece | his coal yy | ORO) | 
| Length of blowholes ................. pamecaselht s OPLON Ne ORO. | | 
| Distance between the anterior end of | | | 
| blowholeste ea O23 earn O00 
| 3 5% posterior end of | 
blowholes ....... 0°073 Sa 0°025 
| eres ape HITE Moa ease onnsoanromtecesvacoocea| OO2 iI) 0°007 
| Length of penis or TUINGED. de sgncoueeeomcoosbeelh OO Keseene 0015 
Diameter of penis at its base........ i) OXORs | 
Number of furrows between the flippers... 48 i 42 
| Distance from penis te the umbilicus ......) 0°38 
From tip of lower jaw to anterior margin 
of umbilicus ...... 5904s REE: Fae a eee ae O51 54:5 
Diameter of the umbilicus cc t OOOu ale pe 0:03 
Breadth of tail-flukes at their base. sceooonsel), O82 7} | 
Distance between the tips of flukes ....... OAR |) 3: 0:20 OMB | 
Distance from notch of flukes to tip of | 
flukes .. sbdedeosnl!|  OFSts) 
Breadth of the body ‘at the flippers See BO tN fe OA elais 
Height of the body at base of flippers ...... oe NY i OPN TPR 
Be 33 at tip of flippers .... 630 13°8 
: : ati iMeleyesmen.. sence At 13°2 | 
3 rs at the dorsal fin ...... eae EWR ORO | 
5 55 at the tail, just in | | 
| front of flukes ...... Bedale vit barn BEG | 
| 


everywhere on the under side. The change from the dark to 
the light colour along the sides was ames ne and no definite 
outline could be seen. There was a more ereyish area just 
behind the eyes on each side. 

No baleen was to be seen on the smaller fcetus, and the whole 
body was of a homogeneous, reddish brown colour. 


Abnormal Specimens. 


A large female specimen with a total length of 14:95 metres, 
captur wel in Durban, November 15th, 1912, was very like the 
fin-whale in colour, Rowethe flippers were white below and the 
colour asymmetrical’ as it is in the fin-whale. The lower jaw and 
the distal part of the upper jaw were white on the right side, while 
those parts on the left side were bluish black. ‘On ‘ae, right 
side there was also a greater number of white baleen-plates ‘shar 


BRYDE’S WHALE. 1089 


on the Jeft. The furrows were bluish black in a belt across the 
under side of the head and backwards to the flippers, but only a 
comparatively small part of the throat on the left side was 
uniformly dark-coloured. No hairs were to be seen on the tip 
of the lower jaw. | This specimen was unusually slender and thin 
and probably was very old and diseased. The stratum of fat was 
only about 3°5 mm. thick; and there were numerous wounds 
distributed over the whole body (Pl. CXIII. fig. 14). These 
wounds were more or less inflamed and partly filled up with 
mortifying fat, and in some of them a Pennella was found. This 
specimen was lying quite motionless on the surface when it was 
observed from the whaling steamer, and was thus very easily 
taken. 

The whalers from Durban told me that on one occasion 
they had caught a whale of this species, which on the whole 
under side of the body was of a colour much resembling that of 
the blue whale (4. musculus); but when it was taken it proved 
to be a “ herring ” whale (B. brydei). 


Parasites. 


Parasites are comparatively rare in this species ; an old female 
(Durban, November 15th, 1912) had, as mentioned above, dis- 
tributed over the whole body a large number of oval-shaped 
wounds, with a length of as much as 10 cm., and a depth of 
2-4 em. (PI. CXIII. fig. 14). Almost ali of these wounds were 
partly filled with mortifying fat and had been caused by a large 
Pennella, specimens of which were found in some of them. . A 
smaller number of these wounds was found on this species at 
Saldanha Bay, but they were very numerous in the few old 
and apparently diseased specimens taken at Porto Alexandre. 

A species of Myxinoid sometimes makes similar wounds in 
Bryde’s whale, but I did not obtain specimens of them because 
they always leave the whale when it is dragged out of the water. 
I do not know whether they are to be found on the whale when 
alive, or only after its death. 


EXPLANATION OF THE PLATES. 
Phare CIX. 


Balenoptera brydei. 
Fig. 1. Side view of male. 
2. Lower surface of male. 
Prat CX, 
Balenoptera brydei. 


3. Male specimen, showing the dorsal side and anterior end of the body. 
Saldanha Bay, 12.9.12. 

4, Same specimen as fig. 3, from the ventral side. 

Female specimen, from the dorsal side. Saldanha Bay, 12.3.13. 


= 
Cee 
ise) 


oN 


1090 THE SECRETARY ON ADDITIONS TO. THE MENAGERIE. 


Prare CXI. 
Balenoptera brydei. 


Fig. 


SH 


Head of a large, but very thin and apparently sick, female specimen, with 
numerous wounds from parasites (Pennella). Durban, 15.11.12. 

. Caudal portion, with the dorsal fin, of a male specimen. 

. Female specimen, from the ventral side. Saldanha Bay, 12.3.13. 


mnt 


Pratre CXII. 


Fig. 9. Bristles of baleen from Balenoptera borealis, captured outside Saldanha 
Bay, —.11.12. 
10. Bristles of baleen from B. brydei, captured outside Saldanha Bay, 
—.11.12. 
11. Baleen from adult specimens of (A) B. borealis, and (B) B. brydei. 


Prate CXIII. 
Balenoptera brydei. 


Fig. 12. Foetus, 0°935 m. in length. Saldanha Bay, 12.3.13. 
13. Foetus, 3°78 m. in length. Saldanha Bay, 12.3.13. 
14. Portion of body of same specimen as fig. 6, showing wounds from parasites 
(Pennella). Durban, 15.11.12. 


EXHIBITIONS AND NOTICES. 
October 28, 1913. 
Prof. KE. A. Mincuty, M.A., F.R.S., F.Z.8. Vice-President, 


in the Chair. 


THe Secrerary read the following report on the additions that 
had been made to the Society’s Menagerie during the months of 
May, June, July, August, and September, 1913 :— 


May. 


The registered additions to the Society’s Menagerie during the 
month of May were 268 in number. Of these, 133 were acquired 
by presentation, 52 by purchase, 31 were received on deposit, 13 in 
exchange, and 39 were born in the Gardens. 

The number of departures during the same period, by death 
and removals, was 185, 

Amongst the additions special attention may be directed to :— 

2 White-bearded Gnus (Connochetes albojubatus), from 
British East Africa, new to the Collection, received in exchange 
on May 19th. 

2 Chimpanzees (Anthropopithecus troglodytes), 1 Anubis Baboon 
(Papio anubis), from West Africa, and several small Monkeys, 
presented by W. O. Danckwerts, Esq., K.C., F.Z.S., on May 2nd, 
28th, and 31st. 

1 Crowned Duiker (Sylvicapra coronata), presented by Capt. 
C. H. Armitage, C.M.G., D.8.0., F.Z.S., on May 4th. 


THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 1091 


1 Poppig’s Woolly Monkey (Lagothrix poppigi) and 1 Ocelot 
(Felis pardalis), from Manaos, presented by Albert Reymond, 
Esq., on May 28th and 20th. 

2 Naked-tailed Mice (Uromys bruijnii), new to the Collection, 
and 2 Sclater’s Cassowaries (Casuarius sclateri), from 8. Dutch 
New Guinea, presented by A. F. R. Wollaston, Esq., and C. Boden 
Kloss, Esq., on May 3st. 

2 Himalayan Yellow-backed Sun-birds (#thopyga seherie) and 
1 Nepalese Bullfinch (Pyrrhula nepalensis), from the Himalayas, 
new to the Collection, purchased on May 29th. 

2 Swainson’s Blue Jays (Aphelocoma sordida), from Mexico, 
new to the Collection, purchased on May 26th. 

1 Chestnut-faced Barn-Owl (Strix castanops), from Tasmania, 
new to the Collection, purchased on May 10th. 

4 Mexican Blue Herons (Ardea herodias lessont) and 2 Northern 
Boatbills (Cancroma zeledoni), from Mexico, new to the Collection, 
received in exchange on May 28th. 

1 Gentoo Penguin (Pygosceles papua), from the Falkland 
Islands, presented by the Rt. Hon. L. Vernon Harcourt, M.P., 
F.Z.S., on May 6th. 

3 Smooth-backed Zonures (Psewdocordylus microlepidotus), from 
S. Africa, new to the Collection, deposited on May 7th. 

1 East African Scorpion (Lepreus tricolor), trom Lake Nyanza, 
new to the Collection, presented by C. B. Blencowe, Esq., on 
May 20th. 


JUNE. 


The registered additions to the Society’s Menagerie during 
the month of June were 295 in number. Of these 144 were 
acquired by presentation, 35 by purchase, 56 were received on 
deposit, 15 in exchange, and 45 were born in the Gardens. 

The number of departures during the same period, by death 
and removals, was 189. 

Amongst the additions special attention may be directed to :— 

1 Pudu Deer (Pudu pudu), from Chili, and 2 Patagonian 
Cavies (Dolichotis magellanicus), from Patagonia, presented by 
Albert Pam, Esq., F.Z.8., on June 2nd. 

3 Pumas (felis concolor), born in the Menagerie on June 13th. 

2 Canadian Beavers (Castor canadensis), from Canada, received 
in exchange on June 5th. 

1 Savanna Sparrow (Passerculus savanna), from North-East 
America, new to the Collection, purchased on June 18th. 

1 Ceylon Mynah (Acridotheres melanosternus), new to the 
Collection, presented by Dr. P. H. Bahr, F.Z.S., on June 29th. 

1 Golden-fronted Woodpecker (Melanerpes flavifrons), from 
Brazil, new to the Collection, purchased on June Oth. 

1 Condor (Sarcorhamphus gryphus), from Chili, presented by 
Albert Pam, Esq., F.Z.S., on June 2nd. 


1092 THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 


2 Crested Screamers (Chauna cristata), bred in the Menagerie 
on June 19th: 

2 Sun Bitterns (Hurypyga helias), from South America, 
purchased on June 16th. 

1 Kagu (Rhinochetus jubatus), from New Caledonia, presented 
by the Marquess of Tavistock, F.Z.8., on June 27th. 

2 Spiny-tailed Skinks (Zgernia depressa), from Australia, new 
to the Collection, purchased on June 10th. 

A collection of Snakes from Sierra Leone including 3 Sooty 
Snakes (Boodon fuliginosus), new to the Collection, presented by 
Guy Aylmer, Hsq., F.Z.S., on June 18th. 

A collection of Snakes from India including 1 Forsten’s Tree- 
Snake (Dipsas jforstenii), new to the Collection, received in 
exchange on June 2nd. 

2 Gopher Frogs (Rana capito), from North America, new to 
the Collection, received in exchange on June 2nd. 


JULY. 


The registered additions to the Society's Menagerie during the 
month of July were 279 in number. Of these 119 were acquired 
by presentation, 46 by purchase, 21 were received on deposit, 37 
in exchange, and 56 were born in the Gardens. 

The number of departures during the same period, by death 
and removals, was 250. 

Among the additions special attention may be directed to :— 

1 Orang-Utan (Simia satyrus), from Borneo, presented by 
Marcus 8. Abrahams, Esq., ¥.Z.S., on July 24th. 

1 Small-clawed Otter (Lutra leptonyx), from India, presented 
by H. R. Taylor, Esq., F.Z.8., on July 25th. 

1 Long-tailed Mouse (Orizomys longicaudata), 2 HKlegant 
Opossums (Marmosa elegans), from Valparaiso, new to the 
Collection, presented by Walter Goodfellow, Esq., F.Z.S., on 
July 8th. 

4 Capybaras (Hydrocherus hydrocherus), from Fray Bentos, 
Uruguay, presented by the Liebig’s Extract of Meat Co., Ltd., 
on July Sth. 

1 Grysbok (Nototragus melanotis), from Constantia, Cape 
Colony, presented by W. Lester, Esq., on July 29th. 

2 Australian Flower-peckers (Dicewm hirundinaceum), from 
Australia, new to the Collection, presented by G. A. Heumann, 
Esq., on July 5th. 

1 Shoebill (Saleniceps rev), from the Sudan, presented by 
Lt.-Gen. Sir Francis R. Wingate, G.C.V.O., K.C.M.G., K.C.B., 
D.S.O., H.M.Z.8., on July 15th. 

3 Seed-Snipe (Thinocorys rumicivorus), from Chili, new to the 
Collection, received in exchange on July 24th. 

2 Spinose Fence-Lizards (Sceloporus spinosus), 1 Graham’s 
Snake (Zamenis grahanz), from N. America, new to the Collection, 
purchased on July 22nd. 


THE SECRETARY ON ADDITIONS TO THE MENAGERIE, 1083 


1 Bird-eating Spider (Citharoscelus kochit) 2, 1 Bird-eating 
Spider (Paraphysa manicata) 3, from Valparaiso, new to the 
Jollection, presented by Walter Goodfellow, Hsq., F.Z.S., on 
July 8th. 

A Collection of Millipedes (Archispirostreptus nitidus), from 
Trinidad, new to the Coliection, purchased on July 11th. 


AUGUST. 


The registered additions to the Society’s Menagerie during the 
month of August were 201 in number. Of these 122 were 
acquired by presentation, 15 by purchase, 14 were received on 
deposit, 7 in exchange, and 45 were born in the Gardens. 

The number of departures during the same period, by death 
and removals, was 231. 

Among the additions special attention may be directed to :— 

1 Fettered Cat (Felis ocreata) and 1 Cheetah (Cynwiwrus 
jubatus), presented by Dr. R. KE. Drake-Brockman, F.Z.8., 
1 Caracal (felis caracal) and 1 Cheetah (Cynelurus jubatus), 
presented by Capt. J. L. Berne, and 2 Cheetahs (Cynelurus 

jubatus), presented by Arnold Hodson, Hsq., from Berbera, on 
* August 8th. 

2 Indian Gazelles (Gazella bennetti), from Karachi, presented by 
A. C. Cooke, F.Z.5., on August 11th. 

1 Barred Warbler (Sylvia nisoria), from Kurope, new to the 
Collection, presented by R. Pearce Page, Esq., F.Z.8., on August 
Sth. 

6 West-African Quail-Finches (Ortygcespiza atricollis), from 
Gambia, new to the Collection, presented by Dr. E. Hopkinson, 
D.S.0O., F.Z.S., on August Ist. 

1 Spot-billed Toucanet (Selenidera maculirostris), bred in the 
Menagerie on August 11th. 

2 Bornean Fireback Pheasants (Lophwra nobilis), bred in the 
Menagerie on August 17th. 

A collection of rare Reptiles from Sierra Leone, including a 
Black-collared Cobra (Waia nigricollis), a Blanding’s Snake 
(Dipsadomorphus blandingii), and 3 Green. Night Snakes 
(Dipsadoboa unicclor), all new to the Collection, presented by 
Guy Aylmer, Esq., F.Z.S., on August 2nd. 

1 Hamadryad (Waia bungarus), from India, received in 
exchange on August Ist. 

6 Pigmy Rattlesnakes (Sistrurws miliarius), born in the 
Menagerie on August 19th. 

4 Miiller’s Xenopus (Xenopus miillert), from West Africa, 
new to the Collection, purchased on August 7th. 

1 Electric Cat-Fish (Malapterurus electricus), from Tropical 
Africa, new to the Collection, purchased on*August 28th. 

1 Scorpion (Bathus hottentotta), from Sierra Leone, new to the 
Collection, presented by Guy Aylmer, Hsq., F.Z.S.,0n August 2nd. 


1094 ON OPOSSUMS AND DEGUS. 


SEPTEMBER, 


The registered additions to the Society’s Menagerie during the 
month of September were 200 in number. Of these 109 were 
acquired by presentation, 21 by purchase, 7 were received on 
deposit, 26 in exchange, and 37 were born in the Gardens. 

The number of departures during the same period, by death 
and removals, was 177. 

Amongst the additions special attention may be directed to :— 

10 Mouflon (Ovis musimon), from Sardinia, presented by 
H.G. The Duke of Bedford, K.G., F.R.8., Pres.Z.8., on September 
16th. 

1 Tigrine Cat (Felis tigrina) and 1 Three-toed Sloth (Bradypus 
tridactylus), from Colombia, presented by W. D. McDougall. Esq., 
on September 15th. 

2 Variegated Agoutis (Dasyprocta variegata), 4 Woolly Opos- 
suns (Philander laniger), and 1 Sooty Amphisbeena (Amphishena 
Juliginosa), from Colombia, presented by Dr. H. G. F. Spurrell, 
E.Z.5., on September 13th. 

1 Azara’s Opossum (Didelphys azarw), 1 Burmeister’s Cariama 
(Cariama burmeistert), and 1 Western Boa (Loa occidentalis), 
from Cordova, presented by Wilfred A. Smithers, Esq.,C.M.Z.S., _ 
on September 16th. 

2 Abyssinian Weavers (Cinnamopteryx rubiginosa), new to the 
Collection, received in exchange on September 7th. 

1 Finch-like Cow-bird (Molothrus fringillarius), from Kastern 
Brazil, new to the Collection, presented by the Marquess of 
Tavistock, F.Z.8., on September 24th. 

1 Brown-headed Parrot (Pwocephalus fuscicapillus), from East 
Africa, presented by Lady D. Prain on September 24th. 

2 Mikado Pheasants (Calophasis mikado), from Mount Arizan, 
Formosa, presented by Mrs. E. J. Johnstone on September 6th, 


The Secretary exhibited a preparation of the alimentary tract 
of a young Elliot’s Pheasant (Phasianus elliott), dissected so as to 
show the gut-pattern in relation to the mesentery, blood-vessels, 
and visceral nerves. The preparation was mounted as a trans- 
pavent object in glycerine jelly, and showed how the various 
morphological conditions of the intestinal tract to which he had 
called attention (Trans. Linn. Soc., Zool. vil. p. 173) might be 
displayed as permanent museum objects. 


Young Woolly Opossums and Degus. 
Mr. R. I. Pocock, F.R.S., F.Z.S., Curator of Mammals, 


exhibited a female and three young specimens of the Woolly 
Opossum (Philander laniger), from Buenaventura in Colombia, 
presented by Dr. H. G. F. Spurrell, F.Z.S., and drew attention 


ON HEEL-PADS IN YOUNG TOUCANET, 1095 


to the rapid growth of the young. When Dr. Spurrell captured 
the mother, the little ones were still in the pouch. Six weeks 
afterwards, when they reached the Gardens, they had left the 
pouch for good, were about one-fourth grown, and were just 
beginning to feed on their own account, though mainly dependent 
upon their mother’s milk. In another six weeks—that is to say, 
three months after capture—they were three-fourths their 
mother’s. size, and apparently quite capable of fending for 
themselves. 


Mr. Pococx also exhibited two young examples of a Degu 
(Octodon degus), born from a pair from Valparaiso, presented by 
Mr. Walter Goodfellow, F.Z.S. The mother had a litter of four, 
but died from inability to give birth to a fifth. The little ones 
were fed by hand, and although two of them quickly died from 
pneumonia, the remaining two throve and promised to do well. 
Like the young of all Hystricomorphous rodents, the Degus are 
of large size at birth and open their eyes within twenty-four 
hours, and are in every respect very precocious as compared with 
young rats. Particular attention was directed to the lateral 
position of the teats in the Degu, a condition which is repeated 
in the Viscacha and Chinchilla and the Coypu. This condition 
is probably an adaptation to the large size of the young, which are 
enabled to suck lying alongside the nother, so that they do not 
interfere with each other. In the Coypu, as first described by 
Owen, the teats are higher up the side than in the Chinchilla and 
the Degu, an arrangement which permits the young of that 
species, it has been alleged, to suck while swimming alongside 
their mother in the water. 


Heel-Pads in young Toucanet. 


Mr. D. Sera-Smiru. F.Z.8., Curator of Birds, exhibited a 
spirit-specimen of a young Toucanet (Selenidera maculirostris) 
(text-fig. 191) which had been hatched in the Society’s Gardens 
in July, and called special attention to the well-developed serrated 
pads on the back of the ankle-joint or heel. This pad had been 
previously observed in the nestlings of Woodpeckers, Wrynecks, 
and Barbets, and it was only to be expected that it would be 
present also in the Toucans. 

The use of this serrated pad, which disappears soon after the 
young bird leaves the nest, is doubtless to enable it to climb up 
the side of the hollow cavity in a tree in which it is hatched— 
these pads forming, as it were, a second set of claws. It is note- 
worthy that no such pads are found in the Parrots, which nest 
in similar situations to those chosen by Barbets and Toucans, but 
which are provided with a hooked beak which aids them in 
climbing. 


1096 ON MELANOCELDUS JOMNSONII. 


Text-fig. 191. 


Nestling of Spot-billed Toucanet (Selenidera maculirostris) showing feather-tracts 
and serrated pads (a) on metatarsal joint. A. Enlarged view of serrated 


pad. 


A Deep-Sea Angler-Pish, MELANOCETUS JOHNSONII. 


Mr. C. Tate Ruean, M.A., F.Z.S., exhibited a specimen of a 
remarkable fish, Melanocetus johnsonit Giinth., that had been 
presented to the British Museum by the Hon. Walter Rothschild, 
F.R.S., and made the following remarks :— 

“This eurlous fish has an enormous mouth armed with slender 


ON A YOUNG MATAMATA TERRAPIN. 1097 


pointed depressible teeth and an extraordinarily distensible 
stomach ; it belongs to the order Pediculati and is a bathy pelagic 
species, uniformly blackish in colour; like the Angler (Lophius 
piscatorius) it has the first ray of the spinous dorsal fin situated 
on the snout and modified into a line and bait; in this ease the 
bait is a luminous bulb. 

«“The species is known from six specimens from the North 
Atlantic, three dredged at considerable depths by the ‘ Travailleur ’ 

‘Talisman,’ ‘ Valdivia’ and ‘ Michael Sars’ expeditions, and 
three in the British Museum collection. The last are from 3 to 
3? inches long (from snout to end of caudal fin) and include the 
type described by Gunther (Proc. Zool. Soc. 1864, p. 301, pl. xxv.) 
from Madeira; this had in its stomacha partly digested Scopeloid 
fish (Lampanyetus crocodilus Risso) about 7 inches long. The 
second specimen was taken at the surface in 31° N. and 37° W., 
and had in its stomach an almost perfect ZL. crocodilus nearly 
8 inches long; this was captured by Captain Charles C . Dixon, 
master of the bark ‘Hlginshire, who also secured the fish 
exhibited to-night at 7 a.m. on August 2/th, 1910, in 40° 19' N. 
and 42° 25' W. By a strange coincidence this specimen also 
contains a LZ. crocodilus, and judging by the anterior part of the 
head, which is still visible, one about 8 inches long; it is curled 
round into a complete circle in the stomach of the Metinoceme 

‘‘Captain Dixon writes that the J/elanocetus was stiil alive 
when captured and that from his observations he believes that it 
could make the ‘lure’ luminous or not at will. The fact that 
it was caught early in the morning is interesting, for it is known 
that at night many fishes that dwell at a depth of 300 fathoms 
or more may approach much nearer the surface; here the large 
Lampanyctus may have been seized and its struggles may have 
brought the captor still further out of its depth and rendered it 
helpless.” 


November 11, 1913. 


Dr. S. F. Harmer, M.A., F.B.S., F.ZS., 
in the Chair. 


Mr. E. G. Boutrnesr, F.Z.8., Curator of Reptiles, exhibited 
a quite young specimen of the Matamarn Terrapin (Chelys fim- 
br iata), the shell of which measured but 5 em. in length. This 
specimen, the smallest yet recorded, differed from the existing 
descriptions of the young of this species in the under surfaces 
being of a bright magenta-red instead of pale yellow. Mr. Bou- 
lenger suggested that as all the young previously described were 
more than double the ‘size of the specimen exhibited, red might 
be the normal coloration of the under surfaces as a juvenile 
livery. 


Text-fig. 192. 


1098 ON A HYBRID BLAGK-WINGED PEAFOWL. 


Hybrid Birds. 


Mr. D. Sers-Srru, F.Z.S., Curator of Birds, exhibited a 
remarkable hybrid (text-fig. 192) between a Black-winged Peacock 
(Pavo nigripennis) and a Domestic Hen (said to be a cross-bred 


game and Leghorn), which had been bred and presented to the 
Society by Mr. R. P. Wheadon of minster. This bird was 
hatched in 1911 and is believed to be only the second example 


Photo. by D. Seth-Smith. 


th its mother, a brown-and-white domestic hen. 


ged Peafowl, wi 


-win 


Hybrid Black 


ON DORSAL GLANDS IN ARMADILLOS. 1099 


of this hybrid ever produced, although hybrids between peafowl 
and guinea-fowl had been recorded. It was mostly white in 
colour, splashed, especially on the neck, with reddish brown, a 
colour derived from the domestic hen, its mother. It ehonwed 
clearly the characters of both parents. 

Mr. Seth-Smith also showed photographs, by means of lantern- 
slides, of a male hybrid Pheasant between Calophasis nikado 
and C. ellioti and of a male Jungle-fowl between Gallus varius 
and G'. gallus. 

The former had been bred in Sussex by Mrs. E. J. Johnstone, 
and kindly presented to the Society. It resembled a very dark- 
coloured Elliot’s Pheasant, with the head and neck, breast, and 
abdomen black. The tail was black, suffused with chestnut and 
crossed with broad white bands. 

The hybrid Gallus, which was one of several bred in the 
Gardens, was very intermediate between the two parent species. 
The comb was shaped like that of G. varius, but was shghtly 
serrated, while the central wattle of G. variws was present 
though reduced in size, as well as the two side wattles of 
G. gallus. The neck hackles, which in G. varius are short and 
rounded, were in the hybrid moderately long as in G. gallus, 
but rounded at the ends, while the colour of these hackles 
as well as of the tail, was violet-purple. This bird appeared to 
be identical with that named Gallus eneus by Temminck. 


Dorsal Glands in Armadillos. 
(‘Text-figures 193-195.) 
Mr. R. I. Pocock, F.B.S., F.Z.S., Curator of Mammals, ex- 


hibited some specimens of Armadillos of the genus Hauphractus* 
(formerly Dasypus) to show the presence of a row of glands upon 
the pelvic portion of the carapace, and remarked :—‘“ In the 
middle line of the pelvie shield there is an unpaired series of two, 
three, or four sears which are ill-defined in the living animal but 
tolerably conspicuous in dried skins and clearly marked when the 
scales are removed from the osseous scutes. ‘Their conspicuousness 
in museum specimens cannot have escaped the notice of syste- 
matic workers, but apparently their significance has never been 
investigated. 

os My attention was originally drawn to them by noticing that 
in newly born Hairy Armadillos (#. villosws) there are two or 
three small holes, one on the fourth, a second on the fifth, and 
sometimes a third on the sixth row of scutes of the shield in 
question. At this early stage the integument 1s unossified and can 
easily be cut with a razor. <A section through the holes in 
question shows that they are the orifices of shallow pits directed 


* According to Thomas, P. Z.S. 1911, p. 141, Dasypus must be transferred to the 
Peba Armadillos, replacing J GeO 


1100 MR. R. I. POCOCK ON 


downwards ant forwards and dipping into the dermis which is 
especially thickened and apparently highly glandular beneath 


Text-fig. 193. 


A. Longitudinal section of the three glands in the newly born 
Hairy Armadillo (Huphractus villosus). 
Ep., epidermis ; D, glandular thickened dermis; O, orifice of pit. 


B. Pelvic carapace of the same showing the glandular orifices 
on the 4th, 5th and 6th belts. 


DORSAL GLANDS IN ARMADILLOS. 1101 


them, the topmost layer of the thickening being marked with a 
few spots of black pigment. The glandular nature of the pits 
was readily ascertained in the case of living adult armadillos of 


Text-fig. 194. 


Photograph of part of the pelvic carapace of an adult living Hairy Armadillo 
showing the three glands on the 4th, 5th, and 6th belts. (The gland-bearing 
scute of the 6th belt was fractured and the pits being filled with secretion 
showed up more clearly than when dry.) 


this species by squeezing the scutes bearing the scars and pressing 
from them drops of fluid with the same objectionable odour as 


Proc. Zoou, Soc.—1913, No. LX XIII. 73 


1102 DORSAL GLANDS IN ARMADILLOS. 


that which characterises these animals. Since the glands, judging 
from external appearances, are equally developed in adults of both 
sexes and appear at birth, there is no reason for thinking their 
function is in any way sexual. Their smell may be sufficiently 
nauseous to protect armadillos from enemies or it may act only 
as a guide enabling these animals to track each other and keep 
together. 


Text-fig. 195. 


Drawing of the central portion of the pelvic carapace of the Six-banded Armadillo 
(Euphractus sexcinctus) with four glandular pits, suggesting the origin of 
the pits from the enlargement and concrescence of the bristle-pores. 


‘‘ When the dorsal carapace of an armadillo is stripped, cleaned 
and dried, the position of the glands is marked on the under side 
of the pelvic shield by a row of bony prominences. Each pro- 
minence represents a gland and is in reality a hollow bony pit 
resulting from the ossification of the dermal layer immediately 
beneath the gland. This at all events is the case in a carapace of 
Euphractus sexcinctus kindly lent to me by Mr. R. EK. Holding. In 
this specimen there are seven free rings behind the scapular 
shield, but the seventh is not so free as those that precede it. 


THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 1103 


Following this are nine rings fused together and constituting the 
pelvic shield. The median glandular pits are observable upon the 
third, fourth, fifth and sixth of these ; and the scutes which carry 
them are distinguished by their shape from those of the rest of 
the pelvic shield. The glandular pit upon the sixth ring is much 
smaller and shallower than the rest and is especially interesting 
as showing the origin of the specialized glandular areas. The 
scutes of the carapace in this speciesare covered with pores, many 
of which are quite small; but a varying number, generally about 
ten and lying in two grooves, are much larger and arranged in two 
longitudinal lines meeting anteriorly and forming a somewhat 
horseshoe-shaped pattern. That these large pores perforate 
the bony scutes may be seen by holding the carapace against 
the light. 

‘“‘ According to de Meijere* these pores are bristle-pores and 
indicate lines of fusion between small and originally separated 
scutes. However that may be, there is no doubt, I think, that 
the glands above described result from the concentration and fusion 
of these rows of pores. The evidence for this is particularly strong 
in the case of the posterior of the four glands in Huphractus 
sexcinctus where the pores may be seen to be concentrated in the 
shallow central depression of the scute that bears them.” 


November 25, 1913. 


Prof, E. W. MacBrips, M.A., D.Sc., F.R.S., Vice-President, 
in the Chair. 


THe SEcrETARY read the following report on the additions 
made to the Society’s Menagerie during the month of October 
1913 :— 

The number of registered additions to the Society’s Menagerie 
during the month of October last was 214. Of these 104 were 
acquired by presentation, 42 by purchase, 16 were received on 
deposit, 38 in exchange, and 14 were born in the Gardens. 

The number of departures during the same period, by deaths 
and removals, was 237. 

’ Amongst the additions special attention may be directed to :— 

3 Entellus Langurs (Presbytis entellus), from North India, and 
1 Wanderoo Macaque (Macacus silenus), from Malabar, presented 
by W. O. Danckwerts, Esq., K.C., F.Z:S., on October 14th and 
24th. 

1 Sclater’s Dog (Canis sclateri), from the Amazons, purchased 
on October 2nd. 

1 Mexican Tree-Porcupine (Coendu mexicanus) and 1 Peba 


* De Meijere, Morphol. Jahrb. xxi. pp. 316, 318 (1894). 
C3" 


1104 MR. R. H. BURNE ON F@TAL SKELETONS. 


Armadillo (Tatusia novem-cincta), from Mexico, purchased on 
October 8th and 30th. 

1 Dorsal Hyrax (Dendrohyrax dorsalis), from Sierra Leone, 
presented by Mrs. A. M. Bettington on October 7th. 

1 Black-backed Hangnest (/cterws mesomelas), from Central 
America, | Loo-Choo Jay (Lalocitta lidthii), from the Loo-Choo 
Islands, and 1 Purple-breasted Parrot (Zriclaria cyanogastra), 
from 8.E. Brazil, all new to the Collection, received in exchange 
on October 27th. 

2 American Harriers (Circus hudsonius) and 1 Mississippi Kite 
(Icterus mississippiensis), from North America, both new to the 
Collection, received in exchange on October 7th. 

3 Pileated Guans (Penelope pileata), 3 Araucuan Guans (Ortalis 
araucuan), and 2 Rufous-necked Rails (Aramides chiricote), from 
Para, all new to the Collection, presented by the Goeldi Museum 
on October 20th. 

2 Brazilian Tortoises (Platemys platycephala), new to the 
Collection, purchased on October 15th. 


Prof. H. Maxweti Lerroy, M.A., F.Z.S., Honorory Curator of 
Insects, briefly described the arrangement of the New Insect 
House, the progress made with the fitting-up of the interior, and 
the installation of and method of displaying the exhibits. 


Mr. R. H. Burne, M.A., F.Z.S., showed some specimens of 
foetal skeletons prepared by the Beale-Schultze method. The 
fetus after being dehydrated is soaked in 1-3 per cent. caustic 
potash, and finally mounted in glycerine. By this procedure the 
soft parts become almost completely transparent, leaving any 
calcareous deposit opaque. 

The specimens shown were fetuses of the Fowl, Horse, and 
Man. In the Fow!] attention was drawn to the three separate 
metatarsals that at an older stage fuse to form the single tarso- 
metatarsus. In the same way in the Horse the ossifying shafts 
of the ulna, fibula, and splint-bones could be clearly distinguished. 

A specimen of the skeleton of a Star-fish (Astertas rubens) 
prepared by the same method was also exhibited. 


No. 124. 


ABSTRACT OF THE PROCEEDINGS 


OF THE 


ZOOLOGICAL SOCIETY OF LONDON.* 
October 28th, 1913. 


Prof. E, A, Miycuin, M.A., F,R.8., F.Z.S., Vice-President, 
in the Chair. 


The Minutes of the last Scientific Meeting were confirmed, 


The Sscretary read a Report on the Additions made to the 
Society's Menagerie during the months May to September, 
ONS: ; 


The Secretary exhibited a preparation of the alimentary tract 
of a young Elliot’s Pheasant (Phastanus elliott), dissected so as to 
show the gut-pattern in relation to the mesentery, blood-vessels, 
and visceral nerves. The preparation was mounted as a trans- 
parent object in glycerine jelly, and showed how the various 
morphological conditions of the intestinal tract to which he had 
called attention (Trans. Linn. Soc., Zool. vill. p, 178) might be 
displayed as permanent museum objects. 


Mr. R. I. Pocock, F.R.S., F.Z.8., Curator of Mammals, 
exhibited a female and three young specimens of the Woolly 
Opossum (Philander laniger), from Buenaventura in Colombia, 
presented by Dr. H. G. F. Spurrell, F.Z.S., and drew attention 
to the rapid growth of the young. When Dr, Spurrell cap- 
tured the mother, the little ones were still in the pouch. 
Six weeks afterwards, when they reached the Gardens, they 
had left the pouch for good, were about one-fourth grown, 
and were just beginning to feed on their own account, though 


* This Abstract is published by the Society at its offices, Zoological Gardens 
Regent’s Park, N.W., on the Tuesday following the date of Meeting to whieh 
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge, 
to all Fellows who subscribe to the Publications ; but it may be obtained on the 
day of publication at the price of Sixpence, or, if desired, sent post -free for 
the sum of Six Shillings per annum, payable in advance, 


38 


mainly dependent upon their mother’s milk. In another six 
weeks—that is to say, three months after capture—they were 
three-fourths their mother’s size, and apparently quite capable of 
fending for themselves, 


Mr. Pocock also exhibited two young examples of a Degu 
(Octodon degus), born from a pair from Valparaiso, presented by 
Mr. Walter Goodfellow, F.Z.S. The mother had a litter of four, 
but died from inability to give birth to a fifth. The little ones 
were fed by hand, and although two of them quickly died from 
pneumonia, the remaining two throve and promised to do well. 
Like the young of all Hystricomorphous rodents, the Degus are 
of large size at birth and open their eyes within twenty-four hours, 
and are in every respect very precocious as compared with young 
vats. Particular attention was directed to the lateral position of 
the teats in the Degu, a condition which is repeated in the Viscacha 
and Chinchilla and the Coypu. This condition is probably an 
adaptation to the large size of the young, which are enabled to 
suck lying alongside the mother, so that they do not interfere 
with each other. In the Coypu, as first described by Owen, the 
teats ave higher up the side than in the Chinchilla and the Degu, 
an arrangement which permits the young of that species, it has 
been alleged, to suck while swimming alongside their mother in 
the water. 


Mr. D. Seru-Smiru, F.Z.S., Curator of Birds, exhibited a 
spirit-specimen of a young Toucanet (Selenidera maculirostris) 
which had been hatched in the Society’s Gardens in July, and 
called special attention to the well-developed serrated pads on 
the back of the ankle-joint or heel. ‘This pad had been pre- 
viously observed in the nestlings of Woodpeckers, Wrynecks, 
and Barbets, and it was only to be expected that it would be 
present also in the Toucans. 

The use of this serrated pad, which disappears soon after the 
young bird leaves the nest, is doubtless to enable it to climb up 
the side of the hollow cavity in a tree in which it is hatched— 
these pads forming, as it were, a second set of claws. It is note- 
worthy that no such pads are found in the Parrots, which nest 
in similar situations to those chosen by Barbets and Toucans, but 
which are provided with a hooked beak which aids them in 
climbing. 


Mr. C. Tare Recan, M.A., F.Z.8., exhibited a specimen of 
Melanocetus johnsonii Giinther, a curious fish having an enormous 
mouth armed with slender, pointed, depressible teeth and an 
extraordinarily distensible stomach. It has the first ray of the 
spinous dorsal fin situated on the snout and modified into a 
line and bait, the latter being a luminous bulb, 


39 


Dr. F. E. Bepparp, M.A., F.R.S., F.Z.S., Prosector to the 
Society, read a paper on the anatomy and systematic arrangement 
of the Cestoidea, in which he described a new genus and species 
of Tapeworms from the Double-striped Thicknee (Wdicnenwus 
bistriatus). 


Dr. F. A. Baruer, M.A., F.R.S., F.Z.8., read a paper entitled 
“The Fossil Crinoids referred to Hy ypocrinus Beyrich.” The 
two specimens of Zypocrinus schneidert Beyr. described by Beyrich 
and Rothpletz respectively are redescribed and refigured. The 
structure of the genus is shown to agree with that of the Devonian 
family Gasterocomide, the content of which is discussed ; but it 
is suggested that in this case and in that of ‘ Lecythiocrinus” 
adamst the distinctive features may have been independently 
acquired. 

The holotype of Hypocrinus piriformis Rothpletz is redescribed 
and refigured, and proved to be no Hypocrinus. It is thought to 
be a highly modified descendant of the Taxocrinide, by way of 
such a genus as Cydonocrinus. The left posterior radial appears 
to have borne a large arm, but the other arms are more or less 
atrophied, and the right posterior radial has almost disappeared. 


A paper on “ Batrachiderpeton lineatum Hancock & Atthey, 
a Coal-Measure Stegocephalian,” communicated by Prof. J. P. 
Hill, D.Sc., F.R.S., F.Z.S., was read by Mr. D. M.S. Watson, 
M.Se. It contained the description of the skull, lower jaw, and 
pectoral girdle of this species, based on a series of specimens in 
the Newcastle Museum, derived from the Low Main Seam of 
Newsham Colliery. 


A paper, communicated by Dr. C. W. Andrews, F.R.S., F.Z.5., 
was received from Mr. R. W. Patmer, M.Sc., entitled ‘‘ The 
Brain and Brain-Case of a Fossil Ungulate of the Genus 
Anoplotherium,” in which a cranium from the Phosphorites 
of Quercy, together with an exceptionally perfect and well- 
marked brain-cast obtained from it, were described from material 
in the British Museum collections. 


40 


The next Meeting of the Society for Scientific Business will 
be held on Tuesday, November 11th, 1913, at half-past Hight 
o'clock p.u., when the following communications will be made :-— 


Exnrerrions AnD Noricrs. 
Sree seattle ctor een 


W. T. Cauman, D.Se.; F.Z.S. 


On Freshwater Decapod Crustacea (Families Potamonide 
and Palemonide) collected in Madagascar by the Hon. Paul 
A. Methuen. 


On a Collection of Reptiles and Batrachians made by 
Dr. H G. F. Spuvrell, F.Z.S., in the Colombian Choco. 
C. Tare Rea@an, M.A., F.Z.S. 
A Revision of the Cyprinodont Fishes of the Subfamily 
Poeciliinee. 
Prof. W. N. Parxer, Ph.D,, F.Z:8. 


Sponges in Waterworks. 


Prof. J. Puayrain McMurricu, C.M.Z.S. 


On Two new Actinians from the Coast of British Columbia. 


The following papers have been received :— 


SORTAN OLSEN. 
On the External Characters and Biology of Bryde’s Whale, 
a new Rorqual from the Coast of South Africa. 
Miss M. V. Lepour, M.Sc. 


A new Species of Trematodes of the Genus Lechriorehis from 
the Dark Green Snake (Zamenis gemonensis). 


T. AH. Wivuers, E.G. 


Ciripedes from the Cenomanian Chalk Marl of Cambridge. 


41 


F. F. Laipnaw, M.A., F.Z.S. 


Contributions to a Study of the Dragon-fly Fauna of Borneo. 
—Part II. The Gomphine and Chlorogomphine. 


Surgeon Josern C. Taompgon, U.S.N. 
Further Contributions to the Anatomy of the Ophidia, 
2 Cir eile) ilravotoneteg Mieke, Lees END US ZAI 


The Peroneal Muscles in Birds. 


Communications intended for the Scientific Meetings should 
be addressed to 
P. CHALMERS MITCHELL, 
Secretary. 
ZOOLOGICAL SoctEery oF Lonpon, 
REGENT’s Park, Lonpon, N.W. 
November Ath, 1913. 


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No. 125. 


ABSTRACT OF THE PROCEEDINGS ~ 


OF THE 


ZOOLOGICAL SOCIETY OF LONDON.* 


November 11th, 1913. 


Dr. 8. F. Harmer, M.A., F.R.S., F.Z.S., in the Chair, 


The Minutes of the last Scientific Meeting were confirmed. 


Mr. R. I. Pococr, F.R.S., F.Z.S., Curator of Mammals, ex- 
hibited a newly-born example of the Hairy Armadillo (Luphractus 
villosus) and the dried shell of a Six-banded Armadillo (Z. sea- 
cinctus), Showing the presence of a series of from two to four 
small orifices in the middle line of the pelvic shield; and explained 
that the orifices are the apertures of shallow invaginations of the 
epidermis dipping down into the dermis, which is specially 
thickened and glandular immediately beneath them, In the 
living animal drops of malodorous fluid can be squeezed from the 
orifices of these glands. 


Mr. D. Seru-Smitu, F.Z.S., Curator of Birds, exhibited a re- 
markable hybrid between a Black-winged Peacock and a domestic 
_hen (cross-bred game and Leghorn) which had been bred by 

Mr. R. P. Wheadon of Ilminster in 1911. This bird was be- 
lieved to be only the second example of this hybrid ever produced, 
although a hybrid between a peafowl and a guinea-fowl had been 
recorded (Hocker, J. f. O. 1870, p. 152). 

The hybrid was mostly white in colour, splashed, especially on 
the neck, with reddish brown, a colour derived from the domestic 
hen, its mother. It showed clearly the characters of both 
parents. 

Mr. Seth-Smith also showed photographs of a hybrid Pheasant 
between Calophasis mikado and C’ elliott, and of a Jungle-fowl 
between Gallus varius and G, gallus. The latter bird, which had 
been bred in the Society’s Gardens, corresponded to the description 
of the bird described by Temminck as Gallus eneus. 


* This Abstract is published by the Society at its offices, Zoological Gardens, 
Regent's Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge, 
to all Fellows who subscribe to the Publications; but it may be obtained on the 
day of publication at the price of Sixpence, or, if desired, sent post-free for 
the sum of Six Shillings per annum, payable in advance, 


Ad 


Mr. E. G. Bounencer, F.Z.8., Curator of Reptiles, exhibited 
a quite young specimen of the Matamata Terrapin (Chelys jim- 
briata), the shell of which measured but 5 cm. in length. This 
specimen, the smallest yet recorded, differed from the existing 
descriptions of the young of this species in the under surfaces 
being of a bright magenta-red instead of pale yellow. Mr. Bou- 
lenger suggested that as all the young previously described were 
more than double the size of the specimen exhibited, red might 
be the normal coloration of the under surfaces as a juvenile 
livery. 


Dr. W. T. Catman, F.Z.S., read a paper on Freshwater Decapod 
Crustacea (families Potamonide and Palemonide) collected in 
Madagascar by the Hon. Paul A. Methuen, in which one new 
species of Potamon and five varietal forms of P. madagascariense 
were described. It is suggested that the River-Crabs of Mada- 
gascar may have had an autochthonous origin from some form 
resembling P, madagascariense. No clear affinities can be traced 
with the Potamonide of Africa or of Peninsular India, but it is 
pointed out that in the present state of knowledge the River- 
Crabs appear to be a hazardous subject for zoogeographical 
speculation. 

Among the Palemonide the most interesting is the species 
’ described by Hilgendorf as “‘ Bithynis ? hildebrandiz,” which is 
figured for the first time. Evidence is brought forward to show 
that the species has no close affinity with the Bithynis gaudichaudi 
of Chili and Peru, and that the single character which led to its 
being referred to Lithynis is a matter of individual variation. 


Mr. G. A. Boutenegsr, F.R.S., F.Z.8., gave an account of a 
collection of Reptiles and Batrachians made by Dr. Spurrell, 
F.Z.8., in the Colombian Choco and presented by him to the 
British Museum. The series of specimens was of great interest, 
and contained several new species. 


Mr. C, Tarr Rucan, M,A., F.Z.S., read a paper entitled 
“A Revision of the Cyprinodont Fishes of the Subfamily 
Peeciliinz.” A number of new genera were defined and several 
new species were described ; the structure of the intromittent 
organ was found to be of great systematic importance, 


A paper received from Prof. W. N. Parker, Ph.D., F.Z8., 
gave an account of some investigations made on a growth of 
Spongilla lacustris in the Cardiff Waterworks system, and described 
the methods adopted to eradicate the sponge from the infected 
areas, 


45 


Prof. J. Puayrarr McMurricu, C.M.Z.S., communicated a 
paper in which he described two new species of Actinians from 
the coast of British Columbia. These specimens probably repre- 
sented stages of a single species, and belonged to a group hitherto 
not recorded from the west coast of America. 


The next Meeting of the Society for Scientific Business will 
be held on Tuesday, November 25th, 1913, at half-past Eight 
o’clock P.m., when the following Conmcniic ations will be made :— 


EXHIBITIONS AND NOTICES. 


RIAN OLSEN. 
On the External Characters and Biology of Bryde’ s Whale, 
a new Rorqual from the Coast of South Africa. 
Miss M. V. Lesour, M.Sc. 


A new Species of Trematodes of the Genus Lechriorchis from 
the Dark Green Snake (Zamenis gemonensis). 


T. H. Wrruers, F.G.S8. 
Cirripedes from the Cenomanian Chalk Marl of Cambridge 


P. Cu CHALMERS Mircnent, M.A. IDs 1h LL.D., emia R.S., PAS. 
ernie Per oneal Maneales = in ae 


The following papers have been received :— 


FE. F. Larpuaw, M.A., F.Z.S. 


Contributions to a Study of the Dragon-fly Fauna of Borneo. 
—Part II. The Gomphine and Chlorogomphine. 


Surgeon JosrpH C. THompson, US.N 


Further Connbutione to ey ree of the Ophidia. 


46 


EU Ue eee 
Sen on an Bieri dovolaped Specimen of Hehinus 
esculentus. 


Gre EN BouLENGER, E.R.S., B.ZS. 


‘An Real ist ee the Reptiles and Batrachians collected 
by the British Ornithologists’ Union Expedition and the 
Wollaston Expedition in Dutch New Guinea. 


C.F. U. Mzex, MSc. LS. F.ZS8. 


“The Possible Comecaon between Spindle-length and Cell- 
volume. 


W ILLIAM Nicout, M.A., 7D) Sc., ., M -D., 8 ZS. 


Ty eines peed Ba om Gittins dying i in the Zoological 
Society’s Gardens during 1911-1912. 


mic ae 7 ce ae a oe Arrangement 
of the Cestoidea.— XII. Further Observations upon the! Genus 
Urocystidium Beddard. 


Communications intended for the Scientific Meetings shoul 
be addressed to 
P. CHALMERS MITCHELL, 
Secretary. 
ZooLoGIcAL Society oF Lonpon, 
Recent’s Park, Lonpoy, N.W. 
November 18th, 1913. 


No. 126. 


ABSTRACT OF THE PROCEEDINGS 


OF THE 


ZOOLOGICAL SOCIETY OF LONDON.* 
November 25th, 1913. 


Prof. E. W. MacBripe, M.A., D.Sc., F.R.S., Vice-President, 
in the Chair. 


The Minutes of the last Scientific Meeting were confirmed. 


The Spcrerary read a Report on the Additions made to the 
Society’s Menagerie during the month of October, 1913. 


Mr. R. H. Burne, M.A., F.Z.S., showed some specimens of feetal 
skeletons prepared by the Beale-Schultze method. The fetus 
after being dehydrated is soaked in 1-3 per cent. caustic potash 
and finally mounted in glycerine. By this procedure the soft 
parts become almost completely transparent, leaving any calcareous 
deposit opaque. 

The specimens shown were feetuses of the Fowl, Horse, and 
Man. In the Fowl attention was drawn to the three separate 
metatarsals that at an older stage fuse to form the single tarso- 
metatarsus. In the same way in the Horse, the ossifying shafts 
of the ulna, fibula, and splint-bones could be clearly distinguished. 

A specimen of the skeleton of a Starfish (Aséerias rubens), 
prepared by the same method, was also exhibited. 


Prof. H. Maxwetu Lerroy, F.Z.8., Hon. Curator of the Insect 
House, briefly described the arrangement of this new building, 
the progress made with the fitting-up of the interior, and the 
installation of and method of displaying the exhibits. 


* This Abstract is published by the Society at its offices, Zoological Gardens, 
Regent’s Park, N.W., on the Tuesday following the date of Meeting to which 
it refers. It will be issued, along with the ‘ Proceedings,’ free of extra charge, 
to all Fellows who subscribe to the Publications ; but it may be obtained on the 
day of publication at the price of Srpence, or, if desired, sent post-free for 
the sum of Six Shillings per annum, payable in advance. 


AS 


A paper received from Mr. Orsan OLseN, and communicated 
by the Secretary, contained the description of a new Rorqual from 
the Coast of South Africa and gave a detailed account of its 
external characters, biology, and distribution. 


Miss Marie V. Lrsour, M.Sc., presented a paper, communi- 
cated by the Secretary, in which she described a new species of 
Trematodes of the genus Lechriorchis, found in the body-cavity 
of a Dark Green Snake (Zamenis gemonensis) that had died in 
the Society’s Gardens. 


Mr. T. H. Wirners, F.G.S., contributed a paper, communi- 
cated by Dr. W. T. Calman, F.Z.8., based upon a large series of 
Cirripede remains from the Cenomanian Chalk Marl in the neigh- 
bourhood of Cambridge. The greater number of the specimens 
are referred to two species of the family Pollicipedide, and add 
materially to our knowledge of the phylogeny of the pedunculated 
Cirvipedes. Both forms are remarkable for their advanced form 
of seutum, in which the umbo is sub-central, and show that the 
transition of the scutal umbo from an apical to a sub-central 
position was acquired independently by unrelated forms in distinct 
lines of development. 


The Srcrerary, Dr. P. Coatmers Mircueti, M.A., F.R.S., 
F.Z.8., read a communication on ‘‘The Peroneal Muscles in 
Birds.” Dr. Mitchell had dissected these muscles in over 300 
birds, and believed that he was able to give a nearly exhaustive 
account of the varieties of form presented by these structures. 
The paper described the peroneal muscles in Chawna chavaria, 
and gave a systematic account of the conditions in the different 
Avian groups which could all be represented as derivatives of 
the Chauna condition by loss of certain portions and increased 
development of other portions. It was diiticult to interpret the 
changes in terms of function, family-relationship appearing to be 
more important in this respect than habit. The paper discussed 
the very few cases of want of conformity between the peroneal 

nuscles and the place assigned in the best-founded systems of 
classification. It also dealt with the use of such anatomical 
characters in systematic classification, and showed that there was 
a definite relation between specialized conditions of the peroneal 
muscles and specialized conditions of the Avian structure 


generally. 


49 


The next Meeting of the Society for Scientific Business will 
be held on Tuesday, February 3rd, 1914, at half-past Hight 
o'clock p.m., when the following communications will be made :— 


EXHIBITIONS AND Noricnus. 


G. A. Boutencsr, F.R.S., F.Z.8. 


An Annotated List of the Reptiles and Batrachians collected 
by the British Ornithologists’ Union Expedition and the 
Wollaston Expedition in Dutch New Guinea. 


Se ae ae cae aa ea Arrangement 
of the Cestoidea.—XII. Further Observations upon the Genus 
Urocystidium Beddard. 


H.G. Prmver, F.R.S., P.Z.5 


ae mi a a Se occurred in the Zoological 
Gardens during 1913. 


The following papers have been received :— 
FF, Larpraw, MA. F.ZS8. — 


Contributions to a Study of the Dragon-fly Fauna of Borneo. 
—Part II. The Gomphine and Chlorogomphine. 


Surgeon Josep C. Tuompson, U.S.N. 


Further Contributions to the Anatomy of the Ophidia. 


H. C. Cuapwick, A.L S. 


Note on an imperfectly - developed Specimen of Hehinus 
esculentus. 


CT Ue Mipies Wleee UNly ein eats. 


Tike a eeae Ge peoween! Spindle- -length and Cell- 
volume. 


Wiiuram Nicotu, M.A., D.Sc., M.D., F.Z.8. 
Bie eae ele 
Tyematode Parasites from Animals dying in the Zoological 
Society’s Gardens during 1911-1912. 


50 
F. J. Mraerrt, M.Se 


The Structure and Life-History of a Tapeworm (Lchthyotenia 
Jilicollis Rud.) parasitic in the Stickleback. 


Communications intended for the Scientific Meetings should 
be addressed to 


P. CHALMERS MITCHELL, 


Secretary. 
ZOOLOGICAL Society or Lonpon, 
Rueent’s Parx, Lonpon, N.W. 
December 2nd, 1913. 


ol. 


53. 


A New rematode of the Genus Lechriorchis from the Dark Green Snake (Zamenis 
gemonensis). By Marin V. Lupour, M.Sc., Assistant Lecturer and Demonstrator in 
Zaolory, Weeds University. (Pl MCLE) site ge te ccs See eee ecw nese rs 
54, Cirripedes from the Cenomanian Chalk Marl of Cambridge. By Tuomas H. Wirusrs, 
BE erses eee (Chal NCO ENV ee RG NCO) Vis) avs etc ictal arene talon «is ciaaee“e'lalwns sista sistaiphers ott oletpiera'e ed 
55, Batrachiderpeton lineatum Hancock & Atthey, a Coal-Measure Stegocephalian. By 
D. M. 8S. Warson, M.Sc., Lecturer on Vertebrate Palaontology in University College, 
Hondon. (Pls. XCVI. & XCVII. and Text-figs. 162-107.) ...... 2.0.02 ...s cece 
56, On two New Actinians from the Coast of British Columbia. By J. Pusyrair 
IMeniCorpnoe Oni Aish = Gab ONALRE icine ad dade an onScMeonooConobocenboOooc Onn 
57. Sponges in Waterworks. By W. N. Parxer, Ph.D., F.Z.8., Professor of Zoology, 
Unrweneiiy- Collage, (Chnthtttcouscoccaocndovduscser on duos acennde syslesaranie whose tenets 
58. A Revision of the Cyprinodont Fishes of the Subfamily Peciliine By C. Tats 
Ruean, M.A., F.Z.S. (Pls. XCIX.-OI. and Text-figs. 168-173.) ........++-.00-00s 
59. On a Collection of Batrachians and Reptiles made by Dr. H. G. F. Spurrell, F.Z.S., in 
the Choco, Colombia. By G. A. Bouruyerr, F.KS., F.ZS8. (Pls. C1L-CVII. 
SUMTER POR ali AiG 5) aie we eicce eee n cinch, oo ale olniote apices 6 a's wha cos siee siete aig rich 
40. The Peroneal Muscles in Birds. By P. Cuaumrers Mircng.y, M.A., D.&c., LL.D., 
F.RBS., F.Z.S8., Secretary to the Society. (Text-figs. 179-190.) ..........e-sseeee 
61. On the External Characters and Biology of Bryde’s Whale (Balenopitera brydei), a 
new Rorqual from the Coast of South Africa. By Mrsan Oxsuy, Zoological 
Laboratory, Christiania University. (Pls. CIX.-CXIIL.).......... 000s. ereens 
CMGI AED Sagiaqoks orien acne aS onan uonciintipn eaten Ae seldtacio Seis Emery rer Ame eo 
Mast OMe OUNCME MELO LNCETS: aan, <ioh eval of crs ciescie s!«)-ele/oidcatlei-oinim sl cleats vin moles ic aie ln'e Gorka : 
brats OF ClonesiiS so on cu dbconn ange 6000 dene UOEOULhe Doub an OO CUMeD SD UOOeES Co Dn de Soe 
Alphabetical List of Contributors .......... -.. ab cis dg OAR Odo SHO AGO Cue soo Lo oon 
EN@wa Generic Donn sisyseicrteec sera) e oteinc toss sicreatiar a toile: « Bia ciel eb eared iol ovate steislaraieicle: shelsiaeie 
iaglae @E SieianinitG INEINES Ses hoo con ogc don dd 50 KOUn Guat Op oo ues HAO sopoodod oo : 
Biirrtve aro ce elas teen proms Gers s: lace scaoe vei olstoe a ereitiare eicletaieiiar ct aneketauas suatsravansavn elapse eilelencvate lesson tonslinne 


Papers (continued). 


The Fossil Crinoids referred to Hypocrinus Beyrich. By F. A. Barnur, M.A., D.£c., 
bors Hee See (Eee sand: NeXtn eS OOM.) cei e's ie wore civisis eaimie wie cele oa als 


. On Freshwater Decapod Crustacea (Families Potamonids and Palemonidz) collected 


in Madagascar by the Hon. Paul A. Methuen. By W. T. Cauman, D.8c., F.Z.8. 
Gi seeNUC Mero oN Cll eancl Metin trom slallly Wractenyoleletorti ere c's e vies siertierace ols e10) ole e/ekelecinmis ols 


Pago 


894 


914 


933 


937 


949 


963 


973 


977 


1019 


1039 


1073 


LIST? OF PLATES, 
1913, Part LV. (pp. 861-1104). 


Plate Page 
XC. 1-6. Hypocrinus schneideri. oe “FL? purcfornst,.ea82 & Oot 
XOI. Potamon methuent....... tro tttntereeetereeeeereee | gry 
XOIL -Palemon hildebrandtii 22, } 
ICME Lechrvonchis weniiisiteteetateyeia ciebevelaliate)ereraiele sipensten tote eee 933 
KCEV.. -Zengmatolepas mocklert wea. +» scls.)ea es oc ote wee ste aera “937° 
XCV. Calantica (Titanolepas) tuberculata ....++....<+.-- Fiete es \ 
Se } BORO ELIGAS OPRT D.C 50 HAG FO DOO iC oie.) Adee 
XOVIII. 1-4. Peachia quinquecapitata. 5-7. Bicidiwm equoree. 
SABA panasicicwmerawere «ati oe ole slo eee oreo 963. 
XOIX. 1,2. Gambusia oligosticta., 3,4. G. wrayi. 5,6. G. gra-\ 
eoliors. “x (Gr, GhOMTNGHUSES * SaobOS oc anne accocnand0s8 os | 
C. 1, 2. Pecilia picta. 3, 4. Peciliopsis isthmensis. 5, 6. | 977 
Phalloceros caudomaculatus. 7, 8. Phalloptychus januarius. f 
CI. 1,2. Limia nigrofasciata. 3, 4. L. heterandria. 5. L. | 
arnoldi. 6. L. caudofasciata. 7. L. ornata............/) 
CII. 1. Bufo hypomelas. 2. Agalychnis calcarifer ........+++» yl 
CIO Agalychnis\spur rena nacelle aha idler tainty = = ateleietcla hina | 
CIV. 1. Dendrobates awrotenia. 2-4. D. tinctorius, var, coctéi. 
5, 6. D. tinctorius, var. Chocensts .s..-...-seereecees 
ean \ OiNOstenmartySPUnnellt Wa carere teietelegsietetntiets etek ee ee eee riOle 
CVIL. 1. Anolis breviceps. 2. Anadia vittata ......06.02..00005 
CVIII. 1. Homalocranium coralliventre. 2. Hlaps microps. 
3. Leptognathus spurrellt .. i. -0secess veers sccccs ose 
CIX. ) 
@X. | Balenoptera brydet 72... 2. ee ce www ces ce ce AMODOS O08 
CXI. | 1073: 
CXII.. 9, 11a. Balenoptera borealis. 10,110. B. brydet ........ | 
(Ch:GUU ny IRA AMMAR OVO» argosuotseAaontobs obodcb on casmqecce y) 
NOTICE. 


The ‘ Proceedings’ for the year are issued in four parts, paged consecutively, 
so that the complete reference is now P. Z. 8. 1913, P- ... The Distribution 


is as follows:— 


Part I. seared in March. 
pie ld Remains June. 
pry EB - September. 
a SEL ee December. 


‘Proceedings,’ 1913, Part III. (pp. 3389-860), were published on 


September Ist, 1913. 


The Abstracts of the ‘Proceedings,’ Nos. 124-126, are 


contained in this Part. 


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