1 Pe he

OF THE

COUNCIL AND OFFICERS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

ESO

COUNCIL.

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

Tue Earu or Atramont, F.S.A. | Srr Epmunp G. Loprr, Bt.

Grorce A. Bou.encer, Esq., | E. G. B. MrEApE-Watpo, Esq.

? » HSQ.,

E.R.S., Vice-President. Vice-President.

Ricoarp H. Burne, Esq., | Prof. Epwarp A. Muncu1n,

M.A. | M.A., Vice-President.

Lt.-Col. Srr R. Havenock- | P. Coatmers MircHer., Esq.,

Cuartes, K.C.V.0.,.M.D. | M.A., D.Sc. Hon. LL.D.,

Aurrep H. Cocks, Esq., M.A. | F.R.S., Secretary.

F. D. Dawrrey Drewirt, Esq., | ALBERT Pam, Esq.

M.A., M.D. | ApriAN D. W. Poxttock, Esa.

CHarLtes Drummonn, KsqQ., | OLDFIELD THOMAS, Ksa.,

Treasurer. F.R.S.

Sir Epwarp Dvuranp, Bt., | A. Trevor-Batrryr, Esq., M.A.

C.B. A. Smrira# Woopwarp, Esq.,

Frepverick GILLerr, Esq. LL.D.,F.R.S., Vice-President.

Srpney F. Harmer, Esq., M.A., | Henry Woopwarp,Esq., LL.D.,

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

PRINCIPAL OFFICERS.

P. Cuaumers Mitrcuett, M.A., D.Sc., Hon,.LL.D., F.R.S.,

Secretary.

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

R. 1. Pocock, F.L.S., Curator of Mammals and Reptiles, and

Resident Superintendent of the Gardens.

D. Seru-Smivu, Curator of Birds and Inspector of Works.

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

F. H. Wareruouse, Librarian.

Jousx Barrow, Accountant,

W. H. Coxe, Ch jef Clerk,

LIST OF CONTENTS,

1910, pp. 589-1033.

Js\jorenll Gs, USING)

Page

The Secretary. Exhibition of a series of photographs taken

in the South Orkneys in 1908 by a correspondent of

SU nie: NUCH GIS Shas bated ore Rohan ce Ce en ee oR 589

Mr. R. I. Pocock, F.L.S., F.Z.8. Exhibition of a living

specimen of a Mole-Rat (Georychus) from Ashanti ...... 589

1. The Caudal Fin of the Teleostomi. By Ricuarp H.

Wuitenovuse, M.Sc., University of Birmingham. (Plates

SRO WVE TMS) | cactoto tates eRnete stoke kai waeeain ace tees soled otites 590

2. Some Notes on Tasmanian Frogs. By T. M. Savacr

UnGiarsta sy (Enlaiber lilly) Sane cece nae eetns s asisn cluster adnate 627

April 19, 1910.

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

Menagerie during the month of March 1910 ............ 634.

Mr. Oldfield Thomas, F.R.S., F.Z.S. | Exhibition and de-

scription of a new Monkey from the Malay Peninsula... 634

Mr. Oldfield Thomas, F.R.S., F.Z.S. Exhibition of a Col-

lection of small Mammals from China, with descriptions

GAN OM PEOEMISt RRR NS enacts seve es ee aNNenK ou dewssniseeeewwes ee

Mr. J. Lewis Bonhote, M.A., F.L.S., F.Z.S. Exhibition of

a yellow variety Of Moris 701008 ......s.scencccverenecnerensns

The Secretary on the Centenary of Philip Henry Gosse,

ERE Senne pea ceenece? | Vays os Senne ene cuneate set an

1. Notes on the Photophores of Decapod Crustacea. By

Srantey Kemp, B.A. (Plates LIT.—LIV.) ..............-

2. On the Varieties of Mus rattus in Egypt; with General

Notes on the Species having reference to Variation and

Heredity. By J. Lewis Bonuorn, M.A., F.LS., F.Z.5.

3. On an example of Posterior Dichotomy in an Aylesbury

Muekdine. (By Gi ABUUuEN, 2. ee eens anne

May 3, 1910.

The Secretary. Exhibition of a photograph of a Giraffe

from:Gallaland. Mast: Atrica, \. o<....5...:seeesn sae eens

Mr. E. E. Austen, F.Z.8. Exhibition of a specimen of a

rare Fly (Cephenomyia ulrichiti Brauer) from East

POURSLa dies oo eae v caine) wc biseas ne Abneinw Vale pene en Ee yee eee RIE

Dr. R. A. O’Brien. Remarks on the habits, illustrated by

lantern-slides, of the North Queensland Green Tree-

Ant (Occophylla smaragdina 2) .......ccscceorsereecssccsaees

Miss Annie Porter and Dr. H. B. Fantham, F.Z.S. Ona

n fa - ‘yt Y .

new Trypanoplasm, 7'rypanoplasma (Cryptobia) dendro-

celi, sp. n., from Dendrocaluim lactewm ..... .ccccceeeeeeeee

1. The Morphology and Life-History of Himeria (Coccidiwm)

avium: » Sporozodn causing a fatal disease among young

Grouse. By H. B. FanruAm, D.Sc. Lond., B.A. Cantab.,

A.R.C.S., F.Z.8., Christ’s College, Cambridge, Assistant

to the Quick Professor of Biology, Cambridge, and

Protozoologist to the Grouse Disease Inquiry. (Plates

AG Ges Ogi bi Re Ao Pattie rete ery etre et toe trie:

Page

639

651

666

669

670

Observations on the Parasitic Protozoa of the Red Grouse

(Lagopus scoticus), with a Note on the Grouse Fly.

By H. B. Fanrnam, D.Sc. Lond., B.A. Cantab., A.R.G.S.,

F.Z.8., Christ’s College, Cambridge, Assistant to the

Quick Professor of Biology, Cambridge, and Proto-

zoologist to the Grouse Disease Inquiry. (Plates

MAE OTA Ae ENS, LU, A Ss Sanat,

. Experimental Studies on Avian Coccidiosis, especially in

relation to young Grouse, Fowls, and Pigeons. By

H. B. Fanrnam, D.Sc. Lond., B.A. Cantab., A.R.CS.,

F.Z.8., Christ’s College, Cambridge, Assistant to the

Quick Professor of Biology in the University of Cam-

bridge; Protozoologist to the Grouse Disease Inquiry.

(OPIEY EULA IN) agate ae ait ea are nes a oa A

. Observations on the Blood of Grouse. By H. B. FanrHam,

D.Sc. Lond., B.A. Cantab., A.R.C.S., F.Z.S., Christ’s

College, Cambridge, Assistant to the Quick Professor

of Biology in the University; Protozoologist to the

Grouse Disease Inquiry. (Plate LXIII.) ...............

. Zoological Results of the Third Tanganyika Expedition,

conducted by Dr. W. A. Cunnington, F.Z.8., 1904—

1905.—Report on the Ostracoda. By Prof. G. O. Sars,

CAME Sam eaves XSW VES ONCRTIT NS oe. Es iio eee aeeen-

. On Tritylodon, and on the Relationships of the Multi-

tuberculata. By R. Broom, M.D., D.Sc., C.M.Z.S.

May 24, 1910.

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

Menagerie during the month of April 1910 ...............

My. Ernest Gibson, F.Z.8S. Exhibition of a skin of Felis

IPE OI MOO MER Chae -cot icc «stem sen melee «ote vamededte ss

1. Observations on the Anatomy and General Biology of

some Members of the Larger Cetacea. By D. G. Litiir,

B.A., Hutchinson Research Student of St. John’s Col-

lege, Cambridge. (Plate LX XIV.) ...........:0sceseeeeee

692

708

wt)

768

769

. Zoological Results of the Third Tanganyika Expedition,

conducted by Dr. W. A. Cunnington, F.Z.8., 1904—

1905.— Report on the Rotifera. By Cuaries F.

RovssEvET, F.R.M.S. (Plate LXAXYV.)...............ce00

3. The Marine Fauna of the Mergui Archipelago, Lower

Burma, collected by Jas. J. Simpson, M.A., B.Se., and

R. N. Rudmose-Brown, D.Se., University of Aberdeen,

February to May 1907.—The Hyproips. By James

Rircuiz, M.A., B.Sc., Natural History Department,

The Royal Scottish Museum. (Plates LXXVI. &

1 001 WD nner Pee pan Sear Ac s)c/cot.cb Go onerar eee enee

ise

. Hydroids from Christmas Island, Indian Ocean, collected

by C. W. Andrews, D.Sc., F.R.S., F.Z.S., in 1908.

By James Rircnie, M.A., B.Sc., Natural History

Department, The Royal Scottish Museum ...............

June 14, 1910.

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

Menagerie during the month of May 1910 ...............

Mr R. I. Pocock, F.LS., F.Z.8. Remarks upon, and

exhibition of two drawings illustrating, an instance

of ant-mimicry by the larve of a species of Mantis;

with a Note on the mimicry of the larva of the

Ceylonese Leaf-Insect (Phylliwm sp. 2) .......:seseereeeees

1. On the Specialised Cutaneous Glands of Ruminants. By

R. I. Pocock, F.LS., F.Z.S., Superintendent of the

Zoological Society’s Gardens <5 .-castecereneemeamemeree nese

2. On a Wapiti and a Muntjac. By R. LypEKKER............

3. On Three African Buffaloes. By R. LYDEKKER............

4, Ona new Antelope and on the Spanish Chamois. By

Prof. A. CaBrEerA, C.M.Z.S.

rrr ree eee ee ee

. The Changes of Plumage in the Red Grouse (Lagopus

scoticus Lath.) in Health and in Disease. By Epwarp

A. Wutson, M.B., F.Z.S., Field-Observer to the Grouse

Page

792

799

837

Disease Inquiry. (Plates LXXVIII.-CL) ............ 1000

Wipe AT List

CONTRIBUTORS,

With References to the several Articles contributed by each.

(1910, pp. 589-1033.)

Page

Austen, Ernest E., F.Z.8.

Exhibition of a specimen of a rare Fly (Cephenomyia

nirichwy braver) trom Wash PLUSSIa, .2 6... .0.-.2-..s00 anne oo 669

Bonuore, J. Lewis, M.A., F.L.S., F.Z.8.

Exhibition of a yellow variety of Mus rattus ............ 638

On the Varieties of Mus rattus in Egypt; with

General Notes on the Species having reference to

Ferretti Loran ea GMO CULV race wis a/0/si >i aldslesiniecic’s ona rasersees ies 651

Broom, Rospert, M.D., D.Sc., C.M.Z.S.

On Tritylodon, and on the Relationships of the

Multituberculata

PCC CC

vill

Page

BULLEN, G. E.

Onan example of Posterior Dichotomy in an Aylesbury

DCM a se aes a's vine ws dus 0 otehne ye Coleen eR ERM ene giay's as a ce eetee 666

Caprera, Prof. ANGEL, C.M.Z.S.

On a new Antelope and on the Spanish Chamois ...... 998

Enautsa, T. M. SAvace.

Some Notes on Tasmanian Frogs. (Plate LI.) ......... 627

Fanruam, Haron B., D.Se. Lond., B.A. Cantab., A.R.CS.,

F.Z.S., Christ’s College, Cambridge, Assistant to

the Quick Professor of Biology, Cambridge, and

Protozoologist to the Grouse Disease Inquiry.

The Morphology and Life-History of Himeria (Cocci-

dium) avium: a Sporozoén causing a fatal disease among

young Grouse. (Plates UuV—-LVIII.).. 0... ee cecseneses: 672

Observations on the Parasitic Protozoa of the Red

Grouse (Lagopus scoticus), with a Note on the Grouse-

Bly... 4(Blaites TRC ST a) ea ssens so anne cnn ieee eaters 692

Experimental Studies on Avian Coccidiosis, especially

in relation to young Grouse, Fowls, and Pigeons. (Plate

GX) ae BF nica spice vc + pie 08 See Le ee eee ental 708

Observations on the Blood of Grouse. (Plate LXII!.). 722

Fantruam, Dr. HArowp B., B.A., F.Z.S,, and Portrer, Miss

ANNIE, B.Sc,

On a new Trypanoplasm, Z'rypanoplasma (Cryptobia)

dendroceli, sp. n., from Dendrocalwm lactewm .......+60+. 670

Gipson, Ernest, F.Z.8.

Exhibition of a skin of Pelis geoffvoyt ccscccceccereeeneenees 769

Page

Kemp, STANLEY, B.A.

Notes on the Photophores of Decapod Crustacea.

(Celene JUIN LAI) Wacontpdncebor spoenacdacnocaaeccoceponeBerne 639

Linuiz, D. G., B.A., Hutchinson Research Student of

St. John’s College, Cambridge.

Observations on the Anatomy and General Biology of

some Members of the Larger Cetacea. (Plate LXXIV.). 769

LyDEKKER, RicHARD.

OnvamWepibieandtay MUMGIACr.. 2... tests cecncses ees once 987

OnglihreceAtieicam) DBUilaloes) «jis seas secceans<vheseaaecrces- 992

MircHett, P. Coatmers, M.A., D.Sc., Hon. LL.D., F.R.S.,

F.Z.8., Secretary to the Society.

Exhibition of a series of photographs taken in the

South Orkneys in 1908 by a correspondent of ‘The

gINWn Comet aceasta atch auc aciiddiie di lecue'saetaenocwiar 589

Report on the Additions to the Society’s Menagerie

@urine the month of March) 1910)... ..scccenc..e0.scenesoeee see 634

On the Centenary of Philip Henry Gosse, F.R.S....... 639

Exhibition of a photograph of a Giraffe from Gallaland,

JORYSth JAMIAIGE), ce.5. coats pb aneobodda cob gee OHO Ber: oda SeE BU See SoocLo Seer 669

Report on the Additions to the Society’s Menagerie

during the month of April 1910 .......... cesses eee eeee ee ees 768

Report on the Additions to the Society's Menagerie

during the month of May 1910............ceceeeeeee eens eeeeee 837

O’Brien, Dr. R. A.

Remarks on the habits, illustrated by lantern-slides,

of the North Queensland Green Tree-Ant (Ocecophylla

SMANGYUUNG 2) crccvevcrccverscnsenenserseenereeneereaeenseeen see ees 669

Pocock, Reerxaup I., F.L.S., F.Z.8., Curator of Mammals

and Reptiles, and Resident Superintendent of the

Gardens.

Exhibition of a living specimen of a Mole-Rat (G@eo-

rychus) from Ashanti

ee ee ry

Remarks upon, and exhibition of two drawings illus-

trating, an instance of ant-mimicry by the larve of a

species of Mantis; with a Note on the mimicry of the

larva of the Ceylonese Leaf-Insect (Phyllium sp. 2)

On the Specialised Cutaneous Glands of Ruminants ...

Porrer, Miss ANNIE, B.Se., and Fanruam, Dr. HAroup B.,

B.A., E.Z.8.

On a new Trypanoplasm, Zrypanoplasma (Cryptobia)

dendroceli, sp. n., from Dendrocelum lactewm

Rircuie, James, M.A., B.Sc., Natural History Department,

The Royal Scottish Museum.

The Marine Fauna of the Mergui Archipelago, Lower

Burma, collected by Jas. J. Simpson, M.A., B.Se., and

R. N. Rudmose-Brown, D.Sc., University of Aberdeen,

February to May 1907.—The Hypromws. (Plates

LXXVI. & LXXVII.)

Hydroids from Christmas Island, Indian Ocean, col-

lected by C. W. Andrews, D.Sc., F.R.S., F.Z.S., in 1908.

RovusseEvet, CHarves F., F.R.M.S.

Zoological Results of the Third Tanganyika Expe-

dition, conducted by Dr. W. A, Cunnington, F.Z.8.,

1904-1905.—Report on the Rotifera. (Plate LXXY.).

Page

589

837

840

Sars, Prof. G. O., C.M.Z.S.

Zoological Results of the Third Tanganyika Expe-

dition, conducted by Dr. W. A. Cunnington, F.Z.S.,

1904—-1905.—Report on the Ostracoda, (Plates LXIV.—

ATE SIMTN re eerie on eta ne cc cide ctieingnsie ceo Oysesnens

Tuomas, OLDFIELD, F.R.S., F.Z.8.

Exhibition and description of a new Monkey from the

Vet enya COMIN Sage ahvaesttalctisthsistaieis seins ois sicisisiscslcle vee sie ene eisislsale

Exhibition of a Collection of small Mammals from

China, with descriptions of new forms.............. Bepatecce

WuiteEnouse, RicuarD H.,M.Sc., University of Birmingham.

The Caudal Fin of the Teleostomi., (Plates XLVII.—L.)

Witson, Epwarp A., M.B., F.Z.S., Field-Observer to the

Grouse Disease Inquiry.

The Changes of Plumage in the Red Grouse (Lagopus

scoticus Lath.) in Health and in Disease. (Plates

Page

634

635

590

HNO NAY TELL (CVE patna ren es asec anen's oe aaGieeakis esac vbeneess 1000

Plate

XLVI.

XLVIIL.

XLIX,

LI.

TUE

LIII.

LIV.

LVII.

VET:

LIX.

LX.

LXL.

TENG:

LXIII.

LXIV.

LXV.

LXVT.

LXVILI.

LXVIII.

LXIX.

LIST OF PLATES.

1910, pp. 589-1033.

Page

Candalefinsion Meleostomiwe esos. . eclle~ + eects 590

TORSTEN TIMOR oe Gk aad eoon coon oCmeu mane oon 627

Photophores of Acanthephyra debilis ...-...-..-.

Photophores of Acanthephyra and Sergestes ...... f cag

Eimeria (Coccidium) avium. (Schizogony) ...... F

Eimeria (Coccidium) avium, (Macrogamete forma- |

AiO IM espn ewedsatey otctaw strc cieti eles ened ya reksurelsysiatrey ar ehees's. 3's es 818 anes

Eimeria (Coccidium) avium. (Microgametes, Spo- ( he

HOMO) dine Wbolls o Sow tie Gaede tee Ren EOE ueee |

Eimeria (Coccidiumy avium. (Sporegony)........

REUL COC TORO OIURUGU ECT a niapete ie eietcn ehciia)s ajo ie viele «'<\ ote) si o\e >

17-28. Leucocytozodn lovate. 29-40. Spirocheta |

lagupodis. 41-42. Hemoproteus mansoni ...... 692

3-50. Trichomonas eberthi, 51-57. Spirocheta |

lovatt. 58-65. Ameba lagopodis.............. J

Aaa COgGIS bs ose adoee coe Sent Hap ennaE 708

Blood cells of birds, chiefly of Grouse............ 722

1-8. Paracypria declivis G. O. Sars, 9-10. P. com-

planata G, O. Sars. 11-12. P. concidea G. O, Sars

Paraeyproa obtusa G.O. Sarvs ........+-e eee eeee

1-6. Paracypria defleva G. O. Sars, 7-11. P. sub- |

epguilaca GOW SATS s,m eee nese nee

1-4, Paracypria reniformis G.O. Sars, 5-9. P. hu- ¢

milis G. O. Sars. 10-15. P. opaca G. O. Sars .. |

1-10. Paracypria claviformis G,. O, Sars. 11-17. |

P, flecuosa G. O. SavTs 6... eee eee eee eee |

1-10. Cypridopsis obliquata G. O. Sars. 11-18.

GesANlG Orn ais xs sivicle vee ve atin we ees

Plate

1DDENG

LXXI,

LXXII.

LXXIII.

LXXIV.

LXXV.

LXXVI.

LXXVII.

LXXVIII.

LXXIX.

LXXX.

LXXXI.

LXXXII.

LXXXIII.

LXXXIV.

LXXXV.

LXXXVI.

LXXXVII.

LXXXVIII.

LAXNXIX.

XC,

XCI.

XCII.

XCIII.

XCIV.

XCV.

XCVI.

XCVIL.

XCVIII.

XCIX.

CI.

{

X1V

Page

1-3. Cypridopsis cunnington G, O. Sars. 4-6. Cr)

pusilla G. O. Sars. 7-9. €. monodonta G. QO. |

ars. 10-12) (Cl serrata Gear. s,s nse

1-3. Cypridopsis bidentata G. O. Sars. 4-6. C. tu- |

midula G. O. Sars. 7-9. C. congenera G. O. Sars. |

LO=12, Cighbai GO aS aren ai gies: > = aa ee

1-3. Zonocypris costata Vavra. 4-7. Z. levis G. r ie

O. Sars. 8-10. Stenocypris perarmata Brady. |

11-14. Heterocypris dubia G. O. Sars.........

1-4. Ilyocypris propingua G. O. Sars. 5-7. I. alta

G. O. Sars. 8-14. Limnicythere obtusata G. O.

Sars. 15-17. Paracypria curta G. O. Sars...... 3

Auditory apparatus of Balenoptera musculus ...... 769

INGtO PS LOFUCTIA A. cieie ie aiais o!e\e > os este Ge eee 792

Hydroids from Mergui Archipelago.............. 799

Pair of Red Grouse in summer with young chicks .)

Male Grouse, black type, in full winter-plumage .. |

Male Grouse, red type, in full winter-plumage ....

Male Grouse, white-spotted bird of the red type

Male Grouse, red type, in full winter-plumage with

a few black-centered feathers of the previous |

AULMM=plUMAe.:. -ycic\e's)- ale eee te tr sis staialc ee |

Male Grouse showing marked beginning of the

autumn-plumage on head and neck............

Male Grouse changing from winter- to autumn-

PUI AGE is ic a ciniecie nie stele hie ten eee ein ane

Female Grouse, black type, in autumn-plumage

Female Grouse, Scottish buff-barred type, in full !

breeding=plumace ir cnn ieiieieMe iis mets sa alerts

Female Grouse, red type, changing from winter- to

SUMMer=PUIUM ATE os egies ieee eee teinreieen eae |

Female Grouse in full summer-plumage ........ 1000

Female Grouse showing bare patch of skin and {

double line of barred feathers on abdomen......

Female Grouse, red type, feathers from flanks ...

Female Grouse, red type, worn upper tail-coverts .

Feet of Red Grouse: (1) New winter-feathers and

nails ;' (2) Full winter-plumage ..............

Feet of Grouse showing stages in moulting of nails. |

Heads of Grouse: (1) Female; (2) Male....,...

Heads of: (1) Female Grouse; (2) Male Grouse ;

and (3) Ptarmigan, showing supra-orbital combs. +

Head of Blackeock, showing supra-orbital comb ., |

Male Grouse showing abnormal erythrism........ |

Female Grouse, Irish buff-barred type

Female Grouse, abnormal yellow variety

Female Grouse, grey variety...........+eeeevees

Female Grouse, grey variety. (Type of Z. persicus

G, R. Gray)

LIST OF TEXT-FIGURES.

1910, pp. 589-1033.

\/. Development of the tail of the Flounder....................

. Curve of hind-foot measurements of Mus ratius....... 2

59. Curve of hind-foot measurements of Mus rattus tectorum......

2, Curve of hind-foot measurements of Acomys cahirinus

. Curve of hind-foot measurements of Mus rattus alevandrinus . .

Curve of hind-foot measurements of Mus norvegicus

Senctenere

63. Aylesbury Duckling with supernumerary legs ..............,

Diagram showing form and relative position of bones composing

supernumerary legs in an Aylesbury Duckling ............

lagram of Life-cycle of Eimeria (Coccidium) avium ........

. Stages in the development of the odeysts of Limerta avium, as

RCeMvIMyiresh) PLEPATALONS) aston ales, oeysisqecrs os ever « es. e+ :

Side view of snout of Tritylodon oneeas Owen, slightly

HSSUO CIE) PRE, Cea e Epo OL NE HOE ns Seeee Oem ate

. Upper view of snout of Trtylodon longevus Owen. ......... :

Dorsal surface of the beak of Balenoptera sibbaldit Guay, ;

SO Wari ps MANS 2) ue We eeiei cusses Sto owoeworns See csc bod one:

. Lateral view of the head of Balenoptera ees Gray, to show

GHEY SITS ocisiay rel otetens! se: NE Etre ctl Os ee ate Te AE Oe

Ventral view of left posterior portion of the skull of Balenoptera

musculus Linneeus. The tympanic bone has been removed . .

Diagrammatic view of the outer and middle ear of Baleno-

{UEDCs ooosdoc0c Mice oie cbG0 0 duldn comUODOO DOM OOE OE saggoae

. Transverse section (diagrammatic) of the pharynx of a

Miyata GOCELE yay seit <i ne dOor sispcaet uehaile Solon cin RiGee? bic

. Transyerse section Geert) of the pharynx of Physeter. .

. Ventral view of anterior extremity of the beak of Balenoptera —

COON GLS WINS TOS Seis Cb Re GO AIEEE Oo oe Ene seer ne

. Dorsal fins of Balenoptera sae Linneus, and Balenoptera

stbbaldu Gray .......... So ab SUR Ee aoe he die Oe toe EEC OE

. Paddles of Balenoptera ee Linneus, and Bulenoptera

siobalda Gray ... 24+... 50 TORRES POUR. Ce Seren anne ws

. View of the left side of the need of Physeter macrocephalus

AGIMNGCUS Misia ccoaeleles os > «© Seeeee “PU OUR Cee oormcee ube

Xvi

. Abnormal hydrothecz of Sertularella quadridens, X 20...... 819

a) RRAISOG ELON MRLGUTNE NPs) 3 RADI G fe faie x gee as aixly boecota hs oh (o ee ae 828

. Portion of hydroclade with hydrothece of Halicornaria hians,

VAR. 1400; Oh. (ECR PeRe Aa ae Cals cee Rass t os ee 836

2, Larva of, West African Mantis in resting attitude and when

CLAWILUIP eens es Rink SO SoNereGte:s eiaeie iss 6 Rene .. 838

. Postcornual glandular swelling on head of male Rupicapra

rupicapra. Also section of foot of a newly born kid ...... 849

. Sections of feet of Oreamnos montanus and Budorcas tibetanus . 852

. Sections of feet of Nemorhedus raddeanus and Capricornis

crispus

. Sections of feet of Ovibos moschatus. Also the lower surface

Of hind#thooktxe she seis Cece oe ee ee LACE . 858

. Inguinal region of young male of Ovis vignet. Also the front

foot of an adult showing the orifice of the pedal gland .... 860

Section of foot of a three days old lamb of Ovis vignez, and the

BATS VOR SOLACE ints tus ps x pie eg A chee wove’ eileen Siete apa ae 860

. Sections of feet of male and female Psewdois nayaur 863

» 00 Seelam

. Section of front foot of Hemitrayus jemlaicus and the same of

Capra hircus (var. dom.)

BSE ROOT NEE EM TE OS 2 . 865

. Preorbital gland of Guevet maawelli, and sections of the same

of Sylvicapra coronata

2. Pedal gland of Guwever manwellt ~ i... ns \scieinle ine oi inner 869

. Sections of feet of Guevet equatorialis and G. melanorheus.... 871

94. Sections of foot of Sylvicapra grimmz, subsp. ?, and of Cephalo-

DRUS MAQUI TONG sveaie a loins nivlsin’n! eset sintefe! aia eiatel ct aie ate eee 872

95. Inguinal region of male of Sylvicapra coronata, and sections of

feet of same ...... wile solee lavets ore sce 0 ib esa Ge ets ea 875

96. Sections of feet and the preorbital gland of Madoqua phillipsit. 877

97. Sections of feet of Dorcotragus megaloti3 ....6.... cece eae 878

98. Pedal and preorbital glands of Raphicerus campestris ........ 880

99, Preorbital gland of Owrebia nigricaudata 9, and section of

front Lo0b Of Same. po) 5 5 s,s oye fee anki wince gueiie tee 883

1C0. Sections of feet, and the inguinal region of Ourebia nigri-

(QTC asao au a a avy 16 ushtia Pee cane ce te Cate D Oe fete .. 804

101, Anterior view of pastern, and a section of the same foot, of

Oreotragus oreotragus. Also section of front foot of Pantholops

ROGGQSONE oa piv 3.0 cant eae bis ene io secon ie Tas eRe nv eee 886

102. Preorbital gland of Gazella dorcas and G, cuviert Q........4. 888

103, Knee gland of G. dorcas, inguinal gland of G. curteri 2, and

inguinal region of G, subgutturosd ...ceceee cere ee ceeuee 890

104.

Hind foot of G, subguttwrosa, sections of front foot of G. bennettii,

of G. cuvieri, and of Antidorcas marsuptalts vo... eeeceenes 892

XVil

105. Sections of the preorbital gland of Antilope cervicapra 3.

Also sections of the front foot of same and of the hind foot of

Page

JD OT ETTORE DCTUIT OO, 8 RCE. BE 8.0.0. Cs UCC ES IEEE 895

106. Sections of preorbital gland and of the feet of Bubalis caama.. 901

107. Sections of feet of Damaliseus horrigum jimela ........ 00006 903

108. Sections of feet and preorbital gland of Connochetes gnu . 904

109, Anterior view and section of front foot of Addax naso-maculatus.

Also section of hind foot of Oryx betsad ... eee ce eee 908

110, Pedal gland and section of preorbital gland of Hippotragus

OER chopaboor onout eo Fo. t10 BION 6.0.0.0 0.0.0 CbIg CiCiO.D COLO RMIT 910

111. Section of foot with pedal gland, and lateral and dorsal views

Oly NOSGION elem CAMLeolUs sr y.ciae ehdeteeks he fede cecalraia ees 912

112. Section of foot of Cervicapra redunca with aborted pedal gland.

Also inguinal region, glands, and nose of thesame ........ 914

118. Sections of the preorbital gland and of front foot of Adenota

kob. Also the inguinal region of same ..........0.0.0008 915

WAS Hind foot of Ainyceros melanipus ,. datees esis sleeves veces 919

115. One of the metatarsal glands of 4pyceros melampus ........ 920

116, Preorbital gland of Tetraceros quadricornis g and inguinal

region of Z'ragelaphus scriptus scriptus S ......000s Soca wee

117. Preorbital gland of Tetraceros quadricornis ....... erotatonercites . 928

118. Inferior and front views and section of hind foot of Tetraceros

CUE 2” Beochogobce pols baud 4 UO NOM Od do OuIn{o 924

119. Inferior view and section of the hind foot of Tetraceros quadri-

cornis §. Also transverse section of the pastern of same,

and transverse section through the region of the false hoofs

of Tragelaphus scriptus Sv. sessccceeeseveseues Boloconae hs!

120. Preorbital gland and nose of Boselaphus tragocamelus 2 . 927

121. Sections of feet of Boselaphus tragocamelus Q .......seeeees 928

122. Anterior aspect of feet of young and adult female Tawro-

tragus oryx, section of front foot of Tragelaphus scriptus

roualeyni?, and inguinal region of female foetal example of

PREGAME GMOS) ga nc0ne oo pARONOOr ROAD Sho. Oa 929

123. Anterior aspect and section of foot of Bos sondaicus three days

old. Also section of foot of calf of Bos taurus ......6+.55. 933

124, Section of feet of Bos bubalus and Bos caffer.......+..eeeeee 93

125. Section of front foot of a prematurely born example of Antilo-

capra americana, and section of hind foot of Giraffa camelo-

pardalts, three weeks Old) (2.0.2. eee cee ce se cet a se ecene 935

126, Lateral aspect of tail and sections of feet of Moschus moschi-

HES G soocssoococbouvscocp oo nb uo KooD DU DOG OODOUD UGE 938

127. Sections of the preorbital ae and of the feet of Cervus

(OBB) TRAYS, 65.5 0.6 6 6 odon DO DOOD DOE OURO OUO COORD 940

128. Section of hind feet of Cervus (Gs vus) elaphus and Cervus

CRUCerrUe OUUTUCEIEN Wine win cles cece se caaeeeeevesaean 941

Proc, Zoou. Soc.—1910. b

XVill

Page

129. Sections of feet of Cervus (Pseudavis) sika ..... 0... c eee 943

130. Sections of feet of Cervus (Panolia) eldi ... 6c... cee eee eee 945

ISL, Sectionsiof featol Rusa unicolor...na.sseeeee +s nl ss es 947

152. Preorbital and metatarsal glands of Avis (Axis) avis. Also

section of hind foot of Avis (Hyelaphus) porcinus ........ 949

133. Sections of feet of Dama dama..ic..cs Geet basescavecssccs 951

134, Anterior aspect and section of hind foot, and section of front

foot of Cervulus muntjac. Also preorbital gland of same .. 953

55. Preorbital gland and sections of feet of Zlaphodus michianus.

Also section of hind foot of Cervulus reevest .........065 955

136. Sections of feet of Capreolus capreolus ...... ccc eee eee ees 957

157% Sections of feet, of Alceialces .,..:.%. esmideiis >» ver ss wae 959

138. Sections of feet of Rangifer tarandus oo. .cc cee cece eee e ees 961

159. Sections of feet and tarsal gland of Mazama nemorivagus 2.

Also preorbital gland and rhinarium of same ............ 965

140. Sections of feet of Mazama bricenit 2, and of hind foot of

Dor celanhus GMenicanusy a. 2 ae eirtseate tin atas 6) + > Ox 6 he 965

141. Sections of feet of Tragulus meminna, of Dicotyles, and of

Phacocherus, and anterior aspect of foot of Tragulus stan-

POY RUS F sisialeientei ds Metts UE RMT ie eae se scot <x 979

142. Section of foot of Lama vicugna, anterior aspect of feet of

Dicotyles collaris and of Phacocherus, and section of dorsal

pland) of Dicotyles:collarts | v.21) .1teeidetedtteieesn sas 2 oa eel 981

143. Antlers of Tibetan Wapiti (Cervus canadensis wardi) ...... 988

144, Skulls of Cervulus reeves (?) and Cervulus bridgemant ...... 990

145. Frontlet and horns of bull ot Bos caffer thierryt .........005 993

146. Head of bull Bos caffer simpsont .......06+ Aotiiaad mike cheers 994

147. Head of cow Bos caffer stmpsont ......eeeeeees 2 Ae pace 995

148. Head of young male Bos caffer cottont (?) ...cccceeecueees 996

149. Whitish-blazed Topi (Damatliscus phalius) from the Uasingishu

De RF RESID) in Si. o.4 Cs SOCCER RRR RE IN 3 OL 999

150. Diagrammatic representation of the growth of Primaries in

the young Grouse .,..4....... Rieti the ys cen oi tae cvs, LOST

xix

NEW GENERIC TERM

PROPOSED IN THE PRESENT VOLUME (pp. 589-1053).

Palacyprial (Ostiacoday) veceuesceswenusescsasscsagsqeeete 752

™“--

TARR SPREE 08 ie

i w

fe te

a pe |

No. 86.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

June 14th, 1910.

Dr. 8. F. Harmer, 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 May 1910.

Mr. R. I. Pocock, F.LS., F.Z.S., exhibited drawings to

illustrate an apparently unrecorded instance of ant-mimicry by

the larve of a species of Mantis from Axim, on the Gold Coast.

Mr. R. I. Pocock, F.L.S., F.Z.S., read a paper “On the

Cutaneous Scent-glands of Ruminants,” based principally upon

work done at the Prosectorium on animals that had died in the

Gardens. The paper was illustrated by lantern-slides showing

the macroscopic structure of the preorbital, pedal, and inguinal

and other specialised glands of most of the genera of this group

of Ungulates. The author pointed out that the structure of the

feet, whether furnished with special glands or not, supplied

valuable data for classifying the genera of Antelopes and Deer ;

and showed that with some modifications, such as the removal of

Tetraceros from the Cephalophine to the Tragelophine, of

Dorcotragus from the Antilopine to the Neotragine, and of

Pantholops, Saiga, and dipyceros from the Antilopine, the sub-

families usually admitted were valid groups. In the case of the

Deer it was interesting to note that Rucervus, Panolia, Elaphurus,

* This Abstract is published by the Society at 3 Hanover Square, London,

W., on the Tuesday following the date of Meeting to which it refers. It will

be issued, 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 aanum, payable in advance.

“eas oie ir

38 ie, if oF) a

ey, ie)

and Sika were closely allied to Cervus, Dama being a totally

distinet type. Ais and Hyelaphus belonged to another group,

while Rangifer, Alces, and Capreolus, as Sir Victor Brooke |

claimed, belonged to the section typified by Dorcelaphus, Mazama,

and other American Deer. ie

Mr. R. LypexKer communicated a paper “On a Wapiti and

a Muntjac,” in which he described two Wapiti antlers from Tibet _

as Cervus canadensis wardi; and a Muntjac from An-wei, China,

as Cervulus bridgemani. The latter was characterised by its dark

blackish-olive colour, the black ears of the female and the yellow

ones of the male, coupled with the relatively wide divergence of

the antler-pcdicles.

Mr. R. LypexKer also presented a paper “ On Three African

Buffaloes.”

Dr. A. Casrera, C.M.Z.S., communicated a paper “On Two

new Antelopes,” in which he described a new spevies of Dama-

liseus from British East Africa and a new Chamois from North

Spain.

Mr. W. R. Oartvie-Grant, F.Z.8., communicated a paper by

Dr. E. A. Wixson, F.Z.8., Field Observer to the Grouse Disease

Committee, entitled ‘Changes of Plumage in the Red Grouse

(Lagopus scoticus) in Health and Disease.”

This Meeting closes the Session 1909-1910. The next Meeting

of the Society for Scientific Business will be held on Tuesday,

November 15th, 1910, at half-past Eight o’clock p.m.

The following Papers have been received :—

}. F. E. Bepparp, M.A., F.R.S., F.Z.8.

On the Alimentary Tract of certain Birds, and on the

Mesenteric Relations of the Intestinal Loops.

2. EK. H. J. Scuuster, F.Z.S.

Cortical-cell Lamination of the Hemispheres of Papio

hamadryas.

3. Prof. Atmex. MEEK, F.Z.S.

Sa a ee a

A Contribution to our Knowledge of the Development of

the Brain and of the Nose of the Crocodile, Crocodilus porosus.

Communications intended for the Scientific Meetings of the

ZOOLOGICAL Socirry oF Lonpon should be addressed to

P. CHALMERS MITCHELL,

Secretary.

3 Hanover, Squarz, Lonpon, W.

June 21st, 1910.

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PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

"| 00LOGICAL soctETY

OF LONDON.

1910.

Pages 589-836.

Part II]. conrtaininc PAPERS READ IN

APRIL anp MAY.

OCTOBER 1910.

PRINTED FOR THE SOCIETY,

SOLD AT THEIR HOUSE IN REGENT’S PARK.

LONDON :

MESSRS. LONGMANS, GREEN, AND CoO.,

PATERNOSTER ROW,

[Price Twelve Shillings. |

LIST OF CONTENTS.

1910, pp. 589-836.

April 5, 1910.

Page

The Secretary. Exhibition of a series of photographs taken in the South Orkneys in 1908

by a correspondent:of “he Dimes sy cteccn caste awewsiet so acce sess as ans 589

Mr. R. I. Pocock, F.L.S., F.Z.8. Exhibition of a living specimen of a Mole-Rat (Georychus)

fromp ASbAmhl >.\. 26 Kancitate ee eames Sie eve vies Ki eke ke le .o e+ 0 » 3 0 yoo er 589

1. The Caudal Fin of the Teleostomi. By Ricnarp H. Wurrenovse, M.Se., University of .

Birmingham,» (Plates LVIEAV,) 25 vin's acs sete ssnvn scveasc- es sce + dente 590

2. Some Notes on Tasmanian Frogs. By T. M. Savacn Eneuisn. (Plate LIL.) ....-....- 627

April 19, 1910.

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

Mar Bh AO 26 one Bie ls cious tnt ss s «.« aceetad Ce hvig SRR Bivens ish oaic/<lp eae er 634

Mr. Oldfield Thomas, F.R.S., F.Z.S. Exhibition and description of a new Monkey from

the Wales Ponnsulae ssi. cislatcysss,'s, acess coscretoeate ercerstnte amet ear eae a tote Lake Seti ST. 634

Mr. Oldfield Thomas, F.R.S., F.Z.S. Exhibition of a Collection of small Mammals from

Ohina, with descriptions of new forms... 05. e ee'eees a> eee sce cas 07 ve es hie 635

Mr. J. Lewis Bonhote, M.A., F.L.S., F.Z.S. Exhibition of a yellow variety of Mus ratius.. 638

The Secretary on the Centenary of Philip Henry Gosse, F.R.S. .......... IS 639

1. Notes on the Photophores of Decapod Crustacea. By Sraniny Kenr, B.A. (Plates

DALE SUV ea inte ta S 10 is,< cate a Wis isl oie So Sie hl msgtal RRM Sister nt W'RIE e's» oc >'s io os 5 Aen 639

On the Varieties of Mus ratius in Egypt; with General Notes on the Species having

reference to Vaiiation and Heredity. By J. Lewis Bonnore, M.A., F.LS., F.Z.S..... 691

3. On an example of Posterior Dichotomy in an Aylesbury Duckling. By G. E. Bunnen.. 666

May 3, 1910.

The Secretary. Exhibition of a photograph of a Giraffe .. 0... ..0+ cece cece cece cee tees 66

Mr. E. E. Austen, F.Z.S. Exhibition of a specimen of a rare Fly (Cephenomyia ulrichii

Braner) OM 9st Lussier sis stamp soe pcs sain 05 r oinisinle Sele teavble's oes sce ste 669

Dr. R. A. O'Brien. Remarks on the habits, illustrated by lantern-slides, of the North

Queensland Green Tree-Ant (Oecophylla smaragdina ?). 1.0 cecceecceeeecsvnccvecee - 669

Contents continued on page 3 of Wrapper.

PROCEEDINGS

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

(April to June, 1910.)

April 5, 1910:

Prof. E. A. Mincut, M.A., Vice-President,

in the Chair.

The Secretary exhibited a series of photographs taken in thie

South Orkneys in 1908 by a correspondent of ‘The Times,’ and

kindly sent to him by the Editor of ‘The Times.’ The photo-

graphs included views of scenery and an interesting series of

studies of living Seals, such as the Sea-Leopard (Stenorhijnchus

leptonyx), Weddell’s Seal (Leptoniychotes weddelli), the White,

or Crab-eating Seal (Lobcdon carcindphdgus), and Ross's Seal

(Ommatophaga rossi).

Mr. R. 1. Pocock, F.L.S., F.Z.8S., exhibited a living specimen

of a Mole-Rat (Georychis) recently presented to the Society by

Capt. C. H. Armitage, who brought it from Ashanti. Mr. Pocock

remarked that the > ihimal possibly belonge ad to the species from

Togoland described by Matschie as @. zechi, and certainly repre-

sented a form never previously exhibited in the Gardens.

Proc. Zoou, Soc.—1910, No. XXXIX. 2

590 MR. R. H. WHITEHOUSE ON THE | Apr. 5

The following papers were read :—

The Caudal Fin of the TeLrosromt.

By Ricuarp H. Warrenouse, M.Sce.,

University of Birmingham.*

[Received February 4, 1910.]

(Plates XLVII.-L.? and Text-figure 57.)

ConTENTS.

1. Introduction, p. 590.

2. Terminology, p. 592.

3. Morphology, p. 593.

4, General Conclusions, p. 620.

5. Additional Note, p. 623.

1. IyrRoDUCTION.

While the study of the paired fins of fishes, and also the dorsal

and anal among the median unpaired fins, has been prosecuted in

ereat detail, a systematic, detailed investigation of the caudal fin

seems to have received much Jess attention. Some authors have

dealt somewhat extensively with the subject, yet these have only

treatect of a comparatively few species; it was thought advisable,

therefore, that such an important organ as the caudal fin of fishes

should receive a broad treatment, and to carry out this purpose,

several scores of different species have been subjected to a detailed

examination of the skeletal parts of the fin. The number treated

of includes species from as many widely different families as it

has been possible to obtain, so that almost every form of tail

likely to occur should be made known. It is impossible to include

every type examined in the present paper, but the subject will be

completed at as early a date as possible.

The Teleostei will receive the fullest treatment, but some

reference will be made to other groups for the sake of presenting

a more complete account of the piscine tail-fin. It has been

impossible, as yet, to investigate the tail of the Dipnoi, but it is

hoped that this group also will be dealt with in due course.

Material for the work has been obtained from various sources,

the largest quantity having been obtained at Naples during my

occupation of the British Association table at the Zoologie: qT

Station in that town, during the six months ended May, 1908,

Other valuable material has: been supplied to me by a number of

friends. I should like to mention here that the whole investiga-

tion is due to the suggestion of the late Professor Bridge, F.R.S.,

under whose superintendence the work was carried out; it was a

pleasure and fortune to have so able a eritie and kindly adviser as

my late revered teacher. I wish also to express my best thanks

to the British Association for appointing me to occupy their table

* Communicated by Dr. P. Cuatmenrs Mirenety, M.A.. F.RS.

+ For explanation of the Plates see p, 625.

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CAUDAL FINS OF TELEOSTOMI.

RE Wittebmse, del. Huth Tsth* London

CAUDAL FINS Of TRLEOS TOML.

a ee ee poy eas ee

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P.Z.S.1910.Pl. XLIX.

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CAUDAL FINS OF TELEO

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RA Whitehouse del.

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CAUDAL FINS OF TELEOSTOMI.

ae , «oS. = a 7” :

1910.] CAUDAL FIN OF THE TELEOSTOMI, 591

at the Naples Zoological Station. The many friends who have

supplied me with material and suggestions also have my best

thanks. For the identification of most of the species I am greatly

indebted to Dr. Lo Bianco of the Naples Zoological Station.

Lastly, I am deeply grateful to the Birmingham Natural History

and Philosophical Society and others, for ‘grants without which

the work could scarcely have been done.

The dissections have been carried out as carefully as possible

under the microscope, and it was found necessary in most cases

to render the specimen transparent by some clearing agent such

as turpentine or xylol, in order to see exactly how the vertebral

column ended, and to clear up any other doubtful points.

In reviewing the literature on the subject of caudal fins, the

very numerous casual references must be omitted for the present.

Perhaps the earliest definite work relating to fishes’ tails is that

of Agassiz in 1833, in which the author recognized two types of

caudal fin, the heterocercal and homocercal; the former term still

retains its original meaning, but the latter was only used to

indicate an externally symmetrical fin as distinguished from the

asymmetrical heterocercal form. In a geological contribution by

McCoy in 1848, the term “ diphycercal” first appeared to desig-

nate the truly symmetrical caudal fin; it is interesting to note

that Cope in 1871 coined the term “isocercal” to mean the same

thing, in ignorance, as that author states, of McCoy’s paper.

Stannius, in a text-book on the Vertebrata, published in 1854,

gives considerable attention to the tail-fin; he refers to a very

lar ge number of Teleosts, but with questionable accuracy. ‘The

“inde interesting feature oi his remarks concerns his belief, which

seems to be true, that the supports of the fin-rays are formed of

arches and radials fused; Lelone is quoted in support of this

view.

But it was in 1859 that Huxley first gave a correct description

and interpretation of the homocercal tail; he exposed the false

symmetry of form in (rasterosteus, and traced its development.

It is unfortunate that the promised account of the development of

the eel’s tail never appeared, for the slight reference made to this

form is of doubtful accuracy, due, no doubt, to only a superficial

examination. Very soon followed (in 1860) an excellent essay

on caudal fins among Ganoids and some ‘eleosts by Koélliker,

accompanied by vaiuable figures which have been reproduced

almost universally since. Lotz followed in 1864 with a paper

containing little new work.

A revival was given to the subject in 1878 by Alex. Agassiz in

a short paper on ‘the development of the tail of the flounder ; his

remarks had the effect of raising the important question df the

true morphology of the homocercal caudal fin. In 1882,

EK. T. Newton read a paper on “ Fishes’ Tails” before the Quekett

Microscopical Club; this consisted of a general review of past

work, together with a description of the ‘tail of the sprat; the

paper included an incorrect figure of the caudal fin of the cod.

39*

592 MR. R. H. WHITEHOUSE ON THE [Apr. 5,

The most comprehensive work of all, however, was published

by Ryder in 1886 ; on the whole, it contained very useful infor-

mation, but his figure of the tail of the young eel is wrong with

respect to the neural arches. Ryder’s suggestions on the origin

of heterocercy are interesting and pr obably correct, though some-

what novel. Ryder’s evolutionary series of the forms of the

caudal fin has come to be acknowledged as the order of appearance

of the different types in time. The next work of importance

dealing with caudal fins of fishes was written in 1895 by Dollo in

discussing the phylogeny of the Dipnoi: basing his conclusions on

eeological evidences, he gives the term ge phyrocerey an extra-

ordinarily wide application; this arises from the idea, strongly

upheld by Dollo, that the present caudal fin in many ‘Teleosts

is really a posterior anal fin, brought into its present position

by complete atrophy of the meal caudal. ~Dollo’s is the last of

important references regarding the tail-fin.

Boulenger, in 1901, in dealing with the fishes of the Congo,

gave a good summary of work done! In 1907, Gregory, in his

work on the classification of Teleostomous fishes, makes continual

references to the caudal fin; but it is difficult to reconcile many

ot his statements: for instance, the caudal fin of MJastacembelus

is referred to as gephyrocercal, ribs: in fact internally it is quite

a generalized homocercal type. As an appendix to the work,

there was given an elaborate scheme of the evolution of the

‘audal fin, which includes several new terms, considered by some

as unnecessary.

2. TERMINOLOGY.

In order to avoid any ambiguity, I have thought it necessary to

modify the meaning of several terms; some have been widened

and others have been more restricted as regards their denotation.

The reasons for such modifications are given in their respective

paragraphs :—

1. Urostyle.—This term is used in the generally accepted sense

denoting the more or less elongated cone-like termination of

the vertebral column, appearing as a prolongation of the last

centrum. It represents the fusion of severai centra.

2. Hypawxial.—This adjective refers to any structure morpho-

logically ventral to the chordal axis.

Epaxial.—This adjective reters to any structure morphologically

dorsal to the chordal axis.

4, Hypwral.—Any hypaxial element having a direct connection

with the chordal axis, and bearing one or more fin-rays

distally, will be called a hypural or hy pural bone. Previously,

this term has been used rather loosely and in such a way as

to leave one in doubtas to its limitation ; the limitation given

above, concerning its direct connection with the chordal

axis—?.e. as a general rule, with centra—as will be seen

later, both widens and narrows the meaning of the term,

1910.) CAUDAL FIN OF THE TELEOSTOMI. 593

5. Hpural.—This term may be regarded as applying to the dorsal

homologue of the ventral hypural, 7. e. any epaxial element,

having direct continuity with the chordal axis, and bearing

one or more fin-rays at its distal end, will be called an epural

orepural bone. It is only after the most careful consideration

that I have decided to change the definition of thisterm. Lt

was established by Huxley in 1859, and appears to have been

used by him to indicate those isolated ray-bearing bones

situated immediately dorsal to the urostyle in Gasterosteus.

The change ‘suggested is based solely on my conclusions

respecting the real morphological value of the several fin-ray

supports, and, reluctant as I am to make any changes in

established nomenclature, I believe the reader will agree that

the subject will be much simplified by the change.

6. Radial.—The term radial is used in its usual sense as synony-

mous with “interspinous bone” and ‘somactid” of some

writers ; 1t implies a more or less dagger-like bone, primarily

having uo direct continuity with the vertebral column, and

supporting distally one or more fin-rays.

7. Last vertebral segment.— This term will refer to the last centrum

plus urostyle, if such exists. The last centrum is never per-

fect, and is often no more than a cone, the apex of which is

directed posteriorly.

3. MorpnuoLtocy or CaupaL Fins.

This section will be descriptive of a selection of caudal fins, and

the order in which they are dealt with corresponds essentially

with the classification given by Sedgwick in his ‘'Text-Book of

Zoology. Owing to lack of time and material, the series is not

as complete as one would wish ; for example, only the Siluride

among the Ostariophysi receive attention, and the Symbranchit

are completely omitted. Except for these sub-orders, however,

the types are fairly representative of the others, at least of the

Teleostei. Almost every example taken presents interesting

features, but owing to the impossibility of including, in the

present paper, figures of all the species treated, only the bavest

description of many forms can be attempted at present.

ACIPENSER sturio. (Chondrostei, Acipenseride.) (Plate XLVIT.

fig. 2.)

The Sturgeon presents a definitely heterocercal tail ; the end of

the caudal axis from the root of the tail-fin is directed upwards,

and the ventral lobe is enlarged, producing a very unsymmetrical

fin. No centra are formed from the notochordal sheath. ‘The

ventral lobe of the fin is supported by a very regular series of

hypurals ; the name hypural has been applied here, since these

structures strictly comply with the detinition. They are of

particular interest, because, in the anterior part of the fin, where

they are most strongly developed, they exhibit a definite segmen-

tation: the largest show four successive segments (/..d., p., u2., and d.,

594 MR. R. H. WHITEHOUSE ON THE Apr. 5

} ;

in the figure); proceeding backwards, as they become smaller,

three segments are shown, then two, anil finally only one can be

traced, “This segmentation is due, I believe, to the formation of the

hypural by the coalescence of radis ul with hemal arch ; anteriorly,

the three segments, the proximal (p.), median (m. ); and distal (d.),

of the nadiale are present, the distal still remaining free; from

before, backwards, the fusion becomes more perfect and finally

ail trace of it is lost This composite nature is a feature which

substantiates the claim of these hypaxial supports to the

title hypural. The dermotrichia constituting the ventral lobe

considerably overlap their endoskeletal supports. Dorsally, the

arches (n.a.) and radials (d.e.7.) remain distinct throughout the

fin and are less regular than the hypaxial elements. The dorsal

radials are almost hidden from view by the forking proximal ends

(removed in the figure) of the stout fulera which i inge the margin

of the dorsal lobe. The whole of the elements at the extr emity of

the tail are hidden by the dermotrichia.

Potyopon spATULA. (Chondrostei, Polyodontide.) (Plate XLVIL.

fig. 3.)

Externally the tail of Polyodon is very deceptive; its symmetry

of form is most marked, and it resembles in outline the deeply

cleft tail of some Teleosts. It is well-known, however, that on

dissection it proves to be an excessively heterocercal type, the

caudal axis being directed upwards and extending to the extremity

of the dorsal lobe. No centra are formed. Almost throughout

the tail-fin the neural arcbes (7.a.) are single elements, and are

somewhat irregular in size, particularly towards the extremity of

the axis. Resting upon these arches are the radials (diew.)s ‘the

first four or five are slender in build, but almost throughout the

fin they resemble the neural arches themselves. They, too, are

very irregular both in disposition and size, and here and there

they show signs of having been more numerous formerly. The

various Bl araeatts at the extremity of the axis are difficult to dis-

tinguish, owing to their small size and tendency to fusion; the

same difficulty is experienced ventrally in this region. The radials

are the immediate supports of the fulera (/.).

Turning to the ventral elements, a very regular series presents

itself, in striking contrast to the dorsal side: there is a regular

series of hypurals, very large at the root of the fin where the large

dermotrichia of the ventral lobe are supported, As in Acipenser,

very many of these hypur: als show distinct segmentation, thus

giving ev idence of bei sing composed of four successive e Jementa the

distal of which, in the case of the first seven supports, is distinet

and separate. The interpretation seems to be that each hypural

is composed of hemal arch and radial fused, the tri-segmental

nature of the latter being retained in several cases. Naturally the

best examples of this are given by the anterior hypurals, where

the supporting elements are larger. The tri-segmental arrange-

ment gradually gives way to a bi- and finally a uni-segmental

1910. ] CAUDAL FIN OF THE TELEOSTOMI. 595

structure in passing from before, backwards. The caudal fin of

Polyodon, then, is obviously heterocercal, but of a more specialized

type than Acipenser, on account of its nearer approach to the

homocereal type.

Ami cALVA. (Holostei.) (Plate XLVII. fig. 4.)

The caudal fin of Amia is externally symmetrical, ov at any

vate the discrepancy in symmetry is very slight indeed, and

negligible. But dissection reveals a very unsymmetrical internal

structure ; at the root of the fin the vertebral column takes a

sudden turn upwards, at a considerable angle. The centra are

well formed and retained throughout the length of the axis, and

consequently no urostyle is present. Nearly the whole of the

dermotrichia are supported by hypurals, a feature which it shares

with the majority of Teleostean fishes; these hypurals are slightly

expanded or somewhat club-shaped at their distal ends, and each

bears a single fin-ray. The last three hypurals belong to the last

centrum, but each of the other centra possesses but one hypural.

The final dorsal element (fig. 4, 2.a'.) is strikingly large, and con-

tinues beyond the vertebral axis for some distance between the

.fin-rays. Kolliker remarks that it is cartilaginous and contains

the spinal cord: he regards it as an incorporation of vertebrae and

neural arches. In the specimens I have examined, the same

structure proved to be bony, and grooved along the dorsal edge to

conduct the spinal cord ; it appears to correspond to a bone, fre-

quently found among the Teleostei, which [ have regarded as a

persistent neural arch, retained, even when its adjacent homologues

are suppressed, as a protection for the delicate termination of the

spinal cord. According to Kolliker, there are three dorsal caudal

radials (d.¢.7.) remaining as supports for the insignificant fin-rays

which constitute the epaxial contribution to the caudal fin; but

these dorsal radials are very poorly developed. Most authorities

agree that the caudal fin of Amia is scarcely correctly described

by being called heterocercal, and therefore it is generally referred

to as hemi-heterocercal. So near, however, does the structure

approach homocerey that hemi-homocereal would probably be a

more suggestive designation : however that may be, the fin may

certainly be considered the most specialized type of heterocercal

forms.

General remarks on the Ganoid Caudal Fin.

It has been thought advisable to omit any detailed references

to other Ganoids than the three considered, for the present, since

they have not been cleared up to my entire satisfaction and

material is lacking. But the Ganoid caudal is characteristically

heterocereal ; truly protocercal forms are absent, and homocercy

is nowhere attained within the group. The three types considered

illustrate how the tendency of specialization is to approach the

homocercal condition ; in Acipenser, the typical heterocercal form,

so characteristic of the Elasmobranchs, is present; Polyodon, by

596 MR. R. H. WHITEHOUSE ON THE [ Apr. 5,

an increase in the size of the ventral lobe reaches a step towards

the externally symmetrical homocercal fin; while Amia proceeds

furthest of all in the homocercal direction, requiring but one single

feature, the possession of a true urostyle, to attain the homocerecal

evade

TELEOSTEI.

MALACOPTERYGII.

Mormyrus KANNUME, (Mormyride.

In general, Mormyrus provides a type of homocercal caudal fin

by no means generalized. The last two or three vertebre are

directed slightly upwards. The urostyle is absent or fused com-

pletely with a hypural bone ; such an adyanced homocercal feature

as this is not consistent with the position of Mormyrus as one of

the least specialized of Teleosts, a position assigned to it according

to general characters. In respect to the hypurals, uncommon

feanieas are shown: in the first place the last centrum presumably

is fused with the last hypural but one, instead of the last as is

usual: again, two hypurals are attached to the penultimate

vertebra and the anterior of these is exceptionally large. It is

important to notice that only two hypuyal bones are associated

with the last vertebral segment—the small number being another

feature of the specialized homocercal caudal. The epaxial

elements are interesting; the neural arch to the last centrum is

strongly developed; it is pierced throughout by the spinal cord

which is continued beyond the arch among the dermotvichia. It

will be remembered that a similar condition of the last neural arch

was found in Ama. The neural arch of the penultimate vertebra

is very much reduced, and the antepenultimate vertebra bears an

epural bone.

ENGRAULIS ENCRASICHOLUS, (Clupeidee.)

The Anchovy provides an illustiation of one of the least speci-

alized of homocercal caudal fins. No centra enter into the

upturned part of the caudal axis. The notochord is prolonged

beyond the long urostyle, but except for the extreme end which

extends among “the dermotrichia, it is protected by several slender

and closely fitting ensheathing bones which it is difficult. to

homologize. Dor sally and ventrally at the end of the notochord

is a delicate car tilage, each, 1t may be, a vestige of once functional

neural or heemal arches : such vestiges are referred to by Ryder

as opisthural cartilages. The hypaxial endoskeletal supports of

the fin-rays are nine in number, seven of which belong to the last

vertebral segment. It is a little difficult to decide whether the

term hypural is correct for all these supports, as some, at any rate,

appear to be ventral caudal radials only; however, it will be

better for the present to call all of them hypurals. This fish

possesses the largest number of hypurals (viz. seven) to the last

1910. ] CAUDAL FIN OF THE TELEOSTOMI, 597

vertebral segment which has been found. The dorsal contribu-

tion tothe caudal fin is comparatively small. Of the last four

vertebre all but the last bear epural bones and the last appears

to have its neural arch greatly reduced. T'he most posterior of

the epaxial fin-rays are supported by two or three poorly developed

dorsal caudal radials; the proximal ends of these radials are

almost in contact with a triangular cartilage applied to the dorsal

side of the notochord near the termination of the urostyle; this

cartilage probably represents a vanishing neural arch.

CLUPEA PILcHARDUS. (Clupeide.) (Plate XLVII. fig. 5.)

The caudal fin of the Pilchard very closely resembles that of the

Anchovy. The urostyle (w.) is long and slender, and the noto-

chord (z.) is produced beyond it, but not beyond the distal ends of

the endoskeletal supports, An opisthural cartilage (0.c.) is

attached to the end of the notochord, and extends between the

dermotrichia. Nine hypural bones are present, seven of which

belong to the last vertebral segment, as in Hngraulis; the

hypurals vary considerably in size, Epaxiallv, the dermotrichia

ave supported by two epurals (ep.) and three dorsal caudal radials

(d.c.r.); the latter are short bones situated between the last

epural and the notochord. Proximally the dorsal radials abut a

triangular bone (.@.')—cartilage in Hngraulis—on the dorsal

side of the urostyle: this triangular bone probably represents one

or more neural arches of vertebra now absorbed into the urostyle.

The last centrum (/.v.) is provided with a large spatulate neural

arch (2.@.) peculiarly constricted at its proximal end,

Comparing the caudal fins of the two Clupeoids here described

and also that of the Sprat as detailed by Newton, it will be seen

that there is a striking similarity throughout; the figure given in

Newton’s paper is that of a young Clupea sprattus, and the

urostyle is seen to be composed of several centra which, no doubt,

are also present in C. pilchardus and Engraulis encrasicholws in

the young, but which are obliterated in the adult All show a

lowly specialized homocercal form, approaching very closely the

Amie type.

NoToPpreRvus BORNEENSIS. (Notopteride.) (Plate XLVII.

fig. 6.)

This fish is provided with a highly specialized caudal fin, and

also presents one or two peculiarities. The caudal is continuous

with the ventral median fin, but the caudal proper may here be

limited to that part composed of dermotrichia which are attached

to the endoskeleton by means of a proximal forking, as distin-

guished from the ball ‘and socket nature of the attachment of the

anterior fin-rays to the radials. Thus the caudal fin is supported

by three hypurals and one epural; hence it is not strictly hypo-

-cercal as Gregory states. No urostyle can be seen in the dissection,

and even clearing fails to give any certain clue as to its exact

outline in the Substance To the last hypural bone, and what

598 MR. R. H. WHITEHOUSE ON THE [Apr. 5,

may be at first considered a urostyle, indicated by a ridge

along the dorsal side of the hypural, has a companion ridge

directed ventralwards; the loss of the urostyle or its fusion

with a hypural bone is an advanced character. A large hypural

bearing four dermotrichia is attached to the penultimate vertebra,

while that of the antepenultimate supports only one. An

interesting feature in this caudal fin is the epaxial fin-ray

support; this consists of a single epural bone broadly expanded

distally, and attached to the penultimate vertebra. Such an

expanded epural is quite unusual in caudal fins, and I know of no

other instance.

General remarks on the Malacopterygqii.

The examples chosen illustrate how varied the caudal fin

structure may be, even in a single sub-order of Teleosts. Two

extreme types are met with: the Clupeoids, by the extension of

notochord beyond the bony axis, by the vestiges of once functional

arches in the form of opisthural cartilages, and by the large number

of hypural bones to the last vertebral segment, provide an

example of a lowly specialized form of caudal fin; while Votopterus,

by the loss of the urostyle and the reduction of the number of

hypurals to the last vertebral segment to a single bone, is an

example of a highly specialized form.

OSTARIOPHYSI,

Cruarias sp.? (Siluride.)

Only a single specimen of this fish has been procurable, but the

caudal fin appears to be quite normal. The tail-fin is fully

differentiated although the dorsal and ventral fins reach nearly to

the end of the body. Internally the skeleton is remarkable in

presenting a very long urostyle, longer than that of any fish

examined, extending posteriorly as far as the distal ends of the

hypurals ; it does not appear to be actually fused to a hypural.

The hypaxial part of the fin is supported by five hypural hones,

four of which are associated with the last centrum and urostyle,

and also by one ventral caudal radial occupying the space between

the first and second hypural bones. The hypural of the penul-

timate vertebra has the appearance of being formed by the

coalescence of a radial anda hemalarch, The epaxial fin-rays are

borne by one epural bone belonging to the last vertebra but one,

and also by a dorsal caudal radial between the epural and the

urostyle. The spinal cord can be traced to the end of the uro-

style. The final centrum possesses the vestige of a neural arch,

through which the spinal cord passes.

Here then is a generalized type of homocercal caudal fin, for in

it are representatives of all possible endoskeletal supports, viz.,

hypurals, ventral caudal radial, and dorsal caudal radial, and

besides, it possesses a well-marked urostyle.

LL: CAUDAL FIN OF THE TELEOSTOMI. 599

Prorosus sp.? (Siluride.) (Plate XLVIT. fig. 7.)

The caudal fin of Plotosus is very interesting and very simple ;

it forms part of a continuous median fin and terminates a gradually

tapering tail. All the specimens examined were from ‘Torres

Straits and were presented to me by the late Professor Bridge.

They were all young, but the skeletal elements were quite fully

established. Before entering into a description of the fin, it will

be instructive to call attention to the remarks made by Mr. Tate

Regan before the Zoological Society on April 28th, 1908, con-

cerning the caudal fin of Cnidoglanis megastomi, also a member of

the Plotosine. He writes :—

“The Siluroid fishes of the subfamily Plotosine have been

generally defined as having two dorsal fins, a short anterior one and

a larger posterior fin continuous with the caudal, similar to the

long anal. The so-called second dorsal proves to be a procurrent

portion of the caudal fin which has extended forward along the

back, and, in the species exhibited, has a base of more than two-

thirds the total length of the fish, and is formed of about 130

rays ; these are supported directly by the neural spines, in striking

contrast to the externally similar anal fin, the rays of which are

attached toa series of interhemals.” Presumably interheemals

are synonymous with radials.

On examining Plotosus, which is a very close relative, I was

pleased to find that the same characters were exhibited in both

cases, or, I believe, practically the same, for although in Plotosus

the rays of this “ procurrent caudal” are not actually attached to

the neural spines but interdigitate between them, they have no

other endoskeletal support, as the anal rays have.

Now, the conclusion to be drawn from the above quotation is

that Mr. Regan would define a caudal fin as that part of the

median fin system which is directly supported by hemal or neural

spines, or in the language adopted in this paper, by hypurals and

epurals alone. It must be pointed out, however, that fin-rays of

true caudal fins are frequently supported by undoubted radials,

in addition to hypurals and epurals, and interdigitating between

these supports; throughout the Gadide this is the case, many

Acanthopterygii show the same thing, as also does Clarias just

considered. But I fully agree with Regan’s remark that the

so-called second dorsal fin is part of the caudal, but for an entirely

different reason. I hope to deal with the definition of the caudal

fin at a future time after my observations have been more extended ;

but for the present, in connection with this discussion I base my

agreement with Regan’s conclusion on the nature of the fin-rays

themselves, and not on their endoskeletal supports. The dermo-

trichia of the caudal fin differ from those of the remainder of

median fins in the method of attachment to their internal

supports ; the caudal fin-rays of Teleosts attach themselves by

means of a forking at their proximal ends, while those of the

dorsal and anal fins are provided with a kind of socket by which

they are attached to the distal segment of the radial. So far as

600 MR. R. H. WHITEHOUSE ON THE [| Apr.5,

T am aware, this difference is universal among Teleosts, but, as

stated above, as yet I do not assert it to be without exception.

The matter is called attention to, in order to make it quite clear

that fin-rays of a caudal fin may be supported directly by un-

doubted radials.

Concerning other features of the caudal fin of Plotosus, the last

dorsal ray is supported by an epural which is attached to the

penultimate vertebra, and which shows the union of two elements,

a radial and a neural arch. Hypaxially is a large hypural firmly

united with the urostyle, and bearing ten dermotrichia. In the

specimen figured the penultimate centrum possesses two ventral

appendages, a hypural supporting one ray, and a hiemal arch.

furiously enough, the next anterior ray is without a support and

seems to anticipate a similar state of affairs ventrally as dorsally :

such, however, is not the case, for each ray preceding is a typical

anal fin-ray supported by a true radial.

Hence the caudal fin of Plotosus is a very highly specialized

organ, totally different from the preceding type, Clarvas.

SYNODONTIS SCHALL. (Siluride.) (Plate XLVIT. fig. 8.)

The last six vertebrae form the basis of support for the deeply

cleft caudal fin in this fish. The urostyle is lost and the last

centrum is firmly fused to the last. two broad hypurals, which

occupy a terminal position. The hypaxial fin-rays are mainly

supported by these two hypurals, but in addition there are five

others, one to each of the preceding five vertebrae ; these two

hypurals afford excellent evidence of the coalescence of radials

and hemal arches in their formation, Epaxially are four epurals,

one to each of the four vertebree preceding the last two; these

epural bones, like the hypurals, show the duplicate nature well ;

it will be seen that the neural arch of the penultimate vertebre

and its corresponding radial have not fused, but a little forward

movement and then the fusion with the neural arch, on the part

of the radial, would produce an exactly similar structure to the

preceding epurals. This caudal fin provides one of the best

illustrations of the fusion of radials with arches in the formation

of epurals and hypurals. Needless to say the fin is a highly

specialized homocercal type.

General remarks on the Siluride.

The single family Siluride, therefore, includes great diversity

in the form of the caudal fin structure, from a very generalized

to a highly specialized type,

APODES.

ANGUILLA VULGARIS. (Murvenide.)

Having obtained a good series of stages, I have been able

to trace the development of the caudal fin in Anguilla from the

Leptocephalus stage to the adult,

1910.) CAUDAL FIN OF THE TELZOSTOMI, 601

Leptocephalus brevirostris. (Plate XLVIII. fig. 9.)

It is perhaps somewhat unfortunate that in the Leptoce) ephalus

all essential skeletal structures are already present; in fact, there

is very little difference between it and the elver four inches in

length. There is very clearly shown a com par: itively long uro-

style slightly directed upwards; this urostyle is not fused with

the hypural bone immediately ventral to it. In my specimen

this hypural, the distal end of which extends beyond the urostyle,

was already a single cartilage, though in older specimens, a very

distinct cleft was frequently to be seen. Three of the four fin-

rays attached to this hypural were bitid distally, though I believe

the number of distally bifurcated rays is a matter of accident.

The penultimate vertebra carries the characteristie U-shaped

hypural of the Apodes, representing two pieces fused proximally.

The antepenultimate vertebra is a Bdoahie structure, and carries

two hemal processes, the posterior of which bears a fin-ray and

shows a definite trace of being formed by the union of a radial

and a hemal arch; this has already been drawn attention to by

Ryder. The double vertebra mentioned is not, as Ryder seems

to suggest, a constant feature, but certainly is very common.

Dorsally, no epural bones are present ; every ray is supported by

aradial. In the only figure of the caudal fin of a young eel I am

aware of, namely that of Rydev’s, the neural arches are not truly

repr esented : in reality they are rectangular plates, equal in

length to fie vertebree to which they ave attached ; even the last

retains its rectangular shape, covering up urostyle and ov erlapping

the last hypural ‘to some extent. The spinal cord can be traced

even among the fin-rays beyond the distal end of the last hypural ;

this fact is evidence of considerable abbreviation.

2. The Elver. (44 ins. or about 12cm. long.) (Plate XLVITI.

fig. 10.)

This stage is next taken as being the last in which the urostyle

can be traced ; the last hypural has already encroached upon it

dorsally, while in specimens five inches long it has vanished, and

the last hypural and the last vertebral segment have become a

single piece. Neural spines have appeared on all the neural arches

except the last two, and a thickening indicative of a spine is seen

there also. The spinal cord does not extend among the fin-rays as

in the Leptocephalus. All other features are much the same

as in the Leptocephalus stage, though naturally more pronounced.

3. Common Hel—adult. (Plate XLVIII. fig. 11.)

After the preceding stage, and sometimes earlier, the last two

neural arches often fuse into one. But the main feature to be

noted in the adult is the complete absorption of the last vertebral

segment, including centrum and urostyle, by the last hypural ; all

efforts to reveal the urostyle by cleaving agents have failed; the

union is complete, hence, what appears in the adult to be hy pural

692 MR. R. H. WHITZHOUSE ON THE [ Apr. 5

only is in reality hypural plus the last vertebral segment. The distal

arms of the U-shaped hypural of the penultimate vertebra often

tend to fuse and so produce an elliptical bone. There is consider-

able variation in the number of fin-rays supported by the various

terminal elements; the terminal dorsal and ventral radial may

or may not support a ray. It is usual for the last two hypurals

to support four rays each, but this too is lable to variation. In

young animals it is quite accidental whether fin-vays bifurcate or

not distally, and it is not safe to conclude, with Ryder, that

bifurcated dermotrichia are caudal proper and others dorsal or

ventral respectively.

ConGER vuLGARIS. (Murenide.) (Plate XLVIII. fig. 12.)

In most respects the caudal fin of the Conger is similar to that

of Anguilla. Only by the aid of clearing agents can the ex ce:

outline of the various elements be traced: : when thus treated,

is seen that the vertebral column proper ends in a long ei:

vertebra, directed posteriorly in a straight line with the vertebral

axis, 7. é. there is no tendency to upturning, and a urostyle cannot

be bad to exist. But all this is invisible without the aid of

cleaving agents, and the last centrum appears to be merely the

proximal end of the three hypurals, which in reality ave firmly

fused with the last centrum. However, any difficulty concerning

the ending of the vertebral axis is removed by a reference to the

young stages: a young conger of 34 inches (9°5 5 cm.) length or

less will show a condition of “things very like those in Anguilla,

viz., 2 urostylar process upw ardly directed, and occupying a

position along the proximal dorsal edge of the last hypural ; in a

specimen 43 inches (or 11:5 em.) long, this could not be seen in

the mere dissection.

Those dermotrichia which may safely be called caudal are

supported by four hypural bones: the first of these is attached to

the penultimate vertebra, instead of the antepenultimate, as in

Anguilla, and the remaining three are all fused to the last

vertebral segment. In this respect, the Conger differs from the

Common Eel in which only one belongs to the last centrum, and

the next two (forming the letter U) to the penultimate vertebra :

this may represent a further shortening of the vertebral axis in

Conger. A word should be said here, calling attention to the

excellent illustration afforded by these hypurals, particularly the

first and second, of their double nature: a line of fusion can be

traced across the hypural dividing the latter into two parts, the

proximal of which is almost exactly similar to the preceding hemal

arches, and the distal part is like a radial.

There are no epural bones. The neural arches closely resemble

those of Anguilla, but are more simple, the anterior ones possessing

a very feeble spine; the last, which is specially long, to protect

the end of the spinal cord, possesses a peculiar bony process

running in an anterio-ventyral direction across the last centrum ;

this is probably a process for the attachment of caudal muscles.

1910.] CAUDAL FIN OF THE TELEOSYOMI. 603

MvuR&NA HELENA. (Mureenidee.)

The caudal fin of the Roman Eel is essentially similar to that of

the Conger, and attention need only be drawn to the peculiarities

presented. The hypural bones are attached to the last centrum.

The spur-like process of the last neural arch, mentioned in Conger,

is more pronounced and fuses with a posteriorly directed process

of the first hypural bone. The posterior neural arches are very

simple, having no spine, and the closely aggregated radials of the

dorsal fin are in close contact with them. The hemal arches very

closely resemble the neural arches : the penultimate vertebra does

not bear a hypural bone.

Myrus vutearis. (Murenide.) (Plate XLVIII. fig. 13.)

The caudal of this fish is again similar to the preceding types.

Only one hypural is attached to the last centrum, which is quite

indistinguishable, appearing only as the proximal end of the

hypural bone: only one hypural may be said to be attached to

the penultimate vertebra, though it is clear that this has been

formed by the coalescence of the distal ends of the arms of the

U-shaped structure in other genera. The hemal arches of the

posterior vertebra are of a horseshoe shape; this may account

for the U-shaped hypural of the penultimate vertebra, which is so

general in the family.

General remarks on the Apodes.

There is a striking uniformity in the structure of the caudal

fin in this sub-order, due possibly to similar habits. In spite of

observations, as early as Huxley’s in 1859, to the contrary, writers

repeatedly refer to the Hels as possessing diphycercal or protocercal

tails. It cannot be too greatly insisted on, that whenever an

upturned urostyle is present, either in the adult or the larva, a

tail is not protocereal or diphycercal. Externally, of course, the

Eels, with their undifferentiated median fin system, do present a

primitive protocercal condition, but dissection proves the terminal

endoskeletal supports to be very unsymmetrical; the broad

hypural bones have no corresponding dorsal homologues. The

caudal fin of the Apodes complies in all its essentials with

the definition of homocerey, which wil! be discussed later; it

cannot be called anything but an advanced form of homocercal

caudal.

While dealing with the Apcdes, it is interesting to note the

references to the ancestral forms. The fossil Urenchelys from the

Chalk beds has a homocereal tail—presumably an externally recog-

nizable homocercal tail—and probably, therefore, much less

specialized a form than the advanced form the modern Apodes

have been shown to possess. Thus all that has happened is a

steady process of specialization, exactly as is to be expected.

604 MR. R, I. WHITEHOUSE ON THE [Apr. 5,

HAPLOMI.

GALAXIAS sp. 2 (Galaxiide.)

Only a single specimen was available for examination; this

revealed : , free urostyle, beyond which the notochord continued

for some ee. but did not reach the dermotrichia. Seven

hypurals are present, five of which are attached to the last

vertebral segment ; the last three are closely applied to each other

and so form a large plate. Dorsally the dermotrichia are supported

by two epural bones and a dorsal caudal radial, the latter being

cartilaginous. Both the epurals and hypurals of the penultimate

and antepenultimate vertebra are very broadly expanded, and

show traces of the inclusion of radials in thei formation. The

last vertebra possesses a small neural arch and the spinal cord can

be traced to the end of the notochord. Such a caudal fin must

be considered a lowly specialized form of homocercy.

No generalizations can be made on the Haplomi until more

species Shave been examined,

HETEROMLI.

Frerasrer Acus. (Fierasteride.)

In this fish the median fins are continuous. At the extreme

end of the tail no fin-rays are present in the fin-fold, so that there

is a gap between the fin-rays of the dorsal and ventral fins. The

vertebral column ends in a perfectly straight line, and, towards

the end, consists of alternate segments of centra and notochord,

the final element being notochord which comes in contact with

the integument. The fin is therefore perfectly symmetrical

externally and internally and moreover shows obvious signs of

degeneration. Ryder concludes that the whole of the true caudal

has been lost, and hence the tail is truly gephyrocercal. It is

interesting to note that the developmental stages do not reveal

any hete rocercal tendencies.

FierasFeR DENTAtUS. (ehiodon dentatus.) (Plate XLVIII.

fig. 14.)

The figure of the tail of this species given by Emery is most

interesting as presenting a typical gephyrocercal tail. ‘The larva

is prov ided with a long filamentous tail-ending, which is eventu-

ally lost in some way, ‘and the adult tail gives the: ‘appearance of

having had the end bitten off; the vertebral column ends ina

vertebra’ ex xactly similar to the preceding ones; the dorsal and

ventral fin-rays attempt to br idge over the gap caused by the loss

of the caudal extremity, but never actually meet. The fin- -rays

of the dorsal and anal fins are supported by very small cartilages

which represent radials, and which are far re snoved from the axis.

I have not yet had an opportunity of examining this candal fin,

and so I reserve any remarks on this rem: arkable form,

1910.] CAUDAL FIN OF THE TELEOSTOMI. 605

General remarks on the Heteromi.

All fishes belonging to this sub-order, whether fossil or living,

appear to be either eel-like or deep-sea forms ; from this it is to be

expected that the tail-fin when present is similar throughout,

i. e. gephyrocercal.

CATOSTEOMI.

GASTEROSTEUS ACULEATUS. (Hemibranchii.)

The caudal fin of this fishy was well described by Huxley in

1859. ‘The vertebral column ends in a long urostyle which is

fused along the dorsal edge of the last hypural bone. There is a

small neural arch to the last centrum, which Huxley figures as

being continuous with it, though there is a line of junction

separating arch and urostyle. The penultimate vertebra possesses

an epural and hypurals ; between the epural and the urostyle is

a dorsal caudal radial (the “ epural” of Huxley).

CENTRISCUS SCOLOPAX. (Hemibranchii.)

The urostyle here is suppressed and the “cone” of the last

vertebra shows only the slightest suggestion of upturning. Two

hypurals are attached and equally well fused to the last centrum.

An epural and hypural are attached to the last vertebra but one,

and both show well that they are composed of a neural and a hemal

arch respectively, together with a radial. A single dorsal caudal

radial is situated between the epural and the last hypural.

Thus the caudal fin of Centriscus is highly specialized in

contrast with the nearly related form Gasterosteus.

Synenataus acus. (Lophobranchii.) (Plate XLVIII. fig. 15.)

The caudal fin of the Pipe-fish presents a perfectly symmetrical

appearance, but morphologically it is quite unsymmetrical since all

the eight dermotrichia are supported by hypaxial elements con-

sisting of two expanded hypural bones. Although the young

stages show a distinct urostyle, this structure is absent in the

adult. There are no complications of any kind and the whole

caudal structure is exceedingly simple. ‘he spinal cord can be

traced to the distal end of the hypural bone. On account of the

presence of a urostyle in the larva and the hypaxial nature of the

fin, the tail must be classed as homocereal; moreover, it is a

highly specialized form of homocercy.

SIPHONOSTOMUS RONDELETH. (Lophobranchii.)

Precisely the same structure is found in this species as in the

preceding type, hence it is unnecessary to repeat the description,

The similarity is no doubt associated with the similarity in habits

of the two fishes.

Proc. Zoon. Soc.—1910, No. XL. 40

606 MR. R. Ht. WHITEHOUSE ON THE [ Apr. 5,

Neroruts oputron. (Lophobranehii.)

This fish has lost its tail-fin probably on account of the

prehensile function of the tail, but other species of Verophis, e. g.

V. equoreus, do possess a very small one, which resembles that of

Syngnathus in miniature.

[LiprocAmpus BREvIROSTRIS. (Lophobranchii.)

The Sea-horse has no caudal fin in the adult, but the larva

possesses a urostyle and hypurals, which degenerate, probably

owing to the prehensile habits.

General remarks on the Catosteomi.

Of the three families of the Catosteomi (Gasterosteide, Cen-

triscidee, and Syngnathide) types of which have been examined,

the Gasterosteide certainly are the least specialized, and the

Syngnathidee the most so, while the Centriscida occupy an inter-

mediate position, as regards the structure of the caudal fin, ‘This

order corresponds exactly with the general classification based on

specialized features as a whole; this bears out what is often seen

elsewhere, that the caudal fin may be useful in classifying fishes

among smaller divisions, though it may not be depended upon as

a broad taxonomic feature.

PERCESOCES.

BeLoxe Acus. (Scombresocidee.)

Only the smallest trace of a urostyle can be seen in the adult.

There are six hypural bones in all, four of which correspond to

the last vertebral segment; the most anterior of these four

possesses a prominent bony process to which muscles are attached.

‘The neural arch to the last vertebra is very large and protects

the spinal cord which can be traced to the distal end of it. The

epaxial fin-supports are lable to some variation : there are some-

times two, and at other times three dorsal caudal radials ; when

two are present, it is on account of the third (the most anterior)

having fused to the neural arch of the antepenultimate vertebr:

and so constituting an epural bone: this point in itself is evidence

as to the way in which epurals may be formed.

The variation exhibited here is probably what Stannius

referred to, in 1854, when he quoted Belone as demonstrating the

fact that the fin-ray supports may be of a double nature, 7. e.

arches plus radials. Lelone, therefore, is interesting as being the

fish in which this feature, so frequently met with elsewhere, was

first seen.

The penultimate vertebra never appears to possess an epural,

as the radial and neural arch are always separate.

On the whole the caudal fin of Belone attains to a fair standard

1910. | CAUDAL FIN OF THE TELEOSTOMT, 607

of specialization, particularly as regards the suppression of the

urostyle; but at the same time the number of hypurals, viz. four,

associated with the last vertebra is indicative of a comparatively

low type of specialization. The proximal ends of the fin-rays

embrace their endoskeletal supports so as to almost exclude them

from view.

AMMODYTES TOBIANUS. (Ammodytide.) (Plate X LIX. fig. 16.)

The last vertebra and last hypural are completely fused and

thus there is no free urostyle. ‘lhe hypaxial supports consist of

five hypurals, four of which ave attached to the last vertebra.

The dorsal elements are somewhat complicated : lying between

the single dorsal caudal radial and the last vertebra and hypural

are four separate bones, through three of which the spinal cord

passes ; these three possibly represent neural arches, while the

fourth may be a dislodged arch, or a spine detached from its

arch. It is interesting to notice that fusion of these four bones

into a single piece would form a large neural arch exactly as in

Belone.

ATHERINA BAYERII. (Atherinide.)

A small but free urostyle is found here ; connected with it and

the last centrum are five of the seven hypural bones supporting

the hypaxial dermotrichia, A. small cartilage, situated between

the distal ends of the first two hypurals, shares in the support of

the anterior rays of the ventral lobe ; this may possibly represent

a radial, but it is doubtful. Dorsally the fin-rays are supported

by two dorsal caudal radials and one epural bone, the latter

attached to the antepenultimate vertebra. The neural arch of the

last vertebra is very large and is composed of three bones: it

affords protection for the end of the spinal cord. Thus A therina

possesses a fairly generalized caudal fin, lacking only a ventral

caudal radial, though even this may possibly be represented by

the small cartilage mentioned.

The development of the caudal fin in this fish is interesting as

being almost identical, in the earlier stages, with that of the

Flounder as figured by Agassiz, and shows very clearly the deep

incision between the original and the permanent caudals.

Muar carrito. (Mugilide.)

The urostyle is lost in the tissue of the last hypural bone ;

three hypurals in all are attached to the last vertebral segment ;

three dorsal caudal radials are present, the last of which is applied

to the dorsal side of the last hypural. The neural arch to the

penultimate vertebra is very much reduced. The anterior fringe

of the ventral lobe of the caudal is supported by small cartilages ;

what these 1epresent it is difficult to determine. ‘lhe fin there-

fore may be classed as an advanced type of homocercy.

40*

608 MR. R, H. WHITEHOUSE ON THE [ Apr. 5,:

General remarks on the Percesoces.

This sub-order presents considerable diversity in caudal fin

structure ; Atherina, with its free urostyle and numerous hypurals

attached to it, possesses a comparatively lowly specialized caudal,

while that of Mugil is quite a specialized form. Arranged in

ascending order of homocercy, the four genera will appear thus :

Atherina, Ammodytes, Belone, Mugil, which arrangement does not

correspond with that of general classification.

ANACANTHINI,

Macruride.—The fishes in this family possess no caudal fin ;

they are deep-sea forms and the body ends in a finely tapering

tail.

Mo ya vuuearis. (Gadide.) (Plate XLIX. fig. 17.)

The Ling possesses a simple type of Gadoid tail. No less than

thirteen vertebre contribute towards the support of the caudal fin ;

all of these vertebrae ave provided with hypural bones (Ay.) and

all, except the last two, with epural bones too; special attention,

therefore, need be directed only to parts connected with these

last two vertebre. The final vertebra possesses no urostyle,

and is firmly fused with the broad last hypural which occupies

a terminal position and bears five dermotrichia, That part of

the last hypural which encroaches upon the dorsal side of the

last centrum is seen in other Gadide, e. g. G. minutus, to be a

secondary extension of this bone, only appearing late in develop-

ment; in young stages the hypural is totally ventral to the

centrum. The hypural bone of the penultimate vertebra is also

broadly expanded and bears three fin-rays. Between this hypural

and the next anterior one is a ventral caudal radial (v.c.7.), in no

way fused with any other skeletal element. Most of the hypural

bones show signs of being formed by the coalescence of hemal

arch and radial, the line of fusion being specially prominent after

clearing agents have been used; the shading in the figure is

intended to show this.

Epaxially, it is interesting to note the great reduction of the

neural arch of the penultimate vertebra ; that of the antepenul-

timate contributes to the formation of an epural bone. Between

this last epural and the dorsal side of the last hypural are two

dorsal caudal radials, each bearing a fin-ray; these radials never

fuse with the neural arch, but always remain perfectly free. It

will thus be seen that the caudal fin of Molva is not symmetrical

internally and that the hypaxial fin-rays outnumber the epaxial ;

it is in every way consistent with homocercy, and its terminal

skeleton with a very highly specialized type of homocercy.

GapicuLus ARGENTATUS. (Gadidee.) (Plate X LIX. fig. 18.)

This small Gadoid possesses a more generalized caudal fin than

does the preceding type; this is due solely to the presence of a

1910.] CAUDAL FIN OF THE 'TELEOSTOMI. 609

large number of radials in addition to the epural and hypural

bones. Except for the fact that eleven vertebre form the basis

cf support for the fin instead of thirteen as in Molva, all remarks

made in reference to the latter genus apply equally to Gadiculus.

The whole interest in this tail-fin lies in the radials; in addition

to the epurals no less than six are present epaxially, and ventrally

three are to be seen alternating with hypurals. Moreover, these

radials may be either quite free, like ordinary radials, or they

may be more or less fused with the neighbouring hypurals or

epurals. This affords one of the strongest cases for the theory

that hypural bones are formed by the coalescence of radials with

hemal arches. In the other median fins two radials are present

between successive vertebral arches; thus probably the hypurals

and epurals of the caudal have already absorbed one radial and

the tendency is to effect a fusion of the second also. There can

be no doubt about these radials being true radials and not a

second neural or hemal arch, since they never reach the centrum

itself and never embrace the spinal cord or heemal vessels at their

proximal ends, as they would do if they were arches. ‘The spinal

cord can be seen to extend to the distal end of the last hypural

bone. This caudal also is therefore a homocercal form.

General remarks on the Gadide.

There seems to be a certain amount of misconception among

writers concerning the caudal fin of the Gadide; one or two

quotations will make this clear. J.T. Cunningham, in ‘ Science

Progress,’ 1897, says :—‘“‘ The structure and development of the

tail described above (heterocercy) occur in the flat fish, but are

entirely wanting in the Gadide. In the latter the tail is per-

manently diphycereal and is composed of dorsal and ventral rays

which are equal in number and size, and, in fact, closely resembles

the tail of the extinct Ceelacanthide. There can be little doubt

that even if the Gadide cannot be directly derived from the latter

family, they are descended from Crossopterygian Ganoids with

diphycercal tails and have never passed through a heteroce pica

condition.” And Boulenger, in the Ann. & Mag. Nat. Hist. x

1902, writes :—‘‘ I have reason to believe that Gadoids must have e

been ‘derived from such a group as the Berycide, through forms

of which the Macruridee with thoracic ventral fins composed of

seven to twelve rays are the nearest known examples, and in

which the caudal fin has entirely vanished. I regard the isocereal

condition of the Gadide as the result of the formation of a new

caudal fin, the homocercal extremity of the vertebral column

having been lost by the direct ancestors of these fishes.” The

same writer in comparing the caudal fin in the Pleuronectide

with that of the Gadide says: (in the Pleuronectid) ‘ the tail,

whenever a caudal fin is well dey eloped, belongs to the homocereal

type (heterocercal in the embryo) with comparatively few rays

(20 or less),”—thus differing from the Gadoids.

610 MR. R. H, WHITSHOUSE ON THE f Apr. a;

Other references might also he quoted, but these will be

sutticient to show what misconceptions are held. Dealing with

Cunningham’s remarks first, a reference to either of the two

figures given will show at a glance that the tail is not “* perman-

ently diphycereal” and is not ‘“ composed of dorsal and ventral

rays equal in number.” In all, eight different species have been

very carefully examined, and any of the following remarks apply

equally to all; there are always more hypaxial fin-rays than

epaxial ; this is due to the fact that a broadly expanded ‘hypural

bone is attached to the last centrum. Thus the tail is a highly

specialized homocercal form.

This author has apparently never been fortunate enough to

secure young forms of Gadide for detailed examination of the

‘audal fin structure or he would never have said that during the

development the heterocercal stage is “ entirely wanting.”

Gadus minutus of one inch length shows an upwardly directed

urostyle along the dorsal edge of the last hypural as clearly as in

the flat-fish; this becomes much reduced in the adult and is

in fact almost completely lost; and since the extremity of the

vertebral axis is almost precisely the same im all adult Gadide,

it is quite reasonable to suppose that the young forms in every

case ave sinilar to G. minutus. Cunningham's error in this

connection Jed him to suppose that the Gadide ‘are descended

from Crossopterygian Ganoids with diphycercal tails and have

never passed through a heterocereal condition.” The latter

statement, however, falls through.

Assuming Cunningham to be correct as regards the diphycercal

condition, Boulenger comes to the conclusion that the Gadide are

descended from a homocereal form, such as the Berycidse, but

that homocercy was entirely lost, and the “ isocercal (i. e. os

diphycereal) condition .... is the result of the formation of :

new caudal fin.” But development proceeds just as in any highly

specialized homocercal form, and hence the caudal fin of the

Gadidee cannot be considered a new thing.

The caudal fin of this family further conforms to advanced

homocerey in the great reduction of the neural arch of the

penultimate vertebra ; in all previous figures of the Gadoid tail,

the last vertebra but one is shown bearing an epural bone ; but

in all the Gadide examined, this has proved not to be the case;

immediately dorsal to the penultimate vertebra there are always

two dorsal caudal radials perfectly free and never fused with any

other part of the neighbouring skeleton. Hypaxially, the same

vertebra always bears a hypur: al bone.

Thus the caudal fin of the Gadidwe is undeniably a specialized

form of homocerey.

ACANTHOPTERYGITI.

A preliminary note will be necessary before entering into

details of the types chosen in this sub-order, In many forms

in which a free urostyle is present, there is a fin-ray support

.. va

1919.] CAUDAL FIN OF THE TELEOSYOMI, 611

which occupies a position exactly in line with the urostyle ;

a difficulty comes about, therefore, in determining whether this

element is hypaxial or epaxial. In shape it closely resembles the

dorsal caudal radials, but the spinal cord runs along its dorsal

edge; it may therefore be considered a dorsal caudal radial which

has moved ventral to the spinal cord; but its present position

below the spinal cord makes it possible also that it is a hypural

bone, or perhaps a ventral caudal radial. For the present,

however, I have decided to call it a hypural in order to simplify

description, but the reader may consign it to any of the three

suggested classes he may consider desirable.

SERRANUS CABRILLA. (Perciformes, Serranide.) (Plate XLIX.

x, 19.)

The vertebral column ends in a_ well-developed and free

urostyle (u.), which is about the length of an ordinary vertebra.

Six hypural bones (Ay.) of varying size are associated with the

last vertebral segment ; the most anterior of these possesses a

long spur-like process, running parallel with the urostyle, and

undoubtedly functioning as an attachment for muscles; the last

hypural is in direct line with the axis of the urostyle. Between

the distal ends of the hypurals of the second and third last

vertebra is a small cartilage (¢c.) which shares the support of the

smaller anterior ventral caudal fin-rays. Dorsal to the urostyle

and extending beyond it, is a slender splint-like bone (7.a.')

protecting the delicate end of the spinal cord; this probably

represents one or more neural arches of vertebre now absorbed

into the urostyle. Independent of this bone, the last vertebra

(/.v.) possesses a neural arch of its own; the neural arch of the

penultimate vertebra (p.v.) is much reduced, and the ante-

penultimate vertebra is provided with an epural bone (ep.).

Three dorsal caudal radials (d.c.r.) support the majority of the

epaxial fin-rays.

This caudal fin, therefore, is a fairly typical form of a com-

paratively lowly specialized homocercal type.

CEPOLA RUBESCENS. (Perciformes, Cepolidz.)

This fish possesses a long tapering form of tail. The urostyle

is prominent though slender; four hypural bones are attached to

the last vertebral segment, the last overlapping the urostyle

slightly. All the dermotrichia of the caudal fin’ ave hypaxial,

though epaxially there is a bone, not actually supporting a ray,

which might be calleda radial. Externally the median fin system

appears to be a continuous one, but closely examined, there is to

be found a very distinct and separate caudal. All the skeletal

structures in the terminal end of the tail are very delicate and

loose, the neural and hemal arches not being actually fused with

the elongated centra. The tail-fin is, however , definitely homo-

cereal,

612 MR. R. H. WHITEHOUSE ON THE [ Apr. 5,

CANTHARUS VULGARIS, (Perciformes, Sparide.)

The urostyle is well marked and free, and four hypurals are

attached to the last vertebral segment; in all there are six

hypurals, the second and third last vertebre each bearing one.

Epaxially, there are three dorsal caudal radials, and one epural

which is attached to the antepenultimate vertebra, The neural

arch of the penultimate vertebra is reduced, Between the last

dorsal radial and the last hypural is a splint-like bone, the

proximal end of which rests on the dorsal end of the urostyle ; it

protects the ending of the spinal cord and is probably a “neural

arch. The fin, therefore, is homocercal.

Box saALpa. (Perciformes, Sparide.) (Plate XLIX. fig. 20.)

The urostyle is well marked and distinct. Corresponding with

the last vertebral segment are three hypurals, two of which are

large, and the most anterior is provided with a ridge for the

attachment of muscles; there are five hypural bones in all ;

between the distal ends of the first two is a small cartilage

bearing a few of the small anterior hypaxial rays; a very small

similar one is present on the anterior side of the first hypural.

Epaxialiy, there are three dorsal caudal radials (d,c.7.), while the

antepenultimate vertebra bears an epural bone; this epural is

particularly interesting from the fact that in an animal 4 em.

long (fig. 20) it is seen to be two distinct and separate pieces—

a neural arch, exactly similar to that of the penultimate vertebra,

and a long cartilaginous radial, the proximal end of which is

slightly applied to the posterior edge of the neural spine. In the

adult, these two elements completely fuse to form an epural.

A neural arch is present on the last centrum.

The caudal fin of Bow is therefore homocercal, and slightly

more specialized than that of Cantharus, on account of the few er

hypurals to the last vertebral segment.

SMARIS ALCEDO. (Perciformes, Mzenidz,)

A fairly long and free urostyle is present, on the ventral side

of which, toge ther with the last centrum, are four fin- vay supports,

three only of which are undoubted hypural bones : the last is

somewhat doubtful. Both the penultimate and antepenultimate

vertebrie bear hypurals, between the distal ends of which is a

Jarge cartilage bearing small fin-ravs. Separated from this car-

tilage only by the distal end of the first hypural are two small

cartils ages, both of which support the very small anterior epaxial

fin-rays; a similar cartilage dorsally performs the same function.

Three definite dorsal caudal radials are present and one epural

bone; neural arches are borne.by each of the last two vertebrie.

\ splint-like bone is closely fitted between the distal ends of the

urostyle and the neural arch of the last centrum; probably it is

a neural arch itself, belonging to some vertebra now absorbed

into the urostyle, Cartilaginous epiphyses are present on most

1916. | CAUDAL FIN OF THE TELEOSTOMI, 613

of the fin-ray supports, and here and there are additional distal

cartilages applied to these epiphyses; they may represent the

persistent distal segments of radials.

This caudal fin is a good intermediate form between the lowly

and the highly specialized homocereal types. i

Lasprus Festivus. (Perciformes, Labride.) (Plate XLIX.

fig. 21.)

No urostyle can be seen in this form, it having completely

fused with the last hypural bone; thus the last hypural, urostyle,

and last centrum (/.v.) become a single bone. The last centrum

bears one other hypural similar in size to the last ; the preceding

centrum also bears a hypural bone. Between the hypural of the

penultimate vertebra and the first of the last centrum is a large

bone (v.¢.7.), Which on account of its not having actually fused

with the centrum, is probably better referred to as a ventral caudal

yadial. Three bony dorsal caudal radials (d.c.7.) are situated

between the neural arch of the antepenultimate vertebra and the

last hypural bone. In an exactly corresponding position dorsally

and ventrally (¢. e. on either side of the distal end of the neural

spine and hemal spine of the antepenultimate vertebra) are

two cartilages (c.) the posterior of which may in some species,

e. g. Labrus turdus, become quite radial-like. The extremity

of the spinal cord is protected along the dorsal side of the last

hypural by a secondary bony development (s.).

Though the structure of this caudal fin approaches closely to

secondary symmetry as regards general form, it is really not

symmetrical owing to the fact that by far the majority of

dermotrichia are hypaxial. By the loss of the urostyle and the

reduction of the number of hypurals to the last vertebral segment

to two, this caudal fin is a considerable advance in homocercy on

the preceding type.

SCOMBER SCOMBER. (Scombriformes, Scombride.) (Plate XLIX.

fig. 22 )

The tail of the Mackerel is somewhat difficult to interpret at

first sight as regards the ending of the vertebral axis; it appears

as though a well-marked urostyle were present extending to the

length of the broad hypural bones. Examined more closely,

doubts will arise as to its true urostylar nature for one or two

reasons :—(1) its end is cartilaginous and it bears dermotrichia ;

(2) its distal end is cut square, a feature no urostyle elsewhere

possesses ; (3) the end of the spinal cord enters slightly into the

tissue of the proximal end of the bone. Another feature worth

notice is that clearing agents do not reveal any indications of a

normal urostyle fused with any other bone, and moreover, the

conical last centrum does not turn upwards as a hook or reduced

urostyle as so many other forms show, but ends in a direct

straight line. The tail of Zrachinus is the only other one

examined which closely resembles Scomber in the termination of

614 MR. R. WL. WHITEHOUSE ON THE [Apr. 9,

the axis, and clearing agents show, in this type, an oblique line

of fusion in the proximal end of this rectangular bone; this

suggests that the distal end is hypural, and the proximal, at least

partly, is urostyle firmly fused with the hypural. It may be so

in Scomber too, but the fact that the spinal cord enters the

proximal end suggests that a neural arch is also involved. And

this seems to be the inter pretation, for it cannot be urostyle

alone, since it is cut square, tipped with cartilage and bears

dermotrichia just as an ordinary hypural does. It seems safe,

therefore, to say that the urostyle is greatly reduced and fused

entirely with the last hypural—in this case a smallone. In all,

then, four hy purals are associated with the last vertebral segment ;

the most anterior being provided with a ridge for the attachment

of muscles. Two dorsal caudal radials are present. The penulti-

mate vertebra possesses a reduced neural arch and a well-developed

hypural, while the antepenultimate vertebra bears an epural and

a hy pural bone. All the hypurals are firmly attached proximally ;

the dermotrichia embrace the terminal endoskeletal supports to

their base ; thus the fin forms a most efficient organ of locomotion,

seen to advantage in the swift swimming Mackerel.

Zeus FABER. (Zeorhombi, Zeide.) (Plate X LIX. fig. 23.)

In this fish, the John Dory, a fairly typical flat-fish form 1s

met with. The urostyle is lost to view; clearing shows it to be

present as a minute hook embedded in the tissue of the last

hypural. The three terminal hypurals (Ay.) are fused with the

last centrum (/.v.) into a single piece, though the outlines of

the hypurals still remain visible. ‘The last two complete centra

each have a well-developed epural (ep.) and hypural bone, and

the former are specially interesting inasmuch as they show clearly

that the epural is formed by a fusion of radial with neural spine,

apparently by means of a secondary ossification. The last centrum

has a small neural arch (7.a.) developed. Of other supports of

dermotrichia there are two dorsal caudal radials (d.c.r.) between

the last epural and hypural, and also a ventral caudal radial

(v.c.7.) immediately anterior to the large terminal hypural plate.

The caudal fin, therefore, is a speci ialized homocereal type.

SoLEA LUTEA. (Zeorhombi, Pleuronectide.) (Plate XLIX.

fig. 24.)

Solea lutea may be taken as possessing a good representative

type of Ple uronectid caudal fin. It has a more delicate build,

but essentially is very similar to that of Zews. A clearing agent

shows the otherwise invisible urostyle to be a mere hook-like

upwardly directed termination of the last centrum. The four

hypurals of this centrum are firmly united, though less so than

in Zeus, and oceupy a terminal position, The penultimate vertebra

has a well-developed epural (ep.) and hypural (Ay.) bone; since

reference has been made by Cole to the corresponding epural and

hypural in the Plaice, it is interesting to compare them in the

1910.] CAUDAL FIN OF THE TELEOSTOMT. 615

two species. In the Liverpool Biological Committee Memoir on

the Plaice, Cole says, on p. 50:—‘‘In the last distinet vertebra

(z.e. the penultimate vertebra) it will be seen that the neural

spine resembles the hemal spine in structure, but that both are

pecuuiar. Hach consists of a partly cartilaginous shaft behind

and a thin laminate portion in front. The posterior shafts so

closely resembie the succeeding epural (7. e. dorsal caudal radial)

and hypural (7. e. ventral caudal radial) bones respectively as to

suggest that an epural above and a hypural below have fused on to

the laminate portions, which latter are undoubtedly similar to and

perhaps iepresent the neural and hemal spines in front. As,

however, we have no positive evidence of such a fusion, the spines

in question are here described as simple neural and hemal

spines.” Though in Solea lutea this point is not so obvious as in

Pleuronectes, there is to be seen a distinct fusion line running

transversely across the proximal part of the epural, so that a

parallelism does occur between the two species. I take it that

Cole reserves committance on the subject on the ground that

actual fusion has not been observed during development ; but it

is scarcely to be expected that such a minute feature as the fusion

of a radial with a neural or hemal spine will be recapitulated in

development, so that lack of embryological evidence would not

be fatal to the theory, But we have seen in the case of Box salpa

that such a fusion can be traced in development ; had Cole been

acquainted with this as well as numerous other examples re-

sembling that of the Plaice, he doubtless would have considered

these epurals and hypurals more than mere neural and hemal

spines.

To complete the description of the caudal fin of Solea, there

are two dorsal caudal radials (d.c.r.) between the epural and

hypural bones; in the corresponding position ventrally there is

a ventral caudal radial (v.c.r.). The last eentrum bears a neural

arch (7.a.) which is not, as may be at first supposed, a urostyle,

since the spinal cord (sp.c.) passes through it, m the fashion of

other neural arches, in its course along the dorsal edge of the

last hypural.

The caudal fin, then, is a homocercal type of an advanced

nature. It may be mentioned that other species of Solea, e. g.

S. monochir and S. impar, differ from S. lutea only in the number

of terminal hypurals and the extent of their collateral fusion.

GoBIUS PAGANELLUS. (Gobiiformes, Gobiide.) (Plate L.

fig. 25.)

The urostyle (wr.) is greatly reduced, and almost non-existent

and fused with the last hypural bone. ‘There are three hypurals,

the last two, the largest, being fused to the last vertebral seg-

ment. A small trace of neural arch is present on the last

centrum and that of the penultimate is also reduced. Dorsally

there are two caudal radials (d.c.r.) and ventrally one (v.c.7.), the

latter lying between the first and second hypurals. The spinal

616 MR. R, H. WHITEHOUSE ON THE [Apr. 5,

cord can be seen to extend along the whole length of the last

hypural.

Cartilage enters very considerably into this caudal fin, Both

the anterior dorsal and anterior ventral fin-rays are supported

by cartilage which is somewhat irregularly disposed, and which

extends inwards even to the vertebral axis. It is the irregular

nature of these cartilages that has led me to attach no importance

to them as homologues of ordinary supporting elements such as

radials, but merely to regard them, both here and frequently

elsewhere, as convenient secondary developments to support the

less important dermotrichia.

I have been fortunate in obtaining a fairly good series of

developmental stages of Gobius, and the larvee show exceedingly

well how a condition in which the urostyle is very much reduced

in the adult, is preceded by stages in which this structure is very

prominent; and how, by a secondary encroachment of the last

hypural bone, the urostyle is incorporated with this hypural.

The caudal fin of Gobius paganellus i Is a type of considerably

advanced homocercy.

ECHENEIS NAUCRATES. (Discocephali.)

A long well-developed urostyle is present, attached to which,

together with the last centrum, are six hypural bones closely

fitting together. A well- developed neural arch is closely applied

to the dorsal edge of the last vertebral segment ; the penultimate

vertebra possesses a hypural and an epural, and immediately pos-

terior to the latter are three dorsal caudal radials, slenderly built.

This fin must therefore be considered a lowly specialized homo-

cercal caudal.

ScorP£NA scRoFA. (Scleroparei, Scorpenide.) (Plate L.

fig. 27.)

The vertebral axis ends in a well-marked and free urostyle.

Three undoubted hypural bones are attached to the last vertebral

segment, while in a direct line with the axis of the urostyle is a

slender vay-bearing bone which may be considered a bhypural or

a radial. Between this bone and the epural (ep.) attached to

the antepenultimate vertebra are three dorsal caudal radials

(d.c.r.). The neural arch of the penultimate vertebra is reduced ;

there is also a slender bone (v.@.') immediately above, and

closely approximated to the urostyle, which protects the delicate

ending of the spinal cord ; it is probably, therefore, a neural arch

of some vertebra now absorbed into the urostyle. Both the

second and third last vertebrae bear a strong hypural bone. This

caudal fin is an excellent example of a form intermediate between

a lowly and a highly specialized type.

TRIGLA LINEATA. (Scleroparei, Triglide.) (Plate L. fig. 28.)

The urostyle here is practically non-existent, and traces of it

can be seen only after the tail has been treated with a cl aring

1910.] CAUDAL FIN OF THE TELEOSTOMI, 617

agent; thus the last vertebral segment and the last hypural

become one solid structure. Three more hypurals are attached

to the last vertebral segment and one is also attached to the

preceding centrum. The whole of the dorsal contribution to

the caudal fin is supported by four caudal radials. The neural

arch to the penultimate centrum is much reduced. Though

classified in the same division of Acanthopterygil as Scorpena,

Trigla provides a striking contrast to this form in the degree of

specialization of the caudal fin, being quite a highly specialized

type.

URranoscopus scaBer. (Jugulares, Trachinide.)

The urostyle is absent or indistinguishably fused with the last

hypural bone. Hypaxially, the last centrum bears three hypural

bones, and epaxially a well-developed neural arch. The pen-

ultimate vertebra possesses a hypural bone, but its neural arch is

greatly reduced; the antepenultimate vertebra bears both a

hypural and epural. Between the epural bone and the neural

arch of the last centrum are three dorsal caudal radials, while

between this same neural arch and the hypural which is fused

with the last centrum is a ray-bearing bone which is probably an

epaxial element, though situated ventral to the spinal cord, since

it is usual for the urostyle to fuse with the last, if any, of the

hypural bones. This caudal fin, therefore, attains to a fairly

high standard of homocercy.

CALLIONYMUS LyRA. (Jugulares, Callionymide.) (Plate L.

fig. 26.)

The caudal fin of Callionymus is a very simple type. No

urostyle is present ; the last vertebral segment has three hypurals

firmly attached to it and to each other, thus forming a single

piece, to which dorsally is also fused the neural arch oF the last

vertebra. The penultimate vertebra bears an epural (ep.) and a

hypural, and one dorsal caudal radial (d.c.7.) is present.

CRISTICEPS ARGENTATUS. (Jugulares, Blenniide.) (Plate L.

fig. 29.)

The urostyle is present only as a small hook embedded in the

tissue of the last hypural bone. Hypaxially, the last vertebral

segment possesses two hypurals of about equal size, while dorsally

it shows traces of two neural arches, through which the spinal

cord passes on its way some distance beyond the end of the axis.

The penultimate vertebra possesses an epural and a hypural ;

posterior to this is a large dorsal caudal radial. Immediately

adjoining the distal end ro the last hypural, and on its dor sal

side, is a small ray-bearing cartilage, and another one on either

side of the dorsal radial. Similar cartilages are to be found

ventrally in a corresponding position. ‘This caudal fin, therefore,

will be regarded as a highly specialized homocercal type.

618 MR. R. H. WHITEHOUSE ON THE [Apr. 5,

OpniruM BARBATUM. (Jugulares, Ophidiide.) (Plate L.

fig. 30.)

All the median fins of this fish are continuous with one another,

and the caudal proper may in this particular case be limited to those

dermotrichia which are not supported by the typical radial or

somactid of the dorsal and anal fins. This confines the caudal

rays to those eight or nine supported by a large terminal hypural,

often deeply and irregularly cleft, which is firmly coalesced with

the last vertebral segment ; the urostyle therefore is reduced to

the merest vestige. A diminished neural arch is to be seen on

the last centrum, and the nerve-cord runs along the whole length

of the dorsal edge of the hypural. ‘Thus the caudal fin itself is

exceedingly simple in form and very highly specialized. The

neural and hemal arches of the penultimate vertebra have a

cartilaginous shaft fused to their posterior side ; in reality, there-

fore, they may be considered epural and hypural respectively,

but bearing no fin-rays.

General remarks on the Acanthopterygian Caudal Lin.

A suflicient number of types from various divisions of the

Acanthopterygii have been dealt with to show how, in a single

sub-order of Teleosts, the structure of the tail- fin may vary.

Even in the same division of the sub-order, vastly different types

are found; for example, in the Perciformes, the genera Serranus,

Bow, Cantharus, and others have features characteristic of the

less specialized homocercal tail, such as a long and free urostyle

and a large number of hypural bones attached to the last vertebral

snommenitise on the other hand, the Wrasses (Labridze) have lost

these lowly specialized features and attained to the rank of high

specialization. The Scleroparei provide a similar contrast,

Scorpena having a much Jess advanced type than Zrigla. It

would be difficult to account for such variations between types

otherwise so closely related, except on physiological grounds, but

it indicates that the tail-fin is not a safe guide, or rather, a

wholly satisfactory character, by which fishes can be classified,

In the main, however, advanced homocercy is exhibited among

Acanthopterygian caudals, except for the Perciformes; this

division possesses the greatest number of less specialized tails

(the Labride are the only family with much exception), but it is,

as a division, the least specialized among the Acanthopterygii.

From the foregoing descriptions it will be seen that the Jugulares

possess the most advanced types of homocercy ; of the six genera

which have been subjected to examination, "viz, Uranoscopus,

Trachinus, Lepadogaster, Blennius, C ‘risticeps, and Ophidium, all

possess advanced features in the cauds ilfin; but the Jugulares are,

as a division, among the most specialized of the Ac: anthopter ‘yeu,

‘The Zeorhombi occupy an intermediate position, and all members

have very similar tails: they have a greatly reduced urostyle

fused with a hypural, and also a large number of hypurals to the

1910.] CAUDAL FIN OF THE TELEOSTOML. 619

last vertebral segment ; thus there is an advanced feature (the

former) and a lower feature (the latter) both present in the same

caudal—in other words, an intermediate condition in specialization

is present, which is consistent with the intermediate position the

dlivision occupies among the Acanthopterygii.

It would, therefore, seem that each individual division has its

own grades of specialization within itself, and in this minor

classification the structure of the tail does correspond with the

general structure of the fish in respect to specialized features in

general. In the Perciformes, the Serranide are among the least

specialized of the several families, and the Labride are at the

other end of the scale; and it has been pointed out that members

of the former family have less specialized tails, by far, than those

of the latter; thus caudal features correspond with specialized

features in general, within this division. Exactly the same

thing occurs among the Seleroparei, where the least specialized

members are the Scorpzenide and one of the most specialized

the Triglide ; the Scorpenide have lowly and the Triglidx

highly specialized caudal fins.

The Scleroparei are a more specialized division than the Perci-

formes, and yet Scorpena, among the former, has a less specialized

tail than that of Zabrus in the latter division. Thus while the

caudal fin may not be depended upon as a feature for classification

in a broad sense, it may be useful within the scope of a single

division,

OPISTHOMTI.

MastTAcEMBELUS sp. ¢ (Plate L. fig. 31.)

This fish possesses a continuous undifferentiated median fin ;

the urostyle is well-marked and free, and equal in length to a

vertebra. The limits of the caudal fin proper are probably best

fixed here by those dermotrichia supported by hypural bones, of

which two are associated with the last vertebral segment, the

anterior one being obviously composed of three fused collaterally :

and one with the penultimate vertebra, All the remaining fin-

rays of the median fin are borne by radials; no epurals are

present. A trace remains of the neural arch to the last vertebra.

Thus the tail is definitely homocercal ; in regard to this, it is

strange that authors refer to it as a gephyrocercal tail. The only

possible foundation for so naming this type is the apparent

symmetry due to the confluent tins; it must, however, be insisted

on that gephyrocercy indicates true secondary symmetry, 7. e. an

abbreviated protocercal condition, It is obvious that such a tail

as that of Mastacembelus cannot be referred to the same category

as that of Mierasfer.

PEDICULATY!.

Lopuius piscatorius. (Plate L. fig. 32.)

This fish may be said to illustrate the extremest type of

advanced homocerey. Practically only one hypural is present,

620 MR. R, H. WHITEHOUSE ON THE [Apr. 5,

though this is somewhat cleft distally ; this structure is the sole

support of the eight caudal rays; the last vertebral segment has

become most completely fused with the hypural. The whole fin

presents a most compact and solid structure. The neural and

hzemal spines form an imbricate series, each overlapping the one

behind.

PLECTOGNATHI.

BauistEes cApriscus. (Sclerodermi.) (Plate L. fig. 33.)

No trace of a urostyle remains, and two large hypural bones

are firmly united with the last centrum. For such an advanced

condition of homocercy an irregularity occurs here, for the pen-

ultimate vertebra possesses an epural bone (ep.) instead of the

usually greatly reduced neural arch; the penultimate vertebra

also bears a hypural bone. Two dorsal caudal radials (d.c.7.) ave

closely wedged between the epural and last hypural, and a ventral

‘audal radial (v.c.7.) is found between the first and second hypural.

The whole fin presents an apparent symmetry.

4, GENERAL CONCLUSIONS.

There is every reason for believing that the most specialized

caudal fins have been evolved from a common primitive type,

which is referred to as protocercal ; modifications of this produced

the heterocercal form, which in turn was followed by the homo-

cercal type.

Prorocercy.—A protocercal fin may be defined as one which is

primitively symmetrical externally and internally, and hence one

which has not undergone any reduction or modification of the

original form. It is very probable that the protocercal tail is not

found in theadult condition among any living fishes. The Cyclo-

stomes are sometimes credited with having a protocercal caudal,

but it may be that it is a degenerate condition, and not a per-

sistent original structure. Since the original protocercal caudal

fin was continuous with the dorsal and anal fins, and thus

undifferentiated, it is not unreasonable to suppose that the con-

tinuous fin was supported similarly throughout, 7. e. by a series

of radials.

It must not be supposed, however, that protocercy is always

associated with a continuous median fin ; during development in

Teleosts the elements of the permanent differentiated median fins

are laid down before heterocercy is established, so that there is

evidence for supposing that differentiation of the median fins

occurred in adult forms before heterocercy was adopted.

Hererocercy.-Confining attention to typical forms, the

heterocercal caudal fin may be defined as one in which the

extremity of the chorda is directed upwards, and which as a

consequence is unsymmetrical, the lower lobe being larger than

the upper, and, moreover, one in which, when centra are formed,

these elements remain distinct to the end of the axis.

1910.] CAUDAL FIN OF THE TELEOSTOMI. 621

Reference to fig. 1 (Pl. XLVII.) will render a full description

unnecessary ; this form may be considered quite typical except in

possessing what may be regarded as an incipient urostyle at the

extreme end of the axis. It should be observed that on the

dorsal side the separation of the radials from the neural arches

anteriorly, and their gradual approach to and ultimate fusion

with them posteriorly, suggests that when the median fins were

continuous there was a eradual transition, from before backwards,

from widely separated radials to fusion of radials and arches.

There is evidence that the same thing occurred ventrally, but the

fusion of radials and arches has taken place much further forward

owing to the fact that the enlarged lobe required a much firmer

base of support; several Kiasmobranchs retain the line of fusion

in the adult (e. g. Galeus and Acanthias).

Homocercy.—The homocereal type of caudal fin is characteristic

of the Teleostei, and is a specialized form of heterocercy. Though

the term is very broad in application, a tail-fin which is externally

symmetrical, but in which the majority of the fin-rays are

supported internally by hypaxial elements, and in which a true

urostyle is present either in the adult or some time during larval

history, may safely be referred to as a homocercal caudal fin.

Special attention has been directed towards ascertaining which

features may be used to determine the degree of specialization

exhibited by the homocercal caudal fin ; these characters may be

enumerated as follows :—

I. Characters indicative of a low degree of specialization :—

(a) Notochord exserted beyond the actual vertebral axis.

(0) Long urostyle, not incorporated with an adjacent hypural

bone.

cartilages.

(¢Z) Large number of hypural bones associated with the last

vertebral segment.

Clupea may be quoted as an example of this type.

Il. Characters indicative of a high degree of specialization :—

(a) Greatly reduced urostyle, fused with a hypural bone.

(6) Reduction of the number of hypurals associated with the

last vertebral segment.

Among the caudal fins examined there is considerable evidence

forthcoming to substantiate the view that hypurals and epurals

are formed by the fusion of vertebral arches and radials, and the

following may be quoted as affording examples :—<Acanthias,

Gateus, Heter odontus, Acipenser, Polyodon, Synodontis, Plotosus,

Anguilla, Conger, Gadus, Gadiculus, Molva, Iotella, Centriscus,

Belone, Box, Zeus, Pleuronectes.

Proc. Zoou, Soc.—1910, No. X LL, 4]

622 MR. R. H. WHITEHOUSE ON THE [Apr. 5,

The figures here reproduced from those of Agassiz are interesting

in showing how, in homocereal tails, the original caudal fin is

replaced by a more anterior fin; I have observed exactly the

same thing in Atherina, and it is highly probable that it is quite

Text-fig. 57.

Development of the tail of the Flounder. After Prof. Alex. Agassiz.

n, notochord ; c, embryonic caudal fin; 2, hypural plates.

a general rule. The deep incision between the original and

permanent caudal is very probably only the vestige of a once

prominent gap between the two fins, and hence the homocerecal

1910. ] CAUDAL FIN OF THE TELEOSTOMI. 625

caudal fin may be considered to be a posterior anal—or, in other

words, a more anterior part of the ventral median fin—brought

into a terminal position by the great abbreviation and upturning

of the chordal axis.

GrpHyrocercy.—A gephyrocercal caudal fin may be defined as

one which is quite symmetrical, externally and internally, but

with symmetry secondarily acquired. Fierasfer dentatus may be

quoted as a typical example. Many forms which are generally

considered gephyrocercal (e. g. Mastacembelus and Anguilla) are

not so, but clearly homocereal. It is worth noticing that there is

no embryological evidence that heterocerey pr eceded the gephyro-

cercal condition.

Many other terms are used in works dealing with the caudal

fin, but the four terms here used—protocercal, heterocercal, homo-

eercal,and gephyrocercal, understood as defined here—will probably

include any type of piscine caudal fin. A common term not

insisted on in the present paper is diphycercy, which implies

perfect symmetry, externally and internally; it tends to be

ambiguous, for symmetry may be either primitive or secondary,

and primitive diphycercy corresponds to protocerey, while

secondary diphycerey corresponds to gephyrocercy. However,

diphycercy may sometimes prove useful to apply to a truly

symmetrical fin about which any doubt may exist as to its

primitive or secondary nature.

Taxonomic Imporrance.—The structure of the caudal fin

cannot be considered a safe criterion for taxonomic purposes,

though it may be useful in this connection among the smaller

divisions, as 1s well illustrated by the Acanthopterygii.

ADDITIONAL Nore.

Since the foregoing paper was written, Professor Dollo has

drawn my attention to his memoir on the fishes of the ‘ Belgica’

Expedition of 1897-99 (see Literature, ref. 11), where he advocates

a double nomenclature for the caudal fin. Briefly, this author

considers it necessary to adopt two terms—(1) physiological, and

(2) morphological. This may be conveniently summarized as

follows :—

I. PuystotocicaL (or ErnotocicaL), which refers to the

external form of the fin in its relation to the mode

of life.

(1) Rhipidocercal, implying an expanded or fan-shaped

form.

(2) Gephyrocercal, implying a tapering form.

JJ. Morpnotocican, which refers to the internal structure

of the fin.

(1) Diphycercal (or Protocercal).

(2) Heterocercal.

(3) Homocercal.

41*

624 MR. R. H. WHITEMOUSE ON THE [Apr. 5,

As a result of some discussion on the subject, Professor Dollo

and myself are of the opinion that the following modification is

advisable :—

I. Puystonoaicay, which refers to external form.

(1) Rhipidocercal, implying an expanded or fan-shaped

form.

(2) Oxycercal, implying a tapering form.

Ii. Morruoroeican, which refers to internal structure.

(1) Protocereal.

(2) Heterocercal.

(3) Homocereal.

(4) Gephyrocercal.

For the use of those who prefer to adopt this double nomen-

clature, a few illustrations may be found useful.

Rhipidocercal homocereal ...... Polyodon.

Rhipidocereal gephyrocercal .... Orthagoriscus.

Oxycercal heterocercal ......... Chlamydoselachus.

Oxycercal homocercal ......... Anguilla, Mastacembelus.

Oxyeercal gephyrocercal ...... ierasfer.

LITERATURE.

“1. Acassiz, A.—‘ On the young stages of some Osseous Fishes:

I. Development of the Tail.” Proc. Amer. Acad. Arts &

Sci. vol. xiii. p. 117, 1878.

2. Ayers, H., & Jackson, C. M.—“ Morphology of the Myxin-

oidei: I. Skeleton and Musculature.” Contrib. from the

Morpholog. Lab. of the University of Cincinnati, March

1898.

3. Boutencer, G. A.— Notes on the Classification of Teleostean

Fishes, 1V.” Ann. & Mag. Nat. Hist. vol. x., 1902.

4, Cambridge Natural History, Vol. vii., Systematic

account of the Teleostei.

5 Les Poissons du Congo. 1901.

6. Cotz, F. J., & Jounsronn, J.—‘‘ Pleuronectes.” Liverpool

Biology Committee, Memoir vii., 1901.

7. Corr, E. D.—Trans. Amer. Phil. Soe. vol. xiv., 1870,

pp. 452-3.

8. —— ‘The Homologies of the Fins of Fishes.” Amer.

Nat. 1890, p. 418.

9. Cunninenam, J. T.—‘ Science Progress,’ 1897.

10. Dotto, L.—‘ Sur la Phylogénie des Dipneustes.” Bulletin

de la Société Belge de Géologie, tome ix., 1895.

"11. Voyage du 8.Y. Belgica. Poissons. Anvers, 1904.

12. Emery, C.—‘ Mierasfer.” Fauna und Flora des Golfes von

Neapel.

* Tt was not until the manuscript was ready for the press that my attention was

dyawn to this excellent work.

T9LOs CAUDAL FIN OF THE TELEOSTOMT, 625

13.

14.

15.

16.

18.

tg:

20.

ai.

22.

Grecory, W. K.—‘“‘ The Orders of Teleostomous Fishes.”

Annals New York Acad. Sci. vol. xvii. pt. ii. no. 3, 1907.

Huxtey, T, H.—“‘On some parts of the Skeleton of Fishes.”

Quart. Journ. Micro. Science, 1859.

Kouirker, A.—Ueber das Ende der Wirbelsiiule der

Ganoiden und einiger Teleostier. Leipzig, 1860.

Lorz, T.—‘* Ueber den Bau der Schwanzwirbelsiiule der

Salmoniden, u.s.w.” Zeit. fiir Wissen. Zoologie, 14, 1864.

McCoy, F.—Annals & Mag. Nat. Hist., 1848.

Newton, EK. T.—‘ Fishes’ Tails.” Jour. Quekett Micros.

Club, vol. i. series i1., p. 79, July 1882.

Reean, C. T.—“ On the systematic position of the Gadoid

or Anacanthine Fishes.” Ann. & Mag. Nat. Hist. x1.

“ Remarks on Cnidoglanis megastomi.” Proc. Zool.

Soc., April 28th, 1908.

Rypver, J. A. “ Evolution of the Fins of Fishes.” Rep. of

the Comm. of Fish & Fisheries, Washington, 1884, pub-

lished 1886.

Wurrensousr, R. H. “The Caudal Fin of Fishes,” Pre-

liminary Paper. Proc. Royal Society, B, vol. 82, 1910.

REFERENCE LETTERING.

B.D. ............ Basi-dorsal.

(eM ao teaneD Te Cartilage supporting fin-rays.

d. ............... Distal segment of radial.

dcr. .......... Dorsal caudal radial.

der. ............. Dermotrichia.

eb. ............ Ensheathing bone protecting notochord.

CDS ee oes sacs Epural.

fas. lostteatiacrsapes Fulcra.

he ...............Hyatus between dorsal and ventral rays.

IPSfigt sop endepecs Jalvorllipitane

[OAM Soke ere Hypural.

SD ee eee een eelnter-corsale

ER pee cect Last vertebra or centrum.

M. ............... Median segment of radial.

Tee NOLOCDOXC.

Mle, viv.coaeeee oNeurallanch:

MiG Lee os Probably a neural arch protecting end of spinal cord.

TMS Wecctec.aatace Neural spine.

OCA ATR, Opisthural cartilage.

Pe eecec0sese.... Proximal segment of radial.

PiU aie Lie. Heese Penultimate vertebra or centrum.

So ccsesccssseee Secondarily developed bony sheath protecting the

spinal cord.

SHA casnagesacss hehe ayRele

ABT silat ve. thet csr Urostyle.

V.07. ........... Wentral caudal radial.

EXPLANATION OF THE PLATES.

Pirate XLVII.

Fig. 1. Caudal fin of Heterodontus phillippi. lustrating a typical Elasmobranch

heterocercal caudal fin. At the anterior end of the ventral lobe will be

seen two radials (v.c.7.) quite distinct from the hemal arches. Dorsally,

at the anterior end of the fin, the radials are’ well separated from the

neural arches. The proximal ends of the horny’ tibres have, throughout

the fin, been removed in order to expose the endoskelet#l supports.

“TI

mee:

17.

18.

19.

20.

ON THE CAUDAL FIN OF THE TELEOSTOMI. [Apr. 5,

Caudal fin of Acipenser sturio. Illustrating the Ganoid heterocercal form.

The proximal ends of the fulcra and dermotrichia have been cut away to

expose the endoskeletal supports.

. Caudal fin of Polyodon spatula. Illustrating a more specialized hetero-

cercal Ganoid caudal. As in the preceding types, the proximal ends of

the dermotric hia and fulcra have been removed.

. Caudal fin of Amia calva. Modified from Kélliker. Illustrating a highly

specialized heterocercal form: usually referred to as hemi-heterocercal.

. Caudal fin of Clupea pilchardus. Mlustrating a lowly specialized homo-

cereal candal.

. Caudal tin of Notopterus borneensis. On the ventral side the caudal is

continuous with the anal fin.

. Caudal tin of Plotosus sp.? Drawn from a young specimen 1} inches

(3°8 cm.) in length. The fin-rays (der.) on the dorsal side illustrate the

* procurrent caudal” of Regan. Note that the first fin-ray anterior to

the hypural of the penultimate vertebra has no radial as a support, and

thus seems to anticipate a similar condition ventrally as is found

dorsally.

. Caudal tin of Synodontis schall. The fusion of radials and arches clearly

shown.

PrateE XLVIII.

. Caudal fin of Leptocephalus brevirostris.

. Caudal fin of Anguilla vulgaris, 43 inches (12 cm.) in length. Of the

10 rays which are trulv caudal, 6 are bifureated distally. Note the

excellent evidence of fusion of radial with hamal spine shown in the first

hypural, the fnsion line being very distinct.

. Caudal tin of Anguilla vulgaris, adult. The urostyle and Jast centrum

have completely coalesced with the last hypural.

. Caudal tin of Conger vulgaris. The figure was drawn from a specimen

cleared in xylol, thus showing the outline of the last vertebra (/.r.),

which otherwise remains invisible. The first and second hypurals show

the fusion of radial and arch extremely well. The dermotrichia are cut

short.

. Candal fin of Myrus vulgaris. Cleared in xylol.

. Caudal tin of Fierasfer (Echiodon) dentatus. From Emery. Illustrates

extreme condition of gephyrocercy. A distinct gap can be seen in the

median line separating the dorsal and ventral dermotrichia.

. Caudal fin of Syngnathus acus. Wlustrates an extremely simple type ot

homocercy, very symmetrical in appearance, though really wholly

hypaxial.

PuatE XLIX.

Caudal fin of Ammodytes tobianus.

Caudal fin of Wolva vulgaris. ‘The dorsal caudal radials (d.c.7.) are quite

free, but slightly overlap the neural arch of the penultimate vertebra,

thus giving, in the figure, the appearance of fusion.

Caudal fin of Gadiculus argentatus. One of the best instances among

Teleostei of the formation of hypurals and epurals by the coalescence of

radials and arches; radials are to be seen in the various stages of fusion.

Caudal tin of Serranus cabrilla. A somewhat loosely constructed skeleton

and a type of homocercy frequently met with among the Acanthopterygii.

Caudal fiz of Box sa/pa. Figured from a specimen 4 cm. long. The

young stage has been chosen on account of the nenral arch of the pen-

ultimate vertebra being separate from the dorsal caudal radial immediately

posterior to it: these two elements fuse in the adult condition; otherwise

the adult form is the sume.

. Caudal fin of Lahrus festivus. The figure gives the appearance after

clearing with xylol.

. Caudal fin of Scomber scomber. Figured as seen after clearing agents were

used; no:e the last centrum shows no sign of upturning at its posterior

end.

3. Caudal fin of Zeus faber.

. Candal fin of Solea lutea. As it appears when cleared in xylol. Note

the spinal cord pierces the neural arch of the last centrum.

1910. ] MR. T. M. S. ENGLISH ON TASMANIAN FROGS. 627

Puate L.

Fig. 25. Caudal fin of Gobius paganellus. Note the large amount of cartilage

entering into the skeleton.

26. Caudal fin of Callionymus lyra. A very advanced form of homocerey with

no complications.

27. Caudal fin of Scorpena serofa. Intermediate between a lowly and a highly

specialized homocercal caudal.

28. Caudal fin of Trigla lineata. Figure drawn from a cleared specimen,

showing well the vestige of a urostyle.

29. Caudal fin of Cristiceps argentatus.

30. Caudal fin of Ophidium barbatum. The caudal proper is confined to the

rays attached to the terminal hypural bone. Note the cartilaginous shaft

fused to the posterior side of the neural and hiemal arches of the pen-

ultimate vertebra. The median fins form a continuous series.

31. Caudal fin of Mastacembelus sp.? Forms part of continuous median fin,

but the caudal proper is wholly hypaxial.

32. Caudal fin of Lophius piscatorius. The extreme case of advanced homo-

cercy—a totally hypaxial fin.

33. Caudal fin of Balistes capriscus.

2. Some Notes on Tasmanian Frogs.

By T. M. Savace ENeuisu”.

[Received March 10, 1910. }

(Plate LI.+)

Taking into consideration its outlying position and its com-

paratively small size, Tasmania is the home of a surprisingly

large number of frogs.

Seven species have come under my own observation, and it is

about these that the following notes were made in the years

1901-3. These species, all of which, I believe, are also found in

the neighbouring State of Victoria, are :—

1. Limnodynastes tasmaniensis.

2. Limnodynastes dorsalis.

3. Crinia signifera.

4. Crinia levis.

5. Pseudophryne bibronii.

6. Hyla ewingit.

7. Hyla aurea.

No others than these seven have come under the notice of

Mr. J. J. Fletcher (Proceedings of the Linnean Society of

N.S. W. 1897, p. 662). He and others, including myself, have

tried in vain to find Crinia tasmaniensis, and Mr. Boulenger

considers that, though this is beyond doubt a well-marked species,

there is reason for thinking that the specimens of it which have

belonged to the British Museum since 1858 may not have come

from Tasmania.

* Communicated by G. A. BounenGer, F.R.S., V.P.Z.S.

+. For explanation of the Plate see p. 634.

62 MR, T. M. 8. ENGLISH ON TASMANIAN FROGS. [ Apr. 5,

Two other frogs, Limnodynastes peronii and Hyla peronii,

have been credited to Tasmania, in both cases on the strength of

single specimens presented to the British Museum by Sir A.

Smith before 1858. Here, again, the evidence of locality seems

doubtful. But in a mountainous and forest-clad country, frogs,

however abundant they may be, are difficult to find, and it is only

« small portion of Tasmania that has been explored for them at

ul; so that it is quite possible that these and perhaps other

species may exist.

The Tasmanian climate is on the whole mild and even, with

regular seasons. Its winter resembles that experienced in the

south of Europe, though the mean temperature of summer is not

higher in Tasmania than it is in the north of France. The

rainfall is distributed over the year, and the droughts of the

Australian mainland are unknown.

This climate allows the frogs of Tasmania to breed at almost

any time; so that it is nearly always possible to find tadpoles in

any suitable pond. But each species seems on the whole to keep

to 1fs own particular season.

Taking the species in order :—

1, Liwopynastes rasmMANrENsIs. (Plate LI. figs. 1, 2.)

| found this frog abundantly in the neighbourhood of Hobart

in the south-east, and towards the north, around Launceston and

Westbury. These places have a smaller rainfall than the average

for the island. On the north-west coast, where, at Ulverstone and

Devonport, there is sufticient rain for tree-ferns near sea-level, I

did not find it.

The stone, or log, under which it makes its home during the day,

is usually in a somewhat moist place, but Limnedynastes tasmant-

ensis is decidedly terrestrial in its habits. I never found a single

individual in the water, and if thrown in it at once makes for the

bank, swimming badly.

A well marked speeimen of this frog is a really beautiful

creature: light brown inclining to yellow, or occasionally pink,

with large and regularly placed spots, which may be either rich

dark brown or green, with or without a lighter edging. From

this coloration it may vary to an almost uniform dark brown,

sometimes looking almost black. The dorsal stripe is generally

bright yellow.

1 found one at Westbury, towards the end of summer (February

1902), whose dorsal stripe was of a brilliant orange, almost red

(cf. colour variations of Hyla ewingit and Crinia signifera from

the same place).

I was not able to determine satisfactorily the note of Zimno-

dynastes tasmaniensis, though a noise like two stones being sharply

struck together, which I repeatedly heard from a heap of loose

rock on the bank of the North Esk, near Launceston, in April,

1901, must have come from either this species or //yla aurea.

Numbers of both were under the stones, buf no other frogs.

1910.] MR. T. M. 8. ENGLISH ON TASMANIAN FROGS, 629

LIMNODYNASTES DORSALIS. (Plate LI. fig. 3.)

Of this species, which is widely distributed over the mainland

of Australia, and is probably common in most parts of Tasmania,

I only secured one adult specimen. This I found at the end of a

burrow in sand under a large stone. It was in a rather dry part

of some Eucalyptus “ bush, 7 but not far from a small pond.

if Tam right in supposing that a loud booming note, which

can be heard ‘considerably more than a quarter of a “milé aw: vy, 1S

due to this species, it is common in the neighbourhood of Hobart,

Launceston, Fingal (in the east of the island), aud on the north-

west coast.

The only other frogs to which this note could be reasonably

attributed would appear to be Limnodynastes tasmaniensis and

Myla aurea; but the former does not seem to occur on the north-

west coast, and the latter is not found near Hobart.

This is by far the most noticeable frog-music in Tasmania, and

when, as is usually the case, a number of frogs are uplifting their

voices together, the effect is that of a somewhat barbaric banjo

performance. These concerts take place in or around some fairly

deep pond, and, however quietly one approaches, according to my

own experience, invariably cease before it is possible to exactly

mark down the performers. I have never been able to notice

any disturbance in the water, and believe that the frogs must be

singing at the entrances of their burrows, and that these are well

concealed under roots, stones, &c.

This frog, on the north-west coast, retired for the winter of 1902

about the middle of April, and was in evidence again towards the

end of August.

At Ulverstone it was spawnmg towards the end of Octeber, the

egg-masses resembling lumps of froth about 6 inches, or rather

less, across. On the 7th of November some of these had fairly

large tadpoles in and under them. These tadpoles were very di ark

in Eales with violet reflections.

Karlier in the year (September) I had found some very large

tadpoles of this species (3 inches long), with hind legs just forming,

and at the same time some quite small ones. On the 10th of

November some of these were showing signs of front legs, and by

December they had become frogs.

3. CRINIA SIGNIFERA. (Plate LI. figs. 4, 5.)

This active and variable little frog was very common wherever

I went in Tasmania, and its cricket-like chirp was to be heard at

almost any time of the year ; so that it does not seem to hibernate.

It is decidedly aquatic, diving to the bottom at the slightest alarm

and remaining motionless.

No kind of water seems to come amiss to it, from a rapid

mountain stream to a muddy puddle. As a rule, however, it

prefers water which is fairly shallow, and so is often obliged to

become terrestrial during the heat of summer, when all the

630 MR. T. M. S. ENGLISH ON TASMANIAN FROGS, [ Apr. 5,

shallow pools are dry. At this time of the year these frogs are

to be found, almost always singly, under stones and in cracks in

the ground, into which their small size allows them to go deep

enough to find a secure and comparatively moist retreat.

They chirp to each other incessantly, by day as well as in the

evening, and though the note of a single one is comparatively

weak the comb ined efforts of the numbers which gather together

in any suitable place can be heard a long way off.

On land these frogs are as active as crickets, and exceedingly

difficult to catch. In the water they swim well.

The colour of Crinia signifera is as a rule dark brown or grey,

but it is most variable. ‘The male is darker than the female and

may be almost black all over, though he is generally lighter under-

neath. The under surface of the female is usually white, with or

without dark spots. While breeding, the male seems always to

have a black throat. There is sometimes a tendency to bright

brown or orange markings ; and in February, 1902, I found one of

these frogs at Westbury which was of a fairly bright orange all

over (cf. colour variations of Limnodynastes iusmaniensis and of

Hyla ewingii from the same place). The young and also the tad-

poles are intensely black.

In January, 1901, I found that most of these frogs at Hobart,

and at Zeehan in the west of the island, had the backs of their

thighs and their hinder parts blood- red, giving them a truly

diabolical appearance. I did not, however, find any of them

coloured in this way except on these two occasions.

This species has a continued chirp, hardly to be distinguished

from that of the mole cricket, which is such a common Tasmanian

insect, or from that of P: seudophryne bibronit.

Crinia signifera has its main spawning season during the

autumn and winter months, from May to August. At this time

of the year these frogs are not nearly so active as usual, though

of course this may be owing to the colder weather.

The male clasps the female round the waist.

The eggs are in masses, spread over stones and weeds at the

bottom of shallow water, and the tadpoles are exceedingly small,

as are the young frogs. ‘These, at first, are less than a quarter of

an inch in length.

4. CriniA L&vis. (Plate LI. fig. 6.)

I found this species abundantly on the north-west coast; also,

when I had got to know where to look for it, at Westbury and near

Launceston. But as it seems to be not only nocturnal but silent,

it is very possibly far more common than it appears to be.

All my specimens were under logs or stones lying on swampy

or at all events fairly moist ground, but, in complete contrast to

Crinia signifera, Crinia levis appears to be altogether terrestrial,

not even going into the water to spawn. Indeed, in life-history

and habits these two frogs seem to have nothing in common,

while Crinia levis closely resembles Pseudophryne bibronii. It is

1910.] MR. T. M. S. ENGLISH ON TASMANIAN FROGS. 631

perhaps a little more active than this last, but still is by no

means difficult to catch.

The colour of this species, in Northern Tasmania, does not

seem to vary much. Above it is light grey, pecomane paler or

white underneath, where it is marbled with dark grey or black.

The “ concealed surface ” at the groin is pink marbled with black.

A breeding pair, near Devonport, had irregular brown blotches

on their backs, while the throat of the larger of the two was dull

yellow.

This frog may perhaps be responsible for some of the chirping

which comes from the haunts of the smaller Tasmanian batrachians,

but of this I was never able to assure myself.

Its breeding habits are peculiar. It does not deposit its eggs

in water, but in a smooth-sided underground chamber, ener

hollowed out or adapted by the parent frogs.

On the 30th of March, 1902, one of these nests contained two

frogs as well as a mass of eggs. (I never found more than two

frogs of this species together : ef. Pseudophryne bibronii.) This

mass, after being put mo water, held together for about a week ;

then the eggs separated. The tadpoles began to free themselves

on the 4th of May. Their colour during their earlier development

was a rather pale brown, but before leaving the egg they became

quite dark. There was no appearance whatever of external gills ;

and while in the eggs the tadpoles did not move at all.

5. PSEUDOPHRYNE BIBRONII. (Plate LI. fig. 7.)

This little toad, the only known representative of the Bufonide

in Tasmania, seems to be widely distributed over the island, for,

though I never saw one of them except in the autumn, when they

gather together for breeding, it was abundant wherever I hap-

pened to be at this time of the year: at Hobart, near Launceston,

and near Devonport, in March, April, and May, 1901, near this

last place in March and April, 1902, and at Perth in March,

1903.

After rain at the end of March, 1901, they became abundant and

noisy near Launceston. I found over thirty packed closely together

under one log.

This species is slow in its movements, and walks rather than

hops. In the water it seems perfectly helpless, and as, like Crinia

levis, it leaves the necessary water for its tadpoles to be provided

by subsequent rain, and frequents dry hilly country, it is probably,

except for a short time as a tadpole, altogether terrestrial.

In colour it is dark brown above, occasionally almost black ;

beneath it is marbled, black and white or grey ; while those parts

of its legs which are usually concealed, except when it is walking,

are brilliant yellow and orange. There are indications of a yellow

vertebral stripe.

The note of Pseadophryne bibronii is a chirp, which I could not

distinguish from that of Crinia signifera. As is the case with this

last frog, the number of singers generally makes up for any

632 MR. T. M. 8S. ENGLISH ON TASMANIAN FROGS. [Apr. 9,

individual want of power, and the resulting concert can be fairly

called noisy.

The breeding habits of the present species are to a large extent

those of Crinia levis, except that it is more social. Fr always

found large companies g gathered together under logs and stones in

places where, after vain, ponds would form. They do not seem

to make nests like Crinia levis, but deposit their eggs under any

convenient stone. (I never found any eggs under a Jog, but this

of course may have been accidental.) The eggs do not seem to

adhere either to each other or to anything else.

On the 20th of April, 1901, I found a collection of these eggs,

which I put into water. The tadpoles began to wriggle at once,

and within twelve hours they were free and swimming about.

6. Hyta Ewine. (Plate LI. figs. 8, 9.)

This frog seems to be abundant in most parts of Tasmania. The

only place from which I might have recorded it, and did not do so,

is Zeehan, and I am almost certain that this is an accidental

omission.

It is on the whole aquatic, seldom going far from water, and

living very generally under stones or logs on the bank of some

pond or stream. When disturbed, it dives in with all speed.

It does not seem to be really arboreal, though it may occasionally

climb up ten feet or so from the evound. I have never heard its

note coming from the top of a tall tr ee, as one hears that of the

European I yla arborea, though this may perhaps be accounted

for by the very slight shelter from either sun or enemies that a

frog would find among the leaves of any full-grown Tasmanian

Eucalyptus.

Where plants are kept in pots on a veranda a colony of these

frogs usually settles, and its members seem to appreciate regular

watering as much as the plants do, generally greeting it with

song.

Except when breeding, //yla ewingii is inactive during the day.

Its regular concerts begin towards sunset, though, as is the case

with many other frogs, its voice is heard when rain is approaching,

or at times when certain loud noises excite it—rapid hammering,

for instance,

It does not seem to hibernate, but is sluggish in cold weather,

Its colour is almost invariably some shade of brown, closely

resembling that of its immediate surroundings.

Mr. J. i Fletcher has found that near the summit of Mount

Koskiusco, in New South Wales, frogs of this species are bright

green. He attributes this to the fact that at this altitude, though

the days may be warm, the nights are almost always frosty ; so

that no insects are abroad then, and the frogs have to hunt for

food by day among green vegetation.

In January, 1902, [ found a specimen at Westbury which had

some bright green patches on its back. The colour of these patches

seemed fixed ; at all events it did not vary with that of the rest

1910.] MR. T. M. S. ENGLISH ON TASMANIAN FROGS. 633

of the frog according to its surroundings (cf. colour variations

of Limnodynustes tasmaniensis and Crinia signifera trom the same

place).

The note of this frog resembles that of Zyla arborea, but is not

so loud,

Hyla ewingii has its regular breeding season in winter. In

July, 1902, it was spawning at both Launceston and Devonport, in

pools and ditches such as would be chosen by Rana temporaria.

Its spawn is in masses, very like those of this last frog but smaller.

The first tadpoles appeared at the end of July. They are very

small, and at first quite black, becoming paler and silvery under-

neath as they develop.

During November the pools they frequent in a normal season

are fast drying up, and the water which remains is packed with

tadpoles. Early in December these pools are mostly dry, and the

tadpoles have either become frogs or perished, the place where

the last water stood being covered by a layer of their dried-up

bodies.

7. HyYLA AUREA.

This, the largest and most conspicuous of the Tasmanian frogs,

seems to be decidedly local. I saw or heard of it only in the

country drained by the North and South Esk rivers, which unite

at Launceston to form the tidal Tamar, and by the Mersey, which

falls into the sea at Devonport. There is only a low watershed

between these two basins. Wherever it is found it seems to be

abundant : near Launceston, Perth, Longford, Westbury, Fingal,

and Devonport; and as it is diurnal, it is’ not likely to be over-

looked if present.

In its habits it resembles Rana esculenta, being essentially

aquatic, and in summer liking to sit in the sun near the water,

into which it dives with a splash when alarmed. It isa powerful

swimmer ; its usual custom of returning at once to the bank after

having dived in, can be well accounted for by the fact that the

rivers it frequents all contain large trout. This frog seems to be

impatient of cold. In 1902, at Devonport, it disappeared after a

few frosty nights at the end of March and the beginning of April,

though the weather was on the whole warm for quite six weeks

after this. Near Launceston I found large numbers of these frogs

under stones during April, 1901. I only once found one climbing ;

it had got about three feet from the ground in a bush.

A well-marked specimen of this species is most gorgeously

coloured : green, gold, blue, and white ; but from this it may vary

to dull brown, and when about to retire for the winter to almost

black.

Beyond an occasional and not very loud croak, I was never able

to identify this frog’s note, though for a considerable part of one

summer my window overlooked a pond full of them.

I was not at any of the places which they frequent during their

breeding-season : this is probably September or the beginning of

634 ON A NEW MONKEY FROM THE MALAY PENINSULA. [Apr. 19,

October. At Westbury, towards the end of January, 1902, tad-

poles of this species were very abundant and were developing their

hind legs. During February they became frogs.

These taclpoles are very fish-like in colour : ereenish olive above,

silvery white beneath. They make a most attractive spinning

bait for large trout.

Hyla aurea is one of the many additions which have been made

of late years to the fauna of New Zealand and to that of New

‘aledonia.

XPLANATION OF PLATE LI.

Figs. 1 & 2. Limnodynastes tasmaniensis, showing different types of colouring.

Vig. 3. Limnodynastes dorsalis.

Crinia signifera, 2 in the breeding season.

Crinia signifera, in late summer.

Crinia levis.

: Pseudophr, yne bibronii, w ith exes showing the development of the tadpole.

» Hyla ewingii, showing its nor mal colour.

Hyla ewingii, from a quarry of brightly coloured sandstone.

GIA

<=)

April 19, 1910.

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

in the Chair.

The Seeretary read the following report on the additions

made to the Society’ s Menagerie during the month of March

1910 :—

The number of registered additions to the Society’s Menagerie

during the month of March last was 100. Of these 30 were

acquired by presentation, 18 by purchase, 35 were received on

deposit, 12 in exchange, and 5 were born in the Gardens.

The number of departures during the same period, by deaths

and removals, was 190.

Amongst the additions special attention may be directed

to :—

One Puma (Felis concolor) 3, from Tucuman, presented by

Alec 8. Waley, Esq., on March 22nd.

Two Musquash (Fiber zibethicus), from North America,

purchased on March 16th.

A new Monkey from the Malay Peninsuia *

On behalf of Mr. Oldfield Thomas, F.R.8., F.Z.8., two specimens

were exhibited of a new Monkey of the genus Presbytis, which

had been obtained by Mr. H. C. Robinson during a recent expedi-

tion to the northern parts of the Malay States.

* Published by permission of the Trustees of the British Museum.

1910. ] ON A COLLECTION OF SMALL MAMMALS FROM CIIINA. 635

It was proposed to be called

PRESBYTIS ROBINSONI Thos. *

Abstract P.Z.S. 1910, p. 25 (April 26).

A white Monkey, with darker underside, hands, and feet.

General colour above creamy white; the hairs white to their

bases, mixed, however, with a small number of blackish hairs.

These dark hairs ecame more numerous on the sides and belly,

where they outnumbered the white ones; the general tone of the

under surface grey, about grey no. 6 or 7 on the belly, darkening

to no. 5 on the throat and sides of neck. Outer side of arms to

wrists whitish like back, inner side and whole of legs with an

equal mixture of dark and light hairs, resulting in about grey

no. 9. Hands and feet greyish brown, with a tinge of bistre in

it. ‘Tail greyish white thr oughout.

Skin of face dark, except Tihe lips, which were white. Long

hairs of occiput directed forwards. Hairs of forehead radiating

from points over the temples with a median crest between them,

but both the whorls and crests differed considerably in the two

specimens. Har-tufts white, considerably surpassed by the long

brownish hairs which grew i om the lower parts of the cheeks.

A female specimen was whiter throughout, the under surface

more nearly whitish, but still darker than the upper surface.

Dimensions of the type, measured in the flesh :—

Head and body 565 mm.; tail 824; hind foot 177; ear 30.

Hab. Ko-Khau, Trang, Nowthern Malay Peninsula.

Type. Adult mele, British Museum, No. 10.10.1.1. No. 1236/10

of the Selangor Museum Register. Original number 3184.

Obtained for Mr. Robinson by a native collector, 10 January,

1910.

This fine Monkey, which was obviously not an albino, differed

conspicuously from all its allies by its white upper and darker

lower surface, all other members of the group having the belly

lighter than the back.

Mr. Thomas had great pleasure in naming it after its discoverer

Mr. H. C. Robinson, Director of Museums, Federated Malay

States, by whose scientific energy our knowledge of Malay

Mammalia had been so lar gely increased.

A Collection of small Mammals from China*

On behalf of Mr. Oldfield Thomas, F.R.S., F.Z.8., a collection

of small mammals obtained by Mr. Malcolm Anderson in Southern

Shen-si for the Duke of Bedford’s exploration of Eastern Asia

was exhibited. Observations were made on the value of this further

contribution by the Society’s President to the National Collection

* The complete account of this new species appears here, but the name and a

preliminary diagnosis were published in the ‘ Abstract,’ No. 83, 1910.—Eprror.

+ Published by permission of the Trustees of the British Museum.

636 MR. OLDFIELD THOMAS ON A COLLECTION [Apr. 19,

and on the resultant increase in our knowledge of geographical

zoology, a subject in whose study the Zoological Society had

always had so large a share.

A complete list of the specimens would be given later, but in

the meantime Mr. Thomas gave the following account of the

new forms* :

Myoris MYOSOTIS ANCILLA Thos.

Abstract P. Z.S. 1910, p. 25 (April 26).

Smaller, with shorter ears, and more drab-coloured than in

true myosotis.

Size rather smaller than in myosotis, not so small as in blytha.

Ears decidedly smaller than in myosotis. General colour above

nearly approaching “ drab” of Ridgway, very different from

the ‘“ wood-brown” of myosotis and its pale Persian subspecies

omari. Head rather greyer. Dark shoulder-patches more

strongly defined than in myosotis, blackish brown, Under

surface as in myosotis.

Skull slightly smaller than in myosotis, and the bull smaller

in correlation with the shorter ears. Protocone apparently as

in myosotis, not as described by Miller in blythi y.

Dimensions of the type, the starred measurements taken in

the flesh :—

Forearm 61 mm. (range 59 te 62).

Head and body *75 mm.; tail *56; hind foot #2 - ear *ile

Skull: greatest length 22°2, basi-sinual length 17, front of

canine.to back of m? 9:2.

Hab. Shang-chou, 8.E. Shen-si. Alt. 3300".

Type. Adult male. B.M. No. 10.5.2.4. Original number 2082,

Collected 27 November, 1909.

This is the eastern representative of the common European

M. myosotis. What its relationship is to the Indian form blythi

remains to be seen when better specimens of the latter are

available for examination. But the type of J/. blythi, received

in the British Museum in the Tomes collection, is decidedly

sinaller.

Microrus (Eornpnomys) NuX Thos.

Abstract P. Z.S. 1910, p. 26 (April 26).

Darker than JZ. inex, with a longer and more distinetly bicolor

tail.

General colour above decidedly darker than in inez, the back

approaching ‘ Prout’s brown” ; under surface also brown, of a

slightly more buffy tone than the upper surface, but not sharply

* The complete account of the new species described in this communication

appears here, but since the names and preliminary diagnoses were published in the

‘Abstract,’ No. 83, 1910, these species are distinguished by the names being under-

lined.—Epiror.

1 a be Biol. Soc. Wash. xiii. p. 155, 1900,

1910.] . OF SMALL MAMMALS FROM CHINA, 637

separated from it, the belly on the average decidedly browner

than in irez. Head rather greyer, Hands and feet dull whitish

or pale brown, averaging rather darker than in inex. Tail

longer than in that Species (average of 17 specimens 38 mm, as

against 33 mm. in 14 inez), bicolor, dark brown or blackish above,

dull whitish below.

Skull agreeing closely with that of J/. inez, except that the

posterior lateral palatal grooves tend to be deeper and more

continuous, and thus to show an approach towards the single

deep depression found on each side in melanogaster. Bulle large,

larger than in either of the other species.

Teeth on the whole most like those of J/. inez, but showing an

approach towards those of melanogaster by the junction of several

of the enamel spaces, closed in the former, open in the latter.

Thus in m’ the normal 2nd and 38rd spaces, separated in inez,

are joined as in melanogaster, and in m, the two spaces next

following the trefoil are frequently joined to each other as in

Pitymys. Small extra internal projection on m' and m* as in

M. wez.

Dimensions of the type, measured in the flesh :—

Head and body 93 mm. ; tail 39; tue foot 16:5; ear 10.

Skull: condylo-basal length 23- 6 mm. greatest breadth 14:3

nasals 7; interorbital breadth 4; alate length 11; mau

65; palatal foramina 4°7; bulle, ‘diagonal horizontal diameter 7 :

upper molar series 5:6.

Hab. of type. Shang-chou; 8.E. Shen-si. 3300’.

Type. Adult male. B. M. No. 10.5.2.79. Original number 2089.

Collected 29 November, 1909. 23 specimens.

The discovery of this interesting species tends to confirm the

reference of IM. inez to Hotheneniys! about which I had some

doubt. For while J/. melanogaster, the type of Mothenomys, has

nearly all its cement-spaces communicating with each other, and

M. inex has them nearly all closed and separate, the present form

differs from Jf. inez exactly in the direction of melanogaster by

the opening of several of the spaces, the great majority of them

being still closed.

Microrus JOHANNES Thos.

Abstract P.Z.8. 1910, p, 26 (April 26).

Near Jf. mandarinus, but rather smaller, with much smaller

skull.

General characters very much as in J. mandarinus. Fur

soft and fine ; hairs of back about 11-13 mm. in length. General

colour above rather browner than Ridgway’s wood-brown,

decidedly paler than in J. mandarinus. Under surface dull

greyish ; the hairs slaty basally, greyish white terminally. Ears

short, projecting but slightly ‘above the fur; pr: actically naked.

Upper surface of hands and feet dull whitish. Tail short,

drabby whitish above, dull white below.

Proc. Zoou. Soc.—1910, No. XLII. 42

638 ON A YELLOW VARIETY OF MUS RATTUS. [ Apr. 19,

Skull broad and flat, very like that of JZ mandarinus on a

smaller scale. Brain-case square, with well-marked angles, but

the ridges little developed. Anterior palatine foramina of about

ejual breadth throughout, those of mandarinus broad in front,

narrowing behind.

Incisors markedly lighter than in mandarinus, their faces

orange to the tips. Molar pattern about as in WM. mandarinus ;

m* with 4 spaces, three outer and three inner salient angles ;

m, with 7 spaces, 5 inner and 4 outer angles. All triangles

separated from each other except in ™,.

Dimensions of the type, measured in the flesh :—

Head and body 95 mm. ; tail 23; hind foot 17; ear 7.

Skull: condylo- basal length 24°6 mm. ; greatest breadth 15:2 ;

nasals 7°0 x 3°1; interorbital breadth 3:5; palatilar length 13:2 ;

diastema 7°1 ; palatal foramina 4°1 ; ape molar series (crowns)

58.

Hab. Mts. 12 miles N.W. of Ko-lan-chow, Shan-si. Alt.

7000!.

Type. Adult male. B.M. No. 9.1.1.178, Original number 1950.

Collected 5 June, 1908.

This Vole was referred to M. mandarinus M.-Edw. in my last

paper *, but by the kindness of Prof. Trouessart I have had an

oppor tunity of re-examining the skull of that species, and I find

it to agree in every detail with two specimens now sent from

Shang- chou, S.E. Shen-si, and to differ equally from the Ko-lan-

chow form, which ther efore needs descr ibing.

Lt, howev er, this identification be accepted, it throws some

doubt on the ‘asserted locality of ‘“‘ Mongolie chinoise” for man-

darinus, and 1 think it not impossible that the type really came

from 8. Shen-si, where David made a small collection, after

getting his Mongolian series.

M. johannes is readily distinguishable from J/. mandarinus

by its paler colour and markedly smaller skull.

Mr. J. Lewis Bonhote, M.A., F.L.S., F.Z.8., exhibited a yellow

variety of Mus rattus. This individual had been bred from two

wild- caught examples of Jus rattus tectorwm trom Egypt. ‘The

long hairs on the upper parts, which are normally black, were

colourless and the shorter body-hairs had grey bases with yellow

tips. So far as he knew, this was the first recorded example of a

yellow rat, which was a colour quite unknown to fanciers and

was never met with by Crampe or Doncaster, who had conducted

colour-breeding experiments on rats. Although it was not defi-

nitely stated, these experiments had probably been carried out with

Mus norvegicus. A careful search through the large series of

Mus rattus in the Museum also failed to bring to light any yellow

forms.

* P.Z, 8. 1908, p. 978.

2. LOLO. PE EE

Huth coll

P Z.S.1910; Pl Ee

Huth, coll.

PHOTOPHORES OF ACANTHEPHYRA (Fig. 1)

AND SERGESTES (Figés.2- 4).

P58. 1910, Piva

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PHOTOPHORES OF ACANTHEPHYRA (Fig.1)

AND SERGESTES (Figs. 2) == 05)

1910.] ON THE PHOTOPHORES OF DECAPOD CRUSTACEA. 639

Centenary of Philip Henry Gosse, F.R.S.

Born April 6, 1810; died Aug. 23, 1888.

The Secretary exhibited the set of the works of Philip Henry

Gosse in the Society’s Library and gave a brief summary of his

contributions to science, upon which he placed a very high value.

The Chairman and Prof. A. Sedgwick, F.R.S., added their testi-

mony to his place in Zoological history; Dr. Henry Woodward,

F.R.S., gave some personal reminiscences; and Mr. Edmund Gosse,

Librarian of the House of Lords, the son and biographer of Gosse,

thanked the Society for their notice of his father’s centenary.

The following papers were read :—

1. Notes on the Photophores of Decapod Crustacea.

By Srantey Kemp, B.A.*

[ Received March 1, 1910.]

(Plates LII.—LIV.7)

Two different methods of producing light are known among

Crustacea. In the vast majority of species possessing this power

the organ is of a glandular nature, excreting drops of oily fluid

which become luminous on reaching the water. The process is

probably one of oxidation, though from the results of Giesbrecht’s

experiments | it would seem that very small quantities of free

oxygen will suffice.

Certain Decapods, such as Polycheles phosphorus, appear to pro-

duce light in this way, while in others, such as Avistews coruscans

and Heterocarpus alphonsi, the excretions from the antennal

glands have been noticed to be brilliantly phosphorescent §.

Both these phenomena seem to be exceedingly rare among

Decapods.

The second method of producing light, and it is only with this

type that the present paper is concerned, is by means of photo-

phores, compound luminous organs which do not excrete a fluid ;

they are in most cases provided with a lens and sometimes also

with a reflector. Very little is known of the chemical processes

which are involved in this type of phosphorescence. ‘The essential

feature, as in the case of the glandular organs, is doubtless one

of oxidation, and it is probable, as Aleock has suggested, that the

oxygen is conveyed to the photophore by means of the blood.

Among Crustacea, photophores are known only in the Euphau-

siacea and in the Decapoda. In the former group they are very

highly specialised and occur in practically all the species known.

* Communicated by Dr. W. T. Catmay, F.Z.S.

+ Vor explanation of the Plates see p. 650.

{ Giesbrecht, Mitth. Zool.-Stat. Neapel, ii. 1895, p. 648.

§ See Alcock, ‘A Naturalist in Indian Seas,’ 1902, pp. mage

640 MR. STANLEY KEMP ON THE [Apr. 19,

Their true function was first demonstrated by Sars *, and their

structure is now well known owing to the work of Chun f and of

Vallentin and Cunningham §.

Photophores occur only in three genera of Decapoda, all of

which belong to the Macrura, or, according to modern systems of

classification, to the Natantia. One of them, Sergestes, belongs

to the Penvidea, and the other two, Acanthephyra and Hoplo-

phorus, to the Caridea. Although, as might be expected, two

wholly different types of structure are found in these two groups,

they have none the less one very striking feature in common—in

both a deep blue pigment is associated with the luminous organ.

This pigmentation is one of the most interesting characteristics

of the photophores of Decapoda, for, except for the fact that it

has once been observed in a Euphausian, such an association

appears to be elsewhere unknown.

The following six species of Decapods possess photophores :—

PENAZIDEA.

Sergestide......... Sergestes challengert Hansen.

Sergestes gloriosus Stebbing.

CARIDEA,

Hoplophoride ... deanthephyra pellucida Perrier.

Acanthephyra debilis A, Milne-EKdwards,

Hoplophorus grimaldii Coutiere.

Hoplophorus sp. juv.

Two other genera, Gennadas and Amalopeneus, belonging to

the family Penzide, may also possess photophores, but it has not

as yet been possible to come to any definite conclusion on the

subject.

The material which I have been able to examine consists of

one species of each of the three genera mentioned above. I am

indebted to Prof. J. Stanley Gardiner for the opportunity of

examining four specimens of Sergestes challengeri and an immature

Hoplophorus from the Indian Ocean. The examples of the former

genus are preserved in formalin and have in consequence retained

some at least of their original pigmentation. <Acanthephyra debilis

occurs in all stages off the West coast of Ireland. This species is

unfortunately far from.common, and although special efforts have

been made, no fresh material has been found during the last

eighteen months. In consequence, it has not been possible to

solve certain problems connected with the pigmentation and in-

nervation of the photophores.

Deep-sea Decapods are almost invariably dead when brought

to the surface, and although in view of recent investigations it

does not seem probable that the vitality of the organism has

* Sars, Report on the ‘Challenger’ Schizopoda, 1885, p. 70.

+ Chun, Bibliotheca Zoologica, Bd. vii. (Heft 19), 1896, p. 191.

t Vallentin & Cunningham, Q. J. Mier. Sei. xxviii. 1888, p. 319,

1910.] PHOTOPHORES OF DECAPOD CRUSTACEA, 641

es direct effect on the production of light, it is none the less the

ase that phosphorescence has actually een observed only in a

aaels species, Sergestes chatlengert. Prof. Gardiner informs me

that the examples of this form which he obtained in the Indian

Ocean were brilliantly phosphorescent on the occasion of their

capture. In the other species the function of the organs has

been deduced from their structure only,

THE PHoTrorHoreEs OF SERGESTES.

The only two species of Sergestes which are known to possess

photophores are S. challengert, which was described by Hansen *

from a single specimen obtained by the ‘ Challenger Expedition

near the Piji Islands, and a closely allied form, gloriosus

Stebbing 7, which was found in South African waters. Hansen

discovered 117 photophores in his mutilated example of JS. chal-

lengert, and estimated that at least 150 would be found in a

perfect. specimen. According to Stebbing’s account an even

greater number exist in S. gloriosus.

It is not necessary to describe the distribution of the photo-

phores in detail. They occur on the lower sides of the eyestalks,

antennules and antennal scales, on the oral appendages, on the

thoracic and abdominal sterna, on the ventral surface of the outer

uropods, and on many of the leg joints. All are so situated that

the light which they produce is thrown directly or obliquely

downwards. In both the species photophores have been described

on the lateral face of the carapace. These, however, at any rate

in S. challengeri, are not external, but are placed in the roof of

the branchial chamber and illuminate the gills from above

(Pl. LIV. fig. 4). To find photophores in such a position as

this is most astonishing, and it is not easy to suggest any theory

which will account for their curious situation.

The photophores are all practically identical in structure and

all are quite immovable, though a few are supported on very

short, thick stalks in order to make them bear more directly

downwards. They vary considerably in size, but even the largest

are much smaller than the organs on the pleopods of Acanth-

ephyra debilis. In a single individual the diameter of the lens

was found to range from -06 to ‘14 mm.

The structure of the organs is illustrated in Pl. LIIT. figs. 2-4

and in Pl. LIV. figs. 2, 3, & 5.

Externally there is a double convex lens. This is made up of

two distinct portions, which are formed from the two layers com-

posing the cuticle. The outer part is double convex, while the

inner, which is closely applied to it, is concavo-convex. In sections

treated with picro-carmine the inner lens often stains to a rather

deeper red tone than the outer, but the complete absence of any

yellow colour indicates that the cuticle of which both are formed

* Hansen, P.Z. 8.1903, p. 72.

+ Stebbing, Marine Invest, in S. Africa, yol. iv. 1905. p. 84,

642 MR. STANLEY KEMP ON THE [Apr. 19,

is not more strongly chitinised than the rest of the exoskeleton.

Both portions are quite transparent in material preserved in weak

formalin, A very delicate investing membrane possibly occurs

on the outer side of the lens, but its presence could not be demon-

strated clearly in any of the sections obtained.

Situated immediately behind the lens and exceeding it in

diameter is the first cellular layer. This is composed of a number

of large wedge-shaped cells which appear to be derived from the

epithelium ; from eight to ten are seen in a median section of

the organ. ‘They are all full of protoplasm and their nuclei

invariably lie close up against the lens. In material preserved in

formalin this layer is seen to be impregnated with a deep blue

pigment,

The second cellular layer is extremely inconspicuous ; it consists

merely of a few flattened nuclei round the outside of the first

layer.

The third layer Hansen very reasonably considers to be a

reflector. In material preserved in formalin it is of a distinctly

yellow colour; it is faintly striated and contains numerous pear-

shaped nuclei which are very regularly arranged with their narrow

apices directed towards the lens.

The fourth and last layer consists of a number of irregularly

disposed cells round the back of the reflector. It is possible that

in life these carried a pigment, but in the preserved specimens

no trace of this remains.

In some instances a nerve-strand communicating with the

photophore was detected, but the exact mode of its entrance could

not be discovered. It is not improbable that it runs round the

edges of the reflector and then turns inwards to supply the first

cellular layer, in much the same way as has been demonstrated

by Chun in the photophores of Euphausians.

One of the two organs placed on the underside of the eyestalk

is situated in closest proximity to the cornea, In this case

(Pl. LIV. fig. 2) the photophore is slightly twisted and is

directed forwards and downwards. It is shut off from the cornea

by a layer of black pigment, and its nerve-supply is not drawn

from any of the optic ganglia, but from a separate strand which

runs up the inferior margin of the stalk.

It will be noticed from Pl. LIII. figs. 2, 3, that the lens

may differ considerably in convexity, and in one case (fig. 4)

it is plano-convex. This photophore is placed at the base of the

exopod of the first maxillipede and is directed forwards; as

Hansen mentions, it is partially, though not entirely, overhung by

the surrounding tissues,

In neither of the two species is anything known of the develop-

ment. ‘The photophores, however, differ slightly in number in

the examples which I have examined, and it is probable that, as

in the case of Acanthephyra debilis, they continue to increase as

the specimen gains in size. Additional organs seem to appear

long after the individual has attained maturity.

a ee

» ~

1910.] PHOTOPHORZS OF DECAPOD CRUSTACEA. 643

A comparison of the foregoing description with the account

and figures which Hansen gives of the structure of these organs

reveals many discrepancies. There can be little doubt that this

is due to the fact that Hansen viewed the organs only in optical

section, 2 method which, even in his hands, has not yielded

satisfactory results.

THe PHOTOPHORES OF ACANTHEPHYRA.

Two species of Acanthephyra are known to possess photophores.

They are first mentioned by Perrier in 1886 * in a form which he

called “ Acanthephyra pellucida A. Milne-Edwards.” There is

unfortunately a good deal of uncertainty regarding the validity

of this species, for it has not been rediscovered in recent years

and Milne-Edwards seems to have never published any deserip-

‘tion. Our knowledge of it is, in consequence, derived solely from

the brief reference in Perrier’s work, and the accuracy of the

account of the distribution of the photophores, which is there

given, is by no means certain *.

Acanthephyra debilis is better known. Coutiére in 1905+

first described the existence of photophores in this species, and in

1906 § he published a more complete account of their number

and distribution.

The photophores in 4. debilis ave not all of similar structure,

as they are in the case of Sergestes, but exist in different degrees

of complexity in different parts of the animal.

The most highly developed organs are twelve in number, and

each is so placed that the light which it produces is thrown

directly downwards. One is situated on the distal and external

aspect of the protopodite of each pleopod, and one behind the

protopodite of each uropod. ‘The structure of these photophores

is illustrated in PI. LIT. fig. 1 and Pl. LIV. fig. 1.

Externally there is a thick concavo-conyex || lens formed from

the cuticle. In adult specimens this measures about ‘24 mm. in

diameter, and during life is of a deep violet-blue colour. The

pigment does not exist as a mere coating, but permeates through-

out the structure of the lens. Sections stained with picro-

carmine show that the lens is made up of three distinct layers.

The inner and outer portions are merely thickenings of the two

cuticular layers which form the normal exoskeleton of Crustacea

and under a high power show the usual striations. The middle

layer, which is also striated, always stains more deeply with carmine

than the others, and, owing to the fact that it sometimes takes

* Perrier, Les explorations sous-marines, 1886.

+ A comparative view of the positions occupied by the photophores in A. pel-

lucida and in A. debilis will be found ia Kemp, Fisheries, Iveland, Sci. Invest., 19U8,

I. [1910], p. 67, where Perrier’s original description is reprinted.

ft Coutiére, Bull. Mus. Océanog. Monaco, no. 48, 1905, p. 7.

§ Coutiére, Bull. Mus. Océanog. Monaco, no. 70, 1908, p. 4.

‘| In a few of the sections obtained the lens is plano-convex, but this [ believe to

be due to distortion.

644 MR, STANLEY KEMP ON THE [Apr. 19,

up a certain amount of yellow colour (due to the picric acid in

the stain), it seems probable that it is formed of a more strongly

chitinised material. ‘The cuticle is thrown in folds on either side

of the lens. Jt is possible that by this structure a lLmited amount

of movement is permitted to the photophore, but no trace of a

controlling muscular apparatus could be found.

Inside the lens there is a series of very large elongate cells

which radiate from a well-defined centre to its inner surface.

They measure from °08 to “10 mm. in length; twenty are usually

visible in a median longitudinal section, consequently at least

three hundred must oceur in the whole organ. The proximal

portions of these cells appear to be w holly dee oid of protoplasmal

contents. The nuclei* are very regular in shape and, as in the

first cellular layer of Sergestes, lie close up against the lens. The

outer end of each is evenly rounded, and a band of cytoplasm

may sometimes be seen between it and the lens; the inner end is

squarely truncate. The nuclei differ curiously in size, for in all

the sections obtained the length is proportional to the distance

from the centre of the lens. This results, in effect, in the forma-

tion of an additional lens, concavo-convex and built entirely of

nuclei, which is placed immediately behind that formed from the

cuticular layer. There can be little doubt that such a provision

as this must have a marked effect on the optical qualities of the

apparatus.

The central part of the organ is oceupied by a number of

minute highly refractive granules which are massed together in

a conical shape round the extremity of the nerve- jin. These

gran ules are quite colourless in every section obtained ; carmine,

picric acid, and hematoxylin are all ‘equally ineffectual in staining

them. The nerve-strand leads straight down the pleopod to the

photophore, and, as may be seen from the figures, 1b expands into

a number of ramifying filaments before it converges to the

granular cone,

Numerous cells with large nuclei are irregularly disposed round

the inside of the photophore. It is possible that these were

pigmented when the animal was living, and served as a screen to

prevent light penetrating inwards, but no confirmation of this

was obtained,

In a freshly caught specimen of Acanthephyra debilis a dark

violet-blue streak is easily seen on each side of the inner wall of

the carapace immediately behind the last pair of thoracic legs.

From the structure which these organs possess it is evident that

they also are photophores, although they are much less highly

specialised than those at the base of the pleopods,

In a transverse section (Pl. LIT. fig. 2) the lens, which is dark

blue in fresh material, is seen to be merely a slight thickening

of the cuticle, and Ete densely staining central layer i 1s entirely

absent. The epithelial cells are gre: ably elongated, as in the

There can be no dowbt that these hodies are nuclei, for by the use of hematoxylin

c FaNcoaas s were demnonstrated in them.

Dare

1910.] PHOTOPHORES OF DECAPOD CRUSTACEA. 645

photophores on the pleopods, but their arrangement is not so

regular, and though the majority of the nuclei are placed near

the lens this is by. no means the case with all. The cells show a

tendency to converge towards a point, but no granules similar to

those in the other photophores were detected. In some sections

a nerve-strand may be seen leading away from the organ and

passing between two muscle-bands.

Other spots and streaks of dark blue pigment to the number of

133 are found in adult examples of 4. debilis. They occur on

the eyestalks, on certain legs, on the branchiostegites and other

parts of the carapace, on the abdominal segments and on the

telson. With the exception of a small number, which occur on

the dorsal aspect of the carapace, abdomen and telson, all ave

situated laterally or ventrally.

Coutiére, while fully realizing that little or no structural

evidence could be advanced in favour of such a theory, has no

hesitation in ascribing a luminous function to the cells under-

lying these pigment- spots . With this view I fully concur, and,

as will be seen later on, | am able to bring forward another fact

which supports this interpretation.

In the only passage in which he remarks on the anatomical

characters of the organs, Coutiére * says: “ Les organes lumineux

de la base des pléopodes paraissent se rapprocher beaucoup de ceux

des Kuphauside, tous les autres semblent étre de simples amas de

cellules a lumiére, disposées sur une seule assise et recouvertes de

pigment.” With the latter part of this sentence [am in entire

agreement. In numerous sections, made through all the more

prominent spots of pigment, the underlying cellular layer presents

no visible difference from that of the adjacent tissue. To this

there is only one exception: a section cut transversely through

the telson near the apex (Pl. LITT. fig. 1) shows that the cells

beneath the dorsal spot of pigment are greatly elongated, though

their nuclei differ in position from those found in the more

elaborate organs.

The brilliant scarlet-red pigment, which is such a notable

characteristic of many deep-sea Decapods, presents features of

special interest in the case of Acanthephyra debilis, for it is quite

undeveloped in the neighbourhood of the luminous organs. This

is particularly well shown in the case of the photophores at the

base of the pleopods. Viewed laterally, these organs would be

quite invisible, being wholly covered by the flaps formed by the

abdominal pleura, were it not that in these parts the red pigment

is entirely absent, leaving the transparent cuticle through which

the light emitted by the photophore may shine as through a

window. The luminous streaks behind the last pair of thoracic

legs are covered by the branchiostegal wall of the carapace, and

in these a precisely similar phenomenon may be observed.

Red pigment is also absent from the vicinity of all the numerous

* Loc. cit. 1906, p. 4, footnote.

646 MR. STANLEY KEMP ON THE [ Apr. 19,

organs Which, when sectioned, show no definite structure. In a

freshly caught specimen each appears as a deep blue spot cireum-

seribed by a belt of colourless tissue. Such a distribution of

pigment seems to afford considerable support to the view,

advanced by Coutiére, that all the blue spots represent luminous

organs.

An examination of young specimens of 4. debilis shows that,

as might have been expected, the complex organs are those which

appear first. The eggs are very large and consequently the young

are liberated in a rather advanced stage. The earliest-known

larva possesses twelve luminous organs, viz., those on the pleopods

and behind the fifth pair of legs. A little later the other com-

pound photophores behind the protopodites of the uropods appear

and simultaneously with them certain of the simple organs. At

every succeeding moult fresh spots of blue pigment appear, until

in the largest individual known they have reached the total of

one hundred and forty-seven *.

Tue PHororyuores or //OPLOPHORUS,

Coutiére (loc. cit. 1905, p. 1) first described these organs in a

species which he named Hoplophorus grimaldii. Thanks to the

kindness of Prof. Gardiner, | have been able to examine an example

of this genus from the Indian Ocean, The specimen is unfortu-

nately immature, but, as might have been expected, the photo-

phores were not found to differ in any essential feature from

those occurring in the closely related form Acanthephyra debdilis.

Although the simpler organs are by no means so numerous as in

that species, all those of a more complex character are present

and occur in the usual positions on the pleopods and uropods, and

behind the base of the last pair of legs.

Couticre mentions that in his specimen of /Hoplophorus, ‘“ con-

serve dans la glycérine formolée,” the luminous organs are slightly

yellowish ; in the example from the Indian Ocean, which was

preserved in spirit, no trace of pigment remains. The photo-

phores are, however, so completely identical in structure with

those occurring in Acanthephyra, that, notwithstanding the fact

that the blue coloration has never been seen in //oplophorus,

there can be little doubt that such a pigment really exists in

living specimens.

Whether the organs occur in all the species or whether, as in

the case of Sergestes and Acanthephyra, they exist only in a

limited number, must be left to future investigators,

The PIGMENTATION OF THE PHOTOPHORES,

It has already been mentioned that a deep blue pigment is, in

life, associated with the photophores of Decapoda, occurring in

* For the order in which these organs arise and their number in specimens of

different age, ». Coutiére, Joc. cit, 1006, and Kemp, Fisheries, Ireland, Sci. Inyest.,

1908, I. {1910},

1910.] PHOLOPHORES OF DECAPOD CRUSTACEA. 647

Sergestes in the first cellular layer and in Acanthephyra in the lens

itself.

There is reason to believe that this pigment is closely allied to,

if not identical with, that found in the Lobster. When the

photophore is placed in absolute alcohol the blue colour soon

becomes bright red, and the same reaction instantaneously appears

when it is boiled in a drop of water. If the lens of Acanthephyra

be dissected out and treated with strong sulphuric or nitric acid

the colour at once changes to red, and immediately afterwards

turns to a dull greenish blue of a much less distinct colour than

that originally present. The greenish-blue tone appears to fade

away a little later, but the concluding stages of the reaction are

somewhat obscured owing to the burning of the tissue by the

acid.

The red pigment which gives the familiar colouring to Vephrops

and to the lobster, when boiled, is known to be one of the lipo-

chromes or fatty pigments, called by Moseley crustaceorubin,

associated with a small quantity of yellow pigment, known as

hepatochrome, which appears to be derived from the liver. The

investigations of Krukenberg * and of Miss Newbigin ft seem to

show that the unstable blue-black pigment or lipochromogen which

occurs in the lobster is a compound of the red hpochrome with a

complex organic base. The blue colour is turned red by any

reagent which alters the form of the proteid, and the red pigment,

extracted and dried, gives with strong acids a brilliant but

evanescent blue reaction.

The photophores are unfortunately so minute that it is not

possible to extract a solution of the pigment; but the reaction

mentioned above, which was obtained by the addition of acid to

the lens of Acanthephyra, furnishes fairly satisfactory evidence of

the nature of the pigment +. The acid breaks up the proteid and

at once converts the blue lipochromogen into the red lipochrome,

and this is immediately followed by the characteristic blue re-

action which this pigment gives in the presence of an acid. The

tissues burn and become brown under the influence of the reagent,

and the rapid evanescence of the blue tint, which is characteristic

in the case of dry extracted pigment, is in consequence somewhat

masked.

It has not been possible to test the blue pigment in the photo-

phores of Sergestes as fully as has been done in the case of

* Krukenberg, Vergleich. Physiol. Studien, IIte Reihe, 3te Abteil., 1882,

pp. 92-107.

+ Newbigin, ‘ Journal of Physiology,’ vol. xxi. 1897, p. 237.

+ The following observations on the red colouring-matter of Acanthephyra may

be mentioned here. An ether extract of the pigment gave a bright yellow solution,

which en evaporation yielded an oily red extract. On the addition of strong nitric

acid a bright, but rapidly evanescent, blue reaction was obtained which was followed

by the separation of the red matter trom the oily yellow pigment, the latter turning

a dull green. This result is practically identical with that obtained by Miss New-

bigin with the extracted pigments of Nephrops. The red colouring which turns

blue under the influence of the acid is the lipochrome, crustaceorubin, while the

oily yellow pigment is hepatochrome.

648 MR, STANLEY KEMP ON THE [ Apr. 19;

Acanthephyra, but from the fact that it tarns red when boiled or

when treated with strong acids it is very probable that it is of

the same nature.

The existence of blue coloration in deep-sea animals is ex-

ceeilingly rare, and its occurrence among Decapoda in close

association with the photophores is almost unique, for among the

EKuphausiacea a similar pigment appears to have been noticed. only

on a single occasion. In November, 1909, a large specimen of

Th ysanopoda acutifrons Holt & Tattersall was caught in a mid-

water net off the West coast of Ireland. This specimen, which

was dlead by the time it reached the deck, was found to possess

patches of deep blue pigment associated with the photophores on

the eyestalks. Casual examination failed to reveal this pigment

in the other photophores, which, however, were of a darker colour

than is usually the case. The specimen was put aside in a dish of

water and when it was again examined, not more than half an hour

later, all trace of the blue pigment had vanished. It is evident

that, even if in this case the blue colouring invariably ocewrs in

the photophores, the phenomenon is one of great rarity among

Kuphausians, for it certainly is not found in J/eganyctiphanes

norvegica or in any of the common N. Atlantic species.

The blue pigment of the photophores of Decapoda is much

more stable than that noticed in 7’. acutifrons. Although rapidly

extracted by alcohol, it will persist for years in specimens preserved

in weak formalin, remaining distinct long after the general red

colouring has dis appeared.

The lens of Acanthephyra, being blue, can necessarily only allow

the emission of blue light and it is not impossible that ‘this. is

also true in the case of Sergestes, where the lens is transparent

and the first cellular layer blue. It seems then that, at least in

the former genus, the production of blue light is a necessity,

but it is impossible to suggest any explanation of this curious

phenomenon,

Photophores have evidently been developed by Crustacea in at

least three separate instances. Those possessed by Acanthephyra

and Hoplophorus ave in structure wholly distinct from those of

Sergestes, while in neither case is there any resemblance to the

very complex organs of the Euphausiacea.

It isa remarkable fact that, whereas in the latter order the

possession of photophores is the general rule (only in Bentheu-

phausia ave they absent), their occurrence in large genera such as

Sergestes and Acanthephyrais limited to a few species only. This

is particularly noteworthy in Sergestes, in which two forms, both

of which are classed among a small group of extremely closely

allied species, exhibit a large number of photophores, whereas

none are to be found in the other members composing the group.

Doflein, in a short but interesting paper *, has summarised the

* Doflein, Sitzungsber. d. Ges. f; Morphol, und Physiol, in Mtinchen, xxii, 1907,

pp. 133-136,

* al Pak i

SiGe PHOTOPHORES OF DECAPOD CRUSTACEA. 649

various suggestions which have been made as to the use of

luminous organs to marine animals. He remarks that they

probably serve different functions in different groups of animals

and classes them in four sections.

i. Attraction of prey (chiefly important in sessile or slowly

moving animals).

ii. Attraction of other individuals of the same species, either

(a) for the formation and maintenance of swarms or (6) to enable

the sexes to find and recognise one another. In this connection

Doflein points out that animals with a complicated system of

photophores always possess highly developed eyes, and refers to

Brauer’s theory that the varying arrangement of photophores

produces light patterns serving as recognition marks, like the

colour-patterns of animals living in daylight.

ii. Protection. The clouds of luminous secretion emitted by

some species may possibly serve the same purpose as the ink of

the cuttlefish, and photophores may also by a sudden flash of light

scare a pursuer. In the fauna of land and shallow water a

brilliant colouring is often assumed as a signal that the species is

distasteful, and some deep-sea animals may, for the same purpose,

exhibit warning lights.

iv. [lumination of objects viewed by the animal. On this

theory it is difficult to account for the ventral and lateral position

of the photophores In many marine animals*. In Crustacea

this is particularly well shown, for the large majority of the

organs illuminate regions which seem altogether out of range of

the eyesight.

It is evident that these suggestions will not account for every

ease which can be found; the photophores in the roof of the

branchial chamber of Sergestes remain inexplicable.

The vast majority of marine animals which possess photophores

live at the surface or at intermediate depths and never occur on

the bottom. No exceptions to this rule have been noticed in the

deep-water fauna of the Irish Atlantic slope, but it seems that

the two Euphausians, Meganyctiphanes norvegica and Nyctiphanes

couchit, ave sometimes found on the bottom in shallow water. On

one or two occasions large numbers of these two species have

been caught off the Irish coast at depths of 40 to 60 fathoms, and

there are indications that the specimens which were obtained in

these hauls were actually living on the sea-floor. The same two

species are frequently obtained over depths of 400-800 fathoms

off the West coast of Ireland, and here they invariably occur in

midwater.

It must be remembered that the ordinary open-mouthed nets,

which are generally employed for bottom work, frequently catch

* Miss Massey informs me that when studying the development of the Cephalopod,

Histioteuthis bonelliana, which when adult possesses photophores all round its

body, she noticed that the organs are developed first on the side which is ventral

when the animal is swimming.

650 ON THE PHOTOPHORES OF DECAPOD CRUSTACEA. [ Apr. 19,

midwater organisms while being hauled, and there is reason to

believe that errors arising from this source exist in many of the

instances in which animals bearing photophores have been

recorded from the bottom.

Many of the higher marine animals live on the sea-floor at

depths to which no ray of sunlight can ever penetrate, and, though

they possess well-developed eyes, are themselves, for the most

part, without any special illuminating apparatus. That light

exists at these depths seems almost certain. It is probably fairly

plentiful in regions thickly populated by Coelenterates, and the

excretions of numerous animals of a more highly organised nature

have been found to be brilliantly phosphorescent. The restriction

of photophores to species living in midwater seems only explicable

on the theory that there is a comparatively plentiful supply of

light on the bottom itself.

The sections of the photophores were made by the paraffin

method in the laboratories of Trinity College, Dublin. Decaleifi-

cation was not attempted, for it was found that the cuticle was

sufficiently soft without it, this, perhaps, being due to the fact

that the specimens were preserved in formalin.

In conclusion, I wish to acknowledge my indebtedness to

De. W. T. Calman for much useful help and criticism, and to

Dr. H. H. Dixon for his valuable assistance and advice in the

preparation of the sections and micro: photographs,

EXPLANATION OF THE PLATES.

Prats LIT.

The figures on this Plate are reproduced from micro-photographs.

Acanthephyra debilis A. M.-Edw,

Fig. 1. A longitudinal section of a photophore from the protopodite of the pleopod.

xX 185. (Compare fig. 1, Pl. LIV.)

2. A transverse section of the photophore behind the base of the last pair of

legs. XX 133.

Prats LITI.

The figures on this Plate are reproduced from micro-photographs.

Acanthephyra debilis A. M.-Edw.

Vig. 1. A transverse section of the telson near the apex, passing through the median

dorsal patch of blue pigment. The section shows the regular arrangement

of nuclei below this area and also the bases of two spines cut trans-

versely. XX 93.

Sergestes challengeri Hansen,

Fig. 2. A median photophore from the thoracie sternum, cut transversely. X 320.

3. Part of the penultimate joint of the second maxillipede, cut longitudinally,

showing a photophore in transverse section. X 247, (Compare fig. 3,

Pi. LIV.)

1. The photophore at the base of the exopod of the first maxillipede in trans-

verse section. X 3800,

1910. ] ON THE VARIETIES OF MUS RATTUS IN EGYPT, 651

Prate LIV.

The figures on this Plate are semi-diagrammatic.

Acanthephyra debilis A. M.-Edw.

Fig.1. A longitudinal section of a photophore from the protopodite of a pleopod.

The cellular layer (c.), which is apparently derived from the epithelium, is

composed of long cells with densely staining nuclei at their outer ends.

The only cytoplasm which is visible lies between the nuclei and the inner

face of the lens. XX 210. (Compare fig. 1, Pl. LII.)

Sergestes challengeri Hansen.

Fig. 2. A longitudinal section of the eye-stalk showing the photophore ( ph.) lying

close to the eye (e.) and separated from it by a curtain of black pigment

(p.). X 160.

3. A photophore from the penultimate joint of the second maxillipede in traus-

verse section. XX 880. (Compare fig. 3, Pl. LITT.)

4. A transverse section of the branchial chamber showing an arthrobranch (a.)

and one of the four photophores ( pf.) which are set in the roof of the

cavity and appear to illuminate the gills from above. X 44.

5, The same photophore on a larger scale. The two layers of the lens (7./. and

o.l,) and the first cellular layer (c.') are formed from the cuticular and

epithelial layers of the inner surface ot the branchiostegite. x 380.

Reference letters :—

a. Arthrobranch. 2.L. Inner layer of lens.

6. Branchiostegite. m.l. Middle layer of lens.

ce. Cellular layer (in Acanthephyra.) n. Nerve.

ce’. First celluiar layer (in Sergestes.) o. Optic ganglia.

ec’. Second cellular layer (in Sergestes). o.c. Outer cuticular layer of branchio-

e. Kye. stegite.

e.l, Epithelial cell-layer. o.l. Outer layer of lens.

g. Cone of minute highly refractive p. Curtain of black pigment between

granules in close connection with photophore and eye.

nerve-strand. ph. Photophore.

z.c. Inner cuticular layer of branchio- 7. Reflector or striated layer.

stegite. s.4. Sheathing layer of cells.

2. On the Varieties of Mus rattus in Egypt; with General

Notes on the Species having reference to Variation and

Heredity. By J. Lewis Bonnorts, M.A., F.L.S., F.Z.S.

[Received December 22, 1909. |

(Text-figures 58-62.)

While spending a few months recently at the Giza Zoological

Gardens, near Cairo, I was enabled through the kindness of the

Director, Capt. 8. 8. Flower, to examine a large number of the

common House Rats of the district. I gladly took advantage of

the opportunities thus offered, as I was convinced that a close

study of this species would throw some light both on the causes

of variation and on the inheritance of the varieties that are

found in such profusion in Jus rattus.

In addition to the rats which were caught in the Gardens,

Dr. Charles Todd, of the Public Health Department, kindly

allowed me to examine and measure all the rats that came into

his hands during the time that I was in Cairo, These rats were

taken in various towns and villages in the Delta by special catchers

employed by the Public Health Department, the result being that

652 MR. J. LEWIS BONHOTE ON THE (Apr. 19,

some 500 rats in all were carefully examined and measured in the

flesh.

Before detailing the results of this examination 1t may perhaps

be as well to review roughly the status and problems con-

cerning this species, which is best known in this country as the

“Old English Black Rat.” Jus rattus is by far the commonest

* ship- ‘rat and has thus become practically cosmopolitan in its

distribution, but when it comes in contact with J/us norvegicus*,

our common rat, a larger, stronger and more pugnacious species,

it has to give way, and owing to this fact it is now extremely

scarce in England.

Tu the East (India and the Malay States) Mus rattus is still

holding its own, although in those localities J. norvegicus is

gradually increasing, and in Egypt, while still very abundant and

in some places the only house-rat, it is slowly but surely being

superseded by the larger and stronger species.

The ready variability of Mus rattus together with the continual

crossing of different forms due to the importation of foreign

varieties on ships, has given rise to such a mass of variation, that

the work of the systematist has been rendered very difficult,

The actual localities from which my specimens came were Ath,

Damankhour, Abu Homos, Giza, Faytim, Foueh, and Cairo. In

all except the last two both Jf. rattus and M. nor vegicus were

found in about equal numbers; at Foueh, however, out of 80

rats only three were MW. nor vegicus. From Cairo itself I did not

see many rats, but all those met with were I. rattus ; and Capt.

Flower tells me that I. norvegicus is not yet known there. As

regards the Giza Gardens, which lie on the opposite side of the

river to Cairo, the latter species has only appeared within the last

eight years and forms now about 30 per cent. of the rats. there.

It has entirely ousted the field-rat (Arvicanthis) which used to

abound in the Gardens.

As regards the proportions of the sexes, both J/. norvegicus and

the two forms of J/. rattus show a slight excess of females, the

actual figures being as follows :—

Percentage of males:in MW. norvegicus..........6008 42 °/,

uA - EPR EECLOTILMN 0 con penne “44 °/,

» aaa) ) A) 2

+ M. 7. alewandrinus ...... 50 ve

The higher percentage of females in ./. norvegicus, if always

present, would tend, apart from other reasons, to cause a higher

vate of increase of this species at the expense of other species.

In both species there is practically no difference in the size of

the sexes, but the males have a wider range of variation.

Thus we have in

M. norvegicus 3’s, hind foot 28-43 mm., 2’s 30-44 mm.

M. rattus 3's, A 26-38 mm.. ?’s 29-38 mm.

* Mus norvegicus Erxl, antedates Mus deewnanus Pall. by which this species is

better known.

1910.] VARIETIES OF MUS RATTUS IN EGYPT. 653

Mus rattus in the Indian Region.

In the Indian Region in particular this species has been

burdened with a tremendous number of specific names, given to

any variety which did not appear to fit in with the existing

descriptions ; and with a view of partially clearing up this tangle of

names, I went into the matter at some length a few years back *.

Briefly summed up my conclusions were as follows :—

If the large towns and seaports, where the rat population is

apparently as cosmopolitan as the human, are put out of con-

sideration it appears :—

(i.) That in most localities three fairly distinct forms are

found ;

(i.) That in any particular locality individuals belonging to the

same form do not show much variation and intermediates

between the forms are comparatively scarce ;

(iii.) In most localities the same three forms are easily recog-

nisable though differing slightly from their corresponding

varieties in other places.

I, therefore, divided the species into three subgroups, re-

presenting the three forms, and distinguished them as follows :-—

(i.) Jalorensis subgroup.

Hairs of underparts white to their bases. Hind foot 30 mm.

(i.) Raufescens subgroup.

Hairs of underparts white with slate-coloured bases. Hind

foot 33 mm,

(ili.) Griseiventer subgroup.

Hairs of underparts entirely slate-coloured or tipped with

fulvous. Hind foot 35 mm.

In addition to the characters given above, the subgroups

appeared, from collectors’ notes, to have slightly different habits,

the last-named being found chiefly in houses, Jalorensis in the

hilly districts, and Rufescens in the date-palins.

On this plan, it became comparatively easy by careful reference

to original descriptions and specimens to refer almost all the

varieties which had been described, to one or other of the sub-

groups, and then, by noting the type locality of each variety, the

mass of names became reduced to a manageable and fairly logical

order. Where two descriptions had been applied to rats from

the same locality, 11 was in most cases at once obvious from those

descriptions that they were not synonyms but referred to in-

dividuals of different subgroups.

Without more material I did not feel justified in making

further comment, but left the work as a foundation to be im-

roved and elaborated should a future opportunity arise.

In 1907, in the Mem. Ind. Mus. i. no. 1, Dr. Hossack has

* “Fasciculi Malayenses,’ i. Zool. pp. 28 ef seqq. (1903).

+ I named the subgroups, not after the oldest species but after the one of which

the original description would leave least doubt as fo the main characters of the

subgroup.

Proc. Zoou. Soc.—1910, No. XLITI. 43

654 MR. J. LEWIS BONHOTE ON THE [Apr. 19,

criticised my paper and comes to the conclusion that my effort

has not cleared up the confusion and that all these rats belong to

one and the same species—J/us rattus.

Dr, Hossack’s remarks, fair and just as they appear to be, fail

from the fact that we were working at the subject from entirely

different points of view—-he as a medical man, anxious to know

the number of species by which the plague might be conveyed—

I, as a zoologist, seeking for a law and order in the apparently

innumerable rani ctice oan among these rats. For Dr. Hos-

sack’s purpose it is sufficient to know that there are but two

species of true rats, usually found in houses—the short-tailed,

heavy-built and practically invariable Mus norvegicus, and the

long, slender, and extremely variable J/us rattus, which includes

all the varieties bearing the mass of names that so confused my

evitic. Had he written to me in the first place, I could have told

him at once that all these long-tailed rats belonged to one very

variable species, and that in the large towns, where all his

material was procured, no order or classification was possible as

the varieties had become hopelessly mixed and crossed, but that

in the country places the varieties were much better defined,

and appar ently bred true. The question of the classification and

study of these varieties is a specialised study for zoologists, and it

is not to be expected that a medical man, studying the plague,

should be able to devote the time necessary for the thrashing out

of such matters.

It would therefore have been better had he applied to reecog-

nised authorities for his information, rather than attempt a task

in a specialised subject outside the scope of his investigations.

IT am glad to have had this opportunity of replying to

Dr. Poseaek, for many people do not yet realize that the study of

geographical forms and minute variations has a ver y great bear-

ing on many large and important zoological problems, but need

wat concern the man engaged on other ¢ ognate investigations, anid

in the present instance ne is sufficient for him to know that all

these varieties of the long-tailed rat belong to one species, J/us

rats.

Mus rattus in Loypt.

In Egypt we find two distinct forms of J/us rattus—the one

with pure white underparts and white feet, which was named

Mus tectorum* by Savi; and the other with slate-grey underparts

and dark feet, which may be known as Jus alewandrinus tT.

Between these two forms many intermediates oceur, the actual

typical forms being comparatively rare }.

* Mus tectorwn Savi, Nuovo Giorn. Pisa, 1825

+ Mus alewandrinus Geott. Deser. de ?Egy te. Hist. Nat. ii. p. 733; Atlas, pl. v.

fig. 1 (1812).

"+ This statement apparently contradicts the remarks on p. 653, but it must be

borne i in mind that those remarks were based on Malay material, obtained in centres

removed from the influence of foreign shipping - whereas in the present case the

specimens came from’ populous centres, contiguous to a large waterway (the Nile

and its branches).

¥ ae a pile

1910. | VARIETIES OF MUS RATTUS IN EGYPT, G55

I have already referred to the fact that in India the three

varieties of MW. rattvs may roughly be considered as hill, tree, and

house forms; but in Egypt, where the cultivated parts frequented

by these rats are low-lying and level plains, only two forms were

found.

I made a number of careful observations in the Giza Gardens,

where both forms are found, to see whether the two varieties were

really different in their fhe Rasy but such evidence as I was able

to get together was very inconclusive. Large numbers of plane

trees grow in the Gardens and on these, often ata considerable

height. from the ground, many heaps of the fruit, looking not

unlike squirrels’ ‘dr ‘eys, were to be found. These were un-

doubtedly brought together by rats, but investigation proved that

they were used merely as convenient platforms on which to feed,

and not as sleeping or breeding places. The seeds of these trees

ave apparently a favourite food, and as it would be impossible for

a rat to devour them 7 situ, they are picked and carried to the

first convenient spot, generally a fork near the main trunk, to be

devoured, and it is on such places that the accumulated débris

forms the apparent nests. On the other hand, the fruit which

had fallen to the ground was frequently taken into a neighbouring

bush and eaten there, so that there is undoubtedly a tendency to

an arboreal life.

On two occasions only did I actually see a rat in a tree, and

neither time were they accurately identified, although one, I am

almost certain, belonged to the white-bellied form (MZ. tectorum).

Both forms were found living in burrows in the ground as well

as in the houses, so that there seems to be but little difference in

habits between the two varieties. The apparent absence of the

third (Indian) subgroup, however, taken in correlation with the

lack of high ground, seems certainly suggestive, but I shail have

more to say on this point later.

In studying the various forms of Mus raéttus I found it con-

venient to classify them tmto four groups.

(i.) The Typical Tectorwm, ia which the hairs of the under-

parts are white to their bases and the upper surface of

the hind feet is also white.

(ii.) The Grey Tectorwm, in which the hairs of the underparts

are white to their bases but the upper surface of the hind

feet shows traces of brown.

Gil.) Zhe Grey Alewandrinus, in which the hairs of the under-

parts have slate-coloured bases and the upper surface of

the hind feet.shows much brown.

(iv.) Zhe Typical Alexwandrinus, a which the hairs of the under-

parts are slate-coloured throughout or nearly so, and the

hind feet entirely brown on the upper surface.

Tn actual shade of colour much variation was shown. Groups

Gil.) and (iv.) were constantly darker than groups (i.) and (ii.),

| 43*

656 MR. J. LEWIS BONHOTE ON THE [Apr. 19,

but in classifying them, reliance was only placed on the characters

noted above, “which allowed no possibility of doubt.

| Mus nor vegicus, on the other hand, proved very constant and

no variation in colour was noticeable. In this species, the hairs

of the underparts have slate-grey bases and the upper surface of

the hind feet is pure white, a combination of characters appar-

ently not found in WW, rattus from Egypt.

Apart from this distinction, JZ. norvegicus is a larger and more

stoutly built animal; the length of the tail never exceeds that of

the head and body, and the ears are shorter and more rounded.

There are also some easily recognisable skull differences, which

need not be gone into here. |

Every specimen was carefully measured in the flesh by myself

with the same ec: ullipers, so that errors from a comparison of

measurements are reduced to a minimum. ‘The measurements

taken were: snout to root of tail, tail, hind foot, ear.

The largest measurements in mm. were as follows :—

Head &

body. Tail. | Har. Hind foot.

Mus norvegicus. 6 ad. No. 250...; 253 233 22 | 43

a i: Gad, No.215...| 225 | 204 | 19 | 41

Mus 1. tectorum. dad. No. 203...) 188 | 232 24 | 30°5

4 45 9 ad. No. 181...) 190 232 24 365

Mus vr. alexandrinus. g ad. No. 290...) 199 | 220 26 35

. 55 Q ad. No. 287... 195 208 26 35

|These measurements are given only as showing the greatest

sizes attained; they are those of individuals and in no sense

average measurements, so that, for instance, it must not be

assumed that the ears of J. alewandrinus are on the average

larger than those of MW. tectorwn. Owing to the difficulty of

accurately estimating the age of any individual, it is impossible to

give average mea surements. |

7 The proportion: ite lengths of the body and tail measurements

were found to vary in beh species, but in J. norvegicus the tail

never exceeded the body length, and in J/, rattus the tail was

always longer than the body length.

The most reliable measurement therefore on which to make

comparisons and the only one which dealt with a struetural

feature and for that reason admitted of least error, was that of the

hind foot. This measurement may therefore be taken as a fair

criterion of general size and as forming a good and reliable index

for comparison.

Curves of this measurement were made, and on referring to

that of Mus rattus (Table L., text-fig. 58), we note that the male

Aor =

1910. ] VARIETIES OF MUS RATLUS IN EGYPT. 657

Text-fig. 58.

Length of Hindfoot in mm.

26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

») DEERE De ee

a a a

BERET

Individuals

ES]

ial

% CSTE es oe ke

= 1

5 TT ie

Pho Se eases

SASS Cae

CTPA TSS

TiO UE ene a

eG SE SLT a a

Taber I.

Curve of hind-foot measurements of Mus ratéus (all forms).

The dotted line represents measurements of the males.

658 MR. J. LEWIS BONHOTE ON THE [Apr. 19,

curve (dotted line) shows three apices at 29, 33, and 35 mm.,

and that the female curve shows an apex at 30 and 35 with a

break in continuity between 32 and 34; that is to say, both sexes

show three apiees, and that therefore a larger number of rats

have hind feet of those three measurements than of the inter-

vening sizes. Now, as these measurements include rats of all

ages, and as there is no reason why there should be more of one

particular age than another, seeing pea the breeding season is

fairly continwous throughout the yea it follows th: at we have

here three ‘limits of erowth,’ or, in Lanee words, that certain

individuals reach their limit of size at these three points, and

consequently that the numbers at these points tend to accumulate

and thus to form the apices in the curve. Those who accept

Mendel’s theory would here maintain, and probably with justice,

that each apex represents a mutation, and that around these

mutations fluctuating variations would occur so that, unless closely

examined, the whole sertes would appear as a case of continuous

variation,

Text-fig. 59.

Length of Hindfoot in mm.

26 27 28:29 203l032 33 3435.36.37 (S832 404i te

of Individuals.

Number

Tasre IT.

Curve of hind-foot measurements of IWus rattus tcctorum {Groups i, & ii.)

1910.] VARIETIES OF MUS RATTUS IN EGYPT. 659

Text-fg. 60.

Length of Hindfoot in mm.

= 27 28 2930 31 32 33.34 35 36 37 38 3940 41 42 43 44

= IS

SRR AR

“S 10 cal

3 See it doe

Sean aaa ee

= BL

A a eT

PNA EENEE

Vane ITT.

Curve of hind-foot measurements of Mus rattus alecandrinus | Groups iii. & iv.).

(N.B.—Tables I. & III. are represented together in Table I.)

aed nf Nt SLL lio | a

Tables IT. and III. (text-figs. 59 & 60) show the curves of

M. 7. tectorum and WM, +. alewandrinus taken separately. In the

first table I have included all the rats which would fall under

groups (i.) and (i1.), and in the second those included in

groups (i11.) and (iv.) (see ate, p. 655).

The tectorwm males show apices at 31 and 35 mm. witha shght

check at 33, and the females at 30 and 35 with a slight check at

32-33. In alewandrinus the males show apices at 29 and 33 with

a slight check at 35, and the females a slight apex at 33 with a

large one at 35. There were no females of alexandrinus below

30, which accounts for the absence of the first apex.

This comparison makes it very clear that the apices in the

general curve (Table I., text-fig. 58, p. 657) are not merely due

to accident, but that the middle one at 33 represents the variety

known as dlevandrinus, and the largest one at 35 yepresents

the white-bellied fectorwn. The fact of the curves in Tables IT.

660 MR. J. LEWIS BONHOTE ON THE [ Apr. 19,

and IIT. showing traces of the other : apices, does not invalidate the

reasoning as groups (ii.) and (iii,) are intermediate forms.

Mus rattus.

Comparison of Indian and Egyptian Forms.

The next question to arise is, what is the significance of the

third and smallest apex at 29 and 30 mm, On reference to my

paper quoted above in which I had divided J/, rattws into three

subgroups, we find the hind-foot measurements as follows :-—

Jalorensis subgroup, hill form. Hind foot 30 mm. (29-31°5).

Rufescens i tree form. - 33 mm.

Griseiventer — ,, house form. + 35 mm.

These measurements, it will be noted, correspond exactly to

the three apices in the eurves of the Egyptian rats; so that we

ave, | think, justified in considering the small apex at 29 to

represent the bill form (J/alorensis subgroup) of the Oriental

Region, but which has become, in the level country of Egypt,

unrec ognisable exter nally.

This is a point which is of extreme importance from a deeper

and more general point of view, for here, in a low-lying country,

we have traces of a variety which in another part of the world

has become the predominant form in the hills. Now, if con-

tinuous variation were the only method of evolution, we should

find that in a level country all variations suitable to the hills

would be eliminated, so that, in course of years, no trace of them

would remain as they would be swamped by the more suitable

forms, but apparently this has not happened in the present

instance. The small-footed variety is still present in Egypt,

permeating the race, and still existing in sufficiently strong

numbers to spring up again quickly and become a dominant form

should it ever find itself under suitable conditions.

Although, perhaps, not absolutely conclusive, this affords con-

siderable evidence that the three subgr oups of J/. rattus, which

I first differentiated in 1903 and which are strongly confirmed

by the present work, probably arose as mutations.

There is yet a further point. One of the characters of the

Jalorensis subgroup (the hill form) is the pure white hairs of the

underparts. In this case it is associated with a small hind foot,

but in Egypt we find it in WV. tectorwm, which has the largest

hind foot of the three forms. . It is therefore evident that the

colour of the underparts is a character which has not always

been inherited in connection with the size of the foot, but may

occur in any one of the subgroups, so that in one country it may

he cha’ aracter istic of the hill form, in another of the tree form, and

SO on *

* | um aware that on the evidence so far brought forward it might be argued

that, if the colour of the underparts is not always found in correlation with the

same sized hind foot, it is a vhavacter ef but little use in distinguishing the sub-

1910.] VARIEITES OF MUS RATTUS IN EGYPT, 661

Comparison of M. vattus and M. norvegicus.

Passing now from J. ratius to MW. norvegicus and comparing

Tables I. (text-fig. 58, p. 657) and IV. (text-fig. 61), we find a

striking and surprising analogy in the curves. As in the case

Text-fig. 61

Length of Hindfoot in mm.

. 28 29 30 3! 32 33 34 35 36 37 38 39 40 41 42 43 44

aia ANSP |

Pee Cee)

ote ic

ToCTRESUEN

Number of Individuals

FEE CA pes

Hie fp zn BO

Tasre LV.

Curve of hind-foot measurements of Ius norvegicus.

of WM. rattus, M. norvegicus also shows three apices at 32, 57, and

39, but in this case there is no colour variation, and therefore no

groups, for being a separate mutation it might be found (if the hind-foot measure-

ment were taken as the determinant character of the subgroup) in two or more of

the subgroups.

The answer to this objection according to our present knowledge is that in

any particular locality we have always found it associated with ore, and only one,

size of hind foot; that is to say, always in the same subgroup as defined by

the size of the foot. Furthermore the breeding experiments, so far as they have

gone, seem to point to the fact that it is inherited in correlation with the size of the

hind foot in the rats from Egypt.

662 MR. J. LEWIS BONHOTE ON THE [Apr. 19,

means of individually distinguishing the different forms or muta-

tions, as we may now, I think, call them.

Here, therefore, in an entirely different species we find the

sume mutations present, and whereas in the case of the one

(/. rattus) we ave enabled to distinguish easily the various forms

owing to the combination of the hind- foot character with a colour

character, yet in the other species (JZ. norvegicus) we are unable

to subdivide it into three forms, owing to the impossibility of

definitely detecting which character is present in any particular

individual. Nevertheless we have cognisance of characters

(mutations) which cannot be recognised by any external inspec-

tion of individuals, although we have proved their presence

beyond a doubt.

Tt would of course be Bu to be too positive, yet it is ex-

tremely unlikely that these rats cend inany way to pair according

to their foot character. If the characters were therefore merely

individual variations, they would undoubtedly tend to be swamped

and an even curve would result. This, however, is not the case.

| For purposes of comparison we have added (Table V., text-

fig. 62) a curve of the hind foot of Acomys cahirinus ; this, it

will be noted, is a perfectly simple curve with only one apex.

As the number of Acomys measured (about 100) was nearly the

same as the number of J. alewandrinus, the comparison is a

perfectly fai: one. |

We have here, therefore, in J/. norvegicus a splendid example

of a Mendelian character (utation) being carried on unimpaired

through generations ; and there can be little doubt that if these

animals found themselves under certain conditions where small

feet, for example, prove of great advantage, very few generations

would suffice for the small-footed race to become the dominant

form.

The Mendelian theory enables us to understand how a small-

footed race could be produced in a very short time, or how, if that

favourable mutation were not present, the species might die out

before the cumulative effects of Natural Selection on continuous

variation would have time to act.

General Notes.

Much work of value still remains to be done by enquiry into

the gradual formation of species, and these rats « fler a wide field

for the study of minute variations. Many groups of the smaller

rodents as, for example, rats of the J/us jerdoni group and many

of the Oriental groups of squirrels, offer similar problems of great

complexity. In the former case we have numbers of very closely

allied forms all inhabiting the same district, and each form has,

auppuwently, its representative in other neighbouring districts.

Systematists of late years have been content to burden each of

these forms with a different name and there to leave the matter.

Such work, necessary as it may be, can be but a- preliminary.

step, and unless the subject is pursted further, the chaos

1910.) VARIETIES OF MUS RATTUS IN EGYPT. 665

brought about by a multitude of names is worse than the old

5 . . .

plan by which one name was made to cover all varieties.

More recently still, the rediscovery of Mendel’s theory has, at

Text-fig. 62.

Length of Hindfoot in mm.

pes SSeneece

Number of Indiviaua/ls

Bis ds

Be eo pee ca

see ae

Saf dai abi

TaBLe V.

Curve of hind-foot measurements of lcomys eahirinus, to show simple curye

for comparison with previous Tables.

664 MR. J. LEWIS BONHOTE ON THE [Apr. 19,

least, given a working hypothesis on which we may attempt to

solve some of the problems of heredity ; and pach J am far

from admitting that Mendel’s law is the sole, or even the chief,

factor governing heredity, it does undoubtedly he us to under-

stand how these closely allied forms can either (1.) exist in the

sume locality as distinct varieties, or (i1.) in another locality

become apparently lost although a closer inspection shows them to

he present, only waiting for a suitable environment in which to

develope,

‘To prove this matter further, it becomes necessary to conduct

Mendelian experiments on these varieties. This I have begun to

do ona small scale. The experiments are, however, as yet in the

initial stages and no definite results can be stated. So far as

they have gone, we find that pure J/. tectorwm mated with pure

A. tectorum have produced pure M. tectorwm. Pure MM. alewan-

drinus when paired together have also bred absolutely true. A

pair of grey JZ, tectorwm (group il.) showing slight traces of brown

on the feet produced young like themselves (g roup il.) and grey

M. alewandrinus (group iii.).

As far as it goes, therefore, this evidence seems to further

confirm the fact that J/us vattus should not be regarded as

a species in which variation has run wild, but rather as a species

containing certain definite mutations, which when mated together

breed perfectly true. From its wandering habits and_ prolific

breeding much intererossing takes place, giving rise to these

innumerable varieties, but no better evidence could be had of the

purity of the gametes than the fact that when the apparently

pure races are mated together (even when captured amongst a

mixed crowd) they breed absolutely true.

Summary.

The main points that have been brought out in this paper are

as follows :—

(1.) Two varieties (and intermediates) of J. ratlus ave found in

Keypt.

(ii.) The hind foot is taken as a standard measurement and

curves of this measurement drawn.

(iii.) These curves show three apices, the first being very

sinall.

(iv.) These three apices correspond with the length of the

hind foot typical of the three subgroups of J/, rattus in the

Oriental Region described by myself some years ago,

(v.) Hence it is argued that these three apices probably repre-

sent mutations, and that the first and smallest apex represents

the Jalorensis subgroup found in the Oriental Region, but which

from some cause or other is practically non-existent in Egypt.

Since, however, a mautation cannot be destroyed, we still

find it in a small percentage of individuals ready under favour-

able circumstances to increase,

1910.] VARIETIES OF MUS RATTUS IN EGYPT. 665

It is pointed out that this may account for the very quick way

in which some species accommodate themselves to altered surround-

ings, since if a favourable mutation is present in the species very

few generations would suffice to make it the dominant form,

(vi.) In M. rattus we found that the length of the hind foot

was also correlated with external differences, and that the white-

bellied J. tectorum had a longer hind foot than the darker

M, alexandrinus.

Although in the Oriental Region the pure white underparts

is characteristic of one of the subgroups, yet in that region it is

correlated with a different sized hind foot to that with which it is

associated in Kgypt. Therefore the length of the hind foot and

the colour of the underparts, although both pure mutations, can

probably be inherited imdependently.

(vul.) An examination of the hind-foot curve in JL. norvegicus

shows also three apices, but in this case there is no colour-

character by which the mutations may be distinguished externally.

The length of the hind foot as a mutation is therefore a character

common to two or more species.

(vili.) It is suggested that this enables us to understand how

several very nearly allied forms of the same species (e. ¢. in Mus

jerdoni and some of the Sciuride) may exist in the same locality

without losing their characteristic differences, however small

those differences may be.

(ix.) Further investigation on these lines is required by means

of experiments in heredity, and so far as the author has.carried

these out the pure varieties of J. tectorum and M. alewandrinus

were found to breed perfectly true.

(x.) Lastly, this paper claims to show that J/us rattus is a species

containing many slight but definite mutations which, as far as

the evidence goes, breed true when paired together, and that the

apparent innumerable and indefinite variations are merely due to

these animals being carried all over the world and mixing together

in the large seaport towns.

In short :—The varieties in I/us rattus appear to run on definite

lines and to have arisen as mutations, they are therefore inherited

on Mendelian lines. Of the three main varieties found in the

Oriental Region only two occur in Egypt, but this paper shows

the presence of the third, though in very small numbers. One of

the characters of these varieties is shown to be present in another

species, J. norvegicus, although it cannot be distinguished in any

particular individual. It is further hinted that many of these so-

called species which are very closely allied have probably arisen as

mutations, and that it is due to this fact that they are able to exist

side by side under precisely the same conditions and yet preserve

their characters intact. It may also be noticed that the colour-

character of the hairs on the underparts is also found in another

species, namely, the house-mouse, I/ws musculus. MM. m. gentilis

has the hairs white to their bases, while J/. m. orientalis has slate-

coloured bases. Both these forms occur in Egypt.

666 MR. G. E. BULLEN ON POSTERIOR [Apr. 19,

3. Onan example of Posterior Dichotomy in an Aylesbury

Duckling. By G. E. Buiien.*

| Received March 14, 1910.)

(Text-figures 63 & 64.)

It is well known that instances of dichotomy and the re-

duplication of limbs occur not unfrequently in domesticated

mammals and birds. Many examples of such malformations

have been recorded hitherto by such authors as_ Forster,

St. Hilaire, Alhfeld, Albers, Ballantyne, His, and Bland Sutton.

The last named, in his work entitled ‘ Evolution and Disease,’

deseribes amongst other ex: vmples of birds having supernumerary

pelvic and other limbs, a chick, which presents the most frequently

observed characters of posterior dichotomy. In the main, the

subject of the present paper is somewhat similar, but with the

addition of certain well-marked deviating char: weters which do not

appear to have been described by any previous author.

Text-fig. 63.

Aylesbury Duckling, with supernumerary legs (reduced).

The specimen under discussion (text-fig. 63), now in the

* Communkated by Joun Tlerkryson, F.L.S., FLZ.S,

1910.) DICHOTOMY IN A DUCKLING. 667

Hertfordshire County Museum, was hatched in July 1909. and

lived for about twenty-four hours. There appear to have been

Text-fig. 64.

Cartilaginous

process.

Pelvis.

Femur:

ie Tibia-Tarsus.

Tarso-Metatarsus.

Diagram showing form and relative position of bones composing supernumerary

legs (twice enlarged).

two individuals with supernumerary legs from the same batch of

eggs, but the second specimen did not pass into the author’s

668 ON POSTERIOR DICHOTOMY IN A DUCKLING. [Apr. 19;

hands. It was stated that the autosite, in both instances, had no

control over the supernumerary legs.

Briefly to summarise the result of the dissection, the following

features were found to exist in the part dichotomised, which

occupied a median position on the ventral aspect, and, as usual

in such cases, was stunted generally in growth. The pelvis was

reduplicated and malformed ; the femora were fused together, and

supported three, and not two stunted, tibie. The fibule and

satellee were entirely absent. The two outside limbs were further

well-developed as described later, whereas the third limb holding

a median position on the base of the confluent femur terminated

with a tarso-metatarsus, reduced to a mere spike-like projection,

covered, however, with scaly integument.

The reduplicated pelvis did not appear to be attached in any

way to the body of the autosite, but was contained entirely within

the integument. Its form, as shown in the illustration (text-

fig. 64), would appear to suggest a tendency toward atavistiec

structure, embodying vs it does certain characters seen in the

iliac arch of Batrachians.

The writer desires merely to place the fact of this apparent

similarity before the reader, without throwing any particular

stress upon it, for as Gegenbaur has insisted, and his opinion has

been backed by subsequent investigators, atavism cannot be ac-

cepted as an explanation of resemblances between forms that are

systematically far distant. It is generally found that striking

instances of supposed atavism are the result of misinterpretation.

The total length of this presumed pelvis from its base to a

point midway between the tips of the two furcations is 10°5 mm.

From paired slight processes occurring at the bases of the

furcations, there arise two unciform cartilaginous processes,

each extending upward in contact with the osseous parts, to a

point about 2 mm. from the apex of each furcation. The free

ends then deflect inward toward the body of the autosite, the

tips being obtusely rounded and in no way connected with the

walls of the abdomen.

The confluent femur occurs as a free jot covered with skin,

and is attached by muscle, and lateral tendons, to the symphysis

yubis, a single caput femoris of minute size occurring directly

hehind toward the ventral aspect of the autosite. The total

iength of this bone is 16 mm. At its lower end it broadens out

to form three distinct series of condyles articulating with the

stunted tibie. The patelle, as already stated, appear to be absent,

and the fibule, if present, are fused with the tibie, What may

perhaps be termed the outer pair of these bones are of even

length, viz. 5 mm., whereas the single centre one is 7 mm.

The two outer tarse-metatarsi are large in proportion to the

tibie, each being 21 mm. in total length and covered with scaly

integument. The hind toe in each is lacking. The right foot

otherwise is normal in structure, but the left lacks the third toe,

its place being taken by a broad elliptical flap of web membrane.

1910.] ON THE NORTH QUEENSLAND GREEN TREE-ANTS, 669

The left tarso-metatarsus and foot, moreover, are directed inward

toward the body of the autosite, presenting a three-quarter

plantar aspect when the specimen is viewed from the front.

In the central limb, as previously stated, the tarso-meta-

tarsus is represented merely as a spike-like projection 6 mm. in

length.

The subject of the present paper is interesting mainly in the

fact that the supernumerary paired limbs are again partially

dichomotised by the presence, in articulation with the confluent

femur, of a third tibia and tarso-metatarsus, the latter imperfectly

formed and stunted.

For assistance in the preparation of this paper my thanks are

due to Mr. John Hopkinson, F.L.S., I’.Z.8., and Mr. E. Leonard

Gill, M.Se.

May 3, 1910.

Dr. A. Smira Woopwarp, F.R.8., Vice-President,

in the Chair,

The Seeretary exhibited a photograph of a Giraffe, kindly lent

him by Mr. Carl Hagenbeck. The animal was an adult female

3:30 metres in height, obtained in Gallaland, East Africa, and

was in the possession of Mr. Hagenbeck. It displayed a

reticulate coloration, in which the lighter areas were reduced to

narrow lines, clearly on the neck and body, but the hind-quarters

were blotched, while the spotting came lower down on the legs

than in the typical Giraffa reticulata. It possibly represented a

type intermediate between G'. reticulata and G'. camelopardalis,

but it was to be remembered that the specimen was a female, and

that females did not exhibit the typical coloration so conspicuously

as old males.

Mr. E. E. Austen, F.Z.S., exhibited a specimen of a rare Fly

(Cephenomyia ulrichii Brauer) from East Prussia, parasitic in the

larval stage in the throat of the Elk (Alces machlis Ogilby), and

remarked on its habits and those of allied species. This specimen

had been recently acquired by the Hon. Charles Rothschild and

presented by him to the British Museum,

Dr. R. A. O’Brien, introduced by Prof. E. A. Minchin, M.A.,

F.Z.S., exhibited a series of lantern-slides illustrating the habits

of the North Queensland Green Tree-Ant (Oecophylla smarag-

dina ®). The Green Tree-Ant is found in the open forest

Proc. Zoou. Soc.—1910, No. XLIV. ™ 44

670 DR. H. B. FANTHAM AND MISS ANNIE PORTER ON | May 3,

country or “ bush,” on the edge of the thick jungle or ‘ serub,”

along the sea-coast of Northern Queensland, It is a very active

pugnacious insect, from half to three-quarters of an inch long,

living wholly or almost wholly in trees. In riding through the

country one is frequently covered with the actively- biting insects

which fall from a bough touched in passing. Fortunately the

bite, though momentarily severe, is not followed by any sub-

sequent irritation.

The nest is built on the bough of a tree and consists of a very

large number of leaves , generally fresh and green. ‘These are

matted together with sh gelatinous material exuded from larve

which the worker ants bri ing up to the site of the projected nest,

where other ants hold the edges of adjacent leaves together.

This process is repeated until the bundle may be several feet in

diameter.

The bridge formed of the bodies of the ants shown on the

screen was re-formed, when broken, by festoons of ants hanging

from the upper leaf until some of them, dropping from the festoon,

joined momentarily with others on top of the leaf and twigs

below. Others quickly joined in strengthening the bridge until

it was about four ants’ width and eight ants’ length. Then the

one leaf was seen to be dragged slowly nearer the other, decreasing

the bridge to five ants’ lensth, and at this stage it remained for

several days, when I left the place.

The ants ‘fon duty” in the bridge over which other ants ran

to and fro carrying their ‘‘ game” were watched carefully for

80 minutes, and none in the centre of the bridge was relieved

during that time—a rather remarkable feat of str ength and

endurance.

On a new Trypanoplasm, 7. dendrocali, sp. n., from

Dendrocelum lactewm.

Dr. H. B. Fantham, F.Z.8., exhibited, on behalf of Miss Annie

Porter, D.Se., and himself, preparations and original drawings of

a new Jr ypanoplasma (Cr yptobia) from the alimentar 'y tract of

Dendrocelum lactewm. In some of the preparations the living

organisms were shown, ‘The parasites measured from 20 ye to 40

long approximately. They were biflagellate, the poster ‘ortincenent

forming a border to the undulating membrane. The blepharo-

plast (kinetonucleus) was large and often somewhat curved. It

was proposed to call the parasite 7’rypanoplasma (Cryptobia)

dendroceli, There was evidence of hereditary infection of the

Planarian by the 7rypanoplasma, for the parasites had been seen

to enter the eggs of the host, and in some cases to divide

longitudinally therein.

Breeding-experiments on Dendroceluwm were in progress in order

to obtain further information concerning the hereditary trans-

mission of the parasite. Dr, Fantham and Miss Porter thought

1910.] A NEW TRYPANOPLASM. 671

it probable that the hereditary method was not the only means

of the transference of the parasite from host to host. They had

seen Trypanoplasms become rounded, the rounded parasites being

probably resting forms which, perhaps, encysted and passed out

from the host in this condition, Living fl: well: ate Trypanoplasms

had also been seen to pass out of the host on a few occasions.

Fresh Dendrocalum lactewm may thus become infected by the

mouth, by ingesting either flagellate or rounded Trypanoplasms

with their food.

It is of interest to note that, while the food of Dendro-

celum im its natural environment consists of animal matter

derived from Annelids, Molluses, and Crustacea together with

plant tissues, yet many of the infected Planarians which were

bred by Dr. Fantham and Miss Porter were kept as compulsory

vegetarians throughout their existence.

So far as is known, this is the first record of the occurrence of

a Trypanoplasma m the Platvhelminthes.

Hitherto Trypanoplasms had been described from the blood of

a number of freshwater fishes, in the alimentary tracts of certain

leeches that feed on the fish, and in the intestine of certain

marine fishes. The genus 7rypanoplasma was created in 1902 by

Laveran and Mesnil, the type-species being 7’. borreli in the blood

of the rudd. Cryptobia helicis of Leidy (Bodo helicis of Diesing),

from the reproductive organs of snails, was stated by Friedrich

(1909) to be a Trypanoplasm. Crawley (1909) claimed that the

generic name Cryptobia had priority over Zrypanoplasma. The

diagnosis of Cryptobia given by Leidy was, however, rather

vague.

There was also the interesting and closely allied genus 7rypan-

ophis, studied by Poche and Keysselitz for flagellates parasitic

in the celenteric cavity of certain Siphonophores. The genus

Trypanophis was merged in that of Zrypanoplasma by some

authorities, for example, Léger.

Dr. Fantham and Miss Porter also desired to record the

occurrence of a Trypanosome-like flagellate which was seen on

rare occasions in the guts of Dendrocalum lactewm and Palycelis

nigra.

The following papers were read :—

44*

672 DR. H. B. FANTHAM ON THE [May 3,

The Morphology and Life-History of Himeria ( Coccidium)

avium: a Sporozoén causing a fatal disease among

young Grouse. By H. B. FaNrHaM, D:Sebonde

B.A. Cantab., A.R.C.8., F.Z.8., Christ’s College, Cam-

bridge, Assistant to the Quick Professor of Biology,

Cambridge, and Protozoologist to the Grouse Disease

Inquiry.

[Received April 5, 1910.]

(Plates LV.-LVII1.* and Text-figure 65.)

CONTENTS.

Page

I. Introduction .......... itis seqgd Sse ROM

Il. The Generic Names Hemera nel Cocctauim adie awe pores 673

TS Miethods wc eniinear weereeete secs bens na daare te sania traagees aan Seen ean

IV. Morgholoer ep anceae aoa ice (El

(a) The young, growing Mearanite 5 Seishin Roles ee eRoheeaeieee Meat aa CHL

(O) Schizomontyaeesteeree co) ss. on cscs ene eee Ree steer en CeO

Macrogametocyte and. macrogamete 2 eaten smg SE OS

Microgametocy te and microgametes ..................... 680

(@) -Mertilasation@rieet rs. jee... teete ee ene ee

(e) Sporogony ...... Wadd baa Og gene Sa SCORER Ble Oh oe etal eee CEES

VY. Summary of the Life EG eiony | RRS eet a Roth asiocdicaes tsi:

WI. The effect of Ei aviwm on the Host ........0 00. ..000.eedeeece essen 686

WATS (Concluding Rem arksiepe cece ss5 0). haces cee eee OS

References towiitieratures,... +25... cb 5..c saaane eee ee CS

xplanationvohebleresmeerste cc... wie socce stk tence eee eee ee eee

: I. Lyrropuction.

The subject of this memoir is a microscopic, protozoal parasite,

which infests the lining epithelium of the alimentary canal of

grouse. It belongs to the Coccidia, a group of parasitic Protozoa

umany of which are known to occur in the digestive tracts of

both Vertebrates and Invertebrates. These minute organisms

reproduce by means of resistant spores, and belong to that class

of the Protozoa known as the Sporozoa. The Coccidia are of

economic importance, inasmuch as they destroy the mucous

membrane of the intestine of the host, thereby setting up

enteritis which is accompanied by diarrhea, and very often

has a fatal effect upon the unfortunate animal harbouring the

parasites, especially if the host be young.

Such a disease—termed coccidiosis—has long been known in

rabbits, and is often fatal, Occasionally coccidiosis occurs im

man. The life-history of a Coccidian parasite is complicated,

There are two phases in the life-cycle :—(i) a multiplicative phase

within the cells of the gut- epithelium of the host, and (ii) a re-

productive phase leading, after a sexual act, to the formation of —

* Por explanation of the Plates see p. 690.

HB.Fantham del.

E.Wilson, Cambridge.

EIMERIA (COCCIDIUM) AVIUM.

(Schizogony,)

PZ.S.1910. PL LVI.

33. 34. 35. 36.

H.B.Fantham del. E.Wilson, Cambridge.

EIMERIA (COCCIDIUM) AVIUM.

(Macrogamete formation.)

7 oe

PZ.S.1910.PLL

HB.Fantham del. \

= EIMERIA (COCCIDIUM) AVIUM.

AW (Microgametes, Sporogony.)

E.Wilson, Cambridge. %

DZS. 1910) PL EVil,

Se a a a

eee ee ee

»Cambridge.

E.Wilson

H.B.Fantham del.

EIMERIA (COCCIDIUM) AVIUM.

(Sporogony.)

1910. | MORPHOLOGY OF EIMERIA AVIUM, ies

resistant spores adapted for life outside the body of the host.

The spores so formed are the means of spreading the parasite

and lead to the infection of fresh hosts. The two phases were

formerly considered to belong to separate parasites; but the

occurrence of alternation of generations in the life-cycle of

Coccidia was first suggested by R. Pfeiffer in 1892, conjugation

was discovered in Coccidia by Schaudinn and Siedlecki (1897),

and the complete life-cycle was demonstrated with a wealth of

morphological and cytological detail in the celebrated memoir of

the late Dr. Schaudinn (1900) on Himeria (Coccidium) schubergi,

parasitic in the gut of the centipede Lithobius forficatus.

Coecidia have been recorded from most of the great groups of

the Metazoa, but very few Coccidian life-histories have been

investigated completely.

Dr. Leiper, while working on helminthiasis in grouse in May

1909, noticed the occurrence of Coccidian cysts in large numbers

in the gut of many grouse-chicks, with concomitant enteritis,

very often proving fatal. The Coccidian cysts are oval and, at

first sight, might easily be mistaken for eggs of worms. Leiper

suggested that coccidiosis was a factor in “ grouse disease,”

especially in young birds. Through the courtesy of Lord Lovat

I was enabled to spend a few days early in June 1909 on a Scotch

moor to investigate coccidiosis in grouse chicks *. Previously we

had noticed, at various times, the occurrence of Coccidian cysts

in adult grouse, but not in large numbers.

I have much pleasure in expressing my thanks to Lord Lovat,

Dr. A. E. Shipley, Dr. E. A. Wilson, Dr. Hammond Smith, and

Dr. Leiper for aiding my researches by procuring material for me,

and to Professor Nuttall, in whose laboratory much of my work

was done. I would also thank Mr. A. 8. Leslie for placing me in

communication with correspondents, and to members of the Grouse

Disease Inquiry and many correspondents I am also indebted.

In this paper I wish to record my researches on the morphology

and life-history of Himeria avium, more especially as it occurs in

the grouse. I would point out that the length of time at my

disposal for these researches has been limited, only one season

being available to me for procuring material, and I have had

several other investigations to consider during the period, so that

T was not able to give undivided attention to the elucidation of

the Protozoa of grouse. However, the complete life-cycle of

Kimeria avium, vesponsible for the dwindling of grouse broods in

spring, is here set forth for the first time, so far as I am aware.

Il. Tue Generic Names Aiversd AND Coccripivum.

Unfortunately, owing to the rule of priority, the generic name

Coccidium (Leuckart 1879) no longer holds, but is replaced by

that of Himeria (A. Schneider 1875). Tam in sympathy with

* The number of grouse chicks dying of coccidiosis on the moors is not easily

stimated, for the chicks die in the heather, and their tiny corpses are rarely found.

674 DR. H. B, FANTHAM ON THE [May 3,

Professor Minchin when he writes in a recent review :—-“‘ We

regret to see the familiar generic name Coccidium replaced by

Limeria; this is one of those many cases where, in onr opinion,

rebellion against the law of priority in nomenclature is not only

lawful but imperative” (‘ Nature, March 3, 1910). Jt would

save much confusion if the question of zoological nomenclature

were settled by an international committee, as ‘has been suggested

by many able workers.

The Coccidia of birds were first recorded in fowls by Silvestrini

and Rivolta (1873), under the name Psorospermium avium.

Subsequently Railliet and Lucet (1891) reeorded Coccidia from

fowls, naming the parasite C. tenellum. I have followed Doflein

(1909) in naming the Coecidia of birds Himeria avinm. The

Coccidian parasites were obtained from grouse chicks, and I have

succeeded in transmitting the Coccidia of grouse to fowl chicks

and to young pigeons.

Til, Meruops.

In this investigation of coccidiosis both fresh and preserved

materials were used. Samples of gut contents, taken from

different regions, were examined fresh, and often these have been

fixed wet with osmic or formalin vapour and stained by Delafield’s

hematoxylin or by Giemsa’s stain. Such smears were sometimes

useful for examining merozoites.

Odcysts, because of the chitinoid and almost impenetrable

character of their walls, had to be examined fresh.

For preserved material, the best fixatives were found to be

Schaudinn’s fluid (corrosive-acetic-aleohol), and Bouin’s fluid

(picro-formol-acetic) to which a few drops of absolute alcohol

were added. Schaudinn’s fluid tends to shrink the tissues, while

Bouin’s fluid requires much washing out. Sections, 5u to 6h

thick, were made of the duodenum and cecum of infected birds,

these parts of the digestive tract being especially examined. The

chief stains used were Delafield’s hematoxylin (either alone or

counterstained with Orange G or eosin), which was found to be

most useful, safvanin and ‘Lichtgriin, iron-hematoxylin (with or

without van Gieson’s picro- fuchsin) and paracarmine. On the

whole the hematoxylins proved of most service,

ITV. Morpnonoey,

On diluting some of the cecal contents or feces of a grouse

chick suffering from coccidiosis, and examining the preparation

mic roscopically, numerous oval cysts are seen (Pl, LVILTI. figs. 61—

66). Sometimes the cysts are also seen in the small intestine

just beyond the duodenum. ‘These cysts may have homogeneous

contents, or, when older, may show four more or less well

differentiated sporocysts within them (figs. 67-70), Each sporo-

cyst, if ingested by another grouse, can develop two active,

motile germs or sporozoites (figs, 71-76), which can penetrate

1910. ] MORPHOLOGY OF EIMERIA AVIUM. 675

the intestinal epithelium—especially of the duodenum—and so

begin a new infection. Though the odcysts and spores are the

most obvious external manifestation of coccidiosis, it is usual

and certainly more convenient to begin the life-cycle of the

Coccidium with the minute sporozoite (Pl. LY. fig. 2), the agent

whereby primary infection is brought about.

(a) The young, growing Parasite.

The sporozoites are minute, faleiform or vermicular bodies

(Pl. LV. fig. 2) capable of fairly rapid movement and possessing

great penetrative powers. They measure from 7p to 10pm in

length. The ends of the sporozoite are rather pointed, the

extremity that moves foremost being slightly more acuminate

than the posterior end. The general body cytoplasm is more or

less homogeneous, exhibiting but very fine granulations. The

nucleus has a definite rounded or oval contour, The chromatin is

evenly distributed throughout the nucleus.

When the sporozoites are liberated from the investing sporocyst

(Pl. LVILI. figs. 74-76), by the action of the pancreatic juice of the

grouse, they are capable of active movement. The usual method

of progression resembles that of the sporozoite or motile tropho-

zoite of a Gregarine. The organism moves forwards with a slow

gliding movement, the forward progression being facilitated by

the secretion of a viscid proteid substance that rapidly hardens.

On the smooth surface thus provided the coccidian sporozoite

glides forward. The track of the sporozoite, as shown by its trail,

can be stained, and the organism then shows the gelatinous or

proteid material issuing from near the posterior region of its

body (Pl. LY. fig. 2). During the gliding movement waves .

travel down the body of the sporozoite, recalling what is seen on

a larger scale in the billowy undulations of the foot of a snail.

On other occasions a more rapid movement of the sporozoite

occurs. The two ends of the organism become approximated and

then rapidly straightened, the effect being to propel the organism

forwards much more quickly than when the gliding movement

alone is used.

The sporozoite thus makes its way to an epithelial cell of the

duodenum and proceeds to penetrate the cell. As it forces its

way inwards (Pl. LV. figs. 3, 4), so the sporozoite curves on itself

(fig. 7) and becomes round and immobile (figs. 5, 6, 8). The

young, rounded parasite (figs. 8-10) is now in the trophic phase

of existence and continues to grow for some time, feeding passively

on the food-materials of the host-cell. During this period the

parasite is called a trophozoite (figs. 8-12 a).

The nucleus of the trophozoite is approximately central in

position, or sometimes to one side, and at first contains scattered

granules of chromatin. It then becomes somewhat vesicular

(figs. 8-11), and gradually the chromatin collects into a central

karyosome, lying within the nuclear sap (tig. 12 @), The position

676 DR. H. B. FANTHAM ON THE [May 3,

of the karyosome, however, is not centrally fixed; it may lie to

one side of the nucleus (figs. 8, 10).

The growth of the trophozoite naturally affects that of the host-

cell. The protoplasm of the latter becomes more and more

tenuous, great hypertrophy of the host-cell occurring. This

condition is maintained for some time and finally a limit is

reached and atrophy sets in, the nucleus of the host-cell then

appearing as a small, often crescentic mass (figs. 10, 14) to one

side of the film (fig. 12) that represents the host-cell. A clear

space often intervenes between the parasite and the enveloping

epithelial film (figs. 9, 11, 12).

The trophozoite, having attained its full size (some 10m to 12

in diameter) within the host-cell, proceeds to divide, and the

result of its division is to increase the number of parasites within

the host. This stage in the existence of Himeria aviwm is known

as the schizont (agamont), and the method of multiplication is

termed schizogony.

(b) Schizogony.

The schizont (fig. 12) is a more or less spherical parasite. At

first it is uninucleate (fig. 12a), but soon its nucleus begins to

fragment (fig. 126). The division of the nucleus of the schizont

is of the nature of multiple fragmentation rather than of a series

of binary fissions of the nucleus and karyosome (figs. 13-18).

The parasite is very small and it is not easy to follow the

cytological details, even under the best and highest powers of

magnification. Some of the portions of chromatin in multiple

fragmentation may sometimes appear connected by thin strands

(fig. 13) for a short period, but the fragments soon travel to

the periphery of the schizont. The small daughter masses of

chromatin, at first homogeneous, gradually show. differentiation,

becoming minutely vesicular with a dot of chromatin usually to

one side (figs. 14-16), but occasionally central. Thus the nuclei

of the future merozoites are forming their karyosomes early.

The daughter nuclei having migrated to the periphery of the

mother tell (fig. 16), the cytoplasm of the schizont concentrates

around them forming small ovoid masses (figs. 18, 19). The

daughter forms so produced are the merozoites (ag gametes) which

measure 64 to 10” length, They gr: dually acquire a vermiform

shape and arrange themselv es around the remains of the proto-

plasm of the mother cell like the segments of an orange or the

staves of a barrel (Pl. LV. figs. 20 a2, Pl. Lively fies. 23, 24).

Owing to this method of grouping, the merozoites are said to be

drranced “en barillet.”. The groups, when ripe, soon break up

and the individual merozoites are liberated, The movements of

the merozoites, when free, resemble those of the sporozoites.

The number of merozoites formed from a single schizont seems

to vary. Eight to fourteen seem to be common numbers, but as

many as twenty have been found.

The merozoites fin: illy are slightly curved vermicules (Pl. LVI.

figs. 23, 24), possessing a nucleus which may be approximately

1910. ] MORPHOLOGY OF EIMERIA AVIUM, 677

central (Pl. LY. figs. 20, 21) or somewhat towards one end

(Pl. LV. fig. 22, Pl. LVI. fig. 23). ‘The nucleus of the merozoite

is small, and the presence of a karyosome is often not very

evident, though there is a small granule of chromatin—repre-

senting the karyosome—usually to one side of the nucleus

(fig. 24). The ends of the merozoites are rather less pointed

than those of the sporozoites, a feature that #. aviwm has in

common with FZ. schubergi as described by Schaudinn.

When the merozoites reach a new host-cell, they enter, become

round, and proceed to grow as trophozoites in the same way as did

their parent organism, and undergo later nuclear fragmentation

ina similar manner. As the result of this, many more merozoites

are produced, and as schizogony may be continued through

several generations, the destruction of the gut epithelium is very

extensive (Pl. LV. fig. 1).

Towards the end of schizogony—especially in the csecum—

relatively smaller schizonts with larger and fewer merozoites

(fig. 25), about five in number on the average, are produced.

These larger merozoites appear to be formed near the end of

infection, in company with large numbers of gametocytes, so far

as evidence is available.

These differences in the schizonts might be taken by some

investigators to be indicative of difference of species—in other

words, that more than one species of Coccidium may occur in the

gut of grouse. I do not state that this is not so, but personally

prefer the view that the differences in the schizonts and

merozoites noted are reflexes of the condition of nutriment of

the parasite. Wenyon (1907) has some interesting observations

on the variations in the schizogony of ZH. falciformis in the

mouse, and states that the variations are due to the nourishment

available for the parasite. Again, the species found in the liver

and gut of the rabbit (Cocetdiwm oviforme and (. perforans) are

now usually united into one species, Himeria stiede.

The merozoites originally produced in the duodenum pass lower

down the gut and reach the ceca. At the ileo-cecal junction,

the epithelium is attacked again, and the merozoites rapidly grow

to schizonts which produce new generations of merozoites, so

that the ceca soon contain very large numbers of the parasites.

Probably coccidiosis set up in the duodenal wall is sufficient to

kill very young chicks, e.g. chicks 8 to 10 days old, while older

chicks dying at the age of about 4 to 6 weeks may have partially

recovered from duodenal coccidiosis, but succumb to coccidiosis

in the cecum (typhlitic coccidiosis). In cases of intense duodenal

coccidiosis, merozoites are found free in the intestinal contents

and even in freshly shed feces.

Sooner or later a limit is reached, both to the power of the

grouse chick to provide nourishment for the parasite, and to

the multiplicative capacity of the parasite itself. In other words,

the host begins to react on the parasite. Consequent on the

now unfavourable environment, the parasite proceeds to form

gametes, in order that its species may be perpetuated, The

678 DR. H. B. FANTHAM ON THE [May 3,

gametocytes or mother cells of the gametes (Pl. LVI. figs. 26-28,

PI.LVIL, fig. 37) ave modified schizonts which are of slow growth,

and therefore can accumulate more reserve food-material in the

form of granules within their cytoplasm. The processes leading

to the formation of the gametes may be termed eametogony,

which we may now consider.

Sexual differentiation is characteristic of the gametes of

Coccidia, and in the case of Himeria avium the differentiation

is apparent in the gametocyte phase. ‘lwo forms of gametocytes

can be distinguished. ‘The first group are intra-cellular parasites

containing large granules of food reserve within their cytoplasm.

These are the macrogametocy tes (Pl. LVL. fig. 26) which give rise

each to one female gamete. On the other hand, the micro-

gametocytes (PI. LVIL. fig. 37) or male progenitors contain a

little reserve food-material in the form of very minute granules,

distributed evenly throughout the body substance. Hach micro-

gametocyte gives rise to many microgametes.

The Macrogametocyte and Macrogamete.

The structure of the macrogametocyte and of the single macro-

gamete that arises from it is very difficult of interpretation

in Himeria avium, on account of the large amount of reserve food-

material contained within the cy toplasm. Further, it is very

difficult to draw the exact dividing line between the macro-

gametocyte and the female gamete, as the one gradually merges

into the other. These forms vary from 11-8u-17-5 in length, by

6u-l1u in breadth, as seen in sections. Many relatively large

granules occur in the cytoplasm of the macrogametocyte. As the

macrogametocyte grows from round to ovoid, these granules

gradually concentrate to form larger, roundish, Deon aaa

staining granules, which are Blpcrairioid (PE vale fig. 29), the

chromatoid granules of many authors. There are also other, non-

basic staining, granules known as plastinoid granules (** granules

plastiques ” of Théloban and Labbe) composed of ebecidiin

(Labbe). These ultimately are large and round (figs. 80-32)

and are refractile in fresh preparations, where they appear yellow

or greyish-green. They tend to shrink in preparations fixed with

sublimate or sublimate acetic (fig. 29). The plastinoid granules

occur between and among the chromatoid granules (figs. 30-82),

In stained preparations * the granules are best seen by staining

with iron-hematoxylin followed by van Gieson’s picro- -fuchsin,

when the chromatoid granules appear blackish, while the plas-

tinoid granules take on a uniform, yellowish hue. The plastinoid

* The reactions of the granules are discussed by Labbé (1896) and by Minchin

(1903),

1910.| MORPHOLOGY OF EIMERIA AVIUM. 679

granules stain with Lichtgriin in marked contrast with the red of

safranin taken up by the ‘chromatoid granules. With Delafield’s

hematoxylin, the chromatoid granules stain intensely (figs. 29,

31, 32), and somewhat misleading appearances result, suggesting

multiplication of the cell.

As the parasite grows, the chromatoid and plastinoid granules

travel towards the periphery (figs. 29-32). The macrogametocyte

at this time encysts within the epithelium, and the chitinoid

material of the cyst seems to be formed from the chromatoid

granules. At any rate, the inner layer of the cyst-wall seems to

take its origin therefrom (figs. 34-36). The formation of the

cyst-wall from the chromatoid granules has been noted by

Simond (1897) and Wasielewski (1904) in the case of Coccidiwm

oviforme (Himeria stiede of Stiles).

The formation of the cyst of Himeria avium takes place while

the organism is still within the epithelium (figs. 34, 35), and

therefore the parasite encysts precociously. The macrogamete at

this stage is ovoid, and the number of chromatoid granules

within it is reduced. The cyst formed is ultimately rather thick-

walled, but a small aperture or micropyle, which may be in a

slight depression, is left for the entry of the microgamete

(Pl. LVII. fig. 47).

When Schaudinn investigated the life cycle of H. schubergi, he

described a process of maturation of the macrogametocyte,

whereby the karyosome of the nucleus was expelled in fragments.

T regret that I am not at all sure as to the fate of the karyosome

of the macrogametocyte of #. avium. Several causes combine to

defeat the attempts made to elucidate this subject. In the first

place, the entire parasite (#. aviwm) is much smaller than

E. schubergi and its karyosome is not nearly so well differentiated

a structure. Further, the presence of the chromatoid granules,

which stain deeply with basic stains, much confuses the structure.

Similar causes prevail in /, oviforme (L. stiede), as I can vouch

from personal observation, and Wasielewski (1904, p. 54)

states that he is unable to follow the maturation process in

L. oviforme.

It may be that #. aviwm is like some other Coccidia (e. g.

lacazei, C. proprium, Adeleaw ovata) where the karyosome is

retained in the gamete and is left behind in the residual proto-

plasm of the odcyst. On a few occasions, I have observed a

small, chromatin-lke granule in the odcyst residuum (PI. LVIT.

fig. 57), and this body may be the karyosome of the macro-

gametocyte. #. avium is distinguished by having a very small

eystal residuum, which, together ‘with the sm allness of the karyo-

some, and indeed of fhe Snile parasite, increases the diffic salty

of investigation. However, at the time of fertilisation, the

distinctness of the karyosome of the macrogamete has disappeared

(Pl. LVI. figs. 29-36), and the nucleus of the macrogamete appears

to contain granules of chromatin which are rather indistinct.

——$— Ee

680 DR. H. B, FANTHAM ON THE [ May 3,

The Microgametocyte and Microgametes.

The microgametocyte * (PI. LVIT. fig. 37) is an ovoid cell about

13 « long and 8 « broad in the specimens that I have seen. It

possesses a central nucleus containing a karyosome. 'The process

of the formation of the microgametes of #. aviwm is as follows :—

The chromatin of the nucleus, largely concentrated in the

karyosome, breaks up (figs. 38-40) into minute granules or

chromidia which pass towards the surface of the cell, where they

appear to form a very fine chromidial network (fig. 38). The

chromidia then collect into groups or patches, arranged in the

form of minute, irregular loops with central hollows (figs. 38, 39).

These chromatic loops form a number of minute, flexible, rod-like

bodies, composed almost entirely of chromatin (figs. 41-45).

These are the young microgametes (figs. 43-46). The adult

microgametes are small parasites, their chromatic bodies measuring

3 py to. 4 in length, possessing a rod-like, somewhat curved body

composed of a core of chromatin which is surrounded by a

tenuous film of cytoplasm, prolonged outwards to form two fine

flagella (figs. 44, 46). The flagellum which I term the posterior

flagellum trails behind the organism and is practically a con-

tinuation of its body ; the other flagellum is at the opposite end

of the body and so is termed the anterior flagellum. In life, the

microgametes are capable of serpentiform movements. Owing

to the minute size of the microgamete (about 3 p to 4 pw long)

it is only with the greatest difficulty that the flagella can be

discerned,

The whole of the microgametocyte is not used in the for-

mation of the microgametes. When separation of the micro-

gametes from their mother cell occurs, a large amount of the

body-substance of the latter remains as a somewhat faint staining

residuum, in which the pale staining remains of the karyosome

(fig. 43, cf. fig. 38) can sometimes be distinguished. The residuum

takes no further part in the vital activities of the parasite.

The microgametes are set free into the lumen of the gut and

proceed to seek out the macrogametes.

(d) Fertilisation.

When the macrogamete has attained its maximum development,

it often lies in the epithelium, near the outer edge of the tissue,

or may even burst through the attenuate wall of the host-cell

and so reach the margin of the lumen of the gut. The minute

but active microgametes (Pl. LVILI. fig. 46) meanwhile have broken

away from the aesiiial protoplasm of the host- cell, and swum out

with vapid lashing movements of their flagella into the gut.

Here they are atti acted, possibly by some chemiotactic subst: ince,

Uninucleate microgametocytes seemed rare in preparations, for the cell rapidly

proceeds to form many microgametes,

1910. } MORPHOLOGY OF EIMERIA AVIUM. 681

towards the macrogametes. The microgametes swarm round the

micropyle of the macrogamete (Pl. LV II. fig. 47*) and several have

been seen trying to enter it simultaneously. The nucleus of the

macrogamete travels upwards nearer the micropyle, and before

long, one microgamete effects an entry (fig. 47 *), appears to bore

its way into the female, reach the nucleus and finally be lost to

view. The macrogamete secretes a plug of protoplasm across the

micropyle, whereby other microgametes are excluded, and the

individuals thus shut out degenerate (fig. 48). Such is the process

of conjugation as seen in the living organism, and the evidence

of stained preparations is fully confirmatory of what has just

been described.

Owing to the presence of granules in the macrogamete it is

extremely difficult to follow the subsequent stages of fertilisation.

Occasionally there are indications of a fertilisation spindle

(figs. 50, 51), but the nature of the material frequently precluded

observation of the same.

The term “fertilisation spindle” is not exactly a happy one,

for the object of that structure is the intimate intermingling of

the chromatin of the uniting gametes.

(e) Sporogony.

After the microgamete has reached the nucleus of the macro-

gamete, fusion occurs and a zygote is produced. The contents of

the zygote at first fill the oval odcyst (Pl. LVI. figs. 49-52) but

gradually they shrink away from the polest. The odcyst itself

may increase slightly in size during the concentration of its

contents, which ultimately form a globular mass, consisting of

cytoplasm rich in fatty matters, within which is a nucleus, usually

centrally placed.

The nucleus of the zygote proceeds to divide directly first into

two (figs. 53, 56) and then into four (fig. 54), the divisions follow-

ing one another very rapidly. The granular protoplasm segments

around the nuclei, and four sporoblasts (fig. 57) are produced,

each sporoblast separating from its neighbours as a small, rounded

body (figs. 55-57). Occasionally odcysts containing two ovoid

masses of protoplasm (fig. 82) are seen, but as a rule the form

containing four sporoblasts is the one found, the four sporoblasts

being formed almost concurrently. The sporoblasts become ovoid

(Pl. LVI. fig. 58; Pl. LVIII. fig. 68) and each gradually secretes

a tough, chitinoid sporocyst, usually differentiated as epispore and

endospore, and so becomes a firm, resistant spore (Pl. LVIII.

figs. 71-76). A minute amount of the cytoplasm of the zygote is

not used in spore formation, but remains within the zygote asa

small cystal residuum.

The sporocysts continue within the odcysts for some time,

* See also Pl. LXII. fig. 3 of my paper on Experimental Coccidiosis (this Journal),

+ In some cases the zygote contents of the oocyst may be slightly nearer one pole

than the other.

682 DR. H. B. FANTHAM ON THE [May 3,

during which period each sporocyst undergoes developmental

changes, leading to the production of actively motile sporo-

zoites. The contents of the spore at first are homogeneous

(Pl. LVILLI. figs. 68-70), but gradually two refractile bodies or

vacuoles appear at either end (fig. 73), and the protoplasm

gradually concentrates into two masses, just internal to each

vacuole. The nucleus is at first central, but divides into two,

and the halves migrate to the opposite poles of the sporocyst

(Pl. LVII. fig. 58). The protoplasmic masses gradually displace

the polar vesicles, so that the two vacuoles move towards the

centre and coalesce (Pl. LVIIT. fig. 73), leaving nearly all the

protoplasm of the sporocyst in two masses, one at each end

(fig. 71). Each of the protoplasmic masses gradually becomes

vermiform, extending along one edge of the spore (figs. 71, 72).

Two vermiform sporozoites are thus formed (figs. 72, 74), some-

times with their more rounded ends placed at opposite ends of

the sporoeyst (téte-béche) (figs. 72, 74, 75), sometimes with the

slightly swollen ends side by side (Pl. LVIT. fig. 59, Pl. LVIIT.

fig. 76), the sporozoites being capable of movement within the

spore just previous to their escape. There is a slight sporal

residuum.

The sporocysts when quite ripe tend to become more pointed

at one end (Pl. LV Ue te. 60, Pl. LVIT figs. 71 fasion

where a slight thickening or small Stieda’s plate (fig. 71) may

appear, which is a point of weakness, for here a rupture may

occur under the action of the digestive juices of the fresh host,

forming a sort of micropyle through which the sporozoites escape.

Partially ruptured sporocysts are sometimes found (Pl. LVII.

figs. 59, 60). In the case of grouse chicks dying from acute

coccidiosis, ripe sporocysts have been found in the cecal walls

themselves, as well as in the cecal contents, though usually

mature sporocysts are found in cecal droppings that have been

exposed.

The odcysts of Himeria aviwm show a fair amount of variation

among themselves. Usually the oocysts are oval (PI. LVIIT.

figs. 65 68, 71, 72, 77, 78), actually measured specimens varying

from 25, to 35 in length and from 14, to 20 in breadth.

Sometimes the odcysts are not oval but subspherical (fig. 70), and

these are from 18 p to 20» in diameter. Somewhat pyriform or

egg-shaped odcysts (fig. 69) are intermediate in size between the

oval and subspherical forms. Morse (1908) noted the occurrence

of both round and oval odcysts when investigating white diarrhea

of fowls, in which coccidiosis played an important part.

Among the odcysts of H. avium certain were tound with

somewhat squarish ends (fig. 78) while others had a_ slight

depression at the apex (fig. 79), but their development was

identical with that of the more common forms. Occasionally,

obcysts with two sporocysts only (figs. 81, 82) were found, but

these were abnormal forms, as was also a parasite (fig. 83) in

isd = So te

Mens

1910.] MORPHOLOGY OF BIMERIA AVIUM, 683

which the cytoplasm extended in a cone or funnel-like fashion to

the edge of the odcyst.

The size and shape of the odcysts are largely a factor of the

space in which the macrogamete develops and the amount of food

available for the parasite. When there are many Himeria present

in any particular region of the gut, the odcysts produced are

relatively small, while where abundance of space and nourishment

are available, the oocysts tend to be large.

From experiments made by feeding birds with Coceidian

odeysts, I conclude that schizogony takes from four to five d: ays.

Uninucleate odcysts mature their sporocysts in two to three day S.

The period for the total life-history of the parasite would be from

eight to ten days.

The larve and imagines of Scatophaga stercoraria, the dung-fly,

ingest the odcysts of BE. avium along with the grouse feces. The

odcysts pass through the bodies of the larvee uninjured, and are

scattered with the excrement, thus serving to disperse the spores

to some extent.

V. SumMMARY OF THE Lire-History or Hyver Avie.

The lite cycle of Himeria avium is complicated, even though

the organism completes its development within one host. The

life-history may be represented diagrammatically as in uorty

fig. 65, A-T (p. 684). Beginning asa sporozoite (Pl. LV. fig. 2 ;

text- fic. 65, A) liberated by the hebion of the pancreatic juice of

the grouse, the parasite rapidly penetrates an epithelial cell of the

duodenum (text-fig. 65, B), and, entering the cell, rounds up

(text-fig. 65, C) and becomes a passive growing trophozoite

(Pl. LV. figs. 3-7; text-fig. 65,D). After a period of rapid

growth, duri ing w hich time the tr ophozoite (figs. 8-11) practically

destroys the cell harbouring it, the parasite enters upon an

asexual, multiplicative phase termed schizogony,

The eclhabamnit is at first uninucleate (fig. 12; text-fig. 65, D)

but the nucleus soon fragments (figs. 12, 13), the daughter mcles

migrate to the periphery (figs. 14-16 ; text. fig. 65, E), cytoplasm

segregates around each (text-fig. 65, B), and the daughter forms

thus produced become meridionally arranged, like the segments of

an orange, the arrangement of the merozoites being ‘‘en barillet”

(ete ve figs. 17- 2, Pl. LVI. figs. 23-25; text- fig. 65, G).

Each merozoite is a small, “vermicular organism, having a

nucleus with a somewhat ill- ileal karyosome usually to one

side (fig. 24). The groups of merozoites break up (text-fig. 65, H),

and the free germs seek out and enter an hitherto uninfected

cell where the. parasite again assumes the trophic phase and then

undergoes division as before. Several successive generations of

schizonts and merozoites are thus produced, resulting in a great

destruction of the gut epithelium of the host. Fin: ly a limit is

684 DR. H. B. FANTHAM ON THE | May 3,

Text-fig. 65,

CVA Mee TO'G Onn)

fo}

Dracram of Life-cycle of Himeria (Coecidium) avium.

D-H represent Schizogony. I-L, Gametogony. N-T’, Sporogony.

Jpithelial host-cells diagrammatically outlined.

A. Sporozoite which penetrates epithelial | EB, Schizont with numerous daughter

cell of the duodenum of the host. nuclei peripherally arranged, differ-

B. Sporozoite curving on itself before | entiating to form merozoites (seen in

becoming rounded within the host | transverse section).

cell. F. Schizont showing further differentia-

C. Young, uninucleate trophozoite, tion of merozoites (seen in oblique

D. Fully grown trophozoite or uni-| section).

nucleate schizont. {

1910.] MORPHOLOGY OF EIMERIA AVIUM, 685

reached to the ability of the host to provide nourishment and to

the multiplicative powers of the parasite, and this results in the

onset of sexual differentiation. Gametogony may occur both in

the duodenum and cecun.

Certain schizonts become considerably modified in one of two

directions. In the first case, food-material accumulates, and a

large, uninucleate food-laden form is produced (figs. 26-30).

This is the macrogametocyte (text-fig. 65,12), destined to give

rise to a single macrogamete (text-fig. 65,J 2). In the second

instance (text-fig. 65,13), nuclear multiplication of a finely

granular parasite (PI. LVII. fig. 37) occurs, and the many

minute nuclei produced reach the periphery of the host-cell

(figs. 838-42), and ultimately the now multinucleate microgameto-

cyte (text- -fig. 65, J ¢) gives rise to a large number of small,

motile micr ogametes (figs. 43-46; text-fig. 65, Ko). The macro-

gamete early on invests itself with a cyst-wall (figs. 35, 36) in

which a thin part or micropyle is left for the entry of the micro-

gamete (figs. 47, 48;

text-fig. 65, K 9 ).

This odcyst-wall

formed while the parasite is within the epithelium.

Fertilisation (figs. 47, 48; text-fig.

65, L) oceurs—the process

Explanation of Text-fig. 65 continued (see opposite).

G. Merozoites arranged

about to issue from host cell. At

one end of the cluster residual proto-

plasm is seen.

H. Free merozoites,

karyosome.

each with a small

“en barillet,”’ |

Fertilisation. The male pronucleus

which entered through a micropyle is

Iving above the female chromatin.

Degenerating microgametes are

shown outside the oGcyst.

N. Odcyst (encysted zygote) with proto-

(?. Young macrogametocyte with plasmic contents filling it completely.

I J chromatoid granules. Nucleus with signs of fertilisation

~) g. Young microgametocyte with spindle.

}

IL fine eranulations.

( 2. Macrogamete with round nucieus

. Obcyst with contents concentrated,

forming a central, spherical mass

and chromatoid granules dis- which has a vacuole in the middle

tributed through the cyto- and the nucleus to one side. Many

plasm. such cysts seen in infected cxeal

J. g. Microgametocyte with nucleus droppings.

divided to form a large number) P. Odeyst wlth four nuclei.

of bent rod-like portions, the | Q. Odcyst with contents segmented to

future microgametes. Remains form four rounded sporoblasts (as

le of karyosome seen m centre. seen in fresh preparations).

{ 2. Macrogamete showing plastinoid | R. Odcyst with four sporoblasts which

eranules, the chromatoid gran-

ules being largely used up in

forming the wall with which the

macrogamete has invested itself,

have grown oval and are becoming

sporocysts; the small cystal residuum

seen to one side.

. Obcyst with four sporocysts in each

K.2 the remaining chromatoid gran- ) which two sporozoites are dif-

a ules being arranged between the ferentiated. The odcyst in this

plastinoid granules. condition ultimately opens, liberating

6. Microgametocyte with many bi- the sporocysts, the upper one of

flagellate microgametes about to

separate from it. Karyosome

left at the centre.

L. Fertilisation. One microgamete is

penetrating the macrogamete, while

other male cells are near the micro-

pyle but will be excluded.

PRoGc. Zoon. Soc. —1910, N

OY ELV.

. Sporocyst

which is seen about to issue from the

oécyst. Slight cystal residuum.

Sporal residuum in each sporocyst.

which has issued from

odcyst. ‘lwo sporozoites are within

it and have assumed the position

most suitable for emergence.

G86 DR. H. B, FANTHAM ON THE | May 3,

has been watched in life—and the micropyle is then closed

(figs. 49-51; text-fig. 65,M). The fertilised odcyst (text-fig. 65, N)

then passes into the lumen of the gut and is voided with the feces

of the grouse. The further development of the odcyst largely

depends on climatic conditions. Under the influence of warmth

and moisture, the contents of the oval odcyst (Pl. LVII. fig. 52,

LVIII. figs. 61, 62, 64) shrink away from the poles and

become a rounded, central mass (figs. 65, 77, 78; text-fig. 65, O).

The nucleus rapidly divides into two (PI. ‘LVI. fig. 56), then four

(fig. 54; text- an 65, P); each nucleus has protoplasm segregate

around it (figs. 55, 67, 70; text-fig. 65, Q), a wall is seer eted, ‘and

the net result is that four sporocysts (figs. 58, 71-76; text-

fig. 65, R) are produced within the odcyst. Within each

sporocyst, two sporozoites gradually differentiate (figs. 58-60,

71-76; text-fig. 65,5), and when the sporocyst (text-fig. 65, T)

is ingested by a new host, the sporozoites creep out of the

sporocyst softened by the pancreatic juice of the new host and

proceed to attack the epithelium of the gut, producing thereby

the primary infection of the bird.

The main differences between Himeria avium (Silvestrini and

Xivolta) and H. schubergi (Schaudinn) may be briefly sum-

marised :—

1. #. avium is smaller than #. schubergi.

2. The merozoites of H. aviwm are arranged ‘en barillet,’

those of £. schubergi ‘en rosace.’

3. Precocious encystment of #. aviwm oceurs before fertili-

sation. This is not the case with /. schubergi.

4. Fertilisation in #. avium is micropylar ; in L. schubergi

a cone of reception is formed by the macrogamete.

. The macrogamete of #. avium contains much more deeply

staining reserve food-material than that of #. schubergi,

thereby increasing the difficulty of minute examination

of the parasite.

. The cysts of #, avium are oval, those of #. schubergi are

round,

VI. Tue Errect oF HiMERIA AVIUM ON THE Host.

The effect of coccidiosis on the grouse may now be considered

briefly, fuller details regarding the symptoms of the disease and

its effects being given in my paper relating to experimental

coccidiosis (P.Z.S. 1910, pp. 708- 722),

External effects.—The chief external evidence of coccidiosis is

the pale colour and great fluidity of the cecal (soft) droppings of

the grouse, the pale tint being due to myriads of odcysts and the

condition being that of diarrhaa, Sportsmen and keepers have

noted diarrhaa as a symptom of “ grouse disease.” A similar

disease in fowls is known among poultry-men as ‘ white

diarrheea.” As the Coccidian parasites cause great denudation

1910. ] MORPiTOLOGY OF EIMERTA AVIUM. 687

of the intestinal epithelium, digestive derangements are brought

about, and consequent on this, malnutrition occurs and the bird

becomes very emaciated and “anemic.” Feathering also is poor

and ragged, leg weakness is fairly common, and a peculi: uw bluish

tint is ‘sometimes seen at the cere, ears and other parts.

Distribution and Kffects on Internal Organs.

Himeria avium appears to be purely a parasite of the gut of

the grouse.and does not affect such gut diverticula as the liver.

The crop and gizzard of infected birds are rarely pavasitised,

though they may contain odcysts in the condition in which they

have been ingested with food. Examination of the duodenum

shows that the sporocysts ingested with the food are attacked by

the pancreatic juice (as I have proved by pancreatic digestion

experiments, using both natural pancreatic juice and trypsin),

and the sporozoites are set free. These invade the tissue of the

duodenum, rapidly become schizonts and multiply, the result

being that the duodenum is often riddled by the parasites,

and consequently inflamed. Both the villi and the erypts of

Lieberktihn are attacked, and the parasites have also been found,

though much more rarely, in the submucosa. Great hypertrophy

followed by atrophy of the epithelial host cell occurs, and the

tissue attacked is often reduced to a finely granular, structureless

mass. Desquamation of the gut is common, and epithelium

containing various developmental stages of the parasite can be

found floating free in the gut contents.

Some of the merozoites formed in the duodenum pass down

the gut, reach the ceca and re-commence their life ie cle there.

Active schizogony and sporogony go on in the ceca *, chiefly in

the epithelium, very rarely in “the submucosa. Often’ the

ceca are as heavily parasitised as the duodenum, whole areas

being completely denuded of the epithelium, especially when

the fertilised odcysts pass outwards into the cecal contents.

The walls of the ceca are often rendered very thin and

tender by the action, direct and indirect, of the parasite. Ripe

odcysts and sporocysts occur in the lumen of the cea of dying

chicks.

Podwyssozki (1890) stated that he found Coccidian odcysts in

the vitellus of eges of fowls, especially in summer. He considered

it possible that the cysts were derived from Coccidia in the

oviduct of the mother, or perhaps from intestinal Coccidia which

had ascended by way of the cloaca. I think that cloacal con-

tamination was the more probable, for I have never seen Coccidia

in the genitalia of adult grouse examined.

A reflex of coccidiosis is seen in the blood of infected birds,

where polymorphonuclear leucocytosis is induced.

* Coccidiosis may sometimes occur along the entire length of the small intestine,

and gametes may be formed far forward, in the duodenum.

45%

688 DR. H. B, FANTHAM ON THE [May 3,

Lesions caused by coccidia in the mucous membrane may

admit bacteria to the circulation of the host. [Cobbett and

Graham-Smith (1910), pp. 24, 2 5|. Rettger (1909) believes that

‘white diarrbea” of fowls in America is due to a bacterium,

while Morse (1908) considers that it is primarily due to coccidiosis.

The discrepancy between the results of these American workers

is thus capable of explanation,

VII. Concitupinc REMARKS.

Himeria avium of grouse is not restricted to this particular

bird, for by administering feeces containing odcysts from diseased

grouse to young fowl chicks and pigeons, I have been able to

reproduce the disease exactly as it occurs in grouse (p. 674).

M‘Fadyean (1894) found coccidiosis in pheas sits while ‘‘ white

diarrhea” of fowls has been the subject. of “arirelie investigation,

particularly recently in America, where Morse (1908) and Hadley

(1909) have worked on the subject. Morse’s account of pre-

ventive measures is very good and he also notes coccidiosis in

many other birds, but the figures of the complete life-cycle of the

parasite are not yet published. Labbe (1886) also has described

coccidiosis in certain marine birds.

Though coccidiosis is peculiarly fatal to grouse chicks during

the first few weeks of their lives, adult grouse also can become

infected, for I have examined an adult bird that probably died of

coccidiosis. Old birds in the chronic condition serve as reservoirs

of odcysts and so may form sources of new infections on the

moors. All infected corpses should be burned, not buried.

1 may add that, while correcting the proofs of this memoir, L

have been able to examine pheasant chicks dying from coccidiosis,

the birds being obtained through the courtesy of Drs. Shipley

and Hanmond. ‘Smith from various parts of England during June

and July 1910.

The economic importance of coccidiosis in birds, especially in

the young, is evident, and I trust that this work, which to the

best of my belief is the first fully illustrated and detailed life-

history of an avian Coccidium, may draw more attention to

a subject both of great scientific interest and of practical

importance,

References to Literature.

Copper, L., and GranAm-Smivra, G.S. (1910).—* An Investigation

of the Pathology of ‘Grouse Disease.” Journ. Hygiene, x.

pp. 1-36, 5 pls.

Dorner, F. (1909).— Lehrbuch der Protozoenkunde. 914 pp.

Fischer, Jena. (Coccidia, pp. 616-653.)

FanrnamM, H. B. (1909).—* Coecidiosis in Grouse Chicks ”

(Prelim. Note). Abstract of Proc. Zool. Soc. Lond., No. 76;

also P. ZS. 1909, pp. 886-7.

1910. ] MORPHOLOGY OF EIMERIA AVIUM. 689

Hanp.ey, P. B. (1909).—* Studies in Avian Coccidiosis. I. White

Diarvhea of Chicks. II. Roup of Fowls.” Centralbl. f.

Bakteriol., Abt. i., Orig., 50, pp. 348-353.

Lass, A. (1896). heverecher ches zoologiques, cytologiques et

biologiques sur les Coccidies.” Arch. Zool. Expér. et Geén.,

ser, 111. t. 4, pp. 517-654, pls. 12-18.

Lasse, A. (1899). Sporozoa ” in Das Tierreich, Lief. 5, pp. 61,

68.

M‘Fapygan, J. (1894).—‘‘Some Observations regarding the

Coccidium oviforme” and ‘ Intestinal Psorospermosis i in the

Pheasant.” Journ. Comp. Path. & Therapeutics, vii.

pp. 131-137.

Merzner, R. (1903).—“ Untersuchungen an Coccidium cuniculi.”

Gch f. Protistenkunde, 1. pp. re 72, pl. i.

Mincuin, E. A. (1903). —‘“ Sporozoa” in Lankester’s “ Treatise on

Zoology,” pt. i, fase. 2. (Coccidridea, pp. 204-238.)

Morse, G. B. (1908). —“* White Diarrhea of Chicks, with Notes

on Coceidiosis in Birds.” Circular 128, U.S. Dept. Agr ‘cult,

Bureau of Animal Industry.

Neumann, L. G., and MacQurxn, J. (1905).—A Treatise on

Parasites aud Parasitic Diseases of Domesticated Animals.

2nd edition, 697 pp. London, Balliére. (See pp. 404-417.)

Perronciro, KE. (1876).—‘‘ Nuove caso di psorospermosi intes-

tinale in una gallina.” Annali Accad. Agricoltura, Tormo,

MOK. 1.

Prererer, R. (1892).—‘‘ Die Coccidienkrankheit der Kaninchen.”

Berlin.

Raiiurer, A. (1895).—Traité de Zoologie médicale et agricole.

2nd Edition. 1303 pp. Paris, Asselin et Houzeau, (See

pp. 142-4.)

Raruirer, A., et Luce, A. (1891).—“ Note sur quelques espéces

de Coccidies encore peu étudiées.” Bull. Soc, Zool. France,

Xv1. pp. 246-250.

Rerreer, L. F. (1909).—“ Further Studies on Fatal Septiceemia

in young Chickens, or ‘White Diarrhea.” Jown. Med.

Research, xxi. pp. 115-123.

Scwaupinn, F. (1900).—* Untersuchungen tiber den Generations-

wechsel bei Coccidien.” Zool. Jahrbiicher, Abt. f. Anat.,

xiil. 2, pp. 197-292, 4 pls.

Stivestrint, A., e Rrvoira, 8. (1873).—“ Psorospermi epizootica

nei gallinacei.” Giorn. d. anat. fisiol. e patol. de animal,

Pisa, v. pp. 42-53.

Srmonp, P. L. (1897).—* L’évolution des Sporozoaires du genre

Coccidium.” Ann. Inst. Pasteur, xi. pp. 545-581, 2 pls.

W asieLEwskt, Th. von (1904).—‘* Studien und Mikrophotograimme

zur Kenntnis der Pathogenen Protozoen.” Heft 1., Coccidia.

118 pp., 7 pls. Leipzig, J. A. Barth.

690 DR. H. B. FANTHAM ON THE [May 3,

EXPLANATION OF PLATES LV.-LVIII.

All figures outlined with Abbé-Zeiss camera lucida, from stained preparations

(sections of infected gut) except where otherwise stated. Zeiss 2 mm. apochromatic

immersion objective and compensating ocular 8 usually employed. The mag-

nification is about 1400 diameters, except where indicated otherwise. Many

parasites are represented lying in portions of the degenerating epitheliuin

snrrounding them.

Puate LY.

Fig.1. Portion of gut (cecum) of infected grouse to show mucous membrane

riddled with parasites. Many stages in life-history of Mimeria avium

shown therein.

. Sporozoite from duodenum, showing its trail stained.

. Sporozoite penetrating epithelium.

. Young parasite just entered epithelium, by fine channel which is still open.

The entermg parasite is by the side of a slightly older (rounded)

parasite or young trophozoite.

, 6. Parasites beginning to become round.

. Sporozoite, curved on itself, preparatory to becoming round. Duodenum.

. Round, young parasite, with relatively yesicular nucleus containing

karyosome.

. Young trophozoite, as yet oval in shape, lying in relatively large space in

the gut-wall.

10,11. Round trophozoites, with large nuclei each containing a karyosome.

Large space surrounding parasite shown in fig. 11.

2. (a) Uninuele: ate, full-grown trophozoite, becoming a schizont.

(6) Schizont in w hich nucleus is dividing. Duodenum.

13. Schizont in process of nuclear fragmentation.

14, 15, 16, 17. Schizonts containing many nuclei which go to the periphery ot

the cell. Very small karyosome differentiating to one side in each

daughter nucleus.

18, 19. Developing merozoites.

20-22. Merozoites arranging themselves “en barillet,” with residual protoplasm

in middle or to one end.

Several smaller and younger parasites shown by the side of the “ bavillet

in fig. 22.

= Co bo

ve) io eS he)

Prater LVI.

Fig. 23, 24. Groups of merozoites, beginning to separate.

Fig. 23 magnification 1400. Fig. 24 magnification 1600.

25. Small schizont, containing tive young merozoites—seen in cross-section.

Cecum.

26. Macrogametocyte, with granules of reserve food-material.

27, 28. Young macrogametocytes.

29. Longitudinal section of macrogamete, stained with Delafield’s haematoxylin,

showing deeply staining chromatoid granules, a few of which are

surrounded by clearer areas of plastin.

30. Transverse section of older macrogamete, stained with iron hematoxylin

and picro-fuchsin, with cyst just formed. Plastinoid granules well

defined; chromatoid granules (largely used in cyst formation) small

and in between the plastinoid granules.

32. Macrogametes, in section, as stained by Delatield’s haematoxylin.

Chromatoid and plastinoid granules alternating. Plastinoid granules

do not stain with hematoxylin. X 1600.

Young macrogametocytes seen in the vicinity.

33, 34. Macrogametes with chromatoid and plastinoid granules, after careful

staining with iron haematoxylin and picro-fuchsin. In fig. 34

chromatoid granules are seen passing to the periphery, to form the

inner wall of the mature macrogamete.

35, 36. Mature macrogametes. Remains of chromatoid granules seen attached

to inner wall of cyst.

1910.] MORPHOLOGY OF EIMERIA AVIUM., 691

Fig.

Prate LVII.

57. Microgametocyte or male mother cell.

38-40. Nuclear fragmentation of microgametocyte. In fig. 38 remains of

parent karyosome seen. In fig. 40 young iicrogametes arranged

along meridians.

41, 42. Young microgametes at periphery of mother cell.

43-45. Separation of the clusters of microgametes.

46. Free microgametes.

47. Fertilisation. One microgamete has gained access to the female pronucleus.

The micropyle has closed, and the microgametes left outside will

degenerate.

48. Fertilised odcyst. Zygote nucleus beginning to form a fertilisation spindle.

Remains of microgametes on outside of cyst.

49. Zygote in odcyst just after fertilisation.

50, 51. Show “ fertilisation spindle ’”’—not very clearly differentiated in Himeria

avium.

52, 53. Longitudinal and transverse sections of odcysts, containing rather a

large amount of reserve material—as judged by staining (hematoxylin).

In fig. 53 the zygote nucleus shows signs of dividing.

54. Odcyst with four nuclei, one ready for each sporoblast. Safranin, Lichtgriin.

55. Small oécyst with four round sporoblasts. Paracarmine.

56. Odcyst, showing segmentation into four sporoblasts. From lumen of cecum

of dying grouse chick.

57. Odcyst, with four ovoid sporocysts, and cystal residuum, containing minute

dot of chromatin (? remains of karyosome of macrogamete). Cut

obliquely.

58. Odcysts with four sporocysts showing commencement of differentiation

(polar separation of nuclei) into sporozoites. Paracarmine.

59, 60. Sporocysts, each with two sporozoites, Dehiscence represented in fig. 60.

Puate LVIII.

Mostly from fresh preparations. > 1000.

61. Zygote nearly filling odcyst. Culture of cwcal contents of grouse containing

such odcysts made and examined for tour days subsequently.

62, 65, 64, 66. Development at beginning of second day.

65. Cyst uninucleate, such as is frequently seen in fresh feces.

67. Odcyst with four sporoblasts, as seen at end of second day, and on third day.

68. Oodcyst with four ovoid sporoblasts, as seen in cultures on the third day.

68-70. illustrate three common forms of odcyst of HB. aviwn—oval, egg-shaped,

and rounded. There is variation among the oécysts.

71. Odcyst with tour sporocysts, in each of which sporozoites are differentiating.

72. Odcyst with four sporocysts, nearly ripe, each containing two sporozoites.

From partially dried cecal droppings, taken from grouse moor.

73. Ripe odcyst debiscing—one sporocyst outside, with sporozoite nuclei at

poles, and central vacuole. Sporocyst inside odcyst shows polar

vacuoles.

74. Free sporocyst, from feces. Sporocyst contains two sporozoites, arranged

“téte béche,” and sporal residuum.

75, 76. Sporocysts, each with two sporozoites.

77. Odcyst, commonly occurring in fresh droppings.

78. Odcyst with somewhat square ends.

79. Odcyst with depression (thin place in cyst-wall) at one end.

80. Zygote in odcyst—zygote nucleus dividing into two. Stained preparation.

Sl. Oocyst containing two precociously separated sporoblasts. There should

be, normally, four sporoblasts. Probably the odcyst represented will

degenerate. Stained preparation.

82. Odeyst with two sporobiasts. Very few of these seen. Possibly abnormal.

The four sporoblasts should be formed almost concurrently in the

normal cyst.

$3. Odcyst showing protoplasmic contents in form of cone-like projection

attached at one point to the cyst. Development retarded, after

keeping three weeks in water.

692 DR. H. B. FANTHAM ON THE [ May 3,

Olservations on the Parasitic Protozoa of the Red

Grouse (Lagopus scoticus), with a Note on the Grouse

Fly. By H. B. Fanrnam, D.Sc. Lond., B.A. Cantab.,

A.R.CS., F.Z.8., Christ’s College, Cambridge, Assistant

to. the Quick Professor of Biology, Cambridge, and

Protozoologist to the Grouse Disease Inquiry.

[Received April 5, 1910.)

(Plates LIX.-LXL.*)

CONTENTS.

Page

Introduction ........ Saeco sar aabecchccnontee aay Mates

A. Parasites found in Ny Blood of Grouse sos cag.auccea ciety femae acieets Ree Samm

(@)) Sporozca——Hieemosporidia, i; 3:..c: osu cesccesbecve-sscoeae eaves (OUE

Ly WeucoctOzoon WOuatie Wasi oi isons 1 oel tens oe deeds ve eee oes

(a) Introductory... Se OHS

(b ) The Relation of the Parasite to its Host Cell... 693

(@) Schizogony RRA See tae a SE

(ec) Concluding Rem: WES. -sacaey ee ga Oa

2. ELEM PrOLEUS MASON ..... 02. cen nese aed pereseteueeney NOU

(5) Spirochwetacea (Proflagellata)) ... ...c.-20.------ cence ssases O97,

Spirocheta lagopodis, sp. 0. se eClanis age ee ee en OT

B. Parasites found in the Alimentary Tract of Grouse .................. 699

(a) ilar) ataeesensrerennrss « wice alsin Gee’ REDE REE REC OSE ee en CO ,

Trichomonas eberthi canes cedudec one semen

(2) Spirochsetaceal(bromacellata) 5....-... ...:29-e:--eecu enna AE

Spurr @La LOUGEIS Nee ah guxes wiew s1d + colon cee eee eee ee ee TOL

Ameba (Entamoeba) lagopodis, sp. a. .....-.....0........ 702

(2) Sporozoa——Gremaninidan eee.c-c..+,---2.--<-ne peru enelen ieee eennna Os

Monocystis sp. ......... esta TALS:

Appendix—Note on the Grouse Fly, Or ‘hee yea na ta gape aR tates 7O4L

References to Waiterditue’ Roemer eee rccs ss csie ces cansat coe scence eaaucean Seem OEs

Explanation ot cPlates’ c.2)jsesemasernindots .lestcasas1 css 'was cbakceeecmai acne eNO

INTRODUCTION.

In the following paper are recorded observations made on the

various parasitic Protozoa found in Grouse examined in connection

with the Grouse Disease Inquiry. The observations were, by

force of circumstances, limited to one season, and so the accounts

of some of the parasites are consequently incomplete. The

parasitic Protozoon of greatest economic importance, and to

which most attention had to be devoted, is Mimeria (Coceidium)

avium, Which is the pathogenic agent of a fatal disease in grouse-

chicks, particularly prevalent in the spring and early summer.

The morphology and_ life- history of Himeria avinm, and the

results of experimental studies on avian coccidiosis are set

* For explanation of the Plates see p. 707.

LEUCOCYTOZOON LOVATI

PZ:S: 1910; Pia

EB Wilson, Cam bridge

E.Wilson, Cambridge

LEUCOCYTOZOON LOVATI. 29-40 SPIROCHAETA LAGOPODIS.

"41-42 HAEMOPROTEUS MANSONI.

aee

{

ia4

: a ;

at ol oe i” y Tela ee te OP @

H.B.Fantham del.

E.Wilson, Cambridge.

43-50 TRICHOMONAS EBERTHI. 51-57 SPIROCHAETA LOVATI.

\ 58-65 AMOEBA LAGOPODIS.

% 4)

baal.

rt?

1910. ] PARASITIC PROTOZOA OF RED GROUSE. 693

forth by me in two other memois*. The remaining Protozoa

found in grouse may be conveniently divided into (2) parasites

of the blood of the bird, (5) parasites of the alimentary tract of

the grouse. So far I have found seven Protozoa parasitic in

grouse, exclusive of the Coccidiwm already mentioned.

(A) PARASITES FOUND IN THE BLOOD OF GROUSE.

Sporozoa-—Hemosporidia.

1. LeveocyrTozoon LOVATI Seligmann & Sambon, 1907.

te}

(Plates LIX., LX. figs. 1-28.)

(a) Lntroductory.

. This parasite was discovered in 1907 by Seligmann and Sambon

in blood-films of grouse. It was found in one bird not suffering

from ‘* grouse-disease.’ Sambon (1908-9) refers again to the

parasite, and states that he found it in five more grouse. Sambon

gives five drawings of the parasite.

Personally, I found this parasite in 1909 in three grouse, all of

which were in splendid condition, though in one case the spleen

was found to be slightly enlarged. The parasites were In no case

numerous. and in one bird only the younger stages of the para-

site were seen. However, in the two remaining birds I had the

good fortune to find the parasites alive in peripheral blood and in

heart-biood taken from birds freshly killed. Observations were

made on the living parasites, both unstained and after colouring

them intra vitam with methylene blue (Pl. LX. figs. 17-22).

The parasites were also seen while making blood- counts of the

avian hosts, when the Lewcocytozodn stained intra vitam with the

methyl-violet of Toison’s fluid.

~ (b) The Relation of the Parasite to its Host Cell.

The nature of the host-cell is controversial, some authorities

considering that it is an erythroblast, others that it 1s a leucocyte,

while the views of some of the observers have changed during the

course of their investigations. The host-cell clearly does not

contain hemoglobin, and no melanin pigment is excr eted by the

parasite. The host-cell is at first round or ovoid, containing a

rather broad and well-marked nucleus, and so the host-cell must

be either a mononuclear leucocyte or an erythroblast. Possibly

both these types of cells may be infected. The nature of the

host-cell is inherently difficult to determine in view of the fact

that all the blood-cells of birds are nucleated. Judging by the

size of the nucleus of the host-cell (Pl. LIX. figs. 1-3), I ine line to

the view that it is a small mononuclear leucocyte or potentially

such. Other workers have acknowledged the resemblance of

the host-cell to a mononuclear leucocyte. However, the matter

can only be detinitely settled by researches on the origin and

i VAS UG KOy pp. 672-691 and 708-722.

694 DR. H. B. FANTUAM ON THE | May 3,

development of the various blood-cells of birds—on which our

present knowledge is meagre and contradictory —running parallel

with the researches on infected birds, studying the young stages

of the Leucocytozoon.

The host-cell of Z. lovati soon becomes drawn out at the ends,

that is, the cell becomes spindle-shaped. This is probably due to

a deformity of the cells brought about by the movements of the

parasite within. Wenyon (1908) considers that some spindle-

shaped cells occur nor mally in the blood of birds and reptiles. I

have very rarely seen spindle-shaped cells in the blood of grouse,

though I have found spindle-shaped cells to be more numerous

in the blood of fowl-chick ks (ef. Fantham, P. Z. 8. 1910, Pl. LXIIT.

fig. 18) dying from coccidiosis. However, I would not suggest,

on present evidence, that coccidiosis is e either directly or indirectly

responsible for the appearance of spindle-cells in the blood.

The presence of Leucocytozoa does not always cause the body

of the host-cell to beeome spindle-shaped. Mathis and Léger

(1909) found that the host-cell of Leucocytozodn eaulleryi of the

‘Tonkin fowl did not have its ends drawn out. They also con-

sidered that the host-cell was probably a mononuclear leucocyte.

Personally, I have seen the spindle or horn-like end of the host-

cell of Z. lovati elongate during the movements of the parasite

within it. Further, both ends usually are elongate (Pl. LX.

figs. 19-22), in other cases one end only becomes deformed

(fig. 18), while in a very few cases the cell contour has been

largely unaffected by the parasite (fig. 17). When both ends

of ‘the host-cell are affected, it not infrequently happens that

the elongation of one end is greater than that of the other

(Pl. Toe figs. 4-6). The shape of the host-cell of different

Leucocytozoa may be merely a reflex of the motility of the

parasite within. It is not wise, therefore, to lay too great stress

at present on the spindle-shape of the host-cells in diagnosing the

Leucocytozoa of birds.

Young Forms.

The parasite in its younger stages is vermiform and like a

hemogregarine (Pl. LIX. figs. 1-3; Pl. LX. figs. 17-22) and is

more refractile in life than the host-cell containing it. Young

forms are by no means common,a feature Z. lovati shares with

L. caulleryi where they have rarely been seen, ad nucleus of

the host-cell is displ aced to one side (Pl. LIX. figs. 1-5) and the

parasite moves very slowly within the cell, slight wave-like move-

ments or pulsations passing from the middle of the parasite

towards either end. The parasite grows in length and breadth

meanwhile (figs. 4-8), ahd the ends of the host-cell become

gradually more and more drawn out (figs. 4-9, 18-22). As

mentioned above, sometimes one end of the containing cell is

more drawn out than the other (figs. 4-6), Free vermicules of

L. lovati have very vavely been seen in my preparations,

1910. ] PARASITIC PROTOZOA OF RED GROUSE. 695

Gametocytes.

The growing parasites gradually become round, and two forms

can be - distinguished to some extent in life (contrast figs. 20

and 22) and more easily after careful staining (figs. 4 4 and 8

from figs. 5-7). In one the protoplasm is relatively hyaline

(figs. 4, 8, 9, 11, 13, 14); im the other form it is granular

more deeply staining and may be slightly alveolar (figs. 5, 7,

10, 12, 15, 16). By analogy with the malarial parasite, these

rounded Leucocytozoa are considered to be gametocytes, the

slightly smaller, hyaline parasites (figs. 11, 13, 14) being charac-

terised as males. while the slightly larger, browder and more

granular Leucocytozoa (figs. 10, 12 iB; 16); are designated females.

The females or mactogametocyte es, which measure 14 p-20 p by

10 »-16 pr, appear to be somewhat more numerous than the m: ales

or microgametocytes, which are about 13 p-17 wp by 6 w-12 wu.

The nuclei of the gametocytes are not well marked, even after

long staining. The nucleus of the microgametocytes (figs. 4, 9, 13)

1s often somewhat larger than that of the macrogametocy te

(ies. 6.10, 12),- 1s) al- defined (figs. 11, 13, 14) in contour, and

frequently has small granules of chromatin scattered within it

(figs. 11,14). The macrogametocyte has a better defined nucleus

(figs. 6, 10, 12), in which a larger eran of chromatin— a karyo-

some—is sometimes present (figs. 5 5, 6, 10). Small chromatoid

granules are often seen at either pole of ie gametocytes (fig. 4, 11,

13), sometimes more numerous at the ends of the micr ogametocy tes

ee 9,11, 13) than at the ends of the macrogametocy tes (figs. 10,

1D) These chromatoid granules are, I think, more probably

derivatives of the nucleus ‘of the host- cell in process of absorption

by the parasite than actual chromidia within the parasite, for

sometimes the chromatoid granules are clearly outside the parasite

(figs. 6, 7

The details of the maturation of the female gametocyte to

form the macrogamete are little known—probably the macro-

gametocyte becomes the macrogamete with little or no cytological

change. Danilewsky (1890), and Sakharoff (1893-5) described

the formation of flagella-like microgametes from the hyaline

microgametocyte in fresh preparations of the blood of certain owls,

crows, and rooks. The microgametes (males) of LZ. ziemanni

were considered by Schaudinn (1904) to be formed normally in

the mid-gut of a mosquito which sucks the blood of the avian host

(an owl). The more recent observers of avian Leucocytozoa have

not usually seen gamete formation in the fresh state, so far as

can be gathered from their published accounts.

Mathis and Léger (1909) have recently made the very in-

teresting statement that there is a periodicity in the occurrence

of the gametocytes of Z. caulleyri in the blood of the fowls they

investigated at Tonkin. The events of the life-cycle of the parasites

in the period intervening between their appearances in the blood

ave unknown, There seem to me to be several alternatives possible :

696 DR. H. B. FANTHAM ON THE [May 3.

either the parasite undergoes a multiplicative stage in some internal

organ (though it is stated that the internal or eans of the fowls

were examined), or a latent phase without meneaee in numbers of

the parasite occurs, or else a re-infection of the host takes place.

Thave found schizogony of Z. lovati (see next section) in the spleen

of the host, but so far neither time nor material have been

available i my observing any possible periodicity in the gameto-

cytes of L. lovati, though | such may occur,

Gametocytes were found to be rare in the peripheral blood,

more numerous in heart-blood, and were also seen in smears of

the liver and spleen of infected grouse.

(d) Schizogony.

Schizogony in the avian Leucocytozoa has not been recorded

before. "However: in my investigations of LZ. lovati, I have

succeeded in finding schizonts of this Leucocytozodn in the spleen

of two infected birds (Pl. LX. figs. 23-28). I saw living

Leucoeytozoa in the heart-blood of these birds, and immediately

made fresh smears of the internal organs. In the spleen

were found rounded or ovoid bodies—the schizonts—with thin

walls, coloured ved with Giemsa’s stain. These schizonts

contained many merozoites (figs. 25, 26).. The capsules of the

schizonts are probably formed, at least in part, by the remains of

the host-cell (fig. 26). The protoplasm of the schizont appeared to

be slightly granular (fig. 26). Some of the schizonts already con-

tained a few nuclei in process of multiplication by amitotic binary

fissions (figs. 28, 24), while other schizonts contained small ver-

micular merozoites (figs. 25, 26) which ultimately escape from the

mother cell (fig. 26) and may be found free in spleen smears

(figs. 27, 28), when their mode of origin and general appearance

are clearly grasped. A small amount of residual protoplasm is

left behind in the thin membranous remains of the schizont

which has just shed its merozoites. ‘The schizonts are from 11 4

to 14 by 8 to 11 ue, and the merozoites are about 7 in length.

The birds whose spleens contained schizonts of ZL. lovati were

not infected with any other Heemoprotozoodn.

Preparations made from the bone-marrow of infected birds,

in which schizogony might also take place, did not exhibit

developmental forms.

(e) Concluding Remarks.

The method whereby Lewcocytozodn lovati is transferred from

grouse to grouse has not yet been shown with certainty. The

vector or second host of ZL. lovati will probably be found in some

blood-sucking insect, which ingests the parasites from one

grouse and transfers them to the next bird from which it obtains

blood; or perhaps there is a cycle of development of the parasite

inside the Arthropod vector.

The grouse-tly (Ornithomyia lagopodis) suggests itself as oa

1910.] PARASITIC PROTOZOA OF RED GROUSE. 697

likely carrier or second host of Levcocytozoon lovati. During my

investigations I have dissected several hundreds of grouse- “Ries.

In the ‘cut contents of a very few of these flies I found unicellular

motile mermicnies which may have been stages in the life-cycle

of LZ. lovati or perhaps of Hemoproteus mansoni. In the grouse-

flies dissected I never found cysts, such as occur in the stomachs

of mosquitoes which have fed eaeabusly on blood infected with

malarial parasites.

As before-mentioned (p. 693) Leucocytozodn lovati, as it has

been found to occur in grouse on the moors, does not appear to be

wu serious agent of disease.

H_#:MOPROTEUS MANSONI Sambon, 1908.

(Plate LX. figs. 41-42.)

This parasite of the red blood corpuscles of grouse was recorded

by Sambon in 1908, though I en not aware that he has yet

ublished a descr iption or fier es * of the parasite.

In blood-films sent to me from Scotland and taken from two

grouse, I found, after staining, a few parasites inside the red

blood corpuscles. Two examples of the organism are drawn in

Pl. LX. figs. 41, 42. They appear to be young forms of the

parasite, one of which showed clearly g granules of melanin pigment.

1 regret that, owing to lack of material, I cannot give a fuller

deser iption of the organism.

It is likely that this heemosporidian is spread from grouse to

grouse by the grouse-fly, Ornithomyia lagopodis, in the gut of

which grouse-blood is found. In the gut-contents of the fly

unicellular vermicules, apparently protozoal organisms, were some-

times seen. F urther, Sambon states that ‘‘in the stomach of one

[grouse-| fly, | discovered a few hemozoin-bearing protozoa, which

were no doubt the odkinetes of the Hoemopr oteus mansoni pre-

viously found in the blood of a grouse from Scotland ” (Sambon

(1909), p. 37).

Spirochetacea (Proflagellata).

SPIROCH_ETA LAGOPODIS, Sp. 0.

(Plate LX. figs. 29-40.)

When examining the blood of grouse for parasites, I have

ecasionally found a few small Spirocheetes exhibiting considerable

morphological ‘variation (Pl. LX. figs. 29-40). As a Spirochete

has hot been previously recorded from the blood of grouse, I have

named it Sprrocheta lagopodis. The Spirochetes were present in

* Since writing the above I find that figures of Hemoproteus mansoni—which

Tam informed are from originals supplied by Dr. Sambon—have been published

recently in Castellani and Chalmers’ * Manual of Tropical Medicine,” fig. 55,

p- 235.

698 DR. I. B. FANTHAM ON THE [May 3,

blood from the heart, liver and spleen of three birds, but were

somewhat rare in each case. On one occasion I saw the para-

sites in life; the remaining preparations were fixed and coloured

with Giemsa’s stain.

Dr. Sambon (1909) has noted the presence of an unnamed

Spirocheete “in smears from the pulp of young feathers, and not

in the general circulation ” of grouse. It is possible that Spiro-

cheta lagopodis and the parasite of feather-pulp may be identical,

but as details of the latter parasite are lacking, | cannot establish

the identity or otherwise. Dr. Sambon also thinks that lice might

act as “alternative hosts” for the Spirochetes. J will return to

the subject of transmission later.

S. lagopodis is trom 10% to 18, long and .is relatively broad

for this class of organism. In life it moves actively in the blood

of the host, spirally boring its way between the corpuscles.

S. lagopodis also often entwines one part of its body about the

other (fig. 31). The organism usually has tapering ends (figs. 29—

AQ).

Though the Spirochzte is small, stained specimens show a fair

amount of structural detail, A narrow undulating membrane

(figs. 29-36) is present, and passes spirally round the body. The

thickened border of the membrane stains bright red with Giemsa’s

stain (figs. 29-34), and even when the membrane is closely con-

tracted round the body of the Spirochzete, the border can be

distinguished as a bright line (figs. 380, 32).

The chromatin of the Spirochete is distributed in the form of

bars throughout the body (figs. 29-35), and at times these

chromatin bars become joined and form a helicoid core (fig. 36)

traversing the entire length of the body. This latter condition

is comparatively rare. A basal granule is present at either end,

and near these granules the membrane i is attached. Longitudinal

division is usually initiated by fission of the basal granules.

Multiplication of S. dagopodis takes place by both longitudinal

and transverse division. In longitudinal division rather thick

Spirochetes divide (fig. 40), the split extending gradually from

one end and the daughter forms diverging more and more until

final separation occurs. ‘Transverse division takes place in

relatively long organisms (fig. 35) and two daughter forms are

produced. The parent organism elongates somewhat during the

division. Very minute forms (fig. 39) about 4 long are the

product of repeated divisions in both directions. Small forms

have been seen in the spleen (figs. 38, 39). By reference to

growth following division, all the variations in size and form

exhibited by S. lagopodis may be explained.

x carding the mode of transmission of S. lagopodis, it seems to

me that the nymphs of Jaoedes ricinus, that sometimes infest

grouse, may be the carriers of the Spirochetes, for in the gut of

nymphs of Jxodes ricinus taken from grouse, I have found Spiro-

chietes which may be S.lagopodis. During my investigations, I have

dissected many Mallophaga, both Nims cameratus and Goniodes

1910.] PARASITIC PROTOZOA OF RUD GROUSE. 699

tetraonis, but have not found Spirochetes, though the sperm of these

lice are deceptively like Spirochetes at first sight. Dr. Sambon

has suggested that lice act as transmitters of the Spivochete that,

he found in feather-pulp. This is possible, and asa matter of fact,

evidence is accumulating and tending to show that the carrying

of disease in a particular host is not restricted to any one species

of Arthropod, but that several organisms differing in genera as

well as in species, may become active agents in the dissemination

of protozoal parasites. '

Among Spirochetes known from the blood of various birds may

be mentioned Spirocheta gallinarwm of fowls, S. anserina of

geese, and a Spirochete cultivated by Tépfer from the blood of

the owl.

S. gallinarum is the pathogenic agent of a fatal disease in fowls,

and is spread from bird to bird by the tick, Argas persicus.

S. lagopodis is much too rare, I think, to be a serious cause of

disease in grouse, although leucocytosis was found in a grouse

infected with S. lagopodis, in the blood of which bird many

vacuolated mononuclear leucocytes occurred. :

(B) PARASITES FOUND IN THE ALIMENTARY TRACT

OF GROUSE.

(a) Flagellata.

TRICHOMONAS EBERTHI.

(Plate LXI. figs. 43-50.)

While examining, on the moors, the cecal contents of grouse—

more especially for Spirochetes—I have sometimes observed a

Flagellate Protozoén moving therein.

The parasite was seen to possess an undulating membrane, the

vibrations of which often regulated the parasite’s active move-

ments ; at other times flagella were seen anteriorly whose active

forward movements dragged the body of the organism onwards.

The structure of the parasite is complex. The organism closely

resembles, both in size and shape, the parasite described recently

(1909) by Martin and Robertson from the cea of the fowl. The

flagellate of the fowl, first seen by Eberth in 1862, was named

Trypanosoma eberthi by Kent (1881), Spirocheta eberthi by Liihe

(1906), while Stein, Leuckart, and Laveran and Mesnil more

correctly considered it to be a Zrichomonas. The nomenclature

of the parasite is discussed at length by Martin and Robertson

(1909). These authors describe both Z'richomonas and Monocer-

comonas forms as well as a Trypanosoma, and state that they

“have cbserved some cases of which the most natural explanation

would be to regard the A [Zrypanosoma], B [ Trichomonas], and

C | Monocercomonas| conditions as stages in one life cycle”; other-

wise the parasites are distinct and there is a mixed infection.

It seems to me that the flagellate which I have seen in the

709 DR. H. B, FANTHAM ON THE | May 3,

cecum of grouse is best named 7'richomonas eberthi, though

Eberth’s original figures published in 1862 do not especially

suggest a typical 7’richomonas. However, the somewhat peculiar

shapes figured by Eberth are perhaps due to the fact that the

organism is very easily deformed.

‘In grouse both typical T'richomonas (P]. LX. figs. 48-46) and

Monocercomonas (fig. 48) forms were seen in life as well as in

fixed and stained preparations (fixed wet with osmic acid and

stained with Delafield’s heematoxylin or Giemsa’s stain). The

bodies of the parasites vary from 8 » to 13 ye by 6 j to 9 p.

Trichomonas eberthi possesses three flagella disposed around

slight cytostomie depression (figs. 43-45), and arising inside

the body close to a chromatin granule, the blepharopk ust or

kinetonucleus. ‘The flagella are often much entangled. There is

an undulating membrane with a flagellar border and short free

Hagellum ; the membrane possesses also a chromatic base line on

the body as seen in some specimens (fig. 45). There is also a

skeletal organella arising at or close to the blepharoplast and

running backwards to the posterior (non-flagellar) end of the

body, where it may project a little (figs. 43-45). This stiff

skeletal rod—which stains blue with Giemsa’s stain—is the

axostyle. A vow of granules or blocks may be seen alongside the

axostyle and base line of the membrane—better marked in some

specimens than others (figs. 43-45). The nucleus is situated

rather nearer the flagellar end of the body close beneath the ble-

pharoplast, and is “fairly large and more or less spherical.

Chromatin granules are seen within it during the resting state,

and a nuclear membrane is present (figs. 43-46).

Trichomonas eberthi divides longitudinally. Although I have

‘arvetully searched for dividing forms, I have only seen them on two

occasions, and then in stained preparations. One of the dividing

forms is shown in fig. 49, but I had not sufficient material to

follow the details of division. The process of division is well

described by Dobell (1909) for Trichomonas batrachorum, and by

Wenyon (1907) for 7". intestinalis in the mouse.

incystment * of 7. eberthi occurs (fig. 50). The organism

evadually becomes oval in outline, and its body substance con-

centrates after gradual absorption of the flagella, undulating

membrane and axostyle (fig. 50). A thin gelatinous cyst is seen

in the specimen drawn (fig. 50). Round, contracte d, uneneysted

forms were also seen ; these ave probably resting forms.

Aflagellate, irregular, non-motile forms with nucleus and axostyle

are occasionally seen, Aieceboid forms are, I believe, the result

cf degeneration.

Sometimes, in stained smears of the cecal contents of grouse

containing V'richomonas, I have seen curious ovoid or figure-of-8

bolies which suggest much deformed specimens of 7'r ‘ichomonas in

which the axostyle acts as a rod around which the flagellar border

* 8. Parisi has recently noted encystment in Trichomonads (Arch. f. Protisten-

kunde, xix. p. 232).

1910.] PARASILIG PROTOZOA OF RED GROUSE, TOL

and strands of the membrane are disposed. Some of these bodies

are not unlike Eberth’s original figures (1862).

Monocercomonas is an ovoid organism possessing four flagella at

the broad anterior end of the body (fig. 48) together with a nucleus

and blepharoplast. Sometimes one of the flagella trails backward

(fig. 47). On one occasion a Jonocercomonas form with a suggestion

of a rudimentary membrane, axostyle and granules was seen

(tig. 47). If such a parasite be normal and not deformed, then it

would be an intermediate form between 7’ ichomonis anc

Monocercomonas (cf, Martin and Robertson’s fig. 8).

(b) Spirochetacea (Proflagellata).

SPIROCH ETA LOVATI, Sp. 0.

(Plate LXI. figs. 51-57.)

When fresh cecal contents of certain grouse of various ages

were examined microscopically, small active organisms have been

seen, moving among the food débris and ‘oun, found in the gut.

These organisms are Spirochetes, and their presence in the cecal

contents of grouse has also been noticed by Dr. Cobbett in adult

grouse, and by Dr. Leiper in a young grouse chick. I have

sometimes observed the Spirochztes in the hinder part of the

intestine as well as in the ceea.

Having had the opportunity of observing this Spirochete,

which I have named Spirocheta lovati, | append a few details

regarding its life-history and structure. I may say that the

parasite does not appear to have a very deleterious effect on its

host.

The organism, as observed in life, occurs in cecal contents,

where it is mingled with semi-fluid food materials, and is often in

company with many minute, rod-like bacteria, both free and

in colonies, and also small hyphal threads, evidently of some

fungus. Because of its associations, the Spirochete is by no

means easy of observation, and much dilution of the cecal contents

to facilitate observation only hastens the death of the Spirochete.

Further, in fixed and stained preparations the Spirochete itself

takes up the stain somewhat faintly.

The movements of the Spirochete are active and resemble those

of the Spirochzetes of Lamellibranchs (oysters, fresh-water mussels,

Tapes) which | have already described (1907, 1908, 1909). ‘* The

motion appears to be resolvable into at least two components :

(3) An undulatory flexion of the body, mainly for progression,

and (ii) a spiral or corkscrew movement of the body as a whole,

due to the winding of the membrane.” 8. Jovati has a small, but

distinct undulating membrane best seen in stained preparations

(Pl. LXI. figs. 51-57). The ends of the organism are pointed, the

length of the body being from 16°5 p to 32°5 u.

In some very narrow forms the relatively long body may be

thrown into many waves (fig. 53), but usually four to seven waves

Proc. Zoon. Soc.—1910, No. XLVI. 46

702 DR. H. B. FANTHAM ON 'THE [ May 3,

oceur along the body (figs. 61, 52,55). The membrane with its

chromatic ‘border is often closely contracted against the body

(figs. 538, 55). The chromatin of the Spirocheete is usually

present in the form of a number of bars (figs. 51-57) which are

probably disposed on a helix as is typical for Spirocheetes.

After examining many specimens of the parasite it was seen that

S. lovati exhibits morphological variation. Thus, the ends of

the body usually taper (figs. 58-57), but occasionally are some-

what roundel (figs. 51, 52). Also, the Spirochietes are of different

lengths and breadths, the results of growth and division.

Multiplication of S. lovati by both longitudinal and transverse

division has been seen in life. Division resembles that of S. re-

currentis and S. duttoni, where a periodicity in the direction of

division occurs (Fantham and Porter, 1909). Broad Spirochzetes

(fig. 51) have been seen to divide longitudinally (fig. 56). Longer

Spirocheetes (fig. 54) often divide transversely (fig. 57), elongating

somewhat as they do so.

The modes of multiplication and the processes of growth are of

the utmost importance in elucidating the morphological variation

of Spirochzetes.

AmacbA (ENTAMG:BA) LAGOPODIS, Sp. 1.

(Plate LXI. figs. 58-65.)

Living Amcebe were seen by Dr. Shipley (1908-9) in fresh

feeces of grouse. I have, on a few occasions, observed Ameebze

in the recently voided feces and in the rectum and intestine of

grouse. ‘The living organism was examined both with and without

staining intra vitam, anid a few stained preparations were made

with Delafield’s hematoxylin after fixation with corrosive-acetic-

alcohol or osmic vapour, but the whole of the material was very

scanty.

In structure the Amoeba exhibits ectoplasm and endoplasm

fairly well differentiated (Pl. LXI. figs. 58-63). The Amoeba

moved slowly by sending out pseudopodia i in the manner usual to

these organisms, the pseudopodia being first chiefly composed of

ec toplasm, Very few pseudopodia were sent out at any one time.

There is a nearly central nucleus, often possessing a karyosome.

Food vacuoles containing partially digested food particles and

bacteria were seen (fig. 61), The gr: emilen endoplasm may contain

a vacuole (figs. 58. 59).

Ameeboid organisms, from the irregularity of their shape, are

not easy to measure precisely. More or less rounded forms may

measure 60 « in diameter (fig. 61) witha eee leus 10 « in diameter.

Other specimens of the parasite (figs. 59, 60, 63) measure from

20 pw to 40 p across, with a nucleus of wh ek 8 jc in diameter.

In preparations stained with Delafield’s haematoxylin the

nucleus was seen to be spherical (figs. 58-61) with a nuclear

membrane on which the chromatin could sometimes be clearly

1910.} PARASITIC PROTOZOA OF RED GRCUSE, 703

seen in the form of granules (fig. 58). The whole nucleus, however,

is somewhat poor in chromatin, though a karyosome may be

present (figs. 58, 60, 63) as before-mentioned.:

Division by binary fission was seen once in life, the process taking

about thirty minutes for completion. The Ameceba when first

noticed was already elongate and the dividing nucleus appeared

as a refractile streak across the body of the parasite. A similar

dividing form (fig. 62) was once seen in a fixed and stained

preparation. ‘The nucleus was drawn out in the form of a long

spindle with a central fibre, which opens out at its extremities

into cone-like expansions, perhaps composed of the remains

of the fibres of the nuclear spindle. The actual ends of the

dividing nucleus were composed of chromatin masses or plates

(fig. 62). . The division of the cytoplasm was delayed somewhat

after that of the nucleus.

Eneystment was rarely seen in life, and the full details could

not be followed. It was noticed sometimes that, followme

division, the daughter individuals proceeded to encyst after a

short interval and so formed small cysts. In fixed preparations

somewhat small uninucleate cysts, about 12 « to 14 » in diameter,

were seen (fig. 64). Larger cysts containing four nuclei were

also found (fig. 65), but the eysts were not numerous. The

larger cysts measured about 20 « in diameter.

Tam sorry that opportunity did not arise for me to make an

extended study of this parasite.

Although Hntameba histelytica (Schaudinn) is the pathogenic

agent of a form of dysentery in man, it is net very likely that

Ameba (Hntameba) lagopodis is the cause of such a serious disease

in grouse, but 4. lagopodis is more like Hntameba coli, which is

usually considered to be a well-nigh harmless parasite in the

human intestine.

Ameba meleagridis (Theobald Smith, 1895), which has been

associated with the disease known as ‘“ blackhead ” in turkeys, is

now generally considered to be a stage in the life-history of a

Coccidium.

(d) Sporozoa—Gregarinida.

MONOCYSTIS sp.

Spores of a Gregarine, almost certainly a species of JMWonocystis,

were occasionally found in the contents of the gut of a few grouse

from the Lowlands of Scotland and the North of England. The

spores present much the same features as those commonly

occurring in the Earthworm, and show the same pseudo-navicellar

appearance. The spores seem to pass through the alimentary

canal of the grouse intact. The grouse acquire the spores

accidentally by way of their food (though earthworms are not

common on many grouse inoors), and the spores have no ill effect

on the birds. Probably the internal heat of the bird’s body aids

in the development of the sporozoites within the sporocysts,

46*

704 DR. H. B, FANTHAM ON THE | May 3,

though the spores are not acted on externally by the digestive

juices of the host.

Uninjured spores of Gregarines have been observed by

L. Pfeiffer in the alimentary tract and faeces of various birds.

APPENDIX.

Nore ON THE GROUSE-FLY, OR V/THOMYIA LAGOPODIS.

The relation of the grouse-fly, Ornithomyia lagopodis, to the

erouse has been to some extent a matter of conjecture. Though

the grouse-flies often ave found clinging to or concealed among

the feathers of the birds, the biology of the fly was uncertain, and

it was not known definitely whether the flies obtained blood from

the grouse or not. Having dissected several hundreds of grouse-

flies in an endeavour to find developmental stages of some of the

Protozoa infecting grouse, and possibly parasites natural to the

fly itself, I append the following notes on the insect, that may be

of interest.

Regarding the food of the grouse-fly, the insect sucks the

blood of the grouse. Blood in all stages of digestion has been

obtained from the gut of the fly. Nucleated red cells of grouse

ton}

blood, which show no sien of digestion, are found in the red

to)

fluid in the crops of recently fed flies taken from grouse, while

on a few occasions leucocytes also have been observed. The

stomach and particularly the intestine contain blood that is semi-

digested and consists almost entively of cell nuclei.

The blood, especially in the fore- part of the gut, has a peculiar

tint, and farcher, retains its fluidity for a very long time. Exam-

ination of the salivary glands of the fly has ‘shown that an

anti-coagulin is secreted by them. I have tested the power of

this anti-coagulin in delaying the clotting of human blood, using

eapillary tubes of blood and amalsion and blowi ing out the éontente

at intervals, controls being carefully kept. Blood mixed with

emulsion of the salivary glands required nine minutes for co-

agulation, while the counrele. using salt solution, had clotted in

six minutes. The intestine of the fly also has anti-coagulin in it.

Another member of the /ippoboscide, the sheep ked, Melophagus

ovinus, also secretes an anti-coagulin. The blood ingested by both

flies is prevented from coagulating, and thereby is kept in a

more suitable condition for dige sehioi and absorption by the fly.

The peculiar colour of the blood in the fore part ‘of the ‘fly’ s

alimentary canal is apparently to be associated with some action

of the anti-coagulin, for an emulsion of the salivary glands of the

fly added to human blood caused the latter to assume the same

peculiar hue as was noticed in the blood from the crop of the

grouse-fly,

A fungus also infests the Malpighian tubes of the fly. The

hyphal threats of the fungus rapidly develop rounded masses of

spores within sporangia, The sporangia completely fill the

1910.] PARASITIC PROTOZOA OF RED GROUSE. 705

Malpighian tubes, the spores finally bursting out as rounded

bodies into the lumen of the gut, and thence pass outside the host.

Melophagus ovinus contains a similar fungus [see Porter (1910)].

Tn connection with the fungus, which is common in the grouse-

flies, I may say that I have never found flagellates present where

the fungus existed, and a similar condition obtains in the sheep-

ked, Melophagus ovinus.

The grouse-fly, examined as a possible vector of the protozoal

parasites of the grouse, yielded rather poor results. However, |

have found protozoal vermicules, some without melanin, others

doubtfully with a little, in the gut of the fly. These might be

stages (cokinetes ?) in the life-history of Leuwcocytozoin lovati ox

Hemoproteus mansoni, or of both. There is also the possibility of

Ornithomyia lagopodis being the transmitter of Spirochetalagopodis,

though J] am inclined to suspect Jxodes ricinus here, but the

whole question of the transmission of the protozoal parasites of

grouse is one demanding much more material and very careful

investigation.

i also kept a number of the pup of the grouse-fly for further

investigation, but unfortunately they have ‘not hatched out yet.

Dz: Shipley (1909, pp. 321-3) has already published an account

of the grouse-fly in which the possible time needed for hatching

is discussed, together with much information regarding the

adult fly.

References to Literature.

Baurour, A. (1908).—Third Report Wellcome Research Labs.,

Khartoum, ‘“Spirochzetosis of Sudanese Fowls,” p. 38

BLANCHARD, R. (1906). —* Spirilles, Spirochetes et auties micro-

poe a corps spiralés.” Semaine méd., Paris, pp. 1-5.

Danttewsky, V. (1890).—‘ Développement des parasites malari-

ques dans les leucocytes des oiseaux (Leucocytozoaires).” Ann.

Inst. Pasteur, iv. pp. 427-431.

Dopett, C. C. (1909).— Researches on the Intestinal Protozoa of

Frogs and Toads.” Quart. Journ. Microsc. Sci., iii. pp. 201-

277, 4 pls.

Dortety, F. (1907).—“ Studien zur Naturgeschichte der Protozoen.

V. Amdbenstudien (4. vespertilio).” Arch. f. Protistenkunde,

Suppl.-Bd. 1. pp. 250-293, 3 pls.

Dutton, J. E., Topp, J. L.,and Tosry, EK. N. (1907).—“ Concerning

certain Parasitic Protozoa observed in Africa.” Ann. Trop.

Med. & Parasitology (Liverpool), i, pp. 287-366, 13. pls.

(Avian Leucocytozoa, pp. 288-297.)

Eperrtn, J. (1862).—‘* Ueber ein neue Infusorium im Darm ver-

schiedener Vogel.” Zeitschr. f. Wissenschaft. Zool. xi.

p. 98.

FantHam, H. B. (1908).—‘ Spirocheta (Trypanosoma) balbianii

(Certes) and Spirechwta anodonte (Keysselitz) ; their move-

ment, structure and affinities.” Quart. Jour, Microse. Sci.,

ln. pp. 1-73, 3 pls.

706 DR. H. B. FANTHAM ON TITE [May 3,

Fanruam, H. B. (1909).—‘ The Spirochetes found in the erys-

talline style of Zapes aureus: a Study in Morphological

Variation.” Parasitology, 1. pp. 892-408, 1 pl.

Fantuam, H. B., and Porrmr, A. (1909).—‘t The Modes of Division

of Spirocheta recurrentis and S duttoni as observed in the

living organisms.” Proc. Roy. Soc. B. 1xxxi. pp. 500-505.

Fantuam, H. B. (1910).—“ The Morphology and Lite- History of

Kimeria (Coccidium) avium: a Sporozoon causing a fatal

ee among young Grouse.” Proc. Zool. Soc. Lond. 1910,

. 672-691, 4 pls.

___lft9109. a4 On the Occurrence of Schizogony in an avian

Leucocytozoon, LZ. lovati, parasitic in the Red Grouse (Lagopus

scoticus).” Ann. ‘Trop. Med. & Parasitology (Liverpool), iy

pp. 255-260, 1 pl.

Kerysseiitz, G., and Mayer, M. (1909).—‘* Ueber ein Leucocyto- .

zoon bei einem ostafrikanischen Perlhuhn (Guttera pucherant

Hartl.).” Arch. f. Protistenkunde, xvi. pp. 237-244, 1 pl.

Leyaprat, C. (1904).-—** Contribution a I’ sage de la Spirillose des

Poules.” Ann. Inst. Pasteur, xviii. pp. 129-149, 1 pl.

Martin, C. H., and Ropertson, M. (1909),.—“ Preliminary Note

on Trypanosoma eberthi (Kent) (=Spirocheta eberthi Lithe)

and some other Parasitic Forms from the Intestine of the

Fowl.” Proc. Roy. Soc. B. )xxxi. pp. 385-391, 1 pl.

Maruis, C., et Liaur, M. (1909).—‘* Leucocytozoon de la Perdrix

du Tonkin.” Ann. Inst. Pasteur, xxi. pp. 740-743, 1 pl.

- (1909).—* Leucocytozoon de la Poule.” C. R. Soe,

Biol. 67, pp. 470-472.

(1909).—“* Recherches sur le Leucocytozoon de la

Poule. Periodicité des formes sexuées dans le sang.” C.R.

Soc. Biol. 67, pp. 688-690.

(1910).—“ Sur un nouveau Leucocytozoon de la

Poule.” C. R. Soe. Biol. 68, pp. 22-24.

(1910),-—“* Leucoeytozoon une Tourterelle (Z'urtur

humilis) et Vune Sarcelle (Querquedula crecea) du Tonkin.”

C. R. Soc. Biol. 68, pp. 118-120.

Mezincescu, D, (1909).—* Leucocytozoon alemannt et Trypano-

somes chez ipevviar (falco nisus).”? C. R. Soe. Biol. 66,

pp. 328-329.

Mincuin, E. A. (1903).—‘ eherozon ” in Lankester’s ‘* Treatise

on Zoology , pt. I, fase. 2.

NEAVE, 8 . (1906). —*A new Hemameeba in the blood of Guinea

Fowl (Vumida ptilorhyncha).” 2nd Rept. Wellcome Research

Labs., Khartoum, pp. 200-202.

Novy, F. G., MacNzrat, W. J., and Torrry, H. N. (1907).—

“The Trypanosomes of Mosquitoes and other Insects.”

Journ. Infect. Diseases, iv. pp. 223-276, 7 pls. (Quotes Tépfer,

259.)

Por’ ne sn, ANNIE (1909).—‘ The Leucocytozoa: Protozoal Par: a

sites of the colourless ae Epnae ‘les of the blood of Vertebrates.

Science Progress, No. 14, pp. 248-266.

1919. ] PARASITIC PROLOZOA OF RED GROUSE. TOT

Porrrer, ANNIE (1910),—* The Structure and Life-History of

Crithidia melophagia (Flu), an Endoparasite of the Sheep-ked,

Melophagus ovinus.” Quart. Journ. Microsc. Sci. lv. pp. 189-

224, 2 pls.

Prowazek, S. von (1906). — “ Morphologische und entwickl-

ungeschichtliche Untersuchungen tiber Hiihnerspirocheten.”

Arb. a. d. Kaiser]. Gesundheitsamte, xxiii, pp. 554-565, 2 pls.

SakHArorF, N. (1893).—‘* Recherches sur les Hématozoaires des

Oiseaux.” Ann. Inst. Pasteur, vil. pp. 801-811.

SamBon, L. W.(1908—9).—“ Remarks on the Avian Hemoprotozoa

of the genus Lewcocytozoon Danilewsky.” Journ. 'Trop. Med.

and Hyg. xi. pp. 245-8, 325-8; xi. pp. 37-8.

Scnaupinn, F. (1903).— Untersuchungen iiber die Fortpflanzung

einiger Rhizopoden (Vorl. Mitteil.)’’ Arb. a. d. Kaiserl.

Gesundheitsamte, xix. p. 547.

Setiemann, C. G., and Samson, L. W. (1907).—“ Preliminary

Note on a Leucocytozoén found in the blood of Red Grouse

(Lagopus scoticus).” Lancet, p. 829 [Sept. 21, 1907].

Surpetey, A. KE. (1909).—“ The Kctoparasites of the Red Grouse

(Lagopus scoticus).” Proc. Zool. Soc. Lond. 1909, pp. 309-

334, pls. 35-47.

Wenyon, C. M. (1907).—* Observations on the Protozoa in the

intestine of Mice.” Arch. f. Protistenkunde, Suppl.-Bd. i.

pp. 169-201, 3 pls.

-—— (1908).—Third Report Wellcome Research Labs., Khartoum.

“ Avian Leucocytozoa,” pp. 157-165, 1 pl.

EXPLANATION OF PLATES LIX.-LXI.

All figures were outlined with the Abbé-Zeiss camera lucida, generally using Zeiss

2 mm. apochromatic immersion objective and compensating oculars 4, 8, 12 (Zeiss).

For a few figures Zeiss DD objective and Huygenian ocular 4 were used.

Prats LIX. figs. 1-16. Leucoeytzoén lovati. (X 1400.)

From preparations, chiefly heart blood, coloured with Giemsa’s stain.

Magnification : 1400 diameters.

Figs. 1-3. Young intracorpuscular parasites.

4-13. Older parasites in which the host cell is drawn out into horns or pro-

cesses at either end, and its nucleus displaced.

4,9, 11, 18. Male gametocytes with hyaline protoplasm.

5, 6,7, 10, 12. Female gametocytes with granular protoplasm. Chromatoid

granules occur around the poles of many of the gametocytes.

14-16. Gametocytes in which much of the host cell is absorbed or discarded,

with only shght remains of host-cell nucleus. Figs. 15, 16 from

specimens in the spleen. Fig. 14 from the liver. Such parasites,

however, are not contined to the internal organs.

Pratr LX,

Figs. 17-28. Leucocytozoon lovati.

17-22. L. lovati as seen in life, stained with methylene blue. Young parasites,

showing formation of tailed processes of host cell. > 1000.

23-28. Schizogony of DL. lovati,as seen in spleen smears from grouse. 1400.

Free merozoites represented in figs. 27, 28.

29-40, Spirocheta lagopodis, trom blood of grouse. 2500.

41,42. Hemoproteus mansoni. Young parasites in erythrocytes of grouse.

< 1800.

708 DR. H. B, FANTHAM ON AVIAN COCCIDIOSIS. | May 3,

Poate LXI.

Figs. 43-50. Trichomonas eberthi and Monocercomonas. X 1409.

43-46. Trichomonas forms from cecum of grouse.

47-48. Monocercomonas forms. Fig. 47 intermediate form.

49. Dividing Trichomonas.

50. Eneysting Trichomonas.

41-57. Spirocheta lovati from the ceca of grouse. XX 1700.

58-65. Amceha (Entameba) lagopodis from intestine and feces of grouse.

58, 59, 61, 62.63. Amcebe. XX 500.

60. Ameeba which has assumed a pyriform shape. 740.

64-65. Encysted ameebe. XX 1000.

3. Experimental Studies on Avian Coceidiosis, especially in

relation to young Grouse, Fowls, and Pigeons. By

H. B. Fanraam, D.Se. Lond., B.A. Cantab., A.R.C.S.,

I'.Z.S., Christ’s College, Cambridge, Assistant to the

(Juick Professor of Biology in the University of Cam-

bridge ; Protozoologist to the Grouse Disease Inquiry.

(Plate LXII.* and text-figure 66.)

[Received April 5, 1910. ]

CONTENTS.

Page

T. Unitroduretronng ob: escesecisny clean ee oe oe Sie eR ete SSS het a

IJ. Experimental Methods saded dca ETRE SEES RNS Ae eee a 0 at ec

Ill. Symptoms of Coccidiosis .............. widen oben ce ee

IV. The condition of the Internal Oreane of Tmtected Bide Ji shee 713

V. Relative resistance of different breeds of Fowls and Pigeons to Coceidiosis ... 716

VI. The Dissemination of Coccidiosis over Tracts of Country ..................6. 717

VII. Duration of Vitality of Coccidian Oécysts REC etre tosmacrite Clue:

VIII. Effects of certain reagents on Coccidian OGcysts ............c:0eeeeeeceseeeees VAD

IX. Concluding Remarks 721

References to oyterature 2... cc.csseaini sinesarg decease ve doaeiteciosstcds teen eeeae neem

Explanation of Plate 722

I. INrropuctrion.

Protozoal parasites are highly specialised animaleules which

live in intimate relation with the hosts they infect. While many

of the Protozoa are capable of living in one host only, there are

others which are under suspicion of infecting several species of

higher organisms as host, and of not being so limited to one host

as was formerly believed. One test for the specifieness of any

protozoal parasite is that it fails to develop in any animal other

than its special, natural host, and merely perishes when inoculated

or otherwise introduced into any other animal. In order to test

the specificness of the coccidian parasite of the grouse, experiments

were made, with the result that the Coccidim pathogenic to young

Por exphination of the Plate see p. 722.

Np,

, abe.

E.Wilson, Cambridge.

H.B.Fantham del.

AVGAN (COC ClDLGsis:

‘hi a ; - o .

1910.] DR. H. B. FANTHAM ON AVIAN GOCCIDIOSIS. 709

erouse and responsible for the dwindling of the broods, particularly

in the spring and early summer, was found to be equally injurious

to healthy young fowls and young pigeons. Healthy grouse chicks

also were experimentally treated by administering food mixed

with small quantities of infected feces from other grouse suffering

from coccidiosis, and fatal results ensued.

The results seb forth in this memoir are those obtained from

one season’s work only, to which my investigations were limited.

II. ExprrtwenraLn Mernops.

The birds used in experimental coccidiosis were grouse chicks,

fowl chicks and young pigeons, all of which were initially healthy.

Coccidian oécysts (Pl. LXIT. figs 4, 5) contaimed in cecal droppings

from infected grouse were nillewed to develop spores to some

extent by being “thinly spread in Petii dishes and covered in order

to prevent complete drying. Under these conditions the spores

(figs. 6, 7) developed in two to three days at summer tempera-

ture. The feces so prepared were then administered to the

healthy young birds.

(A) Grouse Chicks.—As these birds are somewhat wild, even

though bred in captivity, a rapid feeding method was adopted.

Some of the partly died feecal matter was taken up on a spatula

and inserted directly into the mouth of the bird. Rather large

quantities were given as first doses, but if infection was not fairly

vapidly brought about the dose was repeated. The cecal droppings

of treated chicks were collected each morning, for these soft

droppings are more abundant during the night than they are

during the day, and in the early morning there is a better chance

of obtaining excrement free from soil. Examination of feces

collected in the evening was sometimes made.

The grouse chicks were kept on the experimental ground at

Frimley, and the feeding experiments were conducted by Dr. E. A.

Wilson, to whom I tender my best thanks. Samples of the feces

of the birds were sent to me daily.

(B) Fowl Chicks and Young Pigeons. —A method of feeding

similar to that used for grouse chicks was employed with fowl

chicks and young pigeons (squabs). These experiments were con-

ducted by me at Cambridge. The chicks used were from incubated

eggs, the eggs having been care efully cleansed antiseptically before

incubation. Ceecal droppings containing odcysts were administered

directly to the birds. Very small daily doses were used for several

days and then none for a couple of days, and so on. This method

of administering odcysts was quite effective and had the advantage

of reproducing somewhat the condition of wild grouse on the

moors, where intermittent ingestion of odcysts with food or drink

occurs.

One experiment was performed in which a single dose only had

a fatal effect on a fowl] chick.

Droppings containing odeysts in different stages of development

710 DR. H, B, FANTHAM ON AVIAN COCCIDIOSIS, [May 3,

(figs. 4-7) were also used. When the odcysts contained developed

sporocysts the onset of coccidiosis was more rapid.

Again, | have found coccidian odcysts in the water of tarns at

en erouse chicks drink and also in dew collected from the

heather on the moors (fig. 8). Grouse chicks, then, can acquire

coccidian odcysts by way of their drink. To show this method of

infection experimentally, a healthy fowl chick was supplied with

water containing coccidian odcysts. This bird also became

infected with coccidiosis.

Control birds were most carefully kept. These were supplied

with food and drink exactly as were the treated birds, and kept

under the same conditions. Examination of their feces was made

twice daily, and careful search was made for odcysts as the possi-

bility of natural coccidiosis of both fowls and pigeons * was well

recognised and most carefully guarded against. Control birds

invariably were healthy and made more rapid progress in growth

than did the subjects of the experiment

In every instance I most carefully compared the results obtained

experimentally with the cases of natural coccidiosis in grouse

chicks that I investigated at Killin (by the courtesy of Lord

Lovat) and in other cases obtained from Perthshire, Dumfries-

shire and Yorkshire.

Some authors have given the name Coccidiwm cuniculi t to the

parasite of birds, ther éby identifying the coccidium of hirds with

that of rabbits. Having had the opportunity of obtaining Hea

material from rabbits dying rapidly of acute coccidiosis, I fed :

healthy young pigeon’ directly with odcysts of C. cuniculi, At ae

odcysts were voided by the pigeon, then none were found in the

feeces and no symptom associated with coccidiosis appeared at any

time. The first odcysts voided were merely those supplied to the

bird which had passed unchanged through its alimentary tract.

Though this pigeon received several doses of the odcysts of Himeria

(Coccidiwm) cuniculi, it never developed coccidiosis, and the post

mortem examination made immediately after killing the bird

showed a perfectly normal condition of every organ. I consider

that these experiments show conclusively th ne EB. avium and

i}. cuniculi ave distinct species of Himeria and are not identical.

There are also morphological differences between the two, chiefly

of size (2. avium is the smaller).

III. Symproms or Coccrprosts.

The symptoms of grouse suffering from natural coccidiosis and

those of grouse, fowl chicks, and pigeons, im which the disease has

been artificially induced, are identical. The symptoms that have

been noted in the case of the birds examined may now be stated.

* Through the courtesy ofa friend I was able to examine Coccidian odcysts from

a pigeon suffering from natural coccidiosis, On P). LXII. fig. 13 is drawn a cyst of

this Coceidium (CL pfeifferi of Labbé). It is spherical, about 17 4 in diameter.

+ The correct name of this parasite ts Bimeria stiede Lindemann.

> aly

1910. ] DR. I. B. FANTILAM ON AVIAN COCCIDIOSIS. (Alii

Chicks after ingesting coccidian odcysts become far less active

in their movements asa rule, The first noticeable feature is the

drooping of their wings and a habit of constantly looking down-

wards. The birds stand about more than normal birds and their

calls are more plaintive.

While fowl chicks and pigeons appear to mope, their appetite is

increased, and chicks experimentally infected with coccidiosis eat

far more greedily than the control birds. They also drink con-

siderably more. In spite of the increase in the amount of food

consumed, the birds rapidly get thinner, the muscles of the breast

and legs s showing this toa marked degree, Throughout the pro-

gress of the disease the growth of the affected birds is much

retarded.

It was necessary to feed infected young pigeons by hand, for

even when they reached practically adult life they failed to feed

themselves, merely thrusting their heads into the food offered

them, without attempting to swallow any of it.

Several breeds of fowl chicks were used in experimental

coccidiosis and each lost weight steadily till death occurred. The

loss of weight of one pure bred Leghorn chick was very noticeable.

It was first fed with coccidian odcysts when six weeks old. It and

its control bird were then of equal weights (74 0z.).. Two mouths

later the infected chick died, its weight at death being 5 oz.,

while the weight of its control on the same day was 1 lb. 6 oz.

Sample weights of other experimental birds are given below :—

Weight of | Weight of | Difference in

Brrp. Infected Bird.) Control Bird. Weights. Remarks.

Grouse chick A ......... 4 oz. | 52 04. | ldo. (| Dosed once when

| J} 11 days old:

Grouse chick B ......... 42 ,, Viatss 98 ., Killed in extremis

| L whenaged 6 weeks. |

Guowsercuick: | Cameesssce 61 ,, OS ease Died, aged 10 weeks.

| Minorca hen...............) 3 lb. 2 oz. d1b.20z. | 21b.0o0z. | Died, aged 6

| months.

Plymouth Rock cock . .| 4 Ib. 8 oz. 61b.50z. 11b.130z. Killed, at acute

stage of disease.

Cross-bred Leghorn...... 4]b.100z.| 51b.40z. ' 10 oz. Chronic.

Pigeon fa dSb. Geeetcisay 92 oz. 12 oz. | 25 OZ. Died, aged 11 weeks.

Nore.—tThe three fowl chicks were first treated with Coccidian odcysts when aged

3 weeks. Another fowl chick fed with Coccidian oécysts when aged 1 day, died

when 9 days old. The pigeon squab was dosed first when aged 9 days.

Another instance of loss of weight resultant on coccidiosis was

seen in the case of a pure bred female Leghorn chick which was

attacked when seven weeks old by coccidiosis after drinking water

fouled with coccidian odcysts. This bird became a chronic, and

712 DR. H. B. FANTHAM ON AVIAN COCCIDIOSIS. [ May 3,

when adult weighed 4 lb. 3 0z., while its sister bird that acted as

control weighed 5 lb. 4 oz.

Besides loss of weight the infected birds become anzmic.

The cere, comb, and wattles become much paler and the blood-

vessels beneath the wing also look pale. ‘The head appendages

gradually become more and more pale as the disease progresses

and finally acquire a peculiar bluish tinge. This tint also is

shown by the eyelids and ears and the lees are affected though to

a less extent. "The feathers on the bead tend to fall off so “that

the forepart of the head and the region round the bill become

almost bald, and the bird presents a very peculiar appearance,

owing to the bluish colovation. Jeg weakness was present im

several cases.

The plumage of the infected birds is affected im regions other

than the head, and the quills are less rigid than in normal birds.

The feathering of the legs is ragged and the sheen on the neck

and tail-coverts is not so well developed, while the replacement

of nestling down by ordinary feathers is much retarded in diseased

birds.

During the progress of coccidiosis the birds sometimes develop

much mucus and a very offensive “breath,” a smell of sulphuretted

hydrogen being noticea ble. Both sticky mucus and smell disappear

as a rule in a few days, but may recur.

While birds suffering from coccidiosis feed eveedily, internal

digestive troubles occur and the feeces voided by the bigs are very

fluid, the condition being that of diarrhoea. The cxcal droppings

are the more noticeable, and they contain many resistant cysts

(odcysts) (figs. 4-8) of C. avium. Both sportsmen and keepers

have noted that diarrhea is a marked symptom of ‘“ grouse

disease.” Examination of soft droppings daily shows the relative

numbers of odcysts present and may be a rough gauge of the

intensity of the infection*. In ool health the cecal droppings

are of firm consistency and olive-green to brown in colour. When

coccidiosis is slight, the feces become softer and brownish yellow.

in acute cases the excrement is almost fluid and the birds void

sulphur-yellow feces with a heavy, foetid odour, Fatty matters

may be present in the cecal contents.

A day or so before the death of the infected bird the slimy,

mucilaginous discharge recommences, ooze coming from the beak,

nares, ears and eyes. Examination of this liquid by the micro-

scope shows the presence of oval coccidian odcysts (cf. Pl. LAE:

figs. 4-8), all of which show the characteristic cyst-wall within

which is a single uninucleate mass of protoplasm. The mucus

also contains some shed epithelial cells in which occasionally

macro- and micro-gametes may be found. The ooze from the

eyes and beak chiefly contains odcysts which may be due to

resurgitation from the crop just before death. However, it

* It should be noted that in severe cases of duodenal coccidiosis, merozoites may

he found free in the zut-contents and fieces of infected birds when no odeysts are

present,

1910. | DR. HW. B. FANTHAM ON AVIAN COCCIDIOSIS. 715

should be noted that while mucous discharge is common, it is not

an invariable feature of coccidiosis,

Death from coccidiosis appears to be sudden. Some of the

experimental fowl chicks were feeding greedily an hour before

death, though death was almost expected from the great emaciation

and * bluish” appearance of the birds for some days previous to

the actual decease. All infected corpses should be burned, jof

buried.

TV. InvernAat Organs,

Detailed examination of diseased birds shows that coccidiosis of

fowls, pigeons and young grouse is confined chiefly to the digestive

tract, and so is unlike the coccidiosis of the rabbit where both the

liver and the gut may be affected.

Dead chicks have shown oval coccidian ovcysts (text-fig. 66,

p- 714) in the discharge from the nostrils. Scrapings from the

soft palate, trachea and cesophagus of diseased birds have shown

the presence of odcysts embedded in mucilage. Possibly the

odcysts may be regurgitated from the gizzard or intestine.

Odcysts are occasionally present in the crop and gizzard, mixed

with crushed food, having been probably taken up with the food.

The intestine is more highly parasitised than any other part of

the alimentary tract. The odcysts (Pl. LXII. figs. 7, 8, 11, 12),

which show much variation among themselves, are softened by

the pancreatic juice, the four sporocysts (figs. 9, 10) emerge and

from each of them two active, motile sporozoites or germs come out

and proceed to attack the epithelium of the duodenum. Having

penetrated the epithelium (fig. 1) they become round, grow and

produce individuals, destined to divide and give rise to a barrel-

shaped mass of active daughter germs, the merozoites (fig. 1, nz).

These merozoites separate from one another and infect fresh

epithelial cells, the whole mucous membrane being soon badly

infested, and becoming reduced to an almost structureless mass

(fig. 1). Owing to heavy infection in the duodenal epithelium,

death of the young grouse may occur. However, in many cases,

some of the merozoites pass into the lumen of the gut and reach

the ceca, the tissues of which are similarly mutilated. As far

as my experiments go at present, the full period of schizogony

would appear to be from 4 to 5 days, as judged by the appearance

and general moping of the birds.

After several generations of merozoites have been produced,

the power of the host to provide food for the parasite fails, and

consequent on this, the latter begins to make preparation for

extra-corporeal life and produces large, granular forms, which are

female mother cells or macrogametocytes (fig. 1, 2 ) and somewhat

smaller, less granular ones which are microgametocytes (fig. 2, ¢).

Kach macrogametocyte gives rise to one passive macrogamete or

female element (fig. 1), while each microgametocyte gives rise to

many minute, motile microgametes, the male elements (fig. 2).

Fertilisation (fig. 3) occurs through a micropyle left in the wall

714 DR. HH. B. FANTHAM ON AVIAN COCCIDIOSTS. | May 3,

with which the macvogamete invests itself, and the fertilised odeyst

so produced, passes out through the much damaged epithelium

into the lumen of the gut and thence to the outside. The epithe-

lium of the duodenum and eeca is sometimes entirely denuded

by the action of the many parasites that infest it, and the sub-

mucosa also is sometimes infected by the Himeria.

The contents of the odcyst, at first filling the interior (text-

fig. 66, A), gradually contract (text-fig. 66, B) towards the centre,

or occasionally towards one pole, and then divide (text-fig. 66, C)

into four round masses (text-fig. 66, D) known as sporoblasts.

Each sporoblast becomes an oval sporocyst (text-fig. 66, E) while

still within the odcyst, and within each ripe sporocyst two sporo-

zoites or germs are developed. Consequently each odcyst gives

rise to eight sporozoites (text-fig. 66, F).

Text-fig. 66.

Figures A-F,

Stages in the development of the odcysts of Mimeria avium, as seen in

fresh preparations.

A. Odéeyst (encysted zygote) with protoplasm completely filling it.

Bi. Older obcyst with zygote contents forming a central sphere. Many such cysts

are found in infected cea and infected fieces of grouse.

C. Odeyst with four nuclei, about to form sporoblasts.

DPD. Odcyst with four round sporoblasts.

1. Four ovoid sporocysts within odcyst.

F. Fully mature odcyst with four sporoeysts, each containing two sporozoites,

A certain amount of variation, as exhibited by the odcysts of

EL. avium, has been noticed by Morse (1908) in the coccidian para-

site of “ white diarrhoea” of fowls, where the odcysts are round to

oval, and from 12y to 25 in diameter, The o6cysts of #. avium

1910. | DR. H. B. FANTHAM ON AVIAN COCCIDIOSIS. 715

in grouse, which is identical with the parasite found in ‘ white

diarrhea” of fowls, also vary among themselves. Usually the

odcysts are oval (figs. 4-8), but a series of varying sizes and shapes

van readily be found (figs. 8, 11, 12, 14), while round odcysts also

occur (fig. 11). The exact shape and size of the odcyst is a factor

of the space in the cell available for the development of the

macrogamete, and should not be insisted upon as a specific character,

for where many parasites are aggregated together in a limited area

of epithelium, the macrogametes and oocysts are small, while in

areas of the gut but poorly parasitised, large odcysts preponder: ite.

Nutrition of the parasite has obviously a great influence on both

its size and its propagative power.

In some cases the gut-wall is extremely thin and tender; in

other birds this effect is not marked. Inflammatory patches may

be seen at intervals, particularly in the ceca, and the ceca usually

are enlarged. At the ileo-cecal junction, where a recurrence of

schizogony and sporogony occurs, much degenerated epithelium is

present in the gut contents, and this epithelium contains both

schizonts and gametocytes.

The large intestine of chicks infected with coccidiosis sometimes

shows inflammatory patches and blood may be present in the

rectal contents, The rectum itself seems rarely to be attacked

by #. avium, though its contents usually contain odcysts.

The kidneys, spleen, liver, and gall-bladder of birds suffer-

ing from coccidiosis never contained Himeria, though the spleen

and gall-bladder were sometimes enlarged. Examination of

the generative organs has shown no stage of 4. avium so

far, fhough it is possible that eggs may ewonme contaminated

during their passage through the cloaca of the mother, ‘The

young chicks then might be hatched m contact with infectious

material, and so acquire coccidiosis early in life. Eight days old

grouse chicks were the youngest naturally infected chicks that I

examined. ‘lhe period of 8 to 10 days is the one determined

roughly by my experiments as being required for the complete

developmental cycle of #. avium in fowls and pigeons, from the

time of ingestion of the odcysts to the excretion of the maximum

number of odcysts of the second generation.

Many bacteria are present naturally in the gut of the grouse,

and thei active movements can be well seen, especially in the

cecal contents of freshly killed grouse. Examination of sections

of the gut of the grouse, especially of sections stained with iron-

hematoxylin and iron-hematoxylin followed by van Gieson’s stain,

veveals the presence of numerous bacteria which are » present, not

only in the lumen of the gut, but forming a layer along the

striated edge of the colttmnar epithelial cells, and also are found

in lesions left by the outward passage of gametes and merozoites.

These bacteria may have a harmful effect on the tissues invaded,

and there is evidence to show that they gain access to the tissues

very early, the sporozoites and merozoites acting, in fact, as

inoculating needles, whereby the injurious bacteria are passed

into the tissue of the gut, whence, hy way of the blood and lymph,

716 DR. WH. B, PANTHAM ON AVIAN COCCIDIOSIS, | May 5,

they can reach other organs. Further, the denudation of the

epithelium of the gut allows of easy entry of bacterial agents of

infection.

In connection with the action of bacteria in the disease of

fowls known as “* white diarrhea,” there are two opposing views.

Morse (1908), working in America, has investigated white

diarrheea in fowls and other birds and always found intestinal

coceidiosis. Hadley (1909), also working on the subject, found

not only intestinal but also hepatic coccidiosis in fowls suffering

from white diarrhcea. Morse notes the presence of bacteria im

the gut, and thinks that they may gain access to the system

on necount of the denudation of the gut epithelium, Rettger

(1909), on the other hand, considers that white diarrhea is due

entirely to a bacterium, /acterium septicemie gallinarwm or

Bacterium pullorum.

Probably both of these conflicting views are right as faras they

go, but separately they may only partially explain the cause of

“white diarrhea.” Drs. Cobbett and Graham Smith, of the Grouse

Disease Inquiry, have shown (1910) experimentally that Bacteria

may be inoculated by means of coccidia, and find their way into the

internal organs probably by way of the portal vein. ‘The agency

of parasitic worms and coccidia in e: wising lesions of the mucous

membrane through which harmful bacteria may enter, is of far-

reaching importance and probably of wide application in the

elucidation of certain intestinal diseases.

Morse’s paper gives much valuable information regarding

treatment of coccidiosis and brief notes on intestinal coccidiosis of

various birds. Game birds other than grouse are susceptible to

coccidiosis, for M‘Failyean reported coccidiosis in pheasants im

1893-4. At the time of correcting proofs of this article I am

engaged in investigating coccidiosis which is causing the death of

many young pheasants in various parts of England.

The onset of sporogony of #7. avium means either the recover vy or

the death of the infected chick as a rule. When the odcysts pass

out from the body of the host, the epithelium of the gut may be

able to recover itself, when the bird gradually increases in weight

and makes partial or entire recovery. This recovery is sometimes

aided by infiltration of connective tissue into the lesions. If,

however, the infection has been heavy, the epithelium cannot

regenerate itself and the bird becomes exhausted and dies,

V. RELATIVE RESISTANCE OF DIFFERENT BREEDS OF

Fowts AND PiGrons To Coccrptosis.

Young chicks are far more susceptible to coccidiosis than adult

birds* Wi ery young fowl chicks (up to 8 days old) die in a few days

after being fed with coccidian odcysts. Chicks first dosed with

eysts when 14 days old do not suceumb so readily; they may live

* Old birds may become chronics and thus infect the moors. In this connection

infected foster mothers in pheasaut-rearing may be dangerous.

1910.] DR. H. B. FANTHAM ON AVIAN COCCIDIOSIS. 717

for some weeks or some may become chronics, when daily

examination of their feces shows periodic small crops of odcysts.

Older chicks are more likely to recover from coccidiosis. For

example, a grouse chick aged 13 weeks was fed on odeysts, and

when killed 3 weeks later very few coccidia were observed.

Again, a young grouse which had been picked up dead on the

moors in Inverness-shire was received on Sept. 15, 1909. In its

ceca numbers of both coccidian odcysts and Trichostrongylus

pergracilis were found. The bird had probably survived an

attack of coccidiosis and in its weakened condition had fallen

a victim to strongylosis.

Different breeds of fowls have varying powers of resisting

coccidiosis. Pure-bred Leghorn and Minorca chicks died from

the disease in a compavatively short time. Plymouth Rock chicks

were attacked quickly but had greater resisting powers than pure

Leghorns or Minoreas. Cross-bred Leghorns were far more

resistant to the disease than the pure-bred birds.

Regarding pigeons, fantails are more susceptible than the

ordinary pigeons, though squabs of any variety seem easily

attacked and overcome by coccidiosis.

VI. THE DiIssEMINATION OF COCCIDIOSIS OVER

Tracrs or CountTRY:

Coccidiosis outbreaks spread with fair rapidity, but though

many birds in one locality become diseased, yet adjoining estates

may be free. Epizootics of coccidiosis also disappear very

quickly.

Infection has been shown experimentally to be due to the

ingestion of odcysts (cysts), either by way of the food or drink.

Coccidian cysts may occur on the heather and in the tarns and

pools from which the grouse drink, as before mentioned.

When feces of infected chicks are voided, the cecal or soft

droppings are heavily loaded with cysts, and not only do they

foul the ground, heather, and water in their vicinity, but when

dried, the powdery material produced may be disseminated by the

wind, and so oocysts and their contained sporocysts are distributed

over comparatively large tracts of country.

But other methods of producing richer local infections may be

partly due to insects. The agency of insects such as dung-flies

has been observed in nature and also demonstrated experimentally.

Scatophaga stercoraria, the dung-fly, commonly occurs on grouse

moors. The eges of the fly are laid in feeces and hatch out there.

The larve are large and somewhat active. They feed on the

feeces of grouse, which feces may be infected with coccidian cysts.

Dissection of such larve has shown the presence of oocysts within

their guts. When isolated larve were first well washed and then

allowed to defecate on a slide, the preparation of the feeces showed

odcysts when examined microscopically. Some of the pup also

contained coccidian spores in their guts. The freshly hatched

Proc. Zoo. Soc.—1910, No. XLVII. 47

718 DR, 1. B. FANTHAM ON AVIAN COCCIDIOSIS. [May 3,

flies examined rarely showed spores, but as they proceed almost at

once to ingest feces, they rapidly become agents for distributing the

spores. Dung-flies allowed to hatch out in the feces of infected

grouse always contained coccidian odcysts in their alimentary

canal and feeces.

Laboratory experiments were made with the blow-fly, J/usca

vomitoria. Eggs taken from the body of the parent fly were

allowed to hatch out in feces of an infected grouse chick. The

larve greedily ingested the cysts which passed practically unaltered

through their bodies. Some of the larve which pupated were

washed very carefully and then dissected. They contained

coccidian spores. Adults were fed on the infected material and

obeysts were voided in their excrement. The spreading of

young flies, hatched infected, and of older ones that have fed on

infected material, may aid, then, in the dispersal of coccidiosis.

While experimental evidence of the action of Scatophaga ster-

coraria and Musca vomitoria has been afforded in the laboratory,

this is merely confirmatory of what may occur on some moors.

Here on Scots firs, heather and moss, numbers of flies are

found, and dung-flies are known wherever grouse droppings are

to be found, The trail of birds can be tracked to some extent by

the coprophagous flies, while lines of infection are produced by the

birds as they pass down by small paths to their drinking places.

It may be of interest to note that houseflies (JZusca domestica)

have been shown experimentally to ingest coccidian oocysts.

To summarise, wind and rain acting on infected feces are

probably the principal agents in dispersal, while the feeding habits

of Scatophaga and other flies form subsidiary means of spreading

the disease.

VII. Duration or ViraLity or CoccipIAN OdcysTs.

(2) When the Oocysts are kept in water or very moist.

Much moisture is present on many moors, and feces of infected

chicks contain coccidian spores which not only get washed into

the soil, but also into tarns, ete., at which grouse drink. It was,

therefore, of some importance to determine the time required

for the degeneration of the resistant spores of the parasite when

in water.

Coccidian odcysts with undifferentiated contents were placed in

water, kept at about 20° C. (July temperature), the water being

replaced as required, to avoid evaporation effects. Ordinarily, the

odeysts develop sporocysts very rapidly—in 2 to 3 days. In

the case of cysts kept in water, 9 days elapsed before much

change was noted. At the end of that period, a few odcysts

showed differentiated protoplasmic masses, and still fewer showed

four sporocysts. Two days later many more odcysts contained

four sporocysts, and this progressive development continued for

some days. Little signs of degeneration were seen until about the

40th day, when some showed signs of gas bubbles in their interiors.

Others, however, had completed their development, and their four

1910. } DR. H B. FANTHAM ON AVIAN COCCIDIOSIS. 719

sporocysts, apparently unharmed, were set free into the liquid.

By the 50th day, practically all oocysts had either matured or

degenerated, and the sporocysts had begun to degenerate.

From the above experiment, the conclusion i is that the develop-

ment of odcysts and sporocysts is delayed by the presence of much

moisture, but that the power of infection is retained for a long

time by means of the sporocysts.

Very damp air has similar effects.

(B) When Feces are merely kept and allowed to dry on

the outside.

When freshly voided soft droppings of grouse containing

coccidian odcysts are allowed to dry, the odeysts in the surface

layers rapidly develop sporocysts, the inner ones remaining

unaffected.

Feces kept e7 masse in covered dishes for as long as twelve

months have retained the power of infecting birds, as I have

been able to show experimentally. Such material contains un-

differentiated odcysts still, while its outer layers mainly contain

odeysts with four sporocysts within them,

(y) Development under Different Conditions of Temperature.

For experimental purposes, it was sometimes necessary to delay

the development of sporocysts. This was easily done. The oocysts

in feces were transferred to a chamber kept at 10° C., having been

previously kept at 15° C. This change was sufficient to delay all

further development for a considera ble time. Smaller changes of

temperature also arrested the development otf sporocysts, though

‘the effect naturally was not so marked.

Changes of temperature and moisture on the moors may explain

the oceurrence of occasional outbreaks of disease after all disease

seems to have disappeared. Moisture and coolness retard the

development of certain odcysts for a considerable time, during

which period the disease disappears. A return of conditions

favourable to the Coccidium then ensues, rapid development of

sporocysts occurs and a fresh outbreak of disease is reported.

VIII. Errecrs or Certain REAGENTS ON CoccrIDIAN O6cystTs.

Certain experiments were made with a view to finding a means

of destroying odcysts without killing all other forms of life.

This is not an easy matter. While such strong re-agents as

caustic potash will slowly dissolve the odcysts, their applic: ition is

not practicable on grouse moors.

Salé produces plasmolysis in the end, but the process is

rather slow, and the salt is too readily dissolved in dew and rain,

and so merely soaks into the soil *.

_ * However, in this connection see Hammond Smith, “ The Field,” Aug. 20, 1910,

Suppl. p. viii.

47*

720 DR. H. B. FANTHAM ON AVIAN COCCIDIOSIS. [May 3,

Quick-lime destroys the odcysts and sporocysts. It also causes

the faces to cake, thereby preventing scattering of the spores.

It is somewhat doubtful whether lime could be applied on a large

scale. It might be somewhat harmful to the feet of the birds,

apart from the difficulty of distributing it over large areas.

Where the area of infection is small, it is probable that the

application of lime to the soil would be of service. My experi-

ments on a small plot of heather at Cambridge have shown that

small quantities of lime dressing are not detrimental to heather.

Gas lime and slaked lime also are useful, but each is open to the

same objection as quick-lime. Lime in one form or another

certainly seems to have the best and most rapid action on coccidian

odcysts of any reagent that I have tried.

Salicylates—Salicylic acid and sodium salicylate act rather

slowly on coccidian odcysts when mixed with them. Both

chemicals tend to deliquesce, and the feeces mixed with them

remain fluid for a longer period than they otherwise would. The

odcysts become wrinkled and ultimately destroyed, but the con-

tents take longer to degenerate than when lime is used.

Ferrous sulphate.—Copperas or green vitriol is useful to some

extent in destroying coccidian odcysts, but like salicylates it is

somewhat slow in action. A dusting of ferrous sulphate on the

moors would probably be useful, for the combined iron present

might be taken up in small quantites by grouse and, by acting as

a general tonic, might enable the birds to resist coccidiosis the

better if they became attacked. Ferrous sulphate in the pro-

portion of 10 grains to the gallon of drinking-water has been

found of service by the writer in the treatment of coccidiosis in

young fowls and young pheasants. Some keepers and breeders,

at my suggestion, have used catechw in the drinking-water with

success.

Sodium nitrate.—Nitrate of soda mixed with feces destroyed

the contents of the oicysts after some time, but the length of time

required for its effective application would militate against its use

on a large scale.

In my experiments at Cambridge, three portions of infected

feces were mixed with equal quantities of lime, sodium salicylate,

and ferrous sulphate respectively, and were kept in open dishes,

exposed to the action of the weather, The results obtained may

be shortly given.

In the case of lime, the feces rapidly formed a caked mass. In

a fortnight the odcysts were shrunken and wrinkled, and some

showed eracks. At the end of three weeks, the cysts were more

broken up and the contents largely disintegrating, while after a

lapse of two months, there was a difficulty in finding spores at all.

Bacteria were not found after lime treatment, and there was only

a slight feecal odour noticed.

Sodium salicylate added to feces rapidly deliquesced, in fact,

the mixture was quite liquid in less than three hours. After a

1910.] DR. H. B. FANTHAM ON AVIAN COCCIDIO“IS. (PAI

fortnight’s interval, the cysts appeared to be slightly shrunken,

while feecal odour was noticeable. A month later, the obcysts

were more shrunken and a few free sporocysts were found, while

ten weeks after treatment, a slight smell was still perceptible, and

the odcysts present were shr unken and showed oily contents.

Ferrous sulphate (copperas) had much the same effect as sodium

salicylate, but did not deliquesce. Though its action at first

seemed to be rather less effective, it secured the same result

ultimately.

TX. Conciupine REmMARKs.

The ravages of coccidiosis among grouse chicks may be under-

estimated on the moors, since the tiny corpses of the birds lie

hidden among the heather.

While it is relatively easy to take preventive measures in the

case of coccidiosis in fowls, it is most difficult to take active

measures in the case of grouse. The remedy of heather burning

is drastic and coccidian spores, which are present in the tract

burnt, are then destroyed. However, heather is rather slow-

growing, and so heather-burning, while efficacious, is somewhat

restricted in its area of application. Lime dressing is destructive

to spores of coccidia, and could probably be utilised in the case of

limited outbreaks of disease among grouse chicks. The effect

of lime on the heather on the moors should first be carefully

investigated on a large scale (see p. 720). Any condition tending

to raise the general vitality of the birds also makes them much

more resistant to disease. An abundant supply of healthy young

heather, by raising the general standard of health of the birds,

is probably one of the best safeguards against the insidious disease,

coccidiosis.

References to Literature.

Cossett, L.,and GrawAm-Smitu, G.S8. (1910). —“ An Investigation

of the Letina sy of * Grouse Disease.’” Journ. Hygiene, x.

p. 1-86, 5 pls.

Fanruam, H. B. (1910).—* The Morphology and Life History of

Himeria (Coccidiwm) avium; a Sporozodn causing a fatal

disease among young Grouse.” Proc. Zool. Soc. Lond.,

672-691, 4 pls.

Haptey, P. B. (1909).—“ Studies in Avian Coccidiosis. I. White

Diarrhea of Chicks. II. Roup of Fowls.” Centralbl. f.

Bakteriol, Abt. I., Orig., 50, pp. 348-353.

Morss, G. B. (1908). eWhite Diarrhea of Chicks, with Notes on

Coccidiosis in Birds.” Cireular 128. U.S. Dept. Agricult.,

Bureau of Animal Industry.

Rerresr, L. F. (1909).—* Further Studies on Fatal Septicemia

in young Chickens, or ‘White Diarrhea.” Journ. Med.

Research, xxi., pp. 115--123

122 DR. H. B. FANTHAM ON [May 3,

EXPLANATION OF PLATE LXII.

All figures outlined with Abbé camera Incida, using Zeiss 2 mm.

apochromatie lens.

Figs. 1, 2, 3 from stained (hematoxylin) sections of gut of grouse chick infected with

coccidiosis. > 1500 approximately.

Figs. 4-14 from fresh preparations. > 1000.

Figs. 1, 2. Section of gut showing parasites.

par.=parasite.

spz.=sporozoite (infecting germ).

mz.=merozoite (daughter form).

3 =microgametes (male elements) attached to microgametocyte

(gd mother cell).

9 =macrogamete (female), with thin cyst around it.

3. Fertilisation of large macrogamete by minute microgamete. Only one male

element penetrates, the others remain on the outside and degenerate.

4, 5. Odcysts, characteristic of fresh feces.

6, 7. Odcysts with four developing sporoblasts inside, first round, then ovoid.

8. Mature odcyst containing four sporocysts, each with two sporozoites, from

cecal droppings taken from the moors.

9,10. Free sporocysts (spores) from imfected feces. In the sporocyst repre-

sented in fig. 10, which is pointed at one end, the sporozoites are about

to emerge.

11. Round eyst, as found in droppings of grouse or fowl chicks.

12. Somewhat egg-shaped cyst, with four sporoblasts (young spores).

13. Cyst of Coccidiwn (Himeria) pfeifferi (Labbé) trom a pigeon infected

with natural coccidiosis.

14. Cyst from fxces of a pigeon fed on infected grouse feces.

4. Observations on the Blood of Grouse. By H. B.

Fantoam, D.Sc.Lond., B.A.Cantab., A.R:C.8., F.Z.8.,

Christ’s College, Cambridge, Assistant to the Quick

Professor of Biology in the University, Protozoologist

to the Grouse Disease Inquiry.

{ Received April 5, 1910. |

(Plate LXIII.*)

I. [nrropucrion.

The elements of the blood of birds are very different from

those of mammals, and while much is known of the histology

of mammalian blood, the investigation of the blood of birds has

hitherto been very limited, and very little indeed is known of

the subject, Such literature as is available on avian blood is,

unfortunately, largely contradictory, and the few illustrations

relating to the same have not solved entirely the difficulties

connected with the cellular elements of the blood of birds.

The portions of the subject that have been most adequately

dealt with are those relating to certain pathological conditions,

fowl cholera and fowl typhoid, occurring in domestic fowls, and

those relating to normal fowls’ blood. In connection with both

* For explanation of the Plate see p. 731,

bridge

Jeo

1) Ub rh

) WIS On. alm

PZS.1910,P1.LXM,

Ory s

CHIEFLY OF GROT

BLOOD CELLS

ham, del.

i BFa

1910.] THE BLOOD OF GROUSE. 723

these sets of observations there are wide discrepancies between

the results obtained by different investigators, and much con-

fusion has arisen therefrom.

The different results obtained by various investigators are

partly explicable, for the constitution of the blood of ‘birds may

vary among individuals of the same species. Also there are

differences due to age and sex, while the conditions under which

the investigations are made have some influence on the result.

The chief difficulties of manipulation for the investigation of

the blood of such birds as grouse are the rapidity with which

coagulation occurs, and the rapid alteration and disintegration

that often follows the death of the blood elements. The great

‘apidity of the onset of degeneration in the leucocytes is such

that extreme accuracy in counts of these cellular elements is not

easily obtained, and the numbers of erythrocytes and leucocytes

found in different series of counts consequently bear a varying

relation one to another in apparently normal birds, even when

the factors noted in the preceding paragraph have been con-

sidered. Nevertheless, the correlation of the numbers and

condition of the blood-elements, particularly of the leucocytes,

with the occurrence of specific protozoal and helminthic parasites,

renders an examination of the blood a most useful adjunct in

investigating somewhat obscure diseases. Several protozoal para

sites [see Fantham (1910)| are present in grouse, both in ae

blood and in the gut, and the condition i the bird’s blood

can be correlated with the presence of some of the individual

parasites.

II. Meruops.

Blood taken from a wing vein was used in the case of living

birds, of which twelve were thus examined. In the case of freshly

killed grouse, blood taken directly from the heart was employed.

The usual methods and precautions were followed in securing

suitable drops of blood for examination.

Hayem’s fluid was used as a diluent for counting the red cells

of the blood, while Toison’s fluid was used in the case of the

leucocytes,

The Thoma-Zeiss hemocytometer was employed. The disc of

the counting chamber was ruled according to Zappert.

A dilution of at least 200 or 250 was found to be necessary

for counting the red cells, and a dilution of 20 to 25 was absolutely

necessary 1n counting the leucocytes. Sometimes a dilution of

100 was taken in counting the leucocytes, and this greater

dilution was often preferable.

For differential leucocyte counts blood smears were made on

both cover-slips and slides, especially the former.

The stain generally used in making preparations for differential

counting was that of Jenner. Occasionally Giemsa’s stain was

employed.

724 DR. H, B, FANTHAM ON [May 3,

JII. Tae Certs iy true Normat Bxioop or Birps,

ESPECIALLY GROUSE.

The elements present in the blood of normal grouse may be

grouped as:

(a) Erythrocytes.

(>) Leucocytes.

The leucocytes are further subdivided, and may be grouped as

mononuclear, polymorphonuclear, and eosinophile leucocytes, and

lymphocytes. Mast cells, which are markedly basophile, are also

present. Erythroblasts also may be encountered, These forms

may now be considered separately.

(a) Hrythrocytes—These are the most abundant elements in

the blood. The erythrocytes or red blood corpuscles are oval in

contour (Pl. LXIIT, fig. 1). They possess a well-marked oyal

nucleus, centrally placed, The erythrocytes of grouse are about

11°5-12 p long by 7-7:5 p broad. In fresh preparations the

cytoplasm of the ved cells of healthy grouse is practically homo-

geneous, or but very faintly er: anular. Stained preparations give

the same results. It is rare for large granules, chromatoid

granules, or vacuolations to be present in the red cells of healthy

birds. The nucleus stains deeply, and shows a_ well- marked

chromatic meshwork with small net-knots upon it at intervals.

The red cells of various birds are much the same. ‘There may

be slight differences in size and shape. In figs. 2 and 3 are

depicted stained specimens of the red cells of the pigeon and

fowl respectively, from which it will be seen that the erythro-

cytes of the pigeon are slightly narrower than those of the grouse

and fowl, and somewhat leas rounded at the ends.

The number of red cells of grouse varied from 3,600,000 to

5,800,000 per cubic millimetre, averaging 4,800,000 per c.mm.

These figures result from blood-counts of 50 binds.

The number of ved cells is apparently higher in cock grouse

(e. g., 4,400,000 per c.mm.) than in hen grouse (e. g., 3,800,000 per

¢.muny.).

(b) Leucocytes,—The leucocytes of grouse are a subject of much

intrinsic difficulty, and it is impossible to consider the grouping

arrangement given below as more than provisional, The classi-

fication of the leucocytes of birds can be placed on a really sound

basis only when detailed knowledge of the development of all the

elements of the blood of birds has been obtained In spite of

good work by Denys, Dantschakoff, and others, such a complete

developmental investigation of the blood of birds is still wanting.

Owing both to the lack of time at my disposal and to the great

aearcity of guite fresh material, | am unable to attempt such

1910.] THE BLOOD OF GROUSE. 725

an investigation, and therefore have availed myself of the

classification at present adopted by such authorities as Burnett

(1908), modifying their statements in accordance with my own

personal observation and adding a number of details hitherto

overlooked.

1. Lymphocytes (Pl. LXITI. figs. 5, 6).—The lymphocytes are

the smallest of the leucocytes. They are from 5:5 p to 8 p in

diameter approximately. They are smaller than the average red

corpuscles. Each lymphocyte possesses a well-marked round or

oval nucleus which is relatively large and occupies most of et

cell. The cytoplasm of the lymphocyte is small in amount,

basophile, and is reduced to a yim around the nucleus. Phe

lymphocytes have a great tendency to collect together in groups

on blood-smears, and in such aggregations the cytoplasm of the

members away from the periphery of the group is reduced to a

minimum.

The lymphocytes are sometimes subdivided into large and

small varieties. The larger specimens of lymphocytes gradually

merge into the small mononuclears, and the n: aming and classi-

fying of such leucocytes is often a matter of personal opinion.

2. Large Mononuclear Leucocytes (Hyaline cells)—These are

large, approximately circular cells averaging 9 w to 115 p in

diameter (Pl. LXIIT. figs. 7, 8). Ina film stained with Giemsa’s

fluid the mononuclear leucocytes are easily distinguishable not

only from their large size but also from the fact that their cyto-

plasm stains a vivid blue and the nucleus a beautiful purple.

The general cytoplasm of the cell is almost homogeneous in

character.

The nucleus of a mononuclear leucocyte of grouse is large,

occupying about half of the cell, and is often round or oval

(figs. 5, 6). At other times the nucleus is somewhat curved

or indented (fig. 8). The chromatin of the nucleus takes the

form of a fairly dense mass, exhibiting in stained specimens a

uniformly mottled appearance ; it is situated to one side of the

cell.

With Jenner’s stain the nucleus colours a rather deep blue,

while the cytoplasm stains only a faint blue and so is slightly

basophile.

As Burnett (1908, p. 35) writes: ‘One can find all stages

between typical lymphocytes with a small amount of strongly

basophile, coarsely reticular cytoplasm and typical large mono-

nuclears with a much larger amount of faintly basophile, finely

¥eticular cytoplasm.”

3. Polymorphonuclear leucocytes (Burnett) or Crystalloid eosino-

phile cells (Cullen, Warthin).—The apparently polymorphonuclear

leucocytes of birds exhibit marked differences from those of

mammals as regards the contained granules.

These cells in grouse are round, as seen in stained preparations,

and measure from 9 » to 9°5 pw in diameter (figs, 10-12). The

726 DR. HH. B, FANTHAM ON [| May 3,

nucleus consists of two or more lobes and varies in shape—in

other words, is polymorphous. The nucleus is fairly well stained

with Jenner's stain and is coarsely reticular (figs. 10-12). The

cytoplasm stains very faintly.

Embedded in the general cytoplasm are a number of bodies or

granules which stain red with Jenner’s stain and are oxyphilie.

The bodies are generally described as spindle-shaped with

tapering ends, but their outline has also heen compared with that

of a torpedo or cigar (fig. 11), Other leucocytes are found with

thinner inclusions or ‘‘ granules” which are rod-like in shape

(fig. 10).

Cullen (1903) writes: “ As to the nature and origin of these

spindles [inclusions in the cytoplasm of the leucocytes | very little

is known that is definite. They are certainly not artifacts, for

they can be seen in fresh blood as well as in dried specimens.

Dr. Simon is of the opinion that they may be derived from the

second variety [the coarsely eosinophile leucocytes, mentioned

next |, in which granules take the place of spindles; that they

are crystalloids and analogous to similar formations that have

been encountered in certain tissues In man, and notably in the

epithelial cells of the seminal tubules. He thinks that they

result from the granules through loss of water, and as a matter

of fact it is possible to reconvert the erystalloids into granules

in the wet preparation by adding a droplet of a dilute solution of

eosin from the side of the cover-glass. Dr. Simon has also noted

that in certain preparations in which the eosinophilic material

is present in one dense, apparently undifferentiated mass, the

erystalloids separate out upon the application of heat.”

In a few cases, the centre of the spindle bodies is occupied by

a tiny circle or dot, which does not stain as deeply as the rest of

the spindle and is refringent (fig. 12). The presence of this

central dot is noticeable in the case of large spindles.

Asregards the staining properties of the spindle bodies (erystal-

loids), I agree with Burnett that they are not intensely eosino-

philic, but that “in aftinity for stains the [spindle] granules

resemble the polymorphs rather than the eosinophiles.” Cullen,

on the other hand, considers that the crystalloids are more

oxyphilic than the true eosinophile leucocytes. Perhaps the

differences of opinion of the various investigators are explicable

by reference to slight variation in the stains used.

The distribution of the eytoplasmic inclusions within the cell

is dependent on the shape and position of the nucleus.

4. Hosinophile leucocytes (Burnett) or coarsely granular eosino-

phile (oxyphile) cells.—These cells are also present in the blood of

normal grouse (figs. 13-15), but are more numerous in the blood

of birds suffering from helminthiasis, The eosinophile cells are

from 8 x to 10 pin diameter, and have a general resemblance to

the polymorphonuclear leucocytes, only differing in the form of

the inclusions or granules,

1910.] THE BLOOD OF GROUSE. an

The shape of the eosinophile leucocyte is round (figs. 14, 15) or

somewhat quadrilateral (fig. 13). The general cytoplasm is pale

staining and slightly reticulate in str ucture. The nucleus usually

resembles that of the polymorphonuclear leucocyte, and is mostly

bilobed—the lobes being coarsely reticular and usually staining

fairly deeply. ‘The cell- body contains numbers of coarse, oxyphile

(acidophile) granules which stain deeply with eosin. The granules

are often rounded (figs. 13-15) ; they vary in size, some eosinophiles

containing large granules (fig. 13) which may be relatively fewer

in number, while in other cases the granules are small (fig. 14),

and may be relatively more numerous. osinophiles with large

granules are the more prevalent. The contour of the granules is

generally round, but sometimes cells containing somewhat ovoid

granules are seen which stain intensely with eosin.

Occasionally eosinophile cells are found in stained films the

granules of which vary in shape within the same cell, some being

round, some ovoid, and some almost spindle-shaped. Such cells

and their contained granules may be somewhat deformed in the

making of the film, otherwise it is difficult to classify these cells

which are intermediate between crystalloid (polymorphonuclear)

and granular eosinophiles.

Mononuclear eosinophile cells are occasionally seen in the

blood of normal grouse, and perhaps represent an early stage

of development of the polymorphonuclear eosinophiles, Such

mononuclear cells are somewhat smaller than those depicted in

Pl. LXITI. figs. 13-15, and have round nuclei. They are rare,

and are perhaps analogous to the young eosinophile myelocytes

of man.

5. Mast cells (coarsely granular basophile cells) are present in

the blood of normal grouse (figs. 16, 17). They are rare. The

cells are more or less rounded, with a pale staining cytoplasm.

The nucleus is usually rounded (fig. 17) or slightly polymorphous

(fig. 16), and stains blue with Jennevr’s stain—rather more deeply

than similar cells in mammals.

Basophile granules, which vary in size and in number, occur in

the cytoplasm. The granules are usually rounded end stain a

deep purplish colour with Jenner’s stain—in other words the

granules are metachromatic. Mast cells measure from 9, to

10°5 pe in diameter.

(ce) Thrombocytes also occur in the blood of grouse (fig. 4). In

the fresh condition they suggest very narrow “and slightly small

erythrocytes. They are often elliptical, with an oval nucleus

centrally placed.

When stained the cell-body is pale and vacuolated, suggesting

a coarsely reticular cytoplasm. The whole cell is basophile in its

reactions, staining rather faintly blue with Jenner's stain.

Thrombocytes in grouse average 9 yu by 4 pu.

728 DR, H, B. FANTHAM ON [May 3,

These elements may collect in clumps, and show a marked

tendency to degeneration.

Red cells without nuclei were occasionally found in the blood

of grouse, Such non-nucleate cells, however, were very rare.

At times there also appeared to be a number of free nuclei in

the blood of grouse. In this connection we may note that

Warthin (1907) found 16°5 per cent. of degenerated cells in the

blood of normal fowls.

Erythroblasts occur in the blood of grouse in small numbers.

The cells, which are nucleated, are rounder than erythrocytes,

and are devoid of hemoglobin, ‘The nucleus of an erythroblast

is more spherical than that of an erythrocyte. The general

cytoplasm is homogeneous, staining blue with Giemsa’s stain.

The number of leucocytes found per cubic millimetre in the

blood of apparently healthy grouse varied from 22,000 to 50,000,

averaging about 32,000 per c.mm.,

The average results of the differential counts of leucocytes of

apparently healthy grouse may be tabulated thus :—

Tym oy tes! 7 eae eeeeeetraieccs.sc css seem 57 per cent.

Maree Mononuclear saueets..-..s +s. cece ce amet LO sweat

Polymorphonuclears (crystalloid eosinophiles). 20 ,, ,,

Kosinophiles (coarsely granular eosinophiles)... 3 4, ,,

Mastrcells’(basoplilles) aerevss: 0. -+--ceees tutes heey bale

The difficult computations are those of the large mononuclears

and lymphocytes. There are many medium sized mononuclears

in grouse, which different observers would classify differently.

TV. ParuoiocicaAL BLoop oF GROUSE.

The association of altered conditions of the blood with the

presence of protozoal parasites has been noted already, and I have

found certain alterations in the relative proportions of the blood

cells of grouse that harboured Protozoa—Leucocytozoa, Spiro-

chietes and Coccidia—while other differences are to be associated

with helminthiasis. The results may now be summarised :—

(a) When Spirocheeta lagopodis | F antham (1910)] was present in

the blood of grouse, as it was to some extent in two specimens,

the number of mononuclear leucocytes increased and these leuco-

cytes became slightly enlarged and vacuolated (PI. LXIIT. fig. 9).

Levaditi (1901) ‘noticed the presence of vacuolated mononuclear

leucocytes in the blood of fowls infected with Spirocheta galli-

narum, together with mononuclear and polynuclear leucocy tosis.

Balfour (1908) found vacuolated mononuclear leucocytes in the

blood of fowls suffering from spirochetosis in the Soudan. The

infection of grouse with S. /agopodis was prokably not nearly as

1910.] THE BLOOD OF GROUSE. 729

great as that of fowls with S. gallinarwm investigated by Levaditi

and Balfour.

(3) The presence of Leucocytozobn lovati in the blood of grouse

isassociated with mononuclear leucoeytosis. A. differential leuco-

cyte count of the blood of one of the grouse containing L. lovati

gave :

liymphoeytes a: Yics.cctenls 63-0 per cent.

Large mononuclears ......... 2S:0F ys

Polymorphonuclears ......... Ga) Sealer

Hosmmoplilest pe sewaseet sass PO Stats

MASGIC CNSR eccwsen ces fammelhoe TO Ow ae

There was also evidence of polychromatophilia in the red cells

of grouse infected with Lewcocz ytozoon lovati.

Gy) Himeria (Coccidium) aviwm also has its eftect, indirectly,

on the blood. Birds suffering from coccidiosis became anemic.

The paleness of the blood is due to the diminished number of

red cells in the blood. Estimations of the hemoglobin value by

Tallqvist’s scale gave 60-70 in the case of grouse chicks suffering

from coccidiosis, 80-90 for healthy grouse chicks.

There is an increased number of polymorphonuclear leucocytes

(erystalloid eosinophiles). Differential SUEY te counts of two

grouse-chicks dying from coccidiosis gave

Polymorphonuclears ...... 31:5 to 39°5 per cent.

iosmophules™ )eaeesa.s.cecc SO GOMOD TON set

Large mononucleais ......... Se OkGOns a O mans

liyamnpphocy best ssseas he DOL OGoro 0:0) aires

Masti cells vse census 0-6 per cent.

Jn Mammals an increase in the number of polymorphonuclear

leucocytes occurs in inflammation, especially locally. The

increase in polymorphonuclear leucocytes (crystalloid eosino-

philes) is probably to be associated with the intestinal inflamma-

tion caused by the presence of Coccidian parasites.

Similarly in a fowl chick dying of coccidiosis, I obtained the

following differential leucocyte count :

Polymorphonuclears ......... 47 per cent

Hosmurophilesa :.y2va sess PG pS

temo mtn Garceia eee seers tor Oso Mayan

ivanplaeyites” | “saetient.2 ees. DOP dee,

Masticells i..sveee&: nr: it pads.

The blood of this fowl chick * contained basophile spindle-

shaped cells (PI. LXITI. fig. 18).

(0) Helminthiasis is eommon in grouse. The various worms

found in grouse have been well described by Shipley (1909).

* Burnett gives the following differential leucocyte count of normal fowl’s

blood :—Polymorphonuclears 28°8 per cent., eosinophiles 3°3 p. c., large mono-

nuclears 5°5 p. c., lymphocytes 58 p. ¢., mast cells 4°3 p. e.

730 DR. H. B. FANTHAM ON [ May 3,

Grouse which would be considered perfectly healthy by keepers

have often contained a few worms. ‘The causal factor of disease

in adult grouse is the larval stage of Trichostrongylus pergracilis,

as shown by the researches of Dr. Leiper (Abstr. Proc. Zool. Soe.

Lond. No. 80, March 1,1910). Theadult Z'richostrongylus occurs

in the ceca of grouse, and sets up inflammation therein. In the

blood of three adult birds dying on the moors from grouse disease,

I obtained differential leucocyte counts which may be thus sum-

marised :

Hosinophiles ........:.:.....+. 23°5 to 42°0 per cent.

Polymorphonuclears ........- Dito. La OP, Tes

Mononuclears |...:....:-.--- 60) toil0:0 s,s.

Lymphocytes ..........----- AA: Osho p02 OL uy eeieies

Mast icelllsigipee teeter secre: 0-5 per cent.

Contrasting these counts with those given for normal grouse,

the occurrence of eosinophilia is deduced. All three of these

birds, dying of ‘ grouse disease,” had many Hymenolepis microps

in their duodenum in addition to Trichostrongylus in the ceca.

Eosinophilia is often associated with helminthiasis in mammals.

Helminthiasis in grouse results not only in an increase in the

number of eosinophile leucocytes, but also in a diminution of the

number of erythrocytes in the diseased birds. Thus the number

of red cells found in an adult cock grouse dying from hel-

minthiasis was 3,250,000 per cubie millimetre—the number of red

cells for a normal cock grouse being about one million more.

The hemoglobin value estimated by Tallqvist’s scale was 60 for

a grouse suffering from helminthiasis and 80-90 for healthy adult

grouse.

In spite of the incomplete character of this investigation of the

blood of the grouse (due to the lack both of material and time at

my disposal), I think that there are sufficient results contained

herein to emphasise the importance of parallel investigations of

the blood in connection with all animal diseases.

References to Literature.

Apamt, J. G. (1907).—Inflammation, an Introduction to the

Study of Pathology. 240 pp. London: Macmillan & Co.

Buryerr, 8. H. (1908).—The Clinical Pathology of the Blood of

Domesticated Animals. 156 pp., 4 pls. Ithaca, Nei

U.S.A.: Taylor & Carpenter. (Blood of Fowl, Pl. 3.)

(Guiuen, E. K. (1903).— A Morphological Study of the Blood

of certain Fishes and Birds, with special reference to the

Leucocytes of Birds.” 3ull. Johns Hopkins Hesp. xiv.

pp. 352-356.

Dantscuakorr, W. (1908).—‘ Untersuchungen iiber die Ent-

wicklung yon Blut und Bindegewebe bei Végeln.” Archiv

f. Mikroscop. Anatomie, 73, pp. 117-181, 2 pls.

1910. | THE BLOOD OF GROUSE. 731

Denys, J. (1888).—“ La structure de la moelle des os et Ja génese

du Sang chez les Oiseaux.” La Cellule, iv. pp. 203-240,

2 pls.

es HB. (1910).—“* The Morphology and Life-History of

Himeria (Coccidium) avium: a Sporozobn causing a fatal

disease among young Grouse.” Proc. Zool. Soe. Lond. 1910,

pp. 672-691, 4 pls.

Fantuam, H. B. (1910).—* Observations on the Parasitic Proto-

zoa of the Red Grouse (Lagopus scoticws).” Proce. Zool. Soc.

Lond. 1910, pp. 692-708, 3 pls.

Fanruam, H. B. (1910).—‘“ Experimental Studies on Avian

Cozeidiosis.” Proc. Zool. Soe. Lond. 1910, pp. 708-722,

1 pl.

Vee V. A. (1895-6).—* Infectious leukemia in Fowls.” 12th

& 13th Ann. Rept. Bureau Anim. Industry, U.S. Dept.

Agric. p. 185.

NeAVE, SHEFFIELD (1906).—Second Report Wellcome Research

Laboratories at Gordon College, Khartoum, pp. 194-6.

Surpiey, A. H. (1909).—‘* The Threadworms (Nematoda) of the

Red Grouse (Lagopus scoticus).” Proc. Zool. Soc. Lond.

1909, pp. 335-350, 8 pls.

Suipiey, A. KE. (1909).—‘* The Tape-Worms (Cestoda) of the Red

Grouse (Lagopus scoticus).” Proc. Zool. Soe. Lond. 1909,

pp. 3851-365, 5 pls.

Warrain, A. 8S. (1907).—“ Leukemia of the Common Fowl.”

Journ. Infect. Diseases, iv. p. 369.

EXPLANATION OF PLATE LXIII.

All figures were outlined with Abbé-Zeiss camera lucida, using Zeiss 2 mm.

apochromatic immersion objective and compensating ocular 12. Preparations

stained with Jenner’s stain.

The magnification is in all cases approximately 2000 diamcters.

Fig. 1. Erythrocyte of Grouse.

2. Erythrocyte of Pigeon.

3. Erythroc) te of Fowl.

4. Thrombocyte of Grouse.

5, 6, Lymphocytes of Grouse.

7, 8. Mononuclears of Grouse.

9. Mononuclear leucocyte of Grouse showing vacuolations internally.

From a bird suffering from slight spirochztosis.

10-12. Polymorphonuclears (Burnett) or crystalloid eosinophile cells

(Cullen) of Grouse. In fig. 10 the cytoplasmic enclosures are rod-

like, in fig. 11 they are spindle-shaped bodies, while in fig. 12 the

spindle bedies have small clear central areas.

13-15. Eosinophile leucocytes of Grouse (coarsely granular eosinophiles

ot Cullen). The figures represent the variation in the size and shape

of the eosinophile granules. Figs. 13, 15 represent the commonest

forms of eosinophiles.

16, 17. Mast cells (basophiles) of Grouse, showing metachromatic basophile

granules in cytoplasm.

18. Hlongate tailed cell occurring in the blood of a Fowl chick. The chick

was suffering from coccidiosis. Some authors consider such spindle-

shaped or tailed cells to be normal to the blood of birds and reptiles.

Extremely few such cells occur in the blood of normal Grouse.

» a

“7 oe

732 PROF. G. 0. SARS ON | | May 3

Zoological Results of the Third Tanganyika Expedition,

conducted by Dr. W. A. Cunnington, F.Z.8., 1904—

1905.—Report on the Ostracoda. By Prof. G. O. Sars,

C.M.Z.S

{Received February 21, 1910.]

(Plates LXIV.-LXXIIL.*)

Introduction.

In a paper published in the Society's ‘Proceedings’ (1909, p. 31),

T have given an account of the Copepoda found in the samples

taken by De. Cunnington, during the Third Tanganyika Expedi-

tion, from the three large Central African lakes, and I mentioned in

this paper that a number of Ostracoda also occurred in the samples,

some of the species having already been observed by Dr. Cun-

nington, and separated by him in small tubes. I have now

examined the material more closely, and give in the present paper

an account of the species accompanied by carefully drawn figures

of all of them. The number of species determined amounts to

no less than 29 in all, belonging to 7 different genera. Only two

of these species I have been enabled to identify with previously

described forms; all the others seem to be new to science. In

the following pages I give short descriptions of all the species,

with remarks on affinity and occurrence, and at the close of the

paper some general remarks are added.

Description of the Species.

Family CyPRrip&.

Genus 1. PARAcypRIA, gen. n.

Generic Characters.-_Shell compressed, of various form in the

different species, in some cases approaching that in Cypria, in

other cases very different, being, as a rule, very thin and pellucid,

so as to allow the enclosed animal to be rather distinctly traced

through its walls. Valves more or less unequal, the right one

being the smaller and in some cases conspicuously overlapped

by the left both in front and behind. Surface of shell more

generally smooth and only very sparingly hairy at each extremity.

Limbs on the whole built upon the same type as in Cypria (see

the detail-figures given on Pl. LXV. of one of the species).

Caudal vami comparatively short and more or less curved, tapering

distally, dorsal edge smooth; terminal claws of moderate size ;

dorsal seta generally well developed, apical seta very small.

Inferior antennze of male with the third joint distinctly sub-

divided, Copulatory appendages each terminating in a beak-like

extremity consisting of two jaws, one of which is mobile.

* Wor explanation of the Plates ese p. 74

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19 10.] OSTRACODA FROM TANGANYIKA, 733

Bjaculatory tubes each with eight whorls of chitinous stripes, both

extremities conspicuously dilated and encircled by a row of coarset

spiniform processes.

Remarks.—The type of this new genus is the form described

by the present author from a brackish -water lagoon on the

Chatham Islands under the name Paracypris tenuis*. It was,

indeed, only with great doubt that I provisionally referred this

form to the marine genus Paracypris G. O. Sars. I now find

that it is much more closly related to the genus Cypria Zencker,

for which reason I propose to name the new genus Paracypria.

The species from the Chatham Islands is closely allied to some

of the Tanganyika species, as will be shown below. No less than

twelve different species referable to this genus have been found

in the samples.

1. PARACYPRIA DECLIVIS, sp.n. (Plate LXIV. figs. 1-8.)

Specitie Characters.— FemMaue, Shell moderately compressed ;

seen laterally (fig. 1) of oblong cuneiform shape, greatest height

occurring somewhat in frent “of the middle and not attaining

half the. length, anterior extremity obtusely rounded, posterior

much narrower and ending below in a somewhat conical blunt

corner, dorsal margin strongly curved in the middle and sloping

evenly behind to the posterior corner, ventral margin nearly

perfectly straight; seen dorsally (fig. 2) narrow oblong in form,

with the greatest width about equalling one-third of the length,

both extremities acuminate, side-edges nearly parallel in the

middle. Valves only slightly unequal ard very thin and pellucid,

with very slight trace of hairs at the extremities. Surface of

shell quite smooth and shining, without any distinct sculpturing.

Limbs (figs. 4, 5), on the whole, very like those in the type

species ; the slender dactylus of the anterior legs (fig. 4), how-

ever, quite smooth, without any trace of lateral denticles.

Caudal rami (fig. 6) shghtly curved at the base; claws 1ather

strong and perfectly smooth, the distal one much the larger and

exceeding half the length of the ramus; dorsal seta small, but

distinct.

MALE (fig. 3) a little smaller than female, and haying the shell

somewhat less vaulted dorsally. Prehensile palps of posterior

maxill (figs. 7, 8), as usual, somewhat unequal on left and right

sides, and apparently differing a little in shape from those in the

type species.

Length of shell 2 0°91 mm., height 0°42 mm., width 0°30 mm.

Remarks.—This form. is closely allied to ae type species,

exhibiting a very similar shape of the shell. It is, however, of

considerably larger size, and moreover differs in the less com-

pressed shell and in the more robust and quite smooth caudal

claws. Nor is there any trace of the fine denticles found in

P. tenuis on the dactylus of the anterior legs.

* Zool. Jahrbiicher, vol. xxi. pt. 4, p. 404, pl. xx. figs, 173-186. . ¥,

Proc. Zoot. Soc.—1910, No. XLVIII. 48

734 PROF. G, 0, SARS ON | May 3,

Occurrence.—Solitary specimens of this fine species were found

in three different samples from Tanganyika, One of these samples

(88) was taken 7/9/04 at Niamkolo (8. end of lake), another (97)

19/9/04 at about the same place, and the third (138) 13/10/04 at

Sumbu (8.W. of lake).

2. PARACYPRIA COMPLANATA, sp.n. (Plate LXIV. figs. 9, 10.)

Specific Characters—Maur. Shell highly compressed ; seen

laterally (fig. 9) oval reniform in shape, greatest height occurring

about in the middle and equalling half the length, anterior

extremity obliquely rounded and somewhat deflexed, posterior

narrower and obtuse at the end, dorsal margin almost angularly

curved in the middle and sloping behind with a slight curve to

the posterior extremity, ventral margin slightly concave in the

middle; seen dorsally (fig. 10) very narrow, with the greatest

width scarcely exceeding one-quarter of the length. Valves

rather pellucid and somewhat more unequal than in the preceding

species, the right one being conspicuously overlapped by the left

at the anterior extremity below. Surface of shell with scattered

dots and finely hairy at both extremities.

Length of shell 0°96 mm., height 0°49 mm., width 0°24 mm.

Remarks.—This form is of still larger size than the preceding

one, from which it moreover differs conspicuously in the form

and sculpture of the shell. The structure of the several appen-

dages could not be sufficiently examined in the solitary specimen

found, but no essential difference in this respect from that in the

preceding species was to be traced.

Occurrence.—Only a single specimen of this form, a_ fully

grown male, has been secured. It was found in a sample from

Tangs anyika ‘(242) taken 1/3/05 at Kaboge (N.W. shore of the

lake).

3. PARACYPRIA CONOIDEA, Ssp.n. (Plate LXIV. figs. 11, 12.)

Specific Characters—MAtr. Shell rather compressed; seen

laterally (fig. 11) of a narrow, somewhat conoid shape, greatest

height occurring far in front and about equalling half the length,

anterior extremity evenly rounded and much broader than the

posterior, which is conically tapered, though obtuse at the end,

dorsal margin gently curved and sloping evenly behind to the

pee corner, ventral margin nearly straight; seen dorsally

(fig. 12) narrow ‘oblong in fot, with the greatest width about

equalling one-third of the length, both ‘extremities obtusely

acuminate. Valves moderately pellucid and rather unequal, the

right being considerably overlapped by the left along the whole

anterior extremity; lip of the latter somewhat thickened and

giving origin to a row of very delicate hairs. Surface of shell

dotted in a similar manner to that in ?. complanata.

Length of shell 0°49 mm., height 0°24 mm., width 0°15 mm.

Remarks,—This is a rather small species, being scarcely more

1910.] OSTRACODA FROM TANGANYIKA. 735

than half as large as P. complanata. The specimen examined is,

however, evidently fully grown, as proved by the well- developed

ejaculatory tubes shining through the pellucid shell. The shape

of the latter conspicuously differs from that of either of the two

preceding species.

Occurrence.—A. solitary male specimen of this form was found

in a sample from Tanganyika (138), taken 13/10/04 at Sumbu

(S.W. of lake).

4, PARACYPRIA OBTUSA, sp. n. (Plate LXV.)

Specific Characters.—Frmate. Shell moderately compressed ;

seen laterally (fig. 1) of oval or elliptical shape, greatest height

occurring about in the middle and slightly exceeding half the

length, both extremities obtusely rounded and nearly equal,

dorsal margin gently curved with trace of an angle behind the

ocular region, ventral margin very slightly concaved in the

middle; seen dorsally (fig. 2) oblong in shape, greatest wiath

somewhat exceeding one-third of the length and occurring a little

behind the middle, anterior extremity narrower and more

acuminate than the posterior. Valves rather thin and pellucid,

conspicuously unequal, the right one being overlapped by the

left along the whole anterior extremity, as also a little behind;

anterior lips of both valves bordered by a very thin and pellucid

rim. Surface of shell exhibiting a very fine longitudinal stria-

tion, only visible under high magnifying-powers (fig. 4), and

clothed at each extremity with extremely delicate hairs. Inner

coating of valves exhibiting a somewhat irregular reticula-

tion, more conspicuous at the anterior extremity (fig. 1). Limbs

(figs. 5-11) on the whole somewhat more strongly built

than in the type species, but otherwise identical in structure.

Caudal rami (fig. 12) slightly curved, with the claws moderately

strong and less unequal than in P. declivis, the distal one about

equalling half the length of the ramus ; dor. sal seta well developed

and rather slender. —

Mate (fig. 3) of about same size as female, and having the

shell very similar in shape. Sexual characters, manifested in

the structure of the inferior antenna (fig. 13), the palps of the

posterior maxille (figs. 14, 15), the copulatory appendages (fig. 16),

and the ejaculatory tubes (fig. 17), agreeing exactly with those in

other species of this genus.

Length of shell 0°63 mm., height 0°33 mm., width 0-22 mm.

Remarks.—In the general shape and sculpture of the shell,

this form exhibits a certain resemblance to the species of the genus

Cypria Zencker, and, indeed, at first I was inclined to regard it

as a member of that genus. On a closer comparison and exami-

nation of the several appendages, I find it, however, impossible

to separate this form generically from the other species here

recorded as belonging to the genus Paracypria.

Occurrence.—This species seems to be by far the commonest

of the Ostracoda of the Tanganyika Expedition. I have noted it

48*

736 : PROF. G, 0. SARS ON [May 3,

in five of the samples from Tanganyika, and in some of these

it occurred rather abundantly. The localities are as follows:

No. 82, Kituta Bay (S. end of lake), No. 97, Niamkolo

(S. end of lake), No. 138, Sumbu (8.W. of lake), No. 169, Kala

(KE. shore of lake), No. 240, Baraka (N.W. shore). In the last-

named locality it seems to have occurred in great abundance,

and a good many of the specimens captured had also been

separated from. the sample by Dr. Cunnington and placed in a

small tube.

5. PARACYPRIA DEFLEXA, sp.n. (Plate LXVI. figs. 1-6.

I g

Specific Characters.—Frmaur. Shell moderately compressed ;

seen laterally (fig. 1) of oval reniform shape, greatest height

occurring somewhat behind the middle and slightly exceeding half

the length, anterior extremity evenly rounded, posterior scarcely

narrower and obliquely deflexed, terminating in an obtusely

triangular corner, dorsal margin evenly curved with the greatest

curvature behind the middle and sloping rather steeply to the

deflexed hind corner of the shell, ventral margin very slightly

concave in the middle; seen dorsally (fig. 2) oblong-ovate in

form, with the greatest width exceeding oné-third of the length,

anterior extremity somewhat narrower than posterior. Valves

rather unequal, the right being considerably overlapped by the

left both in front and behind, anterior lip of left valve (fig. 4)

exhibiting a somewhat thickened zone crossed bynumerous grooves,

from the bottom of which delicate hairs arise. Surface of shell

smooth and shining, though exhibiting a few small dots similar

to those found in P. complanata and P. conoidea. Muscular

impressions in the centre of each valve (fig. 5) very distinct and

arranged asin the other species of this genus. Caudal rami (fig. 6)

distinctly curved; claws moderately strong and only slightly

unequal, the distal one exceeding half the length of the ramus ;

dorsal seta well developed and rather slender.

Mate (fig. 3) a little smaller than female, but exhibiting a

closely similar shape of the shell.

Length of shell 2 0°75 mm., height 0°40 mm., width 0°26 mm.

Remarks.—In external appearance this form somewhat resembles

a Candona, the shell exhibiting in the lateral aspect a rather

similar shape and having the hind corner deflexed in a similar

manner to that found in the more typical species of that genus.

It is, however, a true Paracypria, as proved by the examination

of the several appendages, which agree closely in structure with

those in P. obtusa.

Occurrence. —Of this species also a considerable number of

specimens has been secured. I have noted it in four of the

Tanganyika samples. In two of these samples (138 and 240) it

occurred vather plentifully in company with P. obtusa. In the

other two (97 and 169) only solitary specimens were found. The

localities from which the samples were taken have been recorded

above.

1910.] OSTRACODA FROM TANGANYIKA, 737

6. PARACYPRIA SUBANGULATA, sp. n. (Plate LX VI. figs. 7-11.)

Specific Characters.—Frmaue. Shell somewhat less compressed

than in the preceding species; seen laterally (fig. 7) broadly oval

in form, greatest height occurring nearly in the middle and

considerably exceeding half the length, anterior extremity

obliquely rounded and subangular below, posterior scarcely

narrower and terminating in a slightly deflexe:l obtuse corner,

dorsal margin forming a rather even curve, ventral almost

straight; seen dorsally (fig. 8) oblong-oval in shape, greatest

width occurring behind the middle and almost attaining half

the length, both extremities somewhat blunted at the end.

Valves less pellucid than in the preceding species and very

unequal, the right one being considerably overlapped by the left

both in front and behind ; : anterior lip of right valve simple and

evenly curved, that of left valve (tig. 10) forming below an

abrupt angular bend and exhibiting inside the hyaline border a

thickened zone crossed by a number of somewhat irregular

branching grooves. Surface of shell smooth and very finely

hairy at each extremity. Caudal rami (fig. 11) considerably

curved; claws of moderate size and slightly unequal, the distal

one about half the length of the ramus; dorsal seta slender and

elongated. Ends of ovarial tubes in this and the two preceding

species deflexed.

Matz (fig. 9) a little smaller than female and having the shell

somewhat less vaulted above.

Length of shell 2 0°86 mm., height 0-48 mm., width 0°39 mm.

Remarks.—This form is undoubtedly nearly related to the two

preceding species, but is of considerably larger size than either

of them, and is, moreover, easily recognized by the more vaulted

shell and especially by the peculiar angular appearance of the

anterior extremity caused by the projecting lip of the left

valve.

Occurrence.—Only three specimens of this form, two females

and one male, have come under my notice. They were found in

a sample from Tanganyika (53) taken 9/8/04 at Niamkolo (S. end

of lake).

7. PARACYPRIA RENIFORMIS, sp.n. (Plate LX VII. figs. 1-4.)

Specitic Characters.—FrMALE. Shell not much compressed ; seen

laterally (fig. 1) oblong reniform in shape, greatest height occur-

ring in the middle and not nearly attaining half the length,

anterior extremity evenly rounded, posterior somewhat narrower

and obtuse at the end, not deflexed, dorsal margin forming a

quite even curve throughout, ventral one rather deeply sinuated

in the middle; seen dorsally (fig. 2) oblong fusiform in shape,

greatest width exceeding one-third of the length and occurring

in the middle, both extremities acuminate. Valves very thin

and pellucid, only slightly unequal; left valve, however, as usual,

overlapping the right a little along the anterior extremity ; lips

738 PROF. G, 0. SARS ON [May 3,

of both valves thin and pellucid. Surface of shell quite smooth,

without any trace of sculpture, but distinctly hairy at both

extremities. Eye very large and conspicuous. Anterior legs

(tig. 3) much more coarsely built than in any of the preceding

species, dactylus very strong, with falciform curve and clothed

inside in its outer part with fine spinules. Caudal rami (fig. 4)

of the structure characteristic of the genus, being slightly curved

at the base and a little tapered distally; claws rather strong and

finely spinulose inside, the distal one about half the length of the

ramus; dorsal seta well developed. Ends of ovarial tubes not

deflexed, but extending upwards along the posterior edges of the

valves (fig. 1).

Length of shell 0°61 mm., height 0°28 mm., width 0-21 mm.

Remarks.—Vhis form is easily distinguishable from any of the

preceding ones by the narrow and pronouncedly reniform shell

and by the coarsely built anterior legs. In the structure of the

other limbs I have failed to detect any essential difference from

those in the preceding species.

Oceurrence.—Only two female specimens of this form have come

under my notice. ‘They were found in a sample from Tanganyika

(169), taken 18/11/04 at Kala (EK. shore of lake).

8. PARACYPRIA HUMILIS, sp.n. (Plate LX VII. figs. 5-9.)

Specific Characters—Fermate. Shell rather compressed ; seen

laterally (fig. 5) of narrow elliptical form, greatest height oceur-

ring about in the middle and not attaining half the length, both

extremities obtusely rounded and nearly equal, dorsal margin

very slightly curved, ventral nearly straight ; seen dorsally (fig. 6)

narrow oblong in form, greatest width only slightly exceeding

one-third of the length and occurring behind the middle, both

extremities somewhat blunted at the end, the anterior narrower

than the posterior. Valves rather pellucid, though not so thin

as in P. reniformis, and only slightly unequal, Surface of shell

smooth, and finely hairy at each extremity. Anterior legs (fig. 8)

resembling in structure those in P. reniformis, being very coarsely

built, with the dactylus strong and abruptly curved at the end.

Caudal rami (fig. 9) markedly curved, claws moderately strong

and quite smooth, dorsal seta very small,

Maus (fig. 7) of about same size as female, and scarcely differing

in the shape of the shell.

Length of shell 0°48 mm., height 0°22 mm., width 0°16 mm.

Remarks.—This is a very small species and may easily be

recognized by the rather regular elliptical shape of the shell. It

is undoubtedly nearest allied to P. reniformis, exhibiting a very

similar strong development of the anterior legs.

Occurrence.—Some few specimens of this form were found in a

sample from Tanganyika (138), taken 13/10/04 at Sumbu (8.W.

of Jake), and a solitary male specimen was found in the same

<ample (169) in which P. reniformes occurred,

1910. ] OSTRACODA FROM TANGANYIKA. 739

9. PARACYPRIA OPACA, sp. n. (Plate LX VII. figs. 10-15.)

Specific Characters.—FrMALE, Shell moderately compressed ;

seen laterally (fig. 10) oval in shape, greatest height occurring

somewhat in front of the middle and exceeding half the length,

both extremities obtusely rounded, the anterior one somewhat

broader than the posterior, dorsal margin almcst straight in the

middle, with slight trace of an angle immediately behind the

ocular region, ventral margin very slightly concave in the

middle; seen dorsally (fig. 11) oblong in shape, greatest width

somewhat exceeding one-third of the length and occurring behind

the middle. Valves rather thick and more or less opaque, so as

not to allow the limbs to be traced through their walls ; left valve,

as usual, a little larger than right and overlapping it along the

whole anterior extremity, anterior lip of this valve considerably

thickened and crossed by a number of somewhat irregular grooves,

some of which divide at the end into fine stripes (fig. 13);

inner duplicatures of both valves rather broad. Surface of shell

of a dull appearance, being sculptured by numerous impressed

dots, larger and more distant in the centre of the valves (see

fig. 12), very densely crowded in the peripheral parts, where they

are partly arranged in curved lines (fig. 10). Anterior legs

(fig. 14) less strongly built than in the two preceding species,

with the dactylus rather slender and only slightly curved.

Caudal rami (fig. 15) almost straight, claws of moderate size,

dorsal seta very small.

Length of shell 0°63 mm., height 0°34 mm., width 0°22 mm.

Remarks.—By the thick opaque shell with its pronounced

sculpture, this form looks very unlike the other species of the

genus. On a closer examination of the several appendages,

however, I have failed to detect any essential difference in their

structure from that of the more typical species, and find it thus

impossible to separate this form generically.

Occurrence.—Some few female specimens of this species were

found in the same sample (138) in which P. humilis occurred.

Solitary specimens were, moreover, found in two other samples

from Tanganyika, the one (82) taken 27/8/04 in Kituta Bay,

the other (97), 19/9/04, at Niamkolo, both localities lying at the

south end of the lake.

10. PARACYPRIA CLAVIFORMIS, sp.n. (Plate LX VIII. figs. 1-10.)

Specific Characters.—Frmaue. Shell highly compressed ; seen

laterally (fig. 1) oblong claviform in shape, greatest height

occurring far in front and not attaining half the length, anterior

extremity obliquely rounded and somewhat deflexed, ‘poster ior

much narrower and exserted below to a triangular posteriorly

pointing corner, dorsal margin subangular in the ocular region

and sloping obliquely behind, being for some extent nearly

straight, ventral margin evenly concave in the middle; seen

dorsally (fig. 2) narrow fusiform in outline, greatest width not

nearly attaining one-third of the length and occurring in the

740 PROF, G. 0, SARS ON [May 3,

middle, both extremities acuminate. Valves very thin and

pellucid, nearly equal and without any visible pilosity. Surface

of shell quite smooth ; inner coating exhibiting a faint reticulation,

more obvious at the anterior extremity. Muscular impressions

in the centre of the valves densely crowded and arranged in the

usual manner (fig. 4). Limbs (figs. 5-9) on the whole built

upon the same type as in the other species. Caudal rami (fig. 4)

unusually short and broad, evenly curved and highly chitinized ;

claws rather thick and scarcely at all curved, their outer part

being clothed inside with fine spinules ; dorsal seta well developed.

Mate (fig. 3) of about same size as female, and exhibiting a

very similar shape of the shell. Sexual characters developed in

the very same manner as in the other species of the genus.

Length of shell 0°58 mm., height 0°28 mm., width 0:17 mm.

Remarks.—Vhis is a very distinct species, exhibiting a rather

characteristic shape of the shell, and also distinguished by the

coarse structure of the caudal rami. It must, however, evidently

be regarded as a member of the present genus, as proved by the

structure of the several limbs.

Occurrence.— Several specimens of this remarkable form, the

greater number of them still immature, were picked up from

the bottom-residue of a sample from Tanganyika (138), taken

13/10/04 at Sumbu (8.W. of lake). It was not found in any of

the other samples.

11. PARACYPRIA FLEXUOSA, sp. n. (Plate LX VIII. figs. 11-17.)

Specific Characters.—FemAvE. Shell highly compressed, seen

laterally (fig. 11) narrow reniform in shape, greatest height

occurring about in the middle and not nearly attaining half the

length, anterior extremity evenly rounded and conspicuously

deflexed, posterior a little narrower and obtuse at the end, dorsal

margin only slightly curved, ventral deeply sinuated in front of

the middle; seen dorsally (fig. 12) narrow oblong, greatest width

scarcely exceeding one-quarter of the length and occurring

somewhat behind the middle. Valves rather pellucid and slightly

unequal, the right one being somewhat overlapped by the left at

the inferior part of both extremities. Surface of shell smooth

and shining, without any visible pilosity, but exhibiting a few

small dots, more distinct in the anterior part. Eyes of smaller

size than in the other species and occurring close to the dorsal

margin. Muscular impressions in the centre of the valves

(fig. 14) only little differmg from those in the other species.

Superior antenne (fig. 15) with the sete: somewhat reduced in

number; natatory sete of the inferior ones apparently likewise

less fully developed than usual. Anterior legs (fig. 16) rather

slender, with the dactylus comparatively short. Caudal rami

(fig. 17) somewhat robust, of equal width throughout, and almost

straight ; claws comparatively short ; dorsal seta very small but

distinct, occurring about in the middle of the ramus.

Mace (fig. 13) agreeing with female both in size and in the

1910.] OSTRACODA FROM TANGANYIKA, 741

shape of the shell, but exhibiting the usual sexual characters,

which seem to be fully in accord with those in the other species

of the genus.

Length of shell 0°41 mm., height 0°16 mm., width 0°11 mm.

Remarks.—\ have been in some doubt about the systematic

place of this form. In the general shape and seulpturing of the

shell, as also in the apparent reduction of the natatory sete of

both pairs of antenne, it exhibits a certain resemblance to the

species of the genus Candonopsis Brady, and indeed at first I was

inclined to refer it to that genus. On a closer examination of

the two solitary specimens found, I now have been induced to

abandon this opimion, and find that this form should more

properly be referred to the genus Paracypria, though constituting

a rather anomalous species.

Occurrence.—Only two specimens of this small Ostracod, a

female and a male, have come under my notice. The female

specimen was found in a sample from Tanganyika (138), taken

13/10/04 at Sumbu (8.W. of lake). The male specimen was

found in another sample from the same lake (109), taken 28/9/04

at Mbete (8. end of lake).

12. PARACYPRIA CURTA, sp.n. (Plate LX XIII. figs. 15-17.)

Specific Characters.—Maue. Shell moderately compressed ; seen

laterally (fig. 15) of a somewhat triangular form, greatest height

occurring a little in front of the middle and exceeding half the

length, anterior extremity rather obliquely rounded, posterior

narrower and obtuse at the end, dorsal margin considerably

arched and slopiug behind with a slight curve to the posterior

corner, ventral margin slightly concave in the middle; seen

dorsally (fig. 16) narrow oblong in form, greatest width some-

what exceeding one-third of the length and occurring behind the

middle, both extremities somewhat blunted at the end. Valves

thin and pellucid, shghtly unequal, and clothed at each extremity

with delicate hairs, more densely crowded anteriorly. Surface of

shell perfectly smooth. Caudal rami (fig. 17) of quite normal

structure, claws rather slender, dorsal seta well developed.

Length of shell 0°55 mm., height 0°30 mm., width 0-20 mm.

Remarks.—This is a well-marked genuine Paracypria, some-

what resembling in shape P. complanata, but of much smaller

size and having the shell considerably more abbreviated.

Oceurrence.—A solitary specimen of this form, a fully grown

male, was found in a sample from Tanganyika (138), taken

at Sumbu (8.W. of lake).

Genus 2. Cypriporsis Brady.

Syn. Candonella Claus.

Cypridopsella Kaufman.

Remarks.—This genus was established in the year 1867 by

Prof. Brady, to comprise the three northern species, C. vidua,

742 PROF. G, 0, SARS ON [May 3,

villosa, and aculeata, and was characterized chiefly by the

imperfect development of the caudal rami. In the year 1896

Messrs. Brady and Norman separated one of these species, viz.

C. vidua, from the other two under a new genus Pionocypris, to

which they also referred the Australian form described by the

present author as Cypridopsis globulus. As pointed out by

Dr. Viavra, these two species, however, are not congeneric,

P. vidua having the caudal rami rudimentary, whereas in the

Australian species they are normally developed, proving it to

belong to the genus Cypretta of Vavra. The genus Pionocypris

therefore must be restricted to P. vidwa and allied European

species (helvetica, obesa, picta). I think we are bound to accept

the arrangement proposed by the British authors, according to

which the type of the genus C ypridopsis is not, as suggested by

Dr. Vavra and Dr. Kaufman, C. vidua, but C. alone w which was

referred by Dr. Vavra to the genus Candonella Claus, by

Dr. Kaufman to his genus Cypridopsella.

A third genus, likewise with rudimentary caudal rami, but

with reduced natatory sete on the inferior antenni, was

established by Brady under the name Potamocypris. The genus

Paracypridopsis of Kaufman is in all probability identical with

that genus. We have thus three northern generic types, all

with rudimentary caudal rami, viz. Cypr idopsis, Pionocypris, and

Potamocypris, to which recently a fourth African type, Zono-

cypris, has been added by Dr. G. W. Miiller. ‘T'wo species of the

last-named genus will be described in the sequel ; the other ten

species of the “ Cypridopsis-group” obtained by the Tanganyika

Expedition I provisionally refer to the genus Cypridopsis, as

restricted by Messrs. Brady and Norman.

13. CyprIDOPSIS OBLIQUATA, Sp. n. (Plate LXIX. figs. 1-10.)

Specific Characters.—Frmaue. Shell moderately tumid; seen

laterally (fig. 1) of oval triangular shape, greatest height occurring

a little behind the middle and exceeding half the length, both

extremities obliquely rounded and nearly equal, dorsal margin

strongly arched in the middle, ventral nearly straight; seen —

dorsally (fig. 2) oblong-ovate in form, greatest width considerably

exceeding one-third of the length and occurring a little behind

the middle, anterior extremity narrower than posterior. Valves

rather pellucid and very conspicuously unequal, the left one being

much the larger and overlapping the right, not only anteriorly, but

also along the greater part of the dorsal face, whereby the shell

assumes a peculiar oblique appearance *. Surface of shell smooth,

but rather densely hairy at each extremity. Muscular impressions

in the centre of the valves (fig. 8) somewhat differing in their

arrangement from those in the genus Paracypria. Kye large and

conspicuous. Superior antenne (fig. 4) of normal structure.

Inferior antenne (fig. 5) rather coarsely built, with the apical

* It may here be noted that in C. villosa, where a similar obliquity of the shell

occurs, it is not the left but the right valve which is the larger.

1910. ] OSTRACODA FROM TANGANYIKA. 743

claws strong and partly denticulate inside; natatory sete

issuing at some distance from the end of the second joint and

extending about as far as the claws. Mandibular palp

(fig. 6) comparatively smaller than in Paracypria, with the last

joint short and thick. Palp and masticatory lobes of anterior

maxille (fig. 7) slender and narrow. Anterior legs (fig. 8) com-

paratively large, with the dactylus strong and evenly curved.

Posterior legs (fig. 9) much smaller, and having the reflexed

apical seta quite short. Caudal rami (fig. 10) reduced to two

small juxtaposed lamella of conical form, each terminating in a

slender seta and having a small lateral hair. Ends of ovarial

tubes, as in most other Cypride, ascending along the posterior

edges of the valves (fig. 1).

Length of shell 0°53 mm., height 0°30 mm., width 0°22 mm.

Remarks.—This form may be easily distinguished from any of

the other species here recorded by the peculiar obliquity of the

shell, the left valve overlapping conspicuously the right dorsally, in

such manner that, in the lateral view of the shell (fig. 1), the line

of junction between the valves cuts off, as it were, on the right

side a narrow marginal area, extending along the greater part of

the dorsal side.

Occurrence.—Two or three female specimens of this form were

found in the same sample (138) from which Paracypria curta

was derived. Moreover, a single specimen, likewise of the female

sex, was found in another sample from Tanganyika (150), taken

29/10/04 in Vua harbour (W. shore of lake).

14. Cypripopsis sINUATA, sp.n. (Plate LXIX. figs. 11-18.)

Specific Characters —FrMALE. Shell less tumid ; seen laterally

(fig. 11) of a somewhat reniform shape, greatest height occurring

a little in front of the middle and scarcely exceeding half the

length, both extremities somewhat obliquely rounded and nearly

equal, dorsal margin evenly curved, ventral distinctly sinuated in

the middle; seen dorsally (fig. 12) oblong-oval in form, greatest

width about equalling two-fifths of the length and occurring

somewhat behind the middle, anterior extremity acuminate,

posterior more obtuse. Valves rather thin and pellucid, slightly

unequal, the left overlapping the right somewhat along the

anterior extremity, and at the lower part of the posterior, but

not dorsally. Surface of shell smooth and finely hairy at both

extremities.

Mate (fig. 13) somewhat smaller than female and _ slightly

ditfering in the shape of the shell, which is more deeply sinuated

below, and has the anterior extremity broader than the posterior

and more blunted. Inferior antenne (fig. 14) of exactly the same

structure as in the female, the third joint not being, as in the males

of Paracypria, subdivided. Prehensile palps of posterior maxille

(figs. 15 & 16) rather dissimilar on right and left sides, that of

right maxilla (fig. 16) having the claw lamella dilated. Copu-

latory appendages (fig. 17) with the terminal piece securiform in

744 _ PROF, G. 0, SARS ON [May 3

shape. Ejaculatory tubes (fig. 18) each with eight verticiis of

chitinons stripes, extremities of tube not dilated.

Length of shel! 9 0°47 mm., height 0°23 mm., width 0°19 mm.

Remarks.—This torm is evidently nearly allied to the preceding

one, but is of smaller size and differs conspicuously in the shape

of the shell, which, moreover, does not exhibit the characteristic

obliquity found in that species,

Occurrence. —Some few specimens of this form were found in

the same sample (138) in which the preceding species occurred,

and solitary specimens were found in two other samples from

Tanganyika, the one (169) taken 18/11/04 at Kala (E. shore of

lake), the other (240) 24/2/05 at Baraka (ON. W. shore).

15. CyPRiIDOPsIS CUNNINGTONI, sp. n. (Plate LXX. figs. 1-3.)

Specific Characters.—FrMAue. Shell rather compressed ; seen

laterally (fig. 1) of a somewhat trigonal form, greatest height

occurring a little in front of the middle and considerably ex-

ceeding half the length, anterior extremity rounded, posterior

much narrower and terminating below in an obtuse corner,

dorsal margin strongly arched in ‘the middle, ventral one slightly

sinuated; seen dorsally (fig. 2) narrow oblong in form, greatest

width about equalling one-third of the length and occurring in

the middle, both extremities acuminate. Vi alves rather thin and

nearly equal, finely hairy at both extremities. Surface of shell

smooth.

Mate (fig. 3) of somewhat larger size than female, and having

the shell more deeply sinu: ated ventrally, with the posterior

extremity narrower and more deflexed.

Length of shell 2 0:56 mm., height 0°32 mm., w idth 0:20 mm.

Remarks.—This is a quite genuine Cypridopsis, resembling

somewhat in shape the type species, C. villosa Jurine, but having

the shell less densely pilose and the valves much less unequal,

without any obvious obliquity of the shell. It is the largest of

the species here recorded.

Occurrence.—Two specimens only of this form, a female and a

male, have come under my notice. They were derived from a

sample (14) taken 17/6/04 in Monkey Bay, Lake Nyasa, and had

been separated by Dr. Cunnington in a small tube. I therefore

find it appropriate to name this species in honour of the dis-

tinguished conductor of the Expedition.

16, CYPRIDOPSIS PUSILLA, sp. n. (Plate LXX. figs. 4-6.)

Specific Characters. —FrMaun. Shell moderately compressed ;

seen laterally (fig. 4) onl in form, greatest height occurring a

little behind the middle and equalling about half ‘the length, both

extremities obtusely rounded and slightly deflexed, nearly equal,

dorsal margin somewhat irregularly curved, ventral almost

straight; seen dorsally (fig. 5) oblong in form, greatest width

about equalling two-fifths of the length and occurring in the

middle, both extremities somewhat blunted at the end. Valves

1910.] OSTRACODA FROM TANGANYIKA, 745

rather thin and pellucid, slightly unequal, and finely hairy at

each extremity. Surface of shell perfectly smooth.

Mats (fig. 6) somewhat smaller than female, but exhibiting a

very similar shape of the shell.

Length of shell 2 0°41 mm., height 0°21 mm., width 0°17 mm.

Remarks.—This form is allied to C. sinwata, but of rather

smaller size, and differs conspicuously in the nearly straight

ventral margin of the shell.

Occurrence.—Some few specimens of this small Ostracod were

found in a sample from Tanganyika (138), taken 13/10/04 at

Sumbu (8.W. of lake).

17, CypRIDOPSIS MONODONTA, sp. n. (Plate LXX. figs. 7-9.)

precue Characters.— MAte. Shell somewhat tumid; seen laterally

(fig. 7) almost semicircular in outline, greatest height occurring

in the middle and nearly attaining two- thirds of the length,

anterior extremity obliquely rounded, posterior blunted and some-

what deflexed, dorsal margin boldly arched, ventral nearly

straight; seen dorsally (fig. 8) subovate in form, greatest width

exceeding half the length and occurring behind the middle, both

extremities somewhat blunted at the end, the posterior one ‘being

the broader. Valves rather unequal, the ‘eft one overlapping the

right conspicuously along the whole anterior extremity, anterior

lip of this valve somewhat deflexed and having below a slight

sinus; posterior lip armed at the lower corner with a small

recurved dentiform process (fig. 9). Surface of shell smooth,

but rather densely hairy at both extremities.

Length of shell 2 0°36 mm., height 0°23 mm., width 0:20 mm.

Remarks.—This is the smallest of all the Tanganyika Ostracoda,

and is, moreover, easily recognized by the short and tumid shell

and by the peculiar dentiform process springing off from the

posterior corner of the left valve, a character which has given rise

to the specific name here pr oposed.

Occurrence. —A. solitary male specimen of this form was found

in the same sample (138) from which C. pusilla was derived.

18. CYPRIDOPSIS SERRATA, sp.n. (Plate LXX. figs. 10-12.)

Specific ee a oe Shell. moderately tumid; seen

laterally (fig. 10) oblong reniform in shape, greatest height

occurring in the middle and about equalling half the length, both

extremities obliquely rounded and somewhat defiexed, dorsal

margin evenly curved, ventral conspicuously sinuated ; seen

dorsally (fig. 11) oblong- ovate in form, greatest width about

equalling two-fifths of the length and occurring behind the middle,

both extremities acuminate. Valves rather unequal, the right

being conspicuously overlapped by the left, both anteriorly and

posteriorly ; each valve exhibiting at the posterior corner a row of

extremely delicate triangular Sonata four on right, three on

left. valve (fig. 12), Surface of shell smooth and finely hairy

at both extremities.

746 PROF. G, 0, SARS ON [May 3,

Length of shell 0°52 mm., height 0°26 mm., width 0°21 mm.

Remarks,—This form may be easily recognized by its com-

paratively narrow reniform ‘shell, and more ‘particularly by the

peculiar delicate serrations occurring on the posterior corners of

both valves, a character from which the specific name here

proposed has been derived.

Occurrence.—Of this species also only a single specimen has

been secured, a fully grown female. It was found in a sample

from Tanganyika (16 39), t taken 18/11/04 at Kala (E. shore of

Jake).

19. CypRripopsis BIDENTATA, sp. n. (Plate LX XI. figs. 1-3.)

Specifie Characters.—Frmate. Shell moderately tumid ; seen

laterally (fig. 1) rounded triangular in shape, greatest height

occurring in the middle and considerably exceeding half the

length, both extremities obliquely rounded and somew hat deflexed,

the posterior one more blunted than the anterior, dorsal margin

boldly arched and sloping more steeply to the anterior than to

the posterior extremity, ventral margin nearly straight; seen

dorsally (fig. 2) sabovate in form, ereatest width ahoub half the

length and occurring behind the: middle, anterior extremity

narrower than posterior. Valves slightly unequal, the left over-

lapping the right along the anterior extremity, as also at the

lower part of the posterior one, hind corner of this valve armed

with two recurved dentiform processes (fig. 3) similar to that

found in C. monodonta. Surface of shell smooth, but rather

densely hairy at each extremity.

Mate of about same size as female and exhibiting a similar

shape and armature of the shell.

Length of shell 0°49 mm., height 0°29 mm.,, width 0°23 mm.

Remarks.—This form somewhat resembles 0. monodonta in the

shape and armature of the shell, but is of considerably larger size,

and is, moreover, distinguished ‘by the presence of two dentiform

processes, instead of a ‘single one, at the posterior corner of the

left valve.

Occurrence.—Two specimens of this form, a male and a female,

were found in a sample from Tanganyika (97), taken 19/9/04 at

Niamkolo (8. end of lake).

20, CYPRIDOPSIS TUMIDULA, sp. n. (Plate LX-XT, figs. 4-6.)

Specific Characters.—FrMaAe. Shell unusually tumid; seen

laterally (fig. 4) of oval reniform shape, greatest height occurring

somewhat in front of the middle and scarcely exceeding half the

length; anterior extremity obliquely rounded, posterior scarcely

narrower and blunted at the end; dorsal margin somewhat

irregularly curved, sloping more steeply in front than behind,

ventral margin distine tly concave in the middle: seen dorsally

(fig. 5) ovate in form, greatest height exceeding half the length and

occurring about in the middle, anterior extremity narrower than

posterior, Valves rather unequal, the left one considerably over-

1910.] OSTRACODA FROM TANGANYIKA, 747

lapping the right along the anterior extremity as also somewhat

along the lower part of the posterior, hind corner of this valve

armed with two delicate triangular teeth, that of right valve

with a single such tooth placed somewhat farther above (fig. 6).

Surface of shell smooth, but rather densely hairy at both ex-

tremities.

Length of shell 0°47 mm., height 0°24 mm., width 0°25 mm.

Remarks.—This form may at once be distinguished from any

of the other species here recorded by the unusually tumid shell,

the width of which even somewhat exceeds the height. The

armature of the hind corner of each valve is also characteristic.

Occurrence.—A._ solitary female specimen of this form was

found in a sample from Tanganyika (138), taken 13/10/04 at

Sumbu (8.W. of lake).

21. CypRIpoPsis CONGENERA, sp.n. (Plate LX XT. figs. 7-9.)

Specific Characters.—FrmMaeE. Shell moderately tumid; seen

laterally (fig. 7) of oval reniform shape, greatest height occurring

about in the middle and slightly exceeding half the length, both ex-

tremities rounded and slightly deflexed, dorsal margin irregularly

curved, with trace of an angle behind the ocular region, ventral

margin very slightly concave in the middle ; seen dor ally (fig. 8)

oblong- -ovate in form, greatest width occurring behind the middle

and not attaining half the length, anterior extremity acuminate,

posterior blunted. Valves slightly unequal, left valve overlapping

the right somewhat along the anterior extremity, but scarcely

behind. Surface of shell smooth and finely hairy at both

extremities.

Mate (fig. 9) a little smaller than female and having the

posterior extremity of the shell somewhat narrower.

Length of shell 0°52 mm. ., height 0°28 mm., width 0°23 mm,

inean ‘ks.—This form is closely allied to C. sinuata, exhibiting

a rather similar shape of the shell. It is, however, of larger size

and has the ventral margin of the shell mock less Sinuated:

Occurrence.—Three specimens of this form, two females and one

male, were found in the same sample from which C. tumidula

vas derived.

2. CYPRIDOPSIS GIBBA, sp.n. (Plate LX XI. figs. 10-12.)

Specific Characters.— FumauE. Shell somewhat compressed ; seen

laterally (fig. 10) of oval trigonal shape, greatest height occurring

about in the middle and “exceeding half the length, anterior

extremity broadly rounded, posterior somewhat narrower and

blunted at the end; dorsal margin forming in the middle an

abrupt angular curve and sloping from thence evenly to each

extremity, ventral margin exhibiting in the middle a distinct

sinus, in front of which is a slight convexity: seen dorsally

(fig. 11) oblong-ovate in shape, greatest width not nearly attaining

half the length and occurring about in the middle, anterior

748 PROF. G. 0. SARS ON {May 3,

extremity acuminate, posterior blunted. Valves rather thin and

pellucid, slightly unequal, the left overlapping the right in front

and partly : ales behind by a narrow hyaline border. Surface of

shell smooth with only very slight traces of pilosity.

MALE (fig. 12) of nearly same size as female and scarcely

differing in the shape of the shell. Sexual characters, however,

easily observable in the dense coils of spermatic tubes occurring

in the posterior part of each valve, and in the translucent ejacu-

latory tubes. The latter with numerous (up to 14) whorls of

densely crowded chitinous stripes. Copulatory appendages of a

sinilar structure to that in the male of C. sinwata, but with the

terminal pieces distinctly bilobed.

Length of shell 0°60 mm., height 0°34 mm., width 0°24 mm.

Remarks. 1 m, like (. cunningtoni, is undoubtedly a

genuine Cypr idopsis. It differs from that species, as also from

the two African species described by Dr. Vavra and Dr. Miiller,

in the shape of the shell and in the imperfect pilosity of the same.

Occurrence.—Some few specimens of this form were found in a

sample from Victoria Nyanza (269), taken 25/4/05 at Bukoba

(W. shore of lake).

Genus 3. Zonocypris Miller.

Remarks.—This genus was established in the year 1898 by

Dr. G. W. Miller to comprise two African species, 7. madagascari-

ensis and Z. elegans, both of which were distinguished by a

very peculiar sculpture of the shell, a character which has given

rise to the generic name proposed by Dr. Miiller. The genus is

nearly allied to Cypridop still more to Pionocypris, but

differs in some points from . h these genera, and can well be

retained. Two species referable to this genus are described

below.

Zonocypnris cosrara Vévra. (Plate LX XII. figs. 1-3.)

Cypridopsis costata Vivra, Die Siisswasser Ostracoden Deutsch-

Ost-Afrikas, p. 10, figs. 1-8.

Syn.: Zonocypris madagascariensis Miiller.

Specific Characters. —F rman. Shell very tumid ; seen laterally

(fig. 1) of a somewhat rhomboidal shape, greatest height occurring

a little in front of the middle and slightly exceeding half the

length, anterior ¢ xtremity rounded, posterior obliquely truncated,

dorsal margin forming a rather even curve, ventral margin slightly

sinuated in the middle; seen dorsally (fig. 2) broadly ‘oviform in

shape, greatest width exceeding two-thirds of the length and

occurring behind the middle, anterior extremity much narrower

than posterior. Valves slightly unequ: al, the left overlapping ibis

right somewhat along the anterior extremity, as also a little a

the upper corner of the posterior one ; anterior lips of both valves

somewhat thickened, but with only slight trace of hairs. Surface

of shell sculptured with very conspicuous elevated ridges arranged

1910.] OSTRACODA FROM TANGANYIKA. 749

ina concentric manner. Inferior antenne rather coarsely built,

with one of the terminal claws knife-shaped and denticulated

along the inner edge (fig. 3). The other limbs resembling in

structure those in Cypridopsis.

Mate of about same size as female, and exhibiting a very

similar shape and sculpture of the shell.

Length of shell 0°60 mm., height 0°32 mm., width 0°39 mm.

Remarks.—I cannot see any essential difference between the

form deseribed by Dr. Miiller as the type of this genus, Z. mada-

gascariensis, and the form recorded by Dr. Vavra as Cypridopsis

costata. Both these forms seem indeed to be identical, and as the

specific name proposed by Dr. Vavra is the oldex one, it ought to

be retained for the present species. ,

Occurrence.—Some few specimens of this form were found in a

sample from Victoria Nyanza (269), taken 25/4/05 at Bukoba

(W. shore of lake).

Distribution. — Ugogo, Hast Africa (Véera), Madagascar

(Miller).

24. ZoNOCYPRIS LEVIS, Sp. n. (Plate LX-XIT. figs. 4-7.)

Specific Characters.—FrmMatn. Shell very tumid ; seen laterally

(fig. 4) of a somewhat trigonal shape, greatest height occurring

about in the middle and considerably exceeding half the length ;

anterior extremity somewhat obliquely rounded, posterior obliquely

truncated below; dorsal margin forming in the middle an abrupt,

almost angular curve, ventral margin rather deeply sinuated in

the middle: seen dorsally (fig. 5) of regular oviform shape,

greatest width equalling two-thir’ ~ ‘the length and occurring

behind the middle, anterior extrCuiicy much narrower than the

posterior. Valves rather thin and _pellucid, nearly equal, and

clothed at each extremity with delicate hairs. Surface of shell

per “ec‘ly smooth, without any trace of the concentric ridges found

in the preceding species. Muscular inipressions in the centre of

each valve (fig. 6) densely crowded and somewhat differing in

form and arrangement from those in the two preceding genera.

Inferior antenn of much the same structure as in Z. costata. one

of the apical claws exhibiting a very similar knife-like shape and

coarse denticulation of the inner edge (fig. 7).

Length of shell 0°58 mm., height 0°33 mm., width 0-37 mim.

remarks.—The above-described form is unquestionably con-

generic with the preceding species, agreeing with it in all

essential structural details. Yet it differs very conspicuously in

even the character from which the present genus has been named,

the surface of the shell being perfectly smooth, without the

slightest trace of any sculpturing. ‘The generic name proposed

by Dr. Miiller, of course, is somewhat inappropriate, since it does

not apply to all the species.

Occurrence.—Some specimens of this form, most of them not

ina good state of preservation, were found together with Z. costata

in the above-mentioned sample from Victoria Nyanzi.

Proc. Zoou, Soc.—1910, No. XLIX. 49

750 PROF, G. 0, SARS ON [May 3,

Genus 4. Srenocypris G. O. Sars.

Remarks.—The type of this genus is S. malcolisoni Brady, of

which a detailed description has been given by the present author

from Australian specimens. The most prominent character dis-

tinguishing this genus is unquestionably the peculiar asymmetry

of the caudal rami, the right one of which is much coarser than

the left and armed along the dorsal edge with a comb-like series of

strong denticles, wanting or only slightly indicated on the left ramus.

This character is found in several ‘other species recently described

by Dr. Vavra and Dr, Miiller, and may thus prove to be a very

good generic criterion. The form recorded by the present author

as Stenocypris chevreuxi, in which such an asymmetry does not

exist, I am now inclined to refer to the genus Herpetocypris, in

spite of the presence in this form of distinct, though very short

natatory setz on the inferior antenne. In all other characters

this form exhibits a much closer relationship to the genus

Herpetocypris than to Stenocypris. A genuine species of the

present genus is described below.

. STENOCYPRIS PERARMATA Brady. (Plate LX XIi. figs, 8-10.)

Wen ae perarmata Brady, Entomostraca collected in Natal

by Mr. J. Gibson, Proc. Zool. Soe, Lond, 1904, vol. 11. p. 126,

pl. viii. figs. 50-57.

Specific Characters.—FrmMaALe. Shell much compressed; seen

laterally (fig. 8) of elongated siliquose shape, greatest height

occurring behind the middle and only slightly exceeding one- third

of the length ; ; anterior extremity somew hat obliquely rounded,

posterior narrower and obtuse at the end, none of the extremities

deflexed ; dorsal margin gently curved, sloping evenly towards the

anterior extremity, more steeply towards the posterior, above

which there is trace of a slight concavity ; vential margin ex-

hibiting in front of the middle a slight sinus bounded anteriorly

(in the oral region) by a small convexity : seen dorsally (fig. 9)

narrow oblong m form, with the greatest width only slightly

exceeding one-quarter of the length, both extremities acuminate.

Valves thin and pellucid, only slightly unequal, the left over-

lapping a little the might along the anterior extremity and also

somewhat behind. Surface of shell smooth, but rather densely

hairy both at the extremities and below. Caudal rami (fig. 10)

exhibiting the structure characteristic of the genus ; right ramus

conspicuously broader than the left and armed on the distal part

of the dorsal edge with a row of about eight coarse teeth, followed

bya number of smaller and somewhat unequal denticles ; dorsal edge

of left ramus with only very small hair-like denticles in its outer-

most part; apical claws of both rami rather strong and somewhat

unequal, the distal one being much the longer, though scarcely

exceeding half the length of the ramus ; both claws coarsely

denticulated along the greater part of the concave edge, seta of

1910.] OSTRACODA FROM TANGANYIKA. 751

dorsal edge wanting, apical seta rather slender, extending almost

as far as tlie distal claw.

Length of shell 1:48 mm., height 0°60 mm., width 0°41 mm,

Remarks.—\ think I am right in identifying the present form

with that described by Brady under the above name, though

some small differences as to the shape of the shell may be found

ou comparing the figures here given with those reprodueed by

Prof. Brady.

Occurrence.—A nearly adult female specimen (that here figured)

of this form had been mounted by Dr, Cunnington on a slide,

together with some other Entomostraca, According to the label,

they were derived from amongst algze taken 23/6/ 04 in Nkata Bay

on the west coast of Lake Nyasa. Another somewhat smaller

specimen of the same species, likewise mounted, was found in

Tanganyika, the exact locality not being stated.

Distribution.— Natal (Brady),

Genus 5. Hererocyeris Claus.

Remarks.—According to Dr. Vavra, a genus with the above

name was established by Claus in. the year 1892, to include

the well-known northern species, Cypris incengruens Ranmdohr.

Dr, Vavra considers this genus as constituting only a subsection

of the genus Cyprinotus “Brady. As, however, both these sub-

sections : are rather well defined and comprise at present each a

number of species, I find it more appropriate to maintain the

genus in Claus’ form. A third closely allied genus is the one

established by the present author as Hemicypris, to include

Cypris pyxidata Moniez and allied specics. The following species

is only provisionally referred to the genus of Claus.

26. HirerocyPRis DuBIA, sp.n. (Plate LX XII. figs. 11-14.)

Specific Characters.—FrMA.e. Shell somewhat compressed ; seen

laterally (fig. 11) of oval reniform shape, greatest height occurring

behind the middle and exeeeding half the length, anterior ex-

tremity narrowly rounded, posterior much broader and obtusely

blunted ; dorsal margin sloping with an even curve to the anterior

extremity, abruptly deflexed behind, ventral margin somewhat

convex behind and ver y slightly sinuated in front of the middle:

seen dorsally (fig. 12) oblong-ov al in form, g greatest width equalling

about two-fifths of the length and occurring in the middle, both

extremities somewhat blunted. Valves very thin and pellucid,

slightly uneaues) the left one overlapping the right a little in

front and also somewhat along the posterior half of the ventral

face; lips of both valves perfectly smooth, without any trace of

the crenulation found in other species of this and allied genera.

Surface of sheil smooth and clethed at both extremities with very.

delicate hairs. Muscular impressions in the centre of each valve

(fig. 13) five in number and arranged in the usual manner

49*

752 PROF. G, 0. SARS ON | May 3,

Caudal rami (fig. 14) rather narrow, nearly straight, and slightly

tapering distally, dorsal edge perfectly smooth; apical claws of

moderate size and somewhat unequal, the distal one the longer and

about equalling in length half the ramus; dorsal seta shorter than

the proximal claw and attached at only a short distance from it,

apical seta very small.

Length of shell 0°86 mm., height 0°48 mm., width 0°36 mm.

Retains .—-In the shape of the shell, as seen laterally, this

form looks rather like the African species described by Dr. Vivra

as Cyprinotus fossulatus. It is, however, of smaller size and far

less tumid, exhibiting, moreover, no trace of the impressed dots

with which the surface of the shell in that species is sculptured,

nor have L been enabled to detect any tubercles on the edges of

the valves. In so far this form differs materially from any of the

other known species. In the structure of the several appendages,

however, as far as these could be examined in the solitary

specimen found, no essential differences could be traced.

Occurrence.—A solitary female specimen of this form was found

in a sample from Victoria Nyanza (250), taken 20/4/05 at Bukoba

(W. shove of lake).

Genus 6. Inyocyrris Brady.

Remarks.—Several northern species of this distinet genus have

been described, and also an Australian species, /. australis G. O.

Sars. In the Ostracod material from the Tanganyika Expedition

two species of this genus are represented, though ‘only by solitary

specimens. I have been unable to identify either of them with

previously deseribed species.

ILYOCYPRIS PROPINQUA, Sp. 1. (Plate LXXITTI. figs. 1-4.)

Specific Characters.—FEMALE. Shell moderately compressed ;

seen Jaterally (fig. 1) of oval reniform shape, ereatest height

occuring far in front and slightly excec ding half the length,

anterior extremity broadly rounded, posterior narrower and

obtusely truncated; dorsal margin nearly straight and obliquely

sloping behind, forming with the posterior margin an obtuse

angle, ventral margin deeply sinuated in the middle: seen dorsally

(fiz. 2) oblong-ovate in form, projecting on each side behind the

mic ile to an Obie protuberane e, another smaller rounded promi-

nence occurring on each side in front of the middle. Valves only

alightly pellue sid and somewhat uneque al, the right overlapping the

left in front by a narrow hyaline rim, inl a somewhat along

the posterior part of the ventral face. Surface of shell sculptured

with densely crowded 1 impr ‘esse(l dots and provided with scattered

stiff hairs, more conspicuous at each extremity. Muscular im-

pressions In the centre of each valve (fig. 3) densely crowded and

somewhat differing in number and arrangement from those inthe

other genera, rauel i} vami (fig. 4) rather narrow and con-

pie suously curved, dorsal edge smooth ; apical claws rather slender

1910.] OSTRACODA FROM TANGANYIKA, 753

and somewhat unequal, the distal one being much the longer and

considerably exceeding half the length of the ramus; seta of

dorsal edge well dev eloped.

Length of shell 0°94 mm., height 0°50 mm., width across the

posterior protuberances 0°40 mm,

Remarks.—This form somewhat resembles the northern species

I. gibba Ramdohr, exhibiting a rather similar shape of the

shell, which, as in that species, is. produced on each side behind

the middle to a well-marked protuberance; but these protuber-

ances are in the present species considerably broader and more

obtuse at the tip, more resembling those in the Chinese species

I. angulata G. O. Sars. The structure of the caudal rami is also

somewhat different.

Oceurrence.—A solitary female specimen of this form was

found in a sample from Lake Nyasa (8), taken 11/6/04 in

Anchorage Bay (8. end of lake).

28. Inyvocypris aura, sp.n. (Plate LX XIII. figs. 5-7.)

Specific Characters. MALE. Sheli less compressed than in the

preceding species, seen laterally (fig. 5) of a somewhat clavate shape,

eveatest height occurring far in front and exceeding three-fifths

of the length, anterior extremity very broad and amply rounded,

posterior much narrower and blunted at the end; dorsal margin

forming in the ocular region an abrupt angular bend and sloping

thence rather steeply behind, joining the posterior margin at

an obtuse angle, ventral margin deeply sinuated in the middle :

seen dorsally (fig. 6) of irregular oblong-ovate form, exhibiting,

as in the pr eceding species, on each side in front of the middle a

rounded tul ereuliform prominence, and farther behind a broad

obtuse protuberance, below which another somewhat smaller

protuberance occurs. Valves rather thin and pellucid, con-

spicuously unequal, the right one overlapping the left in front by

a rather broad hyaline rim, as also somewhat behind. Surface of

shell less distinctly dotted and clothed with scattered delicate

hairs. Caudal rami (fig. 7) comparatively shorter and stouter

than in the preceding species and less curved, apical claws nearly

equal in length, dorsal seta comparatively small.

Length of shell 0:80 mm., height 0°51 mm., width across the

posterior protuberances 0°40 mim.

Remarks.—This form is unquestionably specifically distinct

from the preceding one, differing rather conspicuously both from

this and the other known species in the shape of the shell, as also

in the structure of the caudal rami.

Occurrence.—Of this form also only a solitary specimen has

come under my notice. It had been separated by Dr. Cunnington

in a small tube, and proved, on a closer examination, to be a

nearly adult male. According to the label it was found in a

sample from Tanganyika (169) taken 18/11/04 at Kala (E. shore

of lake).

PROF, G. 0, SARS ON | May 3,

Family CyTHERID &.

Genus 7. LimyicyrHEre Brady.

Remarks.—Of this genus, which exhibits so close a relationship

to true marine Ostracoda, four or five northern species have been

described, all of which are found exclusively in freshwater ponds

or lakes. No species of this genus is, however, as yet known

from the Southern Hemisphere. In the Ostracod material of the

‘Tanganyika Expedition one species referable to this genus is

represented, and another nearly related African species is known

to me, having been found in a sample taken by Dr. Purcell from

a freshwater pond in Cape Colony.

29, LIMNICYTHERE OBTUSATA, sp. n. (Plate LX_XTIT, figs. 8-14.)

Specific Characters.— FEMALE. Shell rather tumid; seen laterally

(fig. 8) of regular oblong-oval shape, greatest height scarcely

attaining half the length, both extremities obtusely rounded and

of about same breadth; dorsal margin straight and horizontal,

forming in front, just above the eye, an abrupt angular bend,

ventral margin very slightly concave in front of the middle:

seen dorsally (fig. 9) irregularly subovate in shape, being con-

spicuously constricted in front of the middle and bulging behind,

greatest width almost attaining two-thirds of the length and

occurring in the posterior part, anterior extremity acuminate,

posterior blunted. Dorsal face of shell somewhat fornicate,

ventral flattened, the greatest convexity of the shell occurring far

below. Valves rather thin and flexible, being very little calcitied

and slightly unequal, the left valve overlapping the right a little

in front, but scarcely behind. Surface of shell of a dull appear-

ance, being distinctly reticulated, and clothed at both extremities

with stiff hairs, more densely crowded along the anterior one.

Muscular impressions in the centre of each valve (fig. 10) four in

number and arranged in a regular vertical series. Kye single,

not, as in the marine Cytheridee, double. Superior antenne

(fig. 11) rather stout, being composed of six well-defined joints,

the first two of which are much the largest and forming together a

geniculate bend ; setze of the terminal part short, partly spiniform.

Inferior antenne (fig. 12) 4-articulate, without any trace of

natatory sete, but with a strong deflexed, biarticulate spine

issuing from the end of the basal joint and extending beyond the

apical claws, the latter comparatively short and only three in

number. Mandibles and maxille of normal structure. The

three pairs of legs (fig. 13) gradually increasing in length and of

comparatively simple structure, each carrying on the end a strong

claw, that of last pair, however, more slender. Caudal lobes

(fig. 14) rather broad, and each provided with four thick and

densely ciliated sete.

Length of shell 0°60 mm., height 0°30 mm., width 0°38 mm.

Remarks.-—This form differs conspicuously in the shape of the

shell from any of the northern species, and more resembles in this

1910.] OSTRACODA FROM TANGANYIKA, 755

respect the South American form recently described by Dr. Daday

as Cypridinella tlosvagi. ‘The structure of the legs and the caudal

lobes is, however, very different, and on the whole in accord with

that found in the species of the present genus.

Occurrence.—Some few female specimens of this form, most of

them not in a good state of preservation, were found in a sample

from Victoria Ny anza (269), taken 25/4/05 at Bukoba (W. shore

of lake).

GENERAL REMARKS.

The occurrence of Ostracoda in the samples taken by

Dr. Cunnington during the Tanganyika Expedition must on the

whole be regarded as a very unexpected circumstance, since these

samples, as a rule, were all taken at the very surface vy the aid

of a fine-meshed tow-net. None of the known freshwater

Ostracoda are, however, limnetic in habits, such as is the case with

many of the Copepoda and Cladocera. They all are true bottom

animals, though some of them are enabled, by the aid of the nata-

tory sete attached to the antenne, to move freely in the water to

some extent. The occurrence of specimens of this group in the

samples may therefore be regarded as quite accidental, chiefly

caused by the circumstance that in some cases the samples have

been taken in quite shallow parts of the lakes, whereby some

parcels of the bottom material have happened to be introduced

in the tow-net*. The great number of species determined is still

more perplexing and could easily lead to the wrong supposition

that the Ostracod material procured was a very large and exten-

sive one. This is, however, by no means the case. The material

is in reality, as to quantity, very small, only one or two of the

species being represented by a tolerably great number of indi-

viduals, all the others only by quite solitary specimens. Although

the present account, therefore, in all probability, only gives a

sight glimpse into the Ostracodous fauna of the three great

Central African lakes, it will, I think, suftice to again emphi Usize

the peculiar faunistie character of Lake Tanganyika, as compared

with the other two lakes. A glance at the annexed table of

distribution will show that the far greater number of species are

derived from that lake, and the contrast between Tanganyika and

the othe two lakes is, in this case, even more striking than in

the case of the Copepoda, no less than 22 species occurring

in Tanganyika, whereas only three species are found in Lake

Nyasa and five species in Victoria Nyanza. It will, moreover, be

seen that at the same time only a single species, Stenocypris per-

armata, occurs in two of the lakes (Tanganyika and Nyasa); in all

other cases the species of the three lakes are different. Two

* Dr. Cunnington has recently called my attention to the fact that several of the

samples, and in particular those which turned out to be most productive in Ostra-

coda, were taken during the night; and he has suggested that their occurrence in

the surface-gatherings may more properly be explained by the very probable assump-

tion that freshwater Ostracoda, like many marine bottom-crustacea, rise to the

surface after dark.

PROF,

G, O. SARS ON

Table of Distribution.

Names of Species.

Parazypria declivis G. O. S............

complanata G. O.

obtusa G.

detlexa G. OLS.

subangulata G. OLS.

reniformis G. O. S.

humilis G.

opaca G. OL S..

claviformis G.

flexuosa G. o

curta G. O. ;

is obliquata @. 0.8.

sinuata G, O.S.

S. weeee

conoidea G. O. S. ........

ORS reat 2

One a:

cunnineton? G. OLS.

pusilla G. O. 8.

monodonta G. O. 8...

VET (EA (OD, Sawant

bidentata G. O. S....

tumidula G. O.S.

congenera G. OS.

| Tanganyika.

+te++tt+++ti+¢e4

t+t+et:

Nyasa.

Victoria.

| Nyanza. |

}

[May 3,

Other

parts of

Africa.

eibba G. O. S.

Zonocypris costata OTUQER Bes Sete : |e

p TEMES (OG) SY) canon sonaeaoe, a oe

Stenocypris perarmata Brady Sie Wek

Heterocypris dubia G.O.S. ............ Mis

Ilyocypris propinqua G. O.S...... e

aS Mh ees (ONS a

Limnicythere obtusata G. O. S.

Py ease

29 species. | 22 lets 33

of the genera here recorded are especially richly represented in

Lake Tanganyika, viz. Paracypria and Cypr idopsis. Of the first-

named genus no “Yess than twelve different species have been

determined, all exclusively peculiar to that lake and exhibiting a

vemarkable speci ialisation as regards the external appearance.

The type of this genus, P. tenwis G. O. S., as mentioned above,

was found in the very same brackish-water lagoon on the Chatham

Islands from which the type of the Copepod genus Schizopera

was derived. The suggestions set forth in my previous paper

about the probable origin of the several species of the last-named

genus occurring in Tanganyik a may therefore also apply to those

of the Ostracod genus here under question. As regards the other

genus, Cupridopsi s, it may be noted here that only two of the

ten species determined exhibit all the characteristic features of

that genus, and these two species do not occur in Tanganyika, one

of them, C. cunningtoni, being derived from Lake Nyasa, the

other, C. gibha, from Victoria Nyanza. The remaining eight

species differ somewhat in the shape of the shell and the mutual

1910.] - OSTRACODA FROM TANGANYIKA. TD

relation of the two valves, as also apparently in the structure of

the ejaculatory tubes in the male, and should perhaps more

properly be combined into a separate new genus, which, in such

cxse, would be exclusively peculiar to Lake Tanganyika.

EXPLANATION OF THE PLATES. 3

Prater LXIV.

Paracypria declivis G. O. Sars.

. Adult female, viewed from right side.

2. Same, dorsal view.

3. Adult male, seen from left side.

4. Anterior leg.

5. Posterior leg.

6. Caudal ramus.

7. Palp of left posterior maxilla of male.

8. Palp of right posterior maxilla of sane.

Paracypria complanata G, O. Sars.

Fig. 9. Adult male, viewed from right side.

bs ’ : oS

10, Same, dorsal view.

Paracypria conoidea G. O. Sars.

Fig.11. Adult male, viewed from right side.

12. Same, dorsal view.

Pirate LXV.

Poaracypria obtusa G. O. Sars.

Fig. 1. Adult female, viewed from right side.

2. Same, dorsal view.

3. Adult male, viewed from left side.

4. Piece of the central part of right valve, showing the finely striated sculpture

ot the shell and the group of muscular impressions.

5. Superior antenna.

6. Interior antenna.

7. Mandible and palp.

8. Anterior maxilla with vibratory plate.

9. Posterior maxilla.

10. Anterior leg.

11. Posterior leg.

12. Caudal ramus.

13. Inferior antenna of male.

14. Palp of right posterior maxilla of same.

15. Palp of lett posterior maxilla of same.

16. Copulatory appendage of same.

17. Ejaculatory tube of same.

Prare LXVI.

Paracypria deflexa GO, Sars.

. Adult female, viewed from right side.

. Same, dorsal view.

. Aduit male, viewed from left side.

. Anterior lips of valves, viewed from right side.

Piece of the central part of right valve, showing the musculav impressions,

Caudal ramus.

Fig.

One cy bo

Paracypria subangulata G, O, Sars.

Fig. 7. Adult female, viewed from right side,

8. Same, dorsal view.

9. Adult male, viewed from left side.

10. Anterior lips of valves, viewed from right side,

11. Caudal ramus.

PROF, G. 0. SARS ON. [May 3,

PratE LXVIT.

Paracypria reniformis G. O. Sars.

Fig. 1. Adult female, viewed from right side.

2. Same, dorsal view.

3. Anterior leg.

4. Caudal ramus.

Paracypria humilis G. O. Sars.

Fig. 5. Adult female, viewed from right side.

6. Same, dorsal view.

7. Adult male, viewed from left side.

8. Anterior leg.

9. Caudal vamus.

Paracypria opaca G, O. Sars.

Fig. 10. Adult female, viewed from right side.

11. Same, dorsal view.

12. Piece from the central part of right valve, showing the sculpture of the

shell and the muscular impressions.

13. Anterior lip of left valve, viewed from the inner side.

14. Anterior lee.

15, Caudal ramus.

Pruate LXVIII.

Paracypria claviformis G, O. Sars.

. Adult female, viewed from right side.

. Same, dorsal view.

. Adult male, viewed from left side.

. Piece from the central part of right valve, showing the muscular

impressions.

. Superior anteina.

. Inferior antenna.

. Anterior maxilla, without the vibratory plate.

Anterior leg.

. Posterior leg.

. Caudal ramus.

Vig.

Sowa

SS ONS OH

Paracypria flexuosa G. O, Sars.

FBig.11. Adult female, viewed from left side.

12. Same, dorsal view.

13. Adult male, viewed from right side.

14. Piece from the central part of vight valve, showing the muscular

impressions.

15. Superior antenna,

16. Anterior leg.

17. Caudal ramus.

Prare LXIX,

Cupridopsis obliquata G, O, Sars,

Fig. 1. Adult female, viewed from right side,

2. Same, dorsal view.

3. Muscular impressions of right valve.

4, Superior antenna.

6. Interior antenna.

6. Mandible and palp,

7. Anterior maxilla, without the vibratory plate,

8. Anterior leg.

9. Posterior leg,

10, Caudal ramus.

1910.]

Vig,

Fig.

Fig. 4

Vig. 7

Fig. 10.

ll.

12.

ane, al

Fig. 4

Fig. 7

Fig. 10

Fig. 1

11.

12.

13.

14.

15.

16.

18.

OSTRACODA FROM TANGANYIKA. 759

Cypridopsis sinuata G. O. Sars.

Adult female, viewed from right side.

Same, dorsal view.

Adult male, viewed from left side.

Inferior antenna of male.

Palp of left posterior maxilla of same.

Right posterior maxilla of same.

Copulatory appendages ot same.

Bjaculatory tube.

PruatTe LXX.

Cypridopsis cunningtoni G. O. Sars.

1. Adult female, viewed trom right side.

. Same, dorsal view.

. Adult male, viewed from left side.

Cypridopsis pusilla G. O. Sars.

. Adult female, viewed from right side.

. Same, dorsal view.

Adult male, viewed from left side.

Cypridopsis monodonta G. O. Sars.

. Adult male, viewed from right side.

. Same, dorsal view.

. Posterior corner of left valve, with the dentiform process.

Cypridopsis serrata G. O. Sars.

Adult female, viewed from right side.

Same, dorsal view.

Posterior extremity of shell, showing the peculiar marginal serrations of

both valves.

Pirate LXXI.

Cypridopsis bidentata G. O. Sars.

. Adult female, viewed from right side.

. Same, dorsal view.

. Posterior corner of left valve, with the two dentiform processes.

Cypridopsis tumidula G. O. Sars.

. Adult female, viewed from right side.

. Same, dorsal view.

Posterior extremity of shell, exhibiting the armattre of both valves,

Cypridopsis congenera G. O, Sars,

. Adult female, viewed from right side.

. Same, dorsal view.

. Adult male, viewed from left side.

Cypridopsis gibba G. O. Sars.

. Adult female, viewed from right side.

. Same, dorsal view.

. Adult male, viewed from left side.

Prate LXXII.

Zonocypris costata Vavra.

. Adult female, viewed from right side.

. Same, dorsal view.

. Extremity of left inferior antenna, showing the peculiai structure of one of

the apical claws.

760 DR. R. BROOM ON 'TRITYLODON | May 3,

Zonocypris levis G. O. Sars.

Fig. 4. Adult female, viewed from right side.

5. Same, dorsal view.

)

—e

Muscular impressions of right valve.

. Extremity of Jeft inferior antenna.

Stenocypris perarmata Brady.

Fig. 8. Adult female, viewed from right side.

9. Same, dorsal view.

10. Caudal rami.

Heterocypris dubia G, O. Sars.

Fig. il. Adult female, viewed from right side.

2. Same, dorsal view.

3. Muscular impressions of right valve.

+ Caudal ramus.

Prare LXNITI.

TIlyocypris propinqua G. O. Sars.

Fig. 1. Adult female, viewed from left side.

2. Same, dorsal view.

5. Muscular impressions of left valve.

4. Caudal ramus.

Ilyocypris alta G. O. Sars.

Fig. 5. Adult male, viewed from left side.

3. Same, dorsal view.

7. Caudal ramus.

Limnicythere obtusata G, O. Sars.

Fig. 8. Adult female, viewed from right side.

9. Same, dorsal view.

10. Muscular impressions of right valve.

11. Superior antenna.

12. Interior antenna.

13. Right series of legs.

14. Caudal lobe.

Paracypria curta G, O, Sars.

Fig. 15. Adult male, viewed from right side.

16. Same, dorsal view.

7. Caudal ramus.

6. On Tritylodon, and on the Relationships of the

Multituberculata. By R. Broom, M.D., D.Se., C.M.Z.S.

{Received March 14, 1910.)

(Text-figures 67 and 68.)

In 1884 Owen deseribed, under the name 7'ritylodon longevus,

avery remarkable imperfect skull submitted to him by Dr. Exton,

of the Bloemfontein Museum. ‘The specimen was stated to have

come from “ Thaba-chou, Basutoland,” and was believed to be a

Triassic mammal allied to the Kuropean Stereognathus. ‘There is

no locality known in Basutoland of the name Thaba-chou, but

there are several mountains called Thaba-tsueu, and it is probably

from one of these that the specimen came, In any case we now

19 10.] AND THE MULTITUBERCULATA. 761

know that the whole of Basutoland is on beds of Stormberg age ;

and as the Stormberg beds are situated above the Molteno beds,

which are known to be Rhetic, and as they contain remains of

true Crocodiles, we may safely assume that they are of Lower

Jurassic age, and not Triassic as was supposed.

The specimen, which consists of the almost perfect snout with

part of the interorbital region, portions of the malar arch, and

the complete palate, was described by Owen at some length, and

his figures have been frequently reproduced. Owen’s opinion

that Tritylodon was a mammal was supported by Cope, Osborn,

Lydekker, Seeley, and others. In 1894, however, Seeley, as the

result of his examination of the Cynodont reptiles, came to the

conclusion that 7'ritylodon was really a slightly modified Cynodont

reptile ; and this view has been followed by a number of later

writers. ,

Text-fig. 67.

~~ mle ee

~—

Side view of Snout of Tritylodon longevus Owen. Nat. size. Slightly restored.

Fr. Frontal. Ju. Jugal. Da. Lachrymal. Mx. Maxilla. Na. Nasal.

Pa. Palatine. Pmx. Premaxilla. Sma. Septomaxilla.

In 1904 I endeavoured to show that none of the arguments

advanced by Seeley in favour of 7'ritylodon being a reptile were

valid, but that in all points Zvritylodon was an undoubted

mammal.

762 DR. R. BROOM ON TRITYLODON [May 3,

While recently in London I made a careful study of the type,

which, unfortunately, is still the only specimen known, and found

that a good deal more could be made out of the original specimen

than was supposed.

In a considerable number of points my conclusions differ from

those of Owen and Seeley. The flat piece of bone on the upper

part of the snout, which both Owen and Seeley believe to be the

frontal, I consider without doubt the upper part of the nasal.

The pair of imperfect bones, which form a median ridge at the

back part of the specimen as preserved, are believed by Owen and

Seeley to be the parietals. I regard them as the anterior halves

of the frontals. The bone which Seeley believes to be the pre-

frontal I hold to be the upper part of the large lachrymal.

Text-fig. 68.

Upper view of snout of Trityloden longevus Owen. 4 nat. size.

Fr. Frontal. Ju. Jugal. Zw. Lachrymal. Ma. Maxilla. Wa. Nasal.

Pmex. Premaxilla. Smex. Septomaxilla.

Owen gives the dental formula as 72, m6. I believe there are

pretty certainly seven molars, and in front of the large incisors

there is a socket which probably hada small anterior incisor. I

therefore conclude that the formula is 73, m7. There is un-

doubted evidence that the lower molars ground with an antero-

posterior motion against the upper. The lower incisors most

prohably passed together between the large upper incisors, which

1910.] AND THE MULITTUBERCULATA. 763

I believe to be 7, and their tips probably rested against the

small 7.

Between the nasals and the premaxillaries is a pair of well-

developed septomaxillaries which have not previously been

observed. The position of these bones is seen in the figures

given of both side and upper views. ‘Lhe septomaxillary com-

pletely separates the premaxillary from the nasal and passes down

into the anterior nasal region. ‘There is no evidence that the

nares have been separated by a complete internasal process of

the premaxillaries.

When the back part of the skull and the lower jaws are restored

there is presented a singularly Rodeut-like appearance.

As regards the habits of Zritylodon, | think it most probable

that it was a root-eating animal, the large tusk-like 2nd incisors

being well adapted for digging out and breaking off the roots,

and the molars with their rows of crescentic tubercles admirably

suited for grinding.

Tritylodon is the oldest Multituberculate mammal the skull

of which is known even partly, and the only other Multituber-

culate of which much besides the lower jaw is known is one of

the latest Kocene types, Piilodus. Fortunately there has recently

been discovered and described by Gidley a good skull with the

lower jaw in position and a number of the more important bones

of the skeleton of Ptilodus gracilis.

From the study of Ptilodus, Gidley comes to the following very

interesting conclusions, which may be stated in his own words :—

“A fact of first importance is that neither in the skull nor

skeleton of the Montana specimen are there any indications of

affinities suggesting the Monotremes, while every character is

marsupial, as shown in the general arrangement and function

of the teeth and the development of the skull and skeleton. The

unequal development of the fore and hind limbs, the character

of the incisors, the form of the palate, and the position of the

cheek-teeth all indicate definite affinities with the Diprotodonts.

At the same time the reduction in numbers of the molars and

the extreme specialisation of the premolars confirms Osborn’s

conclusion regarding the philetic position of the group, namely,

that the Multituberculata may be the last representatives of

a very ancient phylum that became extinct in the early

Tertiary.

“rom the foregoing therefore it appears that the Allotheria

represent an extinct group of wmultituberculate Kutherian

mammals closely related with but not ancestral to the Diprotodont

division of the Marsupialia, with which division they may now

be classed as an Infraorder, or Superfamily, their relationship

dating back to a common ancestry somewhere in the Jurassic or

even to earlier Triassic times, as was suggested by Cope.”

While everyone must be most grateful for the great addition

made by Gidley to our knowledge of the Multitubereulate struc-

ture, I should like to suggest that I think the case is by no

764 DR. R, BROOM ON TRITYLODON [May 3,

means proven that they are at all closely allied to the Diprotodont

Marsupials. There is a little ambiguity i in his language where

he speaks of “ multituberculate euatert ian mammals ” forwiine an

Infraorder of the Marsupialia. Presumably he means “ Meta-

therian.” But it is manifest he regards the Multituberculata as

forming a division of the Marsupialia.

Let us consider in some detail the points that are urged in

favour of this conclusion.

‘“« Every character,” it is said, “is marsupial, as shown in the

general arrangement and function of the teeth and the develop-

ment of the seul and skeleton.” If by arrangement is meant

the occurrence of incisors. premolars, and molars, without canines,

the character is not more marsupial than rodent, and a somewhat

similar arrangement is found in the other Kutherian orders. But

further the number of the molars seems to me opposed to any

close association of the Multituberculates with the Marsupials.

No living marsupial has more than three premolars and almost

every one has four molars. The Multituberculates have apparently

as the normal series four premolars and three molars as in the

Eutheria. The whole seven are seen in dA/lodon, and when the

number is reduced to six as in Ptilodus there seems little doubt

that the loss is in front and not behind. From the position of

the lower teeth I should incline to the view that the large grooved

tooth in Ptiledus and the posterior of the grooved teeth in Plagi-

aulax and Ctenacodon are Ist true molars. But whatever

difference of opinion may be held on the interpretation of the

teeth, there can be little doubt that in structure the molars are

quite unlike anything known among Marsupials.

In the structure of the skull the oe points bearing on the

affinities seem to be the following:-—l. “The malar extends

backward to the glenoid surface, and apparently joms the lachry-

mal bone anteriorly as in the living Marsupials.” Though the

backward extension of the malar is doubtless a marsupii al character

it is not confined to Marsupials, being found in Procavia, Mari-

therium, and other Eutherians. It isa typical character of the

Cynodont reptiles and was doubtless present in all early mammals.

The meeting of the malar and lachrymal is another Cynodont

character which we should expect to find in any early mammal.

“The occiput extends but slightly beyond the posterior root

of the zygoma.” ‘This is a Cynodont character which is not met

with in Marsupials*. Some approach to it is seen in some early

futherians, 3. “ The nasals are relatively large and broad, ex-

panding posteriorly.” Still another Cynodont character which

is retained by most Marsupials, but also by some Kutherians, e. g.

Hystrix. 4, “The premaxillaries are short and widely separated

from the frontals by the ascending portion of the maxilla.” This

character is quite common in Eutherians. 5, “The relatively

broad, high-arched palate is perforated by two pairs of foramina,”

* But is found in Ornithorhynchus,

1910. } AND THE MULYTITUBERCULATA. # (OD

In Cynodonts the palate is relatively narrow and never perforated.

It seems probable that as mastication became established and

the palate becaine greatly widened perforation would be likely to

occur. It is still found in most Marsupials, but it is also met

with in Eutherians, e. g. Hrinacews and Macroscelides. 6. “There

appears to be an alisphenoid canal and a well-developed alisphenoid

bulla.” It is admitted that this region is crushed and broken,

and that the characters cannot be made out clearly. So far as

the figure shows, the condition is very unlike that seen in Mar-

supials, and the structure of the back of the jaw confirms this.

In neither Ptilodus nor Plagiaulax is there any distinct angle,

and in Ctenacodon the angle is very small. This is quite unlike

what is seen in Marsupials, where both in the Diprotodonts and

the Polyprotodonts a well-developed angle curves round a large

alisphenoid bulla. 7. “There is a distinct» notch on the inferior

inner border of the [occipital] condyle, a character also observed

in some of the living diprotodont Mar: supials.” The exact shape

of the occipital condyle depends much on the habits of the animal,

and most of the marsupial peculiarities seem to be paralleled by

some of the higher forms. If we knew the arrangement of the

foramina for the 1xth, xth, xith, and especially sii nerves, we

would have a point of more value in settling the affinities than

anything else connected with the skull. The lower jaw is mar-

supial in having the lower posterior portion inflected, but in the

practical absence of a distinct angle it differs considerably.

The humerus, it is stated, “is distinctly eutherian throughout,

and is very unlike that of any of the living Monotremes.” Though

in general shape it 1s very unlike the humerus of either Hchidna

or Ornithorhynchus and bears a superficial resemblance to that of

Marsupials and Eutherians, the lower end is quite unlike anything

known in any living mammal, and certainly very different from

the ordinary marsupialtype. In fact, the articular portion comes

nearer to the monotreme type than to that of any of the higher

forms.

The pelvis is too imperfect to throw much light on the affinities.

The femur and tibia so far as preserved are certainly not

marsupial in type, being very unlike those of either Diprotodonts

or Polyprotodonts *.

Doubtless all the bones both of the skull and limbs are super-

ficially more like those of Marsupials than Monotremes, but one

is too apt to forget that the only living Monotremes are de-

generate dwarfed digging animals, aud are probably much worse

types of the Prototheria than Yalpa and Chrysochloris ave of

the Eutheria.

Taking all the points into consideration, we find that there is

nothing in the posteranial skeleton that is distinctly marsupial,

* The unequal development of the fore and hind limbs is a point of no weight

in determining affinities. The condition in Macropus is probably a very recent

Re A similar development‘is met with in hopping animals of vavious

orders, e.g. Macroscelides, Pedetes, &c.

}2 60: Zoou. Soc,-—-1910, No. L, 50

766 DR. R. BROOM ON TRITYLOL ON | May 3,

and in the skull only the inflexion of the jaw and the palatal

vacuities. All the other sup posed marsupial characters are

primitive features probably present in all early mammals.

On the other hand, there are one or two characters strongly

opposed to any association of the Multituberculata with Mar-

supials. (1) While there is no evidence of any reduction in the

number of molars, there are never more than three (or according

to Gidley two). In 7'ritylodon we seem to have the full set of

four premolars and three molars. In Clenacodon, Plagiaulax,

and Ptilodus we get a progressive reduction of the premolars but

the molars remain constant. Further, the multituberculate con-

dition is quite unlike anything known in Marsupials. It may

have been an evolution from ordinary tritubercular molars, but

if Triglyphus is really Triassic then the Multituberculates appeared

much before the eaflest-known trituberculate types and may

possibly have been independently evolved from a prototherian

ancestor. (2) The presence of a well-developed septomaxillary in

Tritylodon suggests attinity with the Monotremes and is opposed

to a close rel: ationship with the Marsupials. Both Hchidna and

Ornithorhynchus, as shown by Gaupp, have large septomaxillaries.

No marsupial is known to have a septomaxillary, though a small

one appears to occur in Dasypus, as I have shown. (3) The far

backward position of the glenoid cavity is also against a marsupial

affinity and in favour of the Multituberculates being a more

pr imitive type.

In Upper Triassic time we have numerous Cynodont reptiles,

some so like mammals that it is often difficult to be sure whether

they are mammals or not. We also have the earliest-known

supposed mammals, represented by jaws with very primitive teeth.

These earliest mammals can hardly have been other than Proto-

therians, and as the Multituber culates also appear to arise in the

Triassic and 7ritylodon is certainly not later than Lower Jurassic,

it seems most likely that they are an offshoot from the early

Prototherian group. The Multituberculates can be traced right

on to Eocene times and form a well-marked group with no close

aflinities to any other Jater mammals. The Marsupials of to-day

are apparently a lately evolved group which sprang from a di-

phyodont placental ancestor probably in Cretaceous times. A

Didelphys-like form is perhaps the most primitive type, and from

it have been derived the various Polyprotodont types, including

Conolestes, which for many reasons I have long looked upon as a

slightly moditied Polyprotodont, and later on in Australia have

been evolved the Diprotodonts.

[f we are right in concluding that the Diprotodont marsupials

originated in Australia in Tertiary times from a Polyprotodont

ancestor which was itself derived in later Mesozoic times from a

diphyodont placental, it is difficult to believe that the Multi-

tuberculates which originated in Triassic times can be in any

way nearly related to them, In-the present state of our know-

ledge it seems wisest to leave the Multitulerculata as a distinet

1910.] AND THE MULTITUBERCULATA, 767

independent group with no very near affinities with the living

Monotremes, Marsupials, or Kutherians.

About fifty years ago there was much discussion between Owen

and Falconer and others as to the habits of Plagiaulax, Owen

holding that it was a carnivore, the others that it was a vege-

tarian. Since then everyone who has expressed any opinion on

the subject has sided with Falconer ; and Gidley is also opposed

to Owen. He says :—‘“ The evidence that Ptilodus and Plagiaulax

were not carnivorous in habits seems rather conclusive, but as to

whether they were insectivorous, herbivqrous, or frugivorous

there may still be some differences of opinion. IJ am inclined to

consider them as frugivorous since the incisors were well fitted

for picking small fr its or berri ies, while the large cutting blades

of the lower premolars were admirably adapted to cutting or

slicing the rinds of tough-skinned berries, or to chopping up

fleshy fruits held against the blunt- pointed premolars of the upper

jaw. For masticating the seeds of such small fruits and berries

the multituberculate molars were amply sufficient.”

It has long seemed to me that as with Z'hylacoleo so with

Plagiaulax, Owen was right in regarding it as a carnivore. The

large size of the temporal fosse for the accommodation of powerful

temporal muscles, the small extent of the grinding surface, the

eutéing mechanism of the anterior molars or premolars, pind the

pointed condition and the mode of implanting of the incisors

together with their mode of passing between the upper incisors,

all seem to me to favour a carnivorous habit. An insectivorous

habit, however, seems not improbable, and most small Carnivores

are also partly insectivorous. But a herbivorous habit for such

forms as Plagiaulax or Ptilodus seems to me very improbable,

and a frugivorous habit well-nigh impossible. Péilodus from the

great disparity of its limbs must have been a ground-hopping

animal which was probably quite unable to climb trees. Even if

it could, fruits and berries are only ripe at one or two seasons of

the year, and an animal can only be a satisfactory frugivorous

form if it can fly from place to place like the frugivorous bats or

birds. Further, typical fr ‘ugivorous forms like Pieropus seem to

require neither powerful long incisors nor large cutting molars

or premolars. But the most serious objection to Plagiaale a being

a frugivorous form is the fact that Plagiaulax lived in Jurassic

times, whereas what evidence there is, is against flowering plants

having been on the earth before Lower Cretaceous times. Conifers,

Cycads, and Gingkoes occurred before the Cretaceous, but, so far

as we know, no fruits or berries in the ordinary sense.

It may be urged that the carnivorous mechanism of Plagiaulaw

is of a different type from that of modern carnivorous mamials,

but there is this important point to bear in mind. Present-day

Carnivores almost exclusively feed on other mammals or birds or

in some cases fish. But the carnivorous mammals of Mesozoic

times had to feed probably almost exclusively on reptiles. With

the exception of Zritylodon, which was probably a root-eating

50*

768 THE SECRETARY ON ADDITIONS ''O THE MENAGERIE. | May 24,

animal, it is doubtful if we know of any other herbivorous type

of mammal in Jurassic times. ‘The long-jawed many-toothed

small forms were probably insectivorous, the short-jawed Multi-

tuberculates for the most part carnivorous. The peculiar cutting

premolars and molars of the latter probably were used for cutting

the scaly skins of lizards, and the tubercular molavs for crushing

the bones. Possibly in Eocene times they became extinct owing

to the development of the Creodonts, a better carnivorous type.

« Literature referred io.

R. Owen.—“ On the Skull and Dentition of a Triassic Mammal

(Tritylodon longevus Owen) from South Africa.” Q.J.G.5.

vol. xl. 1884, p. 146.

H. G. Seetey.— The Reputed Mammals from the Karroo Forma-

tion of Cape Colony.” Phil. Trans. 1895, p. 1025.

R. Broom.—‘‘ On the Affinities of Zritylodon.” Tr. S$. Afr. Phil.

Soc. vol. xvi. pt. 1, 1905, p. 73.

J. W. Grotey.—“ Notes on the Fossil Mammalian Genus Ptilodus,

with Descriptions of new Species.” Proc. U.S, Nat. Mus.

vol. xxxvi. No. 1689, 1909, p. 611.

May 24, 1910,

Dr. Henry Woovwarp, 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 April, 1910 :—

The number of registered additions to the Society’s Menagerie

during the month of April last was 249. Of these 148 were

acquired by presentation, 52 by purchase, 26 were received on

deposit, 9 in exchange, and 14 were born in the Gardens.

The number of departures during the same period, by deaths

and removals, was 158.

Amongst the additions special attention may be direeted to :—

1 Lar Gibbon (/Zylobates la), from Siam, presented on April

13th by Lt. Stephen St. L. Moore, R.N.

1 Tayra (Galera barbara) and 1 Kinkajou (Cercoleptes eaudi-

volvulus), from Manaos, presented on April 14th by E. Salis

Schwabe, Esq., C.M.Z.S. ‘

1 Monk-Seal (Monachus albiventer), from Madeira, presented

on April 9th by Godfrey Williams, Esq., F.Z.8.

1 Bay Duiker (Cephalophus dorsalis), 1 Zech’s Mole-Rat

(Georychus zechi), new to the Collection, and 1 Beautiful Wood-

Hawk (Dryotriorchis spectabilis), from Obuassi, S$, Ashanti,

presented ou April Ist by Capt. C. H, Armitage.

PZ. 5: 1910 Pe eae

NTIRLGA 5 Sun dence oe lat

rains =

Bale & Danielssan, ita

AUDITORY APPARATUS OF BALANOPTERA MUSCULUS.

1910.] THE ANATOMY AND BIOLOGY OF THE LARGER CETACEA. 769

1 Tasmanian Wolf (Zhylactnus cynocephalus), from Tasmania,

purchased on April 18th.

2 Bearded Vultures (Gypaétus barbatus), from Europe, pur-

chased on April 9th.

4 Mortier’s Water-Hens (Z'ribonyx mortier?), from Tasmania,

purchased on April 18th.

Mr. Ernest Gibson, F.Z.S8., exhibited a skin of Felis geoffroyi

and made some remarks on the distribution of the animal,

The following papers were read :—

1. Observations on the Anatomy and General Biology of

some Members of the Larger Cetacea. By D. G.

Liuuiz, B.A., Hutchinson Research Student of St. John’s

College, Cambridge *.

[Received March 16, 1910. ]

(Plate LX XIV.t and Text-figures 69-78 )

ContTENTS,

J. Introduction.

Il. The Species captured at Innishkea.

LiL. The Occurrence of Hairs in Whales.

IV. The Auditory Organ of Balenoptera.

V. The Asymmetry of the Odontocete Skull.

VI. Notes on the Species :—

(1) Balenoptera musculus Linneus.

(2) Balenoptera sibbaldii Gray.

(3) Physeter macrocephalus Linziweis.

VII. Miscellaneous Observations :—

(1) Locality of Captures.

(2) Diving Powers.

(3) Copulation, Period of Gestation, and Rate of Breeding.

I. [NrropuctTion.

Within the past seven years six whaling stations have been

established in various outlying parts of the British Isles. Four

stations have been opened on the mainland of Shetland, one in

North Harris in the Hebrides{, and one on the island of South

Innishkea in the west of Ireland. Mr. R. C. Haldane §, of

* Communicated by Dr. Hans Gapow, F.R.S., F.Z.8.

+ For explanation of the Plate see p. 792.

{ For an admirable short account of the history of whaling and of its recent

establishment in Scotland, see two papers by the late Thomas Southwell: Annals of

Scottish Nat. Hist. 1904, vol. xiii. p. 77; and Annals & Magazine of Nat. Hist. ser. 7,

vol. xvi. 1905. See also Lillie, Proc. Cambridge Phil. Soc. vol. xv. pt. iv. p. 347.

§ Rh. C. Haldane, Annals of Scottish Nat. Hist. 1904-1909,

770 MR. D. G. LILLIE ON THE ANATOMY [| May 24,

Lochend, Shetland, availing himself of the opportunities thus

offered, has collected and published wiuch information relating to

the larger Cetacea captured in Scottish waters since the opening

of these whaling stations,

Hitherto, our knowledge of whales has been chiefly derived

from isolated specimens stranded from time to time in various

localities around the coasts of civilised countries. Some of these,

though often in an advanced state of decay, have been examined

by zoologists, and a few observations have been added to the large

though scattered Cetacean literature. The fact that the whaling

industry has been confined to the wildest regions of the earth and

carried on under the severest conditions of physical privation has

prevented other than chance observations of this nature being made.

The establishment of whaling stations near our shores should

give a new impetus to the study of Cetology and induce competent

observers to visit the stations during the whaling seasons. It

may be hoped that new light will be thrown upon the many

obscure problems in the biology of whales before these much

hunted animals become too scarce.

During the spring of 1908 a whaling station was opened in the

west of Ireland, by the Arranmore Whaling Company, on the

island of South Innishkea off the coast of Co. Mayo; and a

certain gentleman, who wishes to remain anonymous, prompted

by considerations such as the above, came forward with a sum of

money for the encouragement of the study of the Irish Cetacea.

This fund, which was added to by others, was placed in the

hands of Dr. 8. F. Harmer, F.R.S., Keeper of Zoology in the

British Museum (Natural History), who was asked to find some-

one to visit the Irish station for as long a period as possible during

the summer of 1909 for the purpose of studying the anatomy, the

specific characters, the general biology, and other questions of

scientific interest relating to the larger Cetacea. I had the good

fortune to be chosen for the work, and accordingly went to

Jnnishkea on the 6th of July 1909 and remained there until the

26th of August, a period of seven weeks.

This visit took place about the middle of the whaling season,

which begins early in May and continues up to the end of

September. The capture of whales last year was very inter-

mittent and depended largely on the weather. During the first

part of my stay at Innishkea the weather was unusually rough

for July and only one whale was caught in four weeks, while in

the last fortnight T saw nearly thirty whales brought to shore.

The huge size of the animals makes any examination of them

very difficult, even at a whaling station with all the necessary

appliances for their dissection at hand. The length of the

specimens seen at Innishkea varied between the limits of 50 and

80 feet in a straight line. Any whale under 40 feet is, according

to the whalers, not worth shooting. The whalers have a definite

method of cutting up a whale which no doubt is best suited to

their purpose, though from the point of view of the anatomist the

1910.] AND BIOLOGY OF THE LARGER CETACEA, 771

process leaves much to be desired. The latter has to make the

best of the operations however, and it is so essential in the whaling

trade to dispose of the whales as rapidly as possible, while the oil

is fresh, that the whalers can hardly be expected to study the

needs of the anatomist. A whole animal 70 feet in length will

often disappear completely in the course of a morning.

When a number of whales are brought in at the same time, as

is often the case, the zoologist is not greatly benefited, for they

are all anchored to buoys at some little distance from the shore,

and brought to the “ flensmg slip” one by one to be stripped of

their whalebone and blubber. The ecarcases are then taken back

to the buoy to wait until they can be further dealt with. This

method of procedure is rendered necessary on account of there

being no room for more than three or four whales on the ‘ flensing

slip” at the same time.

The whales are artificially inflated with air directly they are

killed, and they: are then easily towed home by the whaling

steamer. This inflation is often a source of great imconvenience

to the anatomist, since portions of the viscera are torn away and

shot out of the body-cavity when the body-wall is pierced by the

flensing knife.

Enough has been said to show that to the scientific man a

whaling station does not pretend to olter the advantages of

unlimited time and comfort which are to be found im a laboratory.

But at a station all the largest whales, with the exception of

Lalena mysticetus and Rhachianectes glaucus, can usually be

seen within the space of three months. Very frequently several

individuals of different species can be examined and compared as

regards their external and internal characters. The material is

often sutticiently fresh for histological study, which, on account of

the gigantic size of the animals, should prove of considerable

interest. Moreover, exceptional opportunities are offered to the

naturalist of going out to sea in the whaling steamers. Many

interesting observations upon the habits of the Cetacea could

possibly be made by this means alone.

With regard to the smaller Cetaceans, since they are seldom

killed by man at present, material is ditheult to obtain ; but the

study of these animals is not at the moment so pressing as that of

the rapidly decreasing larger forms.

Before passing to consider the observations resulting from

my stay of seven weeks at Innishkea I wish to express my

thanks to the anonymous donors who made the visit possible, and

to many friends who have helped me to carry out the work.

My special thanks are due to Dr. S. F. Harmer, F.R.S.,

who has kindly read the proof-sheets, for entrusting me with

the research and for his valuable help and advice. I would

also express my hearty thanks to Mr. R. M. Barrington, F.LS.,

for his kindness in many ways and for giving me the benefit

of his local knowledge. I am under obligations to the Rev.

W. 5S. Green, C.B., Chief Inspector of Fisheries for Ireland,

772 MR. D. G, LILLVE ON THE ANATOMY | May 24,

and to Mr. E. W. L. Holt for much kindness shown to me

at Innishkea, I cannot express my thanks too warmly for the

cordial help and hospitality accorded to me by Captain Arff-

Pettersen, Mr. E. Christensen, and others at the Whaling Station.

My sincere thanks are due to Mr, A. K. Shipley, F.R.S., for many

kindnesses. ‘To Dr. W. 'T. Calman I am indebted for the deter-

mination of the Crustacea, Finally, to Dr. H. Gadow, F.R.S., I

am very deeply grateful for his valuable help with regard to

several anatomical points.

IL. Tue Species CApruURED Av [INNISHKEA.

The catch of whales at Innishkea for the opening season of

1908 numbered 77, and was as follows * :—

D Balena biscayensis Gray.

21 Balenoptera musculus Linnieus.

19 Balenoplera sibbaldii Gray.

31 Balenoptera borealis Lesson.

1 Megaptera longimana Rudolphi,

The total number of whales caught during the season of 1909

was 102. These consisted of five species. ‘The numbers and

average length of the individuals are given in the following

table :—

| | | |

‘ote - verage |»; (

Ree Total . Number A erage | Number Average

Species. number or pris. lensth | o¢ Cows, length

killed. ”) of Bale ™ of Cows. |

el b oie 4

ft. ins. } ft. ins.

Baivwena biscayensis Gray ....... ye | 4 | 43 6 | 1 43° 0

Balenoptera musculus Linneus. 56 27 58 3 | 29 60. 0

r } |

Balenoptera sibhaldii Gray. 27. WablBy #))j8d%ep LL 66 6

Balenoptera borealis Lesson .. 9 4 43 0 5 42 0

Physeter macrocephalus Linnieus 5h 5 55 7

Thus the list of species taken off Innishkea is as follows :—

Balena biseayensis Gray,

Balanoptera musculus Linneus,

Balenoptera sibbaldii Gray,

Balenoptera borealis Lesson,

Megaptera longimana Rudolphi,

Physeter macrocephalus Linneus,

* Haldane, Annals of Scottish Nat. Hist, April 1909, p, 63,

kes

1910. | AND BIOLOGY OF THE LARGER CETACEA. 773

and includes all the largest known species of whales with the

exception of Balana mysticetus Lianneus and Rhachianectes glaucus

Cope. Several smaller Cetaceans are seen off the west coast of

Ireland, but they are not killed by the whalers. These include

Balenoptera rostrata Gray, and some members of the subfamily

Ziphiine and the family Delphinide.

The specimens of Balena biscayensis Gray, killed last summer,

were taken during the first fortnight of June. This species is

said by the whalers to leave our shores after the end of June.

Balenoptera musculus Linneus was equally plentiful from the

beginning of May till the end of theseason. Lalenoptera sibbaldir

Gray was taken from the end of June till September. The

captures of Balenoptera borealis Lesson, were restricted to the

last half of May and the first half of June, the last specimen

being caught ten days before the first B. sthbaldii Gray was

taken. his whale is said to leave our shoves upon the arrival

of B. sibbaldii, The Sperm Whales (Physeter macrocephalus

Linnzeus) were captured between the middle of June and the

end of July. If detailed records of the captures of whales

continue to be kept for the Lrish stations * to compare with those

published by Mr. Haldane for the Scotch stations, we may hope,

by this means, to throw some light upon the migrations of these

creatures,

During my stay at Innishkea the following whales were

captured: 21 specimens of Balenoptera musculus Linnzeus, 15 of

which were examined; 20 individuals of 4. sibbaldii Gray, 15

of which were examined; and two male Sperm Whales (Physeter

macrocephalus Linneeus), both of which were examined, The

results obtained from the examination of these whales will now be

described,

ITI. Tur Occurrencr or Harrs In WHALES,

The distribution and significance of the scanty hairs of the

Cetacea do not appear to have been hitherto studied in the detail

they deserve. They have been vaguely referred to as occurring

on the jaws of some adult forms t. Sometimes they have been

found on the foetus only.

In two adult Sperm Whales (Physeter macrocephalus Linnzeus)

seen at Innishkea, no trace of hairs could be found on any part of

the animals even after ¢ vareful searching.

In the case, however, of the Rorquals Balenoptera musculus

Linneus, and 4. sibbaldii Gray, some 15 individuals of each

species were examined, and it was found that a definite distri-

bution of hairs could be made out in each case. Four rows of

straight, white, bristle-like hairs from half an inch to an inch in

ace occur on the dorsal surface of the beak or facial region of

* A second Irish station is apie to be opened for the season 1910.

a Knox, Proc. Roy. Soc. Edinb. i. 1833-4. Eschriclit & Reinhardt, “ Om Nord-

hvalen,” Copenhagen, 1861. Collett, Proc. Zool. Sov. 1886, p. 255. Cunningham,

Proc. Zool. Soc. 1876, p. 680, Clark, Proc. Zool. Soc. 1876. p. GRR.

774 MR. D. G. LILLIE ON THE ANATOMY | May 24,

the head (see text-fig. 69). These consist of two inner rows on

either side of the tieilien ridge which bears the blow-holes or

external nares, and two outer rows following the edges of the

beak from points just behind the blow -holes to its anterior

extremity. The average number of hairs in each row is about

eight. ‘Che hairs in each row occur at irregular intervals, but

they always keep to the above-mentioned lines of distribution

whether there ave more or less than eight hairs forming a row.

Text-fig. 69.

Dorsal surface of the beak of Balenoptera sibbaldii Gray, showing hairs.

On each side of the lower jaw (text-fig. 70) there was a row of some

five or more hairs running from the tip of the mandible along the

middle line of the outer ‘edge of each ramus to a point just in

front of the eye. At the extreme anterior end of the mandible

(see text-fig. 70), over the ligamentous junction of the rami, there

were also two vertical rows of hairs set at right angles to those last

mentioned. These oceur close together and run parallel to each

other from the upper to the under surface of each ramus, The

hairs in these two rows are placed closer together than in the

others, there being generally about fourteen hairs in each row,

The relations of the hairs to the epidermis and adjoining layers

in Balenoptera musculus Linneus, have been figured and described

by G.O. Sars in a paper published in 1865; buat further work

requires to be done in order to determine whether they possess ¢

tactile function or not,

* Says, Vidensk.-Selskabets Porhandlinger for 1866.

19105 AND BIOLOGY OF THD LARGER CETACEA, 775

On looking through the Cetacean literature there appears to be

no record of the occurrence of hairs in adult Odontocetes. It

would seem that the presence of hairs in the adult is restricted to

the Whalebone Whales; and their retention and distribution over

the beak and mandible in these forms may be due to their

possessing a tactile funetion and thus serve to indicate to the

animal the presence of its food. The small size of the Crustacea

which generally form the food of the Mystacoceti make them

difficult to see, and since the olfactory organs of the Cetacea are

very reduced, the occurrence of tactile hairs over the oral region

may be an advantage, as the small food-animals might brush

against them and thus inform the whale when to open its mouth.

Text-fig. 70.

Lateral view of the head of Balenoptera sibbaldii Gray, to show the hairs.

The top of the snout of a whale is, of course, only the prolonged

upper lip, where one would naturally look for vibrissee. In one

of the figures given by Sais in the paper above mentioned, what

appears to be a nerve is seen traversing the blubber at the base

of the hair; which would point to a tactile function, On this

theory, the food of the Odontocetes being of a larger size the

presence of tactile hairs is not so obviously required, and hence

the hairy covermg has entirely vanished in the adult forms of

these whales.

On the other hand it is also conceivable that the vibrisse,

because of their relative stoutness, should be the. last hairs to

disappear; and the fact that they have reached a further stage of

degeneration in the Odontocetes than they have in the Whalebone

Whales may be purely accidental.

IV. Tar Auprrory OrGAN OF BALANOPTERA.

The auditory apparatus in several individuals of Balenoplera

musculus and B. sibbaldii was examined with some difficulty.

The parts of the ear agreed fairly well in both these species with

776 MR. D. G. LILLIE ON THE ANATOMY [May 24,

those parts of the auditory organ, in Balwnoptera rostrata, which

were described by Drs. Carte and Macalister* in their excellent

paper on the anatomy of a member of that species published in

1867. Dr. Dwight f has also described the bones of the auditory

organ of B, musculus at some length in a paper published in 1872.

It will thus be only necessary briefly to recapitulate here the

parts of the organ which are already known and to add some

observations which appear to be new.

The opening of the external auditory meatus on the surface of

the head is a relatively small slit situated at a short distance

behind the eye in a horizontal line with the commissure of the

lips. There is no trace of a pinna, The auditory canal is con-

tinued backwards from this aperture until it reaches as far as the

zygomatic process of the squamosal bone where it turns inwards

and, increasing somewhat in diameter, proceeds along a groove in

the squamosal bone (text-fig. 71, Sq.") which winds round the

posterior border of that bone to reach the tympanic membrane.

The diameter of this tube towards its inner and wider extremity

was about one and a half inches in the two species under con-

sideration,

The tympanic membrane (Pl. LX XIV. fig. 1) seems to have

escaped the notice of previous observers. It is highly moditied and

is a sac-like structure not unlike the finger of a glove. The sac

(Pl. LX XIV. fig. 1,7) is about three inches long and three- -quarters

of an inch in diameter and tapers toa rounded point at the outer

or distal end. The walls of the sac are about one-tenth of an inch

in thickness, and consist chiefly of white fibrous tissue and yellow

elastic tissue. No nerve-cells or fibres and no muscle-fibres have

been found up to the present. From the upper surface of the sac

in the median line a ligament about an inch long and 5 mm. in

diameter projects forwards towards the tympanic cavity. The

ligament is continued along the sac in the opposite direction as a

ridge. The total length of the tympanic sac and ligament is

about four inches. The mouth of the sac opens into the: tympanic

cavity, while the outer portion projects into the external auditory

meatus. The inner extremity of the latter joins the rim of the

sac (Pl. LXXTYV. fig. 1, ,h). The ligamentous process passes under

the junction of the > malleus and inc us, and becomes attached at its

proximal end to the very much reduced manubrium of the malleus.

The malleus is fused to the tympanic bone, as has been already

pointed out by Drs. Carte and Macalister. Thus in Balenoptera

the tympanic membrane has become a sac-like organ, which

projects outwards and is attached by a ligament to a rigid process

of the tympanic bone—this process being the malleus.

In the external auditory meatus of all the individuals examined

at, Innishkea there was a solid plug of wax-like substance of fairly

definite size and shape which does not seem to have been hitherto

described. This plug (Pl. LXXIV. fig. 2) was usually about

* Carte & Macalister, Phil. Trans. 1867, p. 252.

+ Dwight, Memvirs Boston Soc. Nat. Hist. 1872, vol. ii. p. 226,

1910.) AND BIGLOGY OF THE LARGER CETACEA. tar

5 inches long and 13 inches broad at the wider end. It consisted

of a deposit of ear-wax, which formed a coat tothe outer surface of

the tympanic membrane in that region where the latter projected

into the external auditory meatus. The deposit formed a solid

mass, which could easily be detached from the tympanic membrane.

The inner portion of the plug had the form of an oval cup

(Pl. LXXIV. fig. 2, a), measuring about 1} inches across the

broadest part and approximately 1 inch deep. The outer portion,

however, formed a solid thin flattened rod about 4 inches long

and 1 inch broad (Pl. LXXIV. fig. 2,6). The auditory canal

appeared to be full of water, in which the tympanic membrane

was immersed, with the attached plug of wax lying ina horizontal

position, The total length of the membrane and plug thus projecting

into the auditory canal was about 6 inches (Pl. LXXTV. fig. 3).

The Cetacea have a remarkable depression on the base of the

cranium on each side of the median line (text-fig. 71).

Ventral view of left posterior portion of the skull of Balenoptera musculus Linneus

The tympanic bone has been removed.

Fr.=Frontal; Pa.=Palatine; Vo.=Vomer ; 4/.=Alisphenoid ; Pt.=Pterygoid ;

Pt..\=Pterygoid plate of alisphenoid; P#.2= Portion of pterygoid, forming roof

of pterygoid fossa; Sg.=Zygomatic process of squamosal ; Sg.1=Zygomatic

process of squamosal, forming roof of groove in which lies the inner portion of

the external auditory meatus with the plug of ear-wax ; a, anterior pedicle for

attachment of tympanic bone, situated on pro-otic portion of periotic; b, fenestra

ovalis, situated on the labyrinthic segment of periotic; c, posterior pedicle of

tympanic bone, situated on opisthotic portion of periotic ; Op., opisthotic portion

of periotic ; Bo., basioccipital ; Hxo., exoccipital.

In Balenoptera these depressions are bounded posteriorly by

the projecting edge of the exoccipital, externally by the base of the

zygomatic process of the squamosal, on the inner side behind by

778 MR. D. G. LILLIE ON THE ANATOMY [May 24,

the prominent edge of the basioccipital. The anterior portion of

the inner side of this depression and the front of the recess are

bounded by the pterygoid and alisphenoid bones, which are fused

together; the latter also form the roof of the anterior half of the

depression. Thus the anterior portion of the cavity is bounded

on three sides by the pterygoid and externally by the squamosal,

and is known as the pterygoid fossa.

In this recess the united tympanic and periotic* bones lie.

The latter consists of three parts : an anterior or pro-otic, a central

or labyrinthic, and a posterior or opisthotic. The anterior or

pro-otic bone resembles athree-sided pyramid with its apex pointing

forwards and projecting into a cavity in the sqguamosal bone above

the roof of the pterygoid fossa.

On the inferior surface of the pro-otic, at its posterior end, there

is a longitudinally flattened pedicle which unites with the anterior

extremity of the tympanic bone. Just behind this anterior pedicle

of the tympanic—that is, between the pro-otic portion and the

central or labyrinthie—there i is a groove running in a transverse

direction. This furrow forms the roof of the ‘auditor vy meatus.

The central or labyrinthie portion is the smallest of the three

divisions of the periotic, and it contains the vestibule, the cochlea,

and the diminutive semicircular canals. The inferior surface of

this bone is dome-shaped and forms the roof of the tympanic cavity.

The fenestra ovalis is situated in the depression which marks the

outer border of this bone on the under side. The fenestra rotunda

is placed some little distance behind the fenestra ovalis, and is

separated from the latter by a ridge which forms fe poster ior

boundary of the tympanic cavity above. ‘The fenestra rotunda is

thus on the posterior surface of the labyrinthic segment, and

thus outside the tympanic cavity. The superior surface of the

labyrinthie bone is irregular in shape and is turned towards the

cranial cavity. Two foramina open on this surface—the aque-

ductus Fallopii anteriorly and the internal auditory meatus

posteriorly. The aqueductus perforates the bone and opens at

the base of the labyrinthic segment, and is continuous with

a groove in the opisthotie division. At the posterior end of the

labyrinthie the opisthotic segment extends outwards; it is con-

stricted at its origin, but broadens and expands towards _ its

extremity between the squamosal and exoccipital bones almost at

right angles to the pro-otie portion. In fact, the labyrinthie may

be said to lie at the angle formed by the pro-otic and opisthotie.

The tympanic segment is united to the ends of these two bones

by pedicles, which are flattened in the direction of their length.

Thus the posterior pedicle is situated on the inferior surface of

the opisthotic, near its junction with the labyrinthic, and is at

right angles to the anterior ee

The tympanic bone? (PI, LXXIV. fig. 3, 7), which is attached

to the periotic as described above, is usually about five inches long

* Carte & Macalister, Phil. Trans. 1867, pl. vi. fig. 11.

+ Carte & Macalister, /oc. cif. pl. vi. fig. 10; pl. vii. fig. 6.

1910.] AND BIOLOGY OF THE LARGER CETACEA. 779

and shaped like a cowrie shell. It occupies the posterior half of

the depression, into which it fits fairly closely (text-fig. 71, a, ¢),

its outer inferior surface being level with the pr ojecting edge of

the b: asioccipital, the pterygoid fossa lying immediately in front.

The tympanic bone may be said to have an inner and an outer

surface, which meet below, ‘The inner surface is flat, its lower

edge lying near, but not touching, that of the projecting process

of the basioccipital ; ; its upper edge i is smooth and rolled into the

cavity of the bone. The outer side of the tympanic is rounded,

with the convexity pointing outwards. The upper edge of this

outer side forms an irregular extended hp, which gives rise to the

anterior and posterior pedicles. On this lip, immediate ly bebind

the anterior pedicle and at right angles to it, there is a ridge

projecting upwards and nearly touching the periotic. The inner

edge of this ridge is continuous with the processus longus of the

malleus, the mailers thus being fused to the tympanic bone.

This process, which runs parallel ‘to the posterior pedicle, together

with the latter, form the sides of the auditory meatus.

The malleus has been described by Drs. Carte and Macalister”,

and is stated by these attthors to be fused to the tympanic bone

by the handle or manubrium ; whereas Dr. Dwight f regards that

portion of the malleus which has co-ossified with the bulla as the

processus longus or gracilis. The latter would seem to be the

correct interpretation judging from the mode of attachment of

the tympanic membrane. This structure was apparently lost or

had decayed in the specimens dissected by the above-mentioned

writers, so they could only guess at the relationship of the parts.

The manubrium is reduced to a short process, slightly hooked at its

distal end (Pl. LX XIV. fig. 1, 5), on the ventr al surface of which

is attached the ligament of the tympanic membrane. ‘This is the

only attachment of the membrane to the malleus. The rim of

the tympanic membrane sae is joined to the inner extremity of

the external auditory canal. The processus longus or gracilis

(Pl. LXXIV. fig. 1,¢) is well developed, and is fused to the inner

edge of the lip of the tympanic bulla. The incus and stapes have

been described by previous writerst, and are morphologically

similar to those found in other Mammals.

At the anterior end of the tympanic cavity, in front of the

anterior pedicle of the tympanic bone, there is an opening which

communicates with the pterygoid fossa. This may be regarded as

the enlarged inner end of the Eustachian tube. The pterygoid

fossa, which measures from 6 to 8 inches in length, about 4 inches

in width, and 4 inches in depth, is alsoa portion of the Eustachian

tube, for it is limed by the same mucous membrane as the tympanic

cavity and the Kustachian tube proper. The latter isa relatively

narrow canal, about three-quarters of an inch in diameter, which

opens out of the floor of the pterygoid fossa and winds along a

* Carte & Macalister, loc. cit. p. 254.

+ Dwight, Mem. Boston Soc. Nat. Hist. 1872, vol. ii. p. 226.

{ Carte & Macalister, Joc. cit. p. 254; Dwight, loc. cit. p. 226.

780 MR. D, G. LILLIE ON THE ANATOMY | May 24,

groove on the inner side of the pterygoid plate of the alisphenoid

(text-lig.71, Pt.', p.777) to open into the alveolated nasopharyngeal

chamber near the junction of the nares. This tube is about one

foot in length in the larger specimens. The mucous membrane,

which lines the Eustachian tube and is continued into the pterygoid

fossa and tympanic cavity, lies directly on the bony walls of these

cavities and covers over the ossicles.

The cavities of the middle ear are probably filled with air through

the Eustachian tube (text-fig. 72), while the external ear appears to

be filled with water. The pressure of the water upon the tympanic

membrane when the whale dives must be considerable, and it is

Diagrammatic view of the outer and middle ear of Balenoptera.

a=external auditory meatus ; '=plug of ear-wax; c=tympanic membrane attached

to malleus by ligament; @ = imalleus fused to tympanic bone; e = incus;

f = stapes; g = fenestra ovalis; 4 = tympanic cavity; J = pterygoid fossa ;

k = Eustachian tube.

curious that the membrane should be concave on its inner side

instead of on the outer side. The sac-like membrane, being con-

nected by a ligament to the rigid malleus, cannot have a vibrating

function, The animal probably fills the middle ear with air on

coming to the surface to breathe. When the creature dives, this

air is imprisoned by the closing of the Eustachian tube. The walls

of the latter are provided with a strong sphincter muscle. The

air inthe Eustachian tube would be forced into the pterygoid fossa

and tympanic cavity when the tube was compressed, and would

inflate the sac-like tympanic membrane. This air would be

approximately at atmospheric pressure. ‘The increase of pressure

1910.] AND BIOLOGY OF THE LARGER CETACHZA, 781i

produced by the water on the external surface of the membrane

as the animal dived would tend to compress the walls of the sac.

There may be some means by which this increased pressure is

made known to the animal, and thus enable the curiously modified

tympanic membrane to serve as a pressure-gauge; but it is not

possible to assign any function to the structure at present.

The whale probably receives sound-vibrations by means of

vibrating bony surfaces, after the manner of fishes. The tympanic

bulla is a relatively dense and heavy sounding-box, fastened to the

periotic bone by two thin pedicles, so that it could be easily set in

vibration. ‘The bulla is connected with the fenestra ovalis by the

chain of ossicles, the auditory apparatus being thus independent

of the tympanic membrane, which may have some other function,

possibly that of a pressure-gauge.

The deseription of the ear of Balena mysticetus given by Home*

in 1823 seems to correspond to some extent with the above account

of that organ in Balenoptera. The ear of the Odontocetes

appears to be more like that of ordinary Mammals 7.

V. Tue ASYMMETRY OF THE ODONTOCETE SKULL.

It has long been recognized that a want of symmetry exists

in the Odontocete skull, centred round the nasal region; but

authorities seem still to be at a loss to account for this irre-

gularity. Myr. Beddardt sums up the difficulty thus :—‘ It is

easier to say that the asymmetry, being, as it is, chiefly developed

in the regions of the blow-holes, has somethiny to do with these

structures, than to find any adequate reason for connecting the

two.” From an examination of the skull alone an explana-

tion certainly does not present itself very readily. But when

the anatomy of the head and neck of a symmetrical Mysta-

cocete, such as Balenoptera musculus Linneeus, is examined,

and the relations of all the parts studied and compared with those

of an asymmetrical Odontocete, such as Physeter macrocephalus

Linneus, the problem becomes greatly simplified, and a solution

seems to offer itself.

In the Cetacea the arytenoid and epiglottidean cartilages form

a long, rigid, eylindr ical tube, a continuation of the larynx, which

rises up through the floor of the pharynx like a pillar and is thrust

up between the alveolated walls of the nasopharyngeal chamber,

which form the roof of the pharynx in this region, In the larger

whales this pipe is about one foot in length. A somewhat similar

pipe-like epiglottis, connecting the larynx with the posterior nares

also occurs in the Marsupials and Ungulates; but it is less pro-

nounced than in the Cetacea,

In the case of the symmetrical Mystacocetes this tube is

* Tome, * Lectures on Comparative Anatomy’ (1823), Vol. ili. Leot. [X., Vol. iv

Tabs. c. & ci.

+ Hunter, Phil. Trans. 1787, p. 430.

+ Beddard, ‘A Book of Whales,’ p. 49.

Proc. Zoou. Soc.--1910, No. LI. al

782 MR. D. G, LILLIE ON THE ANATOMY | May 24,

situated in the middle of the pharynx. The food of these whales,

consisting as it does of small Crustaceans, can readily enter the

comparatively narrow pharynx and find a passage down the

pharynx on each side of the pipe (see text-fig. 73).

Text-fie. 73.

Transverse section (diagrammatic) of the pharynx of a Mvstacocete.

a= epiglottis; 6 = food-passace.

In the Odontocete Physeter the pharynx is fully four or five

times as large as that of a Lalaenoptera of the same length,

and the pipe-like epiglottis, instead of being placed in "tis

middle of the pharynx, is on the left side close against the left

wall of the latter. The pipe passes up into the nasopharyngeal

chamber, which has alveolated walls; as in Lalenoptera, this

Text-fig. 74.

Transverse section (diagrammatic) of the pharynx of Physeter.

a= epiglottis; 6 = food-passage.

chamber communicates with the posterior nares situated on the

left side of the skull. The comparatively large animals eaten by

this whale, which are always sw: allowed ole! pass down the

spacious pharynx only on the right side of the pipe, which is close

up against the left wall and leaves plenty of room for their transit

(see text-fig. 74). Thus the pharynx of Physeter in the region of

1910. ] AND BIOLOGY OF TIE LARGER CUTACEA., 782

the glottis is divided into a right half for the passage of food and

a lett half for respiration. This arrangement gives an asym-

metrical form to the nasal regions of the skull.

Pouchet and Beauregard*, in their admirable memoir on the

anatomy of the Sperm Whale, refer to the pipe-like epiglottis as

being asymmetrically placed ; but they state that it was situated

on the right side of the pharynx, whereas in both the specimens

of Physeter seen at Innishkea the pipe was undoubtedly on the

left side. These authors appear to have attached no significance

to the position of the epiglottis as bearing upon the asymmetry of

the skull.

VI. Nores on THE SrEctss.

1. BALHNOPTERA MUSCULUS Linnzeus 7.

The length, form, and proportions of all the specimens seen were

in accordance with previous descriptions of the species.

Colour. There appeared to be two colour-types in this species,

distinguished by the colour of the dorsal surface. In the more

common type the colour of the dorsal surface was bluish black ;

in the other variety the dorsal surface was brownish black,

almost sepia. This fact has been noticed by Mr. Haldane, who

attributes the difference in colour to a light-effeect; but it was

sufficiently marked in all the individuals landed during my visit

for them to be placed in either one or other of the two groups.

The asymmetrical colouring of the face was remarkably constant

in all the specimens. On the right side the outer edge of the

beak was white or light grey; on the left side the beak was entirely

bluish black or sepia in colour, according to the variety which the

individual belonged to, The anterior third of the baleen plates

on the right side were invariably white, the remainder on that

side being dark grey or black on the outer halves of the plates

and becoming lighter in colour towards their inner edges. On

the left side all the baleen plates were similar to the posterior

plates of the right side. The bristles were invariably white all

round the mouth, and not black as stated by Mr. Collett $ in his

synopsis of the four northern species of Balenoptera. The lower

jaw was white on the right side, and bluish black or sepia on the

left side. The whole underside of the body, the right lower jaw,

the inner side of the paddles, and the under side of the flukes of

the tail were white in all cases. The roof of the mouth consists

of a broad oval margin of white bristles surrounding a narrow

strip of pink palate, and below, a pink tongue covered with dark

grey mottling. Every specimen had whitish oblong spots dis-

tributed over the dark parts of the body, identical, except in

* Pouchet et Beauregard, Nouvelles Archives du Muséum (Paris), 3 sér. vol. iy

1892, p. 59, pl. 111.

+ Known also as B. physalus Fabricius, B. rorgual Lacépede, Physalus anti-

quorum Gray.

~ Haldane, Annals of Scottish Nat. Hist. 1908, p. 70.

§ Collett, Proc. Zool. Soc. 1886, p. 265.

51*

784 MR. D. G. LILLIE ON THE ANATOMY [May 24,

size, with those fizured and described by Mr, Collett* as occurring

in B. borealis, he spots in B, musculus usually did not exceed

50 mm. in length.

Throat-grooves.—In several individuals of both B. musculus and

B. sibbaldii the skin lining the grooves in the throat-region had a

bright red appearance, It unfortunately did not occur to me at

the time to ascertain the cause of this; but I am inclined to

think that the red colour of the grooves may be due to their

being very highly vascular, and thus helping to aérate the blood.

The throat- -grooves of Balenoptera occupy about half of the

ventral surface of the body, extending from the anterior end of

the mandible to the navel. In B. musculus there are about 100

of these grooves, in the other species about 60. The skin lining

the furrows is extremely elastic. When unextended the grooves

are about an inch in width; when extended they are often

six inches wide. Thus they would give a large aérating surface.

Undoubtedly the principal function of these elastic furrows on

the throat is to inerease the size of the mouth-cavity, so that a

large volume of water containing Crustaceans can be taken in at

each mouthful. The water is strained off through the whalebone

plates, and afterwards the food is swallowed. In Lalena the

mouth itself is very large, owing to the arched form of the skull,

so that its capacity is “naturally ample. Hence the absence of

throat-grooves in the Right Whales. The function of the grooves

as an aérating surface or external gill, if it occurs at all, would

be a secondary one. This theory, however, would stisbis us to

understand the extraordinary powers of remaining under water

attributed to Balenoptera which will be referred to later in this

paper.

Jacobson’s organ.—On the under surface of the snout, about half

way between its extreme tip and the small anterior flamente of

whalebone, there were two small apertures leading into narrow

tubes which ended blindly about two inches from their mouths

(text-fig. 75). These were the ducts of the organ of Jacobson.

In BL. sibhaldii they are only represented by two small depressions.

In most Mammals these organs communicate with the nostrils,

but in an adult B. musculus they are separated from the nostrils

by a distance of some ten feet, and are interesting examples of the

persistence of traces of an organ after it has become obsolete. It

is just possible that the small ducts in Bb. musculus may have

some function, but they were not equally well developed in all the

individuals examined, and in one specimen they were reduced to

mere depressions very much like those of B. sibbaldii.

Testes.—In a member of this species, 51 feet long. the testes

were measured and found to be only 9 inches in length and 3 inches

in diameter. In a specimen 60 feet long the testes were 2 feet

6 inches in length and nearly a foot in diameter, ‘This shows that

male members of this species reach maturity when between 50 and

60 feet long.

* Proc. Zool. Soc. 1886, p. 240, pl. xxvi. fig. 2

1910. ] AND BIOLOGY OF THE LARGER CETACEA. 785

Pewis.—In the Cetacea the penis when not in use is capable of

being entirely withdrawn into the body-cavity. The ventral

surface of the body is thus left free from any protuberance which

would hinder the animal in swimming. The testes remain

permanently inside the body-cavity. The penis of an adult

B. musculus measures from 5 to 6 feet in length when fully

extended ; it is about a foot in diameter at the base, tapering to

a relatively fine point.

Text-fig. 75.

Ventral view of anterior extremity of the beak of Balenoptera musculus Linneus

a = hairy inner surface of baleen plates; & = external openings of the orgaiz

Jacobson.

Food.— An examination was made of the stomach-contents of

nearly all the specimens of this species caught during the months

of July and August, and it was found that only one individual,

killed on July 13th, had fed upon herrings. The alimentary

786 MR. D. G. LILLIE ON THE ANATOMY [May 24,

canals of all the others contained JMeganyctiphanes norvegica

M. Sars (formerly called Vyctiphanes norvegica). Other species

of Euphausiide may have been present but only the above

inentioned was definitely determined. In the pharynx these

Crustaceans were found practically intact ; in the stomach they

were more broken up, and in the intestines they were reduced to

a thick terra-cotta coloured fluid, the red colour being due to a

red pigment commonly found in the Euphausiide.

Futuses.—On July 31st, 1909, two female B. musculus were

killed, one was 63 feet long and contained a feetus 1 foot in length.

The other was 67 feet long and contained a feetus 53 feet in length.

This tends to support the view that the species under consideration

and possibly other species of Salenoptera have uo definite

breeding season.

Parasites.—Vhe external surfaces of all the . musculus

examined were entirely free from parasites with the exception of

the baleen plates. These were very frequently coated with the

adult forms and nauplius larvee of Balenophilus unisetws Auri-

villius. These remarkable Copepods have been found on the

baleen plates of B. borealis and B, sibbaldii*, but this appears to

be the first record of their occurrence on BL. musculus.

2. BALHNOPTERA SIBBALDII Gray (J. latirostris Flower).

The external characters of all the Innishkea specimens were in

agreement with the descriptions of the species given by previous

observer's.

The inside of the mouth, the baleen, bristles, palate and tongue

were entirely black.

The forms of vhe paddles and dorsal fins of B. sihbaldu differ

from those of B. musculus, as is shown by the diagrams (text-

figs. 76 & 77). It would seem that the presence of a dorsal fin is

restricted to the fast-swimming Cetaceans.

The contents of the alimentary canals were examined in the

case of some ten individuals of this species, caught during August,

and found to consist exclusively of JWeganyctiphanes norveyica

in all cases.

A young male member of this species, 63 feet long, was killed

while following the mother and feeding upon her milk. The

specimen had Crustaceans in its stomach, showing that it did not

depend entirely upon the mother for feod. The young of

B. sibbaldii ave said to be between 20 and 30 feet in length at

birth, so, unless the period of lactation is unusually prolonged,

this would point to a comparatively rapid growth and develop-

ment in the larger Cetacea as is maintained by Mr. Haldane f.

[f the period of lactation continued, as is generally supposed, for

one year the animal would increase in size to the extent of over

30 feet in its first year of life.

The external surfaces of all the individuals of this species were

* Collett, Proc. Zool. Soc. 1886, p. 243.

+ Haldane, Annals of Scottish Nat. Hist. April 1905, p. 69.

1910.] AND BIOLOGY OF THE LARGER CETACEA. 787

remarkably free from parasites, the only species found being

Balenophilus unisetus Aurivillius on the baleen plates. No

internal parasites could be found.

Text-fig. 76,

orsal fins of a. Balenoptera musculus Linueus; 6. Balenoptera sibbaldii Gray-

Text-fig. 77.

Paddles of a. Balenoptera musculus Linneus; 6. Balenoptera sibbaldii Gray.

788 MR. D. G. LILLIE ON THE ANATOMY [May 24,

PHyserer MACRocEPHALUS Liuneus *,

Two male Sperm Whales were captured during my visit. Their

general appearance corresponded with the descriptions of previous

observers. But, as there seems to be a little uncertainty concern-

ing the shape of the head of this whale, a sketch is given (text-

fig. 78) of the form of the head in the Innishkea specimens.

The upper part of the animal was black, the ventral surface and

lower jaw a light bluish grey. A white oval patch cecurred in

the middle of the ventral surface, extending from a point just an

front of the navel to the anus.

Text-fig. 78.

View of the left side of the head of Piyseler macrocephalus Litneus,

On the surface of the head of one of the specimens, on the left

side immediately over the position of the nasal bone of the skull,

there was a small groove-like depression about two inches deep

and five inches long. This slit was placed longitudinally and in

a straight line with the blow-hole. It may possibly have been a

vestige of an old nasal opening which was situated further back

than “the present position of the blow-hole.

One of these whales had 24 teeth on each side of the lower jaw.

Six of those on the right and eight on the left side were broken.

No sign of teeth could be found in the upper jaw.

The alimentary canals of the two Sperm Whales contained the

remains of cuttlefish, and the marks of their horny denticulate

suckers could be seen all over the surface of the lower jaw and

extending more than half-way up the head of each whale, from

its anterior extremity to the paddles. The marks of the suckers

varied from faintly dotted circular imprints, formed by the slight

* Probably synonymous with P. catodon Fabricius; P. gibbosus Schieber ;

P. trumpo Gerard; P. polyclystus Couch; Catodon australis Mac Leay; C. colneti

Gray; P. polye. yphus Quoy and Gaimard,

1919.] AND BIOLOGY OF THE LARGER CETACEA, 789

impress of the horny teeth of the suckers upon the epidermis, to

deep circular cuts reaching to the blubber. In many cases the

epidermis had peeled off from within the deeply cut. circles,

exposing the corium. The circles varied in diameter from + inch

or less to 14 inches according to the size of the sucker which formed

them. These marks were noticed by Pouchet and Beauregard *

as occurring on the head of aSperm Whale and were attributed by

them to parasitic Cirripedes which had become detached. But, after

comparing these marks with the horny rings of the suckers taken

from the stomachs and intestines of the two specimens, and taking

into account the fact that the marks only occur on the head, there

can be no doubt that they were made by cuttlefish during their

struggles with the Sperm Whales who had seized them for food.

The marking of the skin of Odontocetes by cuttlefish has been

referred to by Dr. Harmer f.

A group of barnacles was found attached to the fourth anterior

tooth on the right side of the lower jaw of one of these whales.

The cluster consisted of individuals of Conchoderma auritwm,

to one of which was attached a small specimen of C. virgatum.

These were the only external parasites to be seen on the two

Sperm Whales.

The guts of both specimens were infested with internal para-

s.tes. ‘lhe determination of these is in progress.

Several of the whales were fresh enough to have permitted an

examination to be made of their histology and of their gut

Protozoa ; but unfortunately the necessary apparatus for such

work was not at hand at Innishkea. This was to be regretted

since the gut Protozoa of the Cetacea are quite ankenow n and

very little, if anything, has been written upon the histology of

whales.

VII. MiscEnLaANEous OBSERVATIONS.

1. Locality of Captures.

When the station at Innishkea was first opened in the early

summer of 1908, whales were taken at a distance of ten miles from

the island. But during the season of 1909 the steamers had to

go at least sixty miles out to sea before they could find a whale.

It was not possible to obtain the exact latitude and longitude of

the captures, as these were kept as trade secrets by the whalers.

However, it is safe to say that all the whales obtained by this

station during the two seasons of its existence were taken within

a radius of seventy miles, north, south and west of Innishkea.

This can be explained either by their being molested or by an

alteration in the distribution of their food.

During my visit one of the whaling captains discovered about

twenty individuals of Balenoptera musculus occupying the head

* Pouchet et Beawrezard, Nouvelles Archives du Muséum (Paris), 8 sér. vol. i,

1889, p. ¥.

f+ Harmer, Trans, Norfolk and Norwich Naturalists’ Soc. vol. vii. 1901, p. 185.

790 MR. D, G, LILLIE ON THE ANATOMY [May 24,

of an inlet or bay about sixty miles north-west of Innishkea

within the 1000 fathom contour, He hunted this spot for three

weeks, always finding the whales there, and was still doing so

when [ left. I took an opportunity of visiting the place with him

about the middle of August and was impressed with the apparent

regularity in the habits of the animals. Between daybreak and

10 A.M. they were to be seen swimming in this deep inlet and

blowing every ten minutes. After 10 A.M. they disappeared for

the rest of the day, but were all back again in the same place the

next morning. The captain always appeared on the ground at

di ay break, killed one or two and towed them home during the day,

returning to the locality over night.

it as iainieined by the whalers that the Balenoptera spend

several hours near the surface each day blowing every ten minutes

and storing their blood and lungs with air, and that they then go

down and remain below for perhaps eight or twelve hours at a time.

They are said to avoid the heat of the day by remaining under

water during that period and appearing at the surface in the

early morning and evening. On dull cold days they may be at

the surface at any time during the day, though they are most

plentiful at sunrise. Mr. Collett * appears to have been given

the same account by whalers. The explanation of these habits

attributed to the Balenoptera, if they really occur, may be the

rising and sinking of the plankton eaten by the Crustaceans

which form the staple food of these whales. It is more likely

that the mid-day heat may have some effect on the plankton,

and thus affect the whales indirectly, than that there should be

any direct influence.

Diving Powers.

I made enquiries of the whalers on this subject and I was told

that whales, when struck by the harpoon, often dive to the bottom

and sometimes come up with stones adhering to the ventral surface,

This, however, has been denied by other whalers. During my

visit I found on the under surface of one of the specimens of

B. musculus \anded at Innishkea a few broken spines of Echino-

derms, which had pierced the epidermis and were lying between

it and the corium. A harpooned whale is said to have come to

the surface with stones attached to its under surface when killed

at a locality where the depth was given as 100 fathoms on the

chart. If a naturalist were to spend sufficient time on board a

whaling steamer he could probably settle this point, for the whales,

when dead, always turn over on their backs and float with the

ventral surface uppermost, so there would be no difficulty in

detecting any adhering stones if they were present. Assuming

that a harpooned whale does not act very differently as regards

diving powers to one under normal conditions, this would “be a

simple method of ascertaining what those powers are. Of course

* Collett, Proc. Zool. Soc. 1886, p. 2638.

1910. ] AND BIOLOGY OF THE LARGER CETACEA. 791

the objection will be raised that if stones are found on the under

surface, how is one to make sure how long they have been there 2

On this point I was assured by my informant that the stones are

generally so lightly attached that they soon fall off, and a whale

could not travel far with them. If we may assume that this is

correct, when a whale is captured with stones on the ventral

surface it is only necessary to ascertain the depth of the sea at

the locality of capture, by referring to the chart, to ascertain the

depth to which the creature has dived on being harpooned. It is

said by the whalers that when a harpooned whale, after breaking

the harpoon-rope, dives toa depth of 60 fathoms and dies, it comes

to the surface at the end of three days; whereas if it goes below

this depth and dies it never reappears. There seemed to be

unanimous agreement among the whalers upon this point.

3. Copulation, Period of Gestation and Rate of Breeding.

The Balenoptera are said by whalers to copulate at the surface

of the sea. The pair swim towards each other and turn slightly

on their sides so that their ventral surfaces face oneanother. The

male makes several dashes at the female to insert the penis.

When the pair first rush together the long axes of their bodies

ave parallel with the surface of the sea; but they curve up

vertically at the end of the act. After copulation the male is said

to be exhausted and easily caught.

As regards the period of gestation and the rate of breeding

among Cetaceans, it is difficult to see how any definite information

can be obtained on these and similar subjects, which are of

interest to the cetologist and of considerable importance com-

mercially, unless individuals are kept in a confined place for

purposes of observation. This would perhaps not be such an im-

possible undertaking as it would at first appear. Of the manv

deep sounds or straits, through which the tides pass, on the

western coasts of Ireland and Scotland, one could doubtless be

found which could be converted into an aquarium for whales. To

do this it would only be necessary to place barriers at the mouths

of the strait in order to imprison the animal and yet allow of the

ingress and egress of the tide and food-supply. The animals

would have to be caught by the Japanese method of capturing

whales, which consists of throwing a large rope-net over the animal

and towing it to the shore alive. Balena biscayensis Gray, has

been taken by this method and possibly other species also *.

A Whalebone Whale imprisoned in a strait could probably be

kept alive on the organisms swept in by each tide.

Until some such scheme as this is brought to pass we must

continue to remain in ignorance, or be content with vague

speculations, concerning many points in the biology of the larger

Cetacea.

* K. Mobius, Sitzungsberichte der Akademie der Wissenschaften zu Berlin, lii.

1893.

792 MR. C, F. ROUSSELET ON | May 24,

EXPLANATION OF PLATE LXXIV.

“

Fig. 1. Posterior view of the right tympanic membrane of Balenoptera musculus

Linneus, showing its ‘attachment to the malleus.

a. Head of malleus with the two articular surfaces for the incus.

6. Manubrium of malleus,

c. Processus longus of malleus.

d. Portion of the lip of the tympanic bone to which the processus longus

of the malleus ts fused.

». Ligament of tympanic membrane.

J. Sac-like tympanic membrane.

. Portion of the mucous membrane lining the tympanic cavity.

. Wall of external auditory canal which has been cut near its junction

with the rim of the tympanic membrane-sac.

Fig 2. Plug of ear-wax from the external auditory meatus of Balenoptera musculus

Linneus. A. Dorsal view of plug. B. Posterior view.

a. Cup-like portion for the reception of the tympanic membrane.

b. Flattened distal portion.

Fig. 3. Ventral view of the left tympanie bone and inner portion of the external

auditory meatus of Balenoptera musculus Linneus.

a. Tuner portion of external auditory meatus with the ventral wall

removed to show the tympanic membrane and plug.

b. Plug of ear-wax.

ce. Tympanic membrane.

d. Tympanie bone.

Zoological Results of the Third Tanganyika Expedition,

conducted by Dr. W. A. Cunnington, F.Z.S., 1904—

1905.—Report on the Rotifera. By CHarzes I. Rous-

SELET, F.R.M.S.*

[Received April 7, 1910.]

(Plate LXXV.Tt)

Amongst the collections brought back by Dr. W. A. Cunnington

from the great inland lakes of Central Africa, visited during this

Expedition, were a number of tubes containing fine surface

Plankton nettings which I have searched over for Rotifera.

The gatherings were not specially made with a view to collect

these creatures, and being, moreover, made only from a boat in

the open water and never among the vegetation near the shore,

these circumstances may account for the comparatively poor

results obtained.

The Collection is, however, interesting and important from the

fact that, with the exception of 8 species (excluding doubtful

ones) collected in and near the Victoria Nyanza by Dr. Stuhlmann

in 1891 and deseribed by Dr. Ant. Collin (1) in 1896, and 14

additional species (again excluding the doubtful ones) collected in

the Victoria Nyanza by Dr. Borgert in 1904, and deseribed by

Prof, E. von Daday (2) i in 1907, no previous records of Rotifera

* Communicated by Dr. W. A. Cunninaton, F.ZS8.

+ For explanation of the Plate see p. 799.

PZ.Se 1910; Pl Exe:

F R.Drxon -Nurrant, del. ad. Nar London Stereoscopic Co. imp.

NOTOPS LOFUANA RovsseEter.

PZ.S.1910,PLLXXVI. 4

James Ritcuim, del. London Stereoacopic Co. imp.

a HYDROIDS FROM MERGUI ARCHIPELAGO. , |

PZ.S. 1910. PL LXXVi

JAMES Rircuise ‘ del ; London Stereoscopic Co. imp.

HYDROIDS FROM MERGUI ARCHIPELAGO.

1910.) ROTIFERA FROM TANGANYIKA, 793

from the Central African Region, and particularly from Lakes

Tanganyika and Nyasa, have ‘been made.

The tubes examined by me came from three lakes :—

1. Lake Tanganyika, including the Lofu river.

2. Lake Nyasa.

3. Victoria Nyanza.

From these I obtained a total number of 37 species, of which 11

had already been recorded from the Victoria Nyanza and adjacent

region. Most of these species have a wide distribution and are

known from various parts of the world, and only one species,

from the Lofu river, can be described as new.

In order to make this record of Central African Rotifera

complete, I will preface my results with the list of species found

by the above mentioned authors, Dr. Collin and Prof. v. Daday,

in and about Victoria Nyanza, excluding the doubtful species not

identified :—

Rotifer vulgaris Ehrenbg. (C.)

Philodina emini Collin. (C.)

Triarthra longiseta Ehrenbg. (D.)

Asplanchna brightwelli Gosse. (D.)

Notops macrourus Barr. & Dad. (D.)

Copeus copeus Khrenbg. (C.)

Tetramastix opoliensis Zach. (D.)

Euchlanis longicaudata Collin. (C.)

(Which is identical with Muchlanis propatula Gosse.)

Monostyla bulla Gosse. (D.)

Noteus stuhlmanni Collin. (C.)

Brachionus rubens Ehrenbg. (C.; D.)

angularis Gosse. (D.)

& bakerit Khrenbg. (D.)

forficula Wierz. (D.)

falcatus Zach. (1).)

pala Khrenbg. (D.)

militaris Khrenbg. (D.)

a tetracanthus Collin = B. caudatus Barr. & Dad.

(CS: )

Schizocerca diversicornis var. homoceros Daday. (D.)

Anurea aculeata Ehrenbg. (D.)

. Ma var. was ga Khrenbg. (C.)

re cochlearis Gosse. (D.)

Pedalion mirum Hudson. (D.)

LAKE TANGANYIKA.

From this lake a large number of tubes were available, collected

from October 1904 to February 1905, all fine tow-nettings made

some distance from the shore and near the following localities :—

Mbete, southern end of lake; Komba Bay, south-western corner ;

Kala, a short distance up the eastern shore; Kasawa. on the west

794 MR. C. F, ROUSSELET ON [May 24,

coast; Vua harbour, on the west coast ; Bismarkburg (Kasanga),

on the east ccast; Maswa, on the east coast.

All these samples contained vast quantities of microscopic algze

and diatoms ; but though I searched most diligently for many days

and weeks, I succeeded in finding only 10 species of Rotifera, and

very few specimens of these, ne nothing new, This paucity of

Rotatorian life is certainly most rem: arkable for such a vast lake ;

but it is probable that, had collections been made amongst the

vegetation near the shore, a greater number and variety might

have been obtained. I understand that the water of this lake

was formerly somewhat brackish, though the actual salinity was

never ascertained, and also that the Cladocera are totally absent,

whilst the Copepods are abundant.

Rotifera found in Lake Tanganyika,

(eistes (sp %) (tubes only, on Pliumatella).

Polyarthra platyptera Ehvenbg.

Synchata (sp. ¢) (badly preserved and wholly contracted),

Asplanchna intermedia Hudson,

Rattulus stylatus Gosse.

Cathypna luna Ehrenbg.

Monostyla bulla Gosse.

es. Seat Some

Salpina brevispina Khrenbg.

Anurea aculeata Ehrenbg.

Brachionus pala Ehyenhg. (very small variety).

Of these, the only species which was found in some abundance

and from various parts of the lake is the small variety of pe i

onus pala, measuring only 205 pin total length of the lorica. The

European specimens ust ily measure 325 fs ae some exc autiondllt

large specimens found in Ormsby Broad, in Norfolk, attain a size

of 442 ll

Special interest attaches to the presence in Tanganyika of

Asplanchna intermedia, as 1 was not able to record any member

of this genus from South Africa (3).

Loru River.

The Lofu river is a large tributary of Lake Tanganyika,

ente rng itat its extreme Souihe western corner. | giv @ a separate

list. of species found in this river, as the Rotatorian fauna shows 2

marked increase in numbers and variety as compared with those

found in the lake.

Although only two small tubes of material were collected by

the tow-net, | obtained therefrom 23 species, one of which is new

to science Sarl altogether different from anything known; the

specimen is figured on Plate LAX VY, and described below under

the name of Wotops lofuana, sp.n. The two gatherings were

made in October 1904 about a mile and a half from the mouth of

the river, so that it is quite clear that a far greater number of

1910. ] ROLIFERA FROM TANGANYIKA. 795

Rotifers are carried into the lake by this river than can afterwards

be found in its water.

Rotifera found in the Lofu Liver.

(“eistes mucicola Kellicott.

Limmias (sp. %) (tubes only).

Callidina (sp. ?) (wholly contracted),

Syncheta oblonga Khrenbg.

Notops lofuana, sp. n., Rousselet.

Proales daphnicola Thompson.

Rattulus longiseta Schrank.

sf gracilis Tessin.

a bicristatus Gosse.

* macerus Gosse.

Euchlanis hyalina Yudson.

¥ oropha Gosse.

5 propatula Gosse,

Monostyla lunaris Khrenbe.

Cathypna leontina Turner.

Bs ungulata Gosse.

= luna Khrenbe.

Salpina macracantha Gosse.

Pterodina trilobata Shephard.

Noteus quadricorms Khrenbg.

A: Hy var. brevispinus Daday.

Brachionus militaris Ehrenbg.

Anurea aculeata var. valga Ehrenbg.

Norops LOFUANA, sp. n. (Plate LX XV. figs. 1-3.)

Amongst the Rotifera found in the Lofu river there is one new

species which appears so different from any known kind, that I find

difficulty in assigning it to any known genus. Unfortunately

the only specimen available is not sufficiently well preserved to

enable one to be sure that the integument is not to some extent

distorted. Its general shape and appearance suggests a Votops, or

Gastropus, and in the absence of a fuller. knowledge of its anatomy

I have provisionally placed it in this genus under the above name.

The figs. 1 and 2 of Plate LXXV. give an exact outline of its

appearance in dorsal and lateral view, and I am indebted to my

friend Mr. F. R. Dixon-Nuttall for these accurate drawings which

he has made from the mounted specimen. Fig. 3 represents the

jaws, which I have drawn under great difficulties and as accurately

as was possible under the circumstances.

The anterior half of the body is stout, somewhat compressed

dorso-ventrally, presenting a wide semicircular front with broad

lateral lobes. ‘The oral region appears to project prominently on

the antero-ventral side; but it is quite possible that this appearance

may be due to distortion, as the whole animal is not very well

preserved. The posterior half of the body, which shows a deep

796 MR, C,. F, ROUSSELET ON [May 24,

constriction behind the great frontal lobes, is globose in shape,

high and rounded dorsally, contracting sharply ‘postero- ventrally

for the issue of a short tubular foot, apparently one-jointed, and

carrying two fairly thick toes. The integument is firm but not

hardened into a lorica. Two red front: al eyes are present, wide

apart, and situated near the middle of the lateral lobes.

The jaws are shown in fig. 3; I must state, however, that,

owing to a granular deposit below the integument, which I was

unable to clear or dissolve with potassium hydrate, this figure

must be taken as only approximately correct. I have unfortunately

been unable to obtain a clear view of the various parts, but the

figure represents as much as | have been able, though imperfectly,

to make out of the structure. The rami are broad triangular

plates resting on a long straight fulerum, which appears to be

surrounded by a CO: arsely striated muscle ; 5 the mallei consist of an

uncus with four teeth of which the anterior tooth is the largest

and curved, and a narrow manubrium directed outward.

The very imperfect state of preservation of the specimen did not

enable me to make any observations regarding the internal organs,

the stomach, intestine, ovary, glands, muscles, &e., being wholly

dissolved into a granular mass. A short tubular dorsal antenna,

however, is distinct on the dorsal side behind the middle of the

head.*

Total length, including foot, 204 py (;4= in.).

Widest part, across frontal lobes, 108 jx (s4s in.).

A considerable number of Plankton collections were made in

this lake in June 1904, and I examined many tubes full of fine

Alge and Diatoms ; but only in two of these, from Monkey Bay

and Anchorage Bay, situated at the southern end of the lake,

could J find a very few Rotifera, six in all, as follows :—

Rotifera found in Lake Nyasa.

Diurella stylata Kyfert.

Rattulus stylatus Gosse.

Brachionus pala var. dorcas Gosse,

- militaris Ehrenbg.

Notholea labis Gosse.

Anurea aculeata var, valga Ehrenbg.

* Soon after writing the above description I received from Mr. H. K, Harring of

Washington some drawings of Rotifera collected in that ne ighbourhood, and amongst *

these was this same extraordinary species, which he had found in some abundance

near Washington, U.S.A. ‘The extended animal shows a hammer-shi uped head with

two large circular ciliated lobes projecting outward, a tubular buceal region, and two

ved eyes situated very wide apart on the ciliary lobes ; Lee the general shape

of the animal is well represented on Plate LXXV. No doubt Mr. “Harring will

publish his drawings and description of this very st ange animal which will

peeatiily require a hew genus, if not a new family, to receive it.

1910.] ROTIFERA FROM TANGANYIKA. 797

Victoria NYANZA.

Only a very few collections from this lake were available ; they

were made near the shore at Bukoba on the west coast im April

1905, and from these [ obtained the following 8 species of Rotitera,

all of which are already known from this lake :—

Monostyla bulla Gosse.

Brachionus bakerit Khrenbe.

faleatus Zach.

forficula Wierz.

- caudatus Barr. & Dad.

Schizocerca diversicornis var. homoceros Daday.

Anurea aculeata var. valga Ehrenbg.

os cochlearis Gosse.

Tn this list the presence of a number of rare species of Brachi-

onus is notable, a fact already noted by Prof. v. Daday. It is

evident tome that a more thorough exploration of the micro-fauna

of this vast and shallow lake would reveal a very considerable

Rotatorian fauna.

Excluding the few species which could not be identified, the

above lists contain 37 species of Rotifera collected by Dr. Cun-

nington in the great lake regions of Central Africa, of which

11 species were already known by the work of Dr. Collin and

Prof. v. Daday, and 21 of these species were also previously

recorded by me from South Africa, whilst one species is new to

science.

LAKE ALBERT.

Having also received for examination a very small tube of

Plankton material collected with the tow-net in July 1907 in

Lake Albert by Dr. R. T. Leiper, the Surgeon accompanying

an expedition sent to this region by the Egyptian Survey

Department, I will here record the three species of Rotifera

found therein :—

Monostyla bulla Gosse.

Noteus quadricornis Khrenbg.

Lrachionus bidentata Anderson.

The last mentioned species appears to be very rare, having beer

found only once since its first discovery in Calcutta by Anderson

in 1891, namely, in Bulgaria by Dr. Konsuloff (who sent me some

specimens for identification) about two years ago.

Proc. Zoon. Soc.—1910, No, LIT.

798 ON ROTIFERA FROM TANGANYIKA.

DisrripuTion oF Rorrrera 1x Cenrrat Arrica, being a list of

all the species known from this region at the present time.

CEcistes mucicola Kellicott ...............0.0+-0+ee2+

Rotifer vulgaris Ehrenbg.

PiwlOuwanenarins COMM wtererewc ee tesnreley= scr 6e es veel Cees

Polyarthra platyptera Ehy enbg. ee dite iats ts x

Triarthra longiseta Whrenbg. ...........-... 220000000] oe

Syncheta oblonga Ehrenbg. ............ +00 000000eee 0 se

Asplanchna intermedia Hudson ............ 0.0... +++ x

. brightwelli Gosse .. HE Recs eisl Vie

Notops macrourus Barr. & Meek oe

55 fey 32 SOLED, soa-nceea tee SOR EBC UBSOnER Ra See

Proales daphnicola Thompson { Se crn Sad RBH NBaBehEHC

Copeus copeus Ehrenbg.

Tetramastix opoliensis Zach. Sobcobesoeeecee||) cbc

Rattulus longiseta Schrank ...............00-.2s000 ee

3 stylatus Gosse pe ofan ih she: 2 See

33 IGT GLAS MUOERID Mens os oy cieen coe = «0s «oleae eee

a BUCTUSUAERBI(RORSD. nasy-t- vacinasiecsns + ceeiecer ne

a macerus Gosse

Diurella stylata Eyfert ieee dh aero 3

Euchlanis hyalina Hudson ...........0000eevee eee 50

p DOD GOSKO Sree e coe wean susnssicsy as -inns eke

x GEIR OY PER OUAT ELE 0) CN, Senter 5059 CEOOQOERIRSAE ADs | Sc

Monostyla lunaris enone Behe ah vines svsinas soot awe

e bulla Gosse . hE ERO nee ALeS@¢

Cathypna leontina Turner. vente

: ungulata Gosse... sl ae

- luna Ehrenbg. rel Me:

Salpina macracantha Gosse .. kal oy

a brevispina Ehrenbg. At Sx2

Pterodina trilobata Shephard .. ‘ae

Noteus quadricornis Ehrenbg. : art

Bs var. brevispinus Daday ere

* stuhImanni Collin aN A aoe Fe)

Brachionus pala Ehrenbg. Ree rtesirh cu dortel wow

5 » var. dorcas cee hes

3 DOPAC DR Uasccesncotesccisesees ees

3 Saloatus Za. <....... eevee neces

ee Sorficula Wierz.

Anurea aculeata Ehrenbg.

on be var. valga Ehren.

A cochlearis Gosse .......

Notholea lahis Gosse ............0c0 00s

Pedalion mirum Fudson ...........

Schicacer ca diversicornis var. homoceros s Daday t

| Lake Tanganyika. |

5 caudatus Barr. & Dad. ............ .. ie

5 angularis Gosse Berets er

se rubens Ehrenbg.

militaris Bhrenbg. es

bidentata Anderson .........

Be SOK Me IK OKs PGS HE EK IG sa

x | Lofu River.

1 ORE

| Lake Nyasa.

> KKXX

| Victoria Nyanza.

XX:

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[May 24,

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1910.] ON HYDROIDS FROM MERGUI ARCHIPELAGO.

BIBLIOGRAPHY.

1. Cott, A.—‘“ Rotatorien, Gastrotrichen und Entozoen Ost-

Afrikas.” Deutsch-Ost-Afrika, 1896, vol. iv. no. 16.

2. Dapay, Prof. E. von.—‘ Plancton-Tiere aus dem Victoria

Nyanza.” Zool. Jahrbiicher, Bd. 25, 1907, pp. 245-262.

3. Rovusseter, C. F.—‘ Contribution to our Knowledge of the

Rotifera of South Africa.” Journ. Royal Mier. Soc. 1906,

pp. 393-414.

EXPLANATION OF PLATE LXXV.

Fig. 1. Notops lofuana Rousselet, dorsal view. X 360.

2 lateral view. X 360.

thejaws. % about 700.

33 bb} 99

”? bb} ”

3. The Marine Fauna of the Mergui Archipelago, Lower

Burma, collected by Jas. J. Simpson, M.A., B.Se., and

R. N. Rudmose-Brown, D.Se., University of Aberdeen,

February to May 1907.—The Hyprorms. By James

Rircaie, M.A., B.Se., Natural History Department, The

Royal Scottish Museum *.

[Received April 28, 1910. }

(Plates LXRX VI. & LXXVILF, and Text-fig. 79.)

The Hydroids were represented in the collections brought

together in the Mergui Archipelago by Dr. John Anderson in

1882, and described in the Journal of the Linnean Society for 1889,

by a meagre total of six species, two of which were regarded by

Hincks as new. The reasonable anticipation that the careful

collecting of Dr. Brown and Mr. Simpson would add many species

to the Hydroid fauna of this region of the Indian Ocean has been

amply fulfilled; for the present collection, confined to a littoral

area, contains representatives of thirty species, only three of which

(Campanularia raridentata and Idia pristis, both widely distributed

species, and Aglaophenia crispata, a synonym of Lytocarpus pen-

narius, fide Billard, 1909, p. 329) were included in Hincks’s list.

As only two of the species, beimg described as new to science,

must be regarded as peculiar to the Archipelago, the collection is

well fitted to show the relationship which the Hydroid fauna

bears to that of neighbouring seas. Leaving out of account

an undetermined species of Plumalaria, nineteen of the Mergui

species have already been recorded from the Indian Ocean,

the majority of these occurring off Madagascar and the eastern

shores of <Africa. The remaining ten species, indicated by

* Communicated by R. KirKparrick, ¥.Z.S.

¥ For explanation of the Plates see p. 825.

800 MR. J. RITCHIE ON HYDROIDS | May 24,

asterisks in the following list, have thus been added to the

fauna of the Indian Ocean. As to the wider relationships :—

Three species are peculiar to the Indian Ocean (Corydendriwm

sessile, Hebella crateroides, Calycella oligista), leaving twenty-

six to be accounted for. Of these, twelve are so widely

distributed that their occurrence, emphasizing the normal nature

of the fauna, can give no indication of special affinity: three

being found in the Atlantic and North Pacifie Oceans and in Aus-

tralasian seas (Opercularella lacerta, Plumularia setacea, Antenella

secundar ia) 5 two in the Atlantic and North Pacific Oceans

(Perigonimus repens, Campanulariararidentata) ; and seven in the

Atlantic Ocean and in Malay-Australian seas (Pennaria disticha,

Halecium tenellum, Campanularia corrugata, Hebella calearata,

Sertularella polyzonias, Idia pristis, Diphasia digitalis), Of the

remaining fourteen, four have been found only in the Atlantic

Ocean (Ludendrium attenuatum, Cuspidella costata, Lafoéa serrata,

Lafoéa venusta), seven in the Malay-Australian area (Hudendrium

generalis, Cladocoryne haddoni, Halecium simplex, Thyroscyphus

vitiensis, Sertularella cylindrica, Sertularella quadridens, Sertularia

turbinata), one in the North Pacitic Ocean (7hyroscyphus regularis),

and two in both the last-named aveas (Lytocarpus pennarius and

L. pheeniceus).

The closer relationship is obviously with the fauna of the

Pacifie Ocean (represented by ten distinctive species), and in

particular with the Malay-Australian portion of that ocean

(represented by seven species). The significance of the apparent

Atlantic atlinity is minimised when it is understood that of

the four species common to it and the Mergui Archipelago, one

(Ludendrium attenuatum) is identified with much doubt, while

the remaining three are very minute epizoie species, the presence

of which in other regions may conceivably have been overlooked.

It is remarkable ‘that of the thirty species recorded, seven-

teen were found climbing upon larger Hydroids and one upon

a Polyzoon, the majority being minute, habitual epizoa, be-

longing in the main to the families Halecide, Campanularide,

Campanulinide, and Lafoéide. The examination of this collection,

and of others, leaves with me the distinct impression that the

epizoic Hydroids are not distributed indiscriminately upon all

types of Hydroid host. Thus, in general, the members of the

family Plumularide remain comparatively free from extraneous

growths—a fact to be correlated, perhaps, with their possession

of nems itophores ; while colonies, belonging especially to the

family Sertularide, are occasionally so overgrown that the

structures of the host are much obscured, Even distinet pre-

ferences for certain species may be observed. ‘Taking, for each

epizoic species in this collection, the total numbers of stations at

which it was found, and adding those numbers for all the epizoic

species, we find that epizoic species were found at a sum total of

forty-five stations. At thirty-two of these /dia pristis was either

the only host, or one of the hosts where more than one species

1910.] FROM MERGUI ARCHIPELAGO. 801

was infested; Zhyroscyphus vitiensis at nine; Sertularella quad-

ridens at four; Corydendrium sessile, Hudendrium attenuatum,

Lytocarpus pheniceus each at two; and Diphasia digitalis and

Plumularia setacea each at one. This result bears out the

impression gained from handling the collection, that /dia

pristis is remarkably subject to infestation by extraneous Hydroid

colonies, Frequently its hydrothecee were all but obscured by

the attendant growths of Hydroids and Polyzoa, and this

although the state of the hydranths and of the coenosare generally,

indicated that the host colony was in a perfectly healthy condition.

The occurrence, in Lafoéa venusta (2), of what appears to be

a step towards a more intimate symbiosis; of a canaliculated

ccenosare in the stem and branches of Sertularella quadridens ;

of the hitherto undescribed gonosomes of Halecium simplex and

Thyroscyphus regularis, are worthy of notice.

Finally, I would express my thanks to Dr. R. N. Rudmose-

Brown and Mr. J. J. Simpson for entrusting this collection to

me for examination,

List of Species.

I. Gymnoblastea.

Family CLavip&.

*Corydendrium sessile, sp. n., p. 802.

Family Bov@artnvILLip2.

*Perigonimus repens Wright, p. 804.

Family EupENDRID&.

we udendrium attenuatuwm Allman (?), p. 804.

a generalis Lendenfeld, p. 805,

Family Corynipm.

Cladocoryne haddoni Kirkpatrick, p. 805.

‘Family Preynariw2.

Pennaria disticha Goldfuss, var. australis Bale, p. 806.

II. Calyptoblastea.

Family Havecirpa.

* Halecium simplex Pictet, p. 807.

53 tenellum Hincks (2), p. 808.

Family CAMPANULARID.

Campanularia corrugata Thornely, p. 809.

raridentata Alder, p. 809.

Hebella calearata A. Agassiz, p. 810.

» erateroides Ritchie, p. 810.

Thyroscyphus regularis Jaderholm, p. 811.

F vitiensis Marktanner, p. 812.

802 MR. J. RITCHIE ON HYDROIDS [May 24,

Family CAMPANULINID2,

* Opercularella lacerta Johnston, p. 812.

* Calycella oligista, sp. n., p. 813.

Cuspidella costata, Hincks, p. 814.

Family Laro&m.

Lafoéta serrata Clarke, p. 815.

* venusta Allman (2), p. 815

Family SERTULARIDA. ;

* Sertularella cylindrica Bale, var. pusilla, n., p. 817.

polyzonias L., var. cornuta Ritchie(?), p. 818.

=f quadridens Bale, p. 818.

Tdia pristis Lamouroux, p. 820.

Diphasia digitalis Busk, p. 821.

Sertularia turbinata Lamouroux, p. 821.

Family PLUMULARID&.

Plumularia setacea Linn. (?), p. 822.

fl sp. indet., p. 822.

Antenella secundaria Gmelin, p. 822.

Lytocarpus pennarius Linn., p. 822.

5 phaniceus Busk, p. 823.

* indicates a species recorded fer the first time from the Indian Ocean.

SysreMAric Discusston,

I. GYMNOBLASTEA,

Family CLAVID 4.

CoryYDENDRIUM SESSILE, sp. n. (Plate LX XVI. figs. 1 & 2.)

TrorpHosomE.—Colony small, the lar vest of the three collected

being only 37 mm. in height. The ‘stem is strongly fascicled,

1:5 mm. thick towards the base, and straight. It may divide

into main branches, which are beset by numerous smaller, almost

equal branchlets, about 1 mm. in length. These spring from

two opposite sides, and lie roughly in one plane. ‘The branchlets

on any one side are almost equidistant from each other, but their

positions bear no regular relation to those of the opposing series.

‘They taper very slightly towards the base, but there is no trace

of ringing nor of wrinkling.

The majority of the hydvanths spring from tubes on the anterior

surtace of the branchlets, although a few are also scattered on the

main branches. The hydranth tubes do not become free from

the common fascicle, except rarely and for an extremely short

distance; nor are special hydrophore-like portions developed.

Thus the hydranth projects simply from the open mouth of a

tube, the adcauline wall of which is adnate. The bydranth tubes

1910.] FROM MERGUI ARCHIPELAGO. 803

are arranged in two series, the polyps of one series alternating

with those of the other. The aperture faces outwards and

upwards, is slightly elliptical in shape, and has an even margin,

indicating the boundary between the thicker perisare and the

place where the chitinous coat becomes so thin and delicate that

it follows the movements of the polyp. The perisare throughout

is covered by a thin coating of extraneous material—diatoms,

sand-grains, fragments of sponge-spicules, and such like,

The hydranths are much contracted, and in this state exhibit

a pyriform body with well-developed hypostome. The tentacles

appear to number about forty.

GoNosoME unobserved.

Colour, in alcohol, grey.

Dimensions :—

Diameter of branchlets immediately above origin 0:31-0°38 mm.

3 Inycipain Png emer. 85 hs debacle scinsran 0°25-0°31 mm.

Eigdmamba; lengthy ) caraamens sts. dads as catia 0:52-0:98 mm.

is CiamCterg prea: feces oseesweteta os 0-21—0°42 mm.

Nematocysts on tentacles, length ................:. D pl.

is we cera thy. sss. ce foci 3 ties 3 pe.

Locality. Three small colonies, with hydrorhiza embedded in a

sponge, were found at St. 35, between Warden Island, Howe

Island, and Lyall Island, 15 to 20 fathoms, rock and sand.

So little is known of the variation liable to oceur in the indi-

vidual species of this genus, that it is difficult to fix on definite

specific characters. In this case, I have relied mainly on the

structure of the tubes from which the hydranths project. In

the majority of the species of Corydendriwm so far described

‘these are free for a considerable distance below the hydranth ;

but here the tubes are adnate up to the end. This gives to the

pane a dumpy appearance, unlike the lax erowth of other

ecies. In this respect it approaches most closely y C. corrugatum

Nene (1905, p . 941), which is to be distinguished, however, by

its distinct hy drophore- like structures,” ‘which are ‘ usually

distinctly corrugated with deep irregular annulations ” ; by the

presence of a well marked annular constriction near the origin of

the branches ; and by the large size (5 inches) of the colonies.

The species, Soleniopsis dendrifor mas, described by me in 1907

(1907, p. 494) as representing a new genus, under the erroneous

supposition that Corydendriwm gave rise to gonophores with free

medus: ad been unable to refer to the m agnificent description

of Weismann (1883),—obviously belongs to the genus Cory-

dendrium, as Dr. Stechow has recently pointed out ( 1909, p. 9).

Corydendri iwm dendriformis differs from C. parasiticum in

possessing thick, definite stems, with branchlets arranged in

pseudo-pinnate fashion, and in having the free portion of tlie

tubes from which the hydranths project much shorter than those

of C. parasiticum.

804 MR. J. RITCHIE ON HYDROIDS [May 24,

Family BOUGAINVILLID&.

PpeRIGONIMUS REPENS Wright, 1858.

Rare examples of an epizoie species occur, which cannot be

specifically distinguished from British specimens of P. repens,

the simple lax habit of which they exactly assume. There

are differences between the dimensions of the Mergui and of

British examples; for while the former are smaller in height

and in the proportions of their hydranths and hydrocaulus, in

respect of the nematocysts in the tentacles the order is reversed.

The comparative table which follows shows at a glance the size-

relations of the two forms. About twelve tentacles crown each

hydranth,

No trace of gonosome was observed.

Dimensions :— Mergui Typical Scottish

specimen, example *.

Height of colony ........... a ise : 3 mm, 6 mm.

Diameter of hydvocaulus ......... 0-04 mm. 0-07 mm.

Hydranth; lon ethe eye seenese n> «> 0°17-0:24mm. 0°'24-0°34mm.

= greatest breadth ...... 0:08-0:13 mm. 0:14-0°15 mm.

Nematocysts of tentacles, length DD pe. 4D p.

> a breadth 3p. 2°2 p.

Locality. Rare colonies epizoic on Corydendrium sessile, trom

St. 35, between Warden Island, Howe Island, and Lyall Island,

15 to 20 fathoms, rock and sand.

The present record adds LP. repens to the fauna of the Indian

Ocean. It has already been noted from the eastern and western

sides of the North, and the western side of the South Atlantic

Ocean, from the Mediterranean Sea, and from the eastern and

western (Japan: Stechow, 1909, p. 25) sides of the Pacific Ocean.

Family EUDENDRIDA.

Eupenprium ATrENUATUM Allman (?), 1877.

Many poor colonies, lacking any trace of hydranth or gono-

some, | refer, with uncertainty, to this species, on account of

resemblance in the skeleton. ‘The Mergui specimens attain a

somewhat greater length (3 inches) than the original examples ;

but the delicate, very slender, non-fascicled stems, with their few

branches, and their short hydranth-bearing ramules lying m one

plane and set alternately at regular intervals of about 1 mm., are

very similar in both cases. Three or four rather irregular annu-

lations mark the base of each ramule, and occasionally a few odd

rings occur irregularly on the ramules and on the stem itself,

The regular and close alternation of the hydranth-bearing ramules

seems the most evident character of an indefinite species, though

a somewhat similar arrangement is observed in 2, maldivense

Borradaile (1905, p. 838).

* Slide of specimen from Loch Carron, 60 fathoms, in my collection.

1910.] FROM MERGUI ARCHIPELAGO, 805

Dimensions :—

Stem) diameter) eens eeass.ci.iev2 — O°20=0'24 mm.

Hydranth-bearing ramules, diameter. 0:11—0°16 mm.

Localities. Many colonies from St. 22, Hastings Harbour, 3 to

20 fathoms and shore, rock and sand ; anda few poor colonies, paler

in colour, from each of Stations 23, Five Islands, 8 to 12 fathoms,

rock and sand, and mud; and 25, Gregory Group and Crichton

Island, 4 to 14 fathoms, stones and broken shells, and rock.

Recorded originally by Allman from 8.8.W. of Tortugas, at a

depth of 60 fathoms, this doubtful species forms an addition to

the fauna of Indian seas.

EUDENDRIUM GENERALIS Lendenfeld, 1885.

The colonies belonging to this species are mature, but are

only about 1 em, in height. They bear well-developed female

gonophores, oval in shape, “which spring in a whorl of four or five

from the base of a hydranth, as in Kirkpatrick’s description and

figures (1890, p. 607, pl. xv. fig. 2).

“The tr ophosome of this species is hard to distinguish from that

of #. capillare. The more slender character of the branches of

the latter, relied on in part by Kirkpatrick, is of little value,

since a considerable amount of variation occurs in both species.

On the other hand, the branching of H. capillare is more profuse,

and irregular; also in the Australian species there is, in the

smaller branches, distinct alternation, while the whole colony

appears more rigid than the lax growths of H. capillare.

Dimensions :—

Shen, diam Chorumesmee eee es: PERE DC ek oe POOH,

Hydranth-bearing ramule, diameter ......... 0:09 mm.

Locality. Creeping in small quantity upon Hudendrium attenw-

atum % and Idia pristis, both from St. 22, Hastings Harbour,

3 to 20 fathoms and shore, rock and sand.

Until the present record from the Indian Ocean, this species

has been found only in the Australian region: Port Phillip, near

low water (Lendenfeld, 1885, p. 351); Torres Straits—20 miles

N.N.W. of Warrior Island, BA fathoms; Murray Island, 15 to

20 fathoms (Kirkpatrick, 1890, p- 607).

Family CoRYNID&.

CLADOCORYNE HADDONT Kirkpatrick, 1890.

Except in size (for they are even more diminutive than those

recorded by Kirkpatrick and Jiderholm), the Mergui specimens

agree with the description and figures of the former, the double

wrinkle at the base of the stalk being very evident. On occasion,

one or two additional wrinkles may occur at the base of the stalks

and, more seldom, on their middle portions; but in the majority

of cases these rings are of little significance, since they belong to a

new stalk regenerated after the destruction of the old one. Here,

806 MR, J. RITCHIE ON HYDROIDS [May 24,

as in simple Campanularian hydroids, regeneration is not content

with simply adding a portion to the old stalk, but a complete new

stalk is reproduced, springing from within the truncated end of

the old one.

Dimensions :—

Stalls, denieths; eibeeteeeee weet ssc .seeewe 0°43-0°98 mm.

sy M@IRMIRGEI TT Peer ees. scl os ceade 0:06-0:08 mm.

Stolon, diameter ...... Mee eich enact 2a 0-10 mm.

Hydranth, lengthen ents. ssc fos. 0. see 0°34—0°36 mm.

i ere: atest Meee hn! ss)t. seek 0-11-0°21 mm.

Cnidoblast threads, armed with barbs such as Kirkpatrick has

figured, accompany several of the hydranths. They are remark-

able for their large size, the barbed portion being 6 p in diameter

from tip to tip of the barbs.

Localities. Rare hydranths on Thyroscyphus vitiensis and on

Idia pristis from St. 1, east of Tavoy Island and Port Owen,

4 to 12 fathoms, sand and broken shells, and mud. A solitary

hydranth on Idi pristis and a few on Corydendrium sessile from

St. 35, between Warden Island, Howe Island, and Lyall Island,

15 to 20 fathoms, rock and sand.

The species has been previously recorded only from Murray

Island, Torres Straits, 15 to 20 fathoms (Kirkpatrick, 1890, p. 606) ;

and, in Indian seas, from Paumben, 1 to 3 fathoms (Jiiderholm,

1903, p. 268).

Family PENNARIDA,

* PENNARIA DISTICHA Goldfuss, 1820. Var. AusTRALIS Bale, 1884.

(=Pennaria cavolinii Ehrenberg, 1832.)

A few well preserved colonies, the largest 30 mm. high,

represent this species. The ramules from which the hydranths

project increase considerably in diameter upwards from their

origin, where they bear three or four annulations, to the distal

end, which is smooth. The number of filiform tentacles varies

from nine to twelve, but there is much variation in the number

of the capitate tentacles. The numbers depend, to some extent,

upon the age of the hydranth, for the tentacles are fewest in

those hydranths which, springing from ramules, lie towards the

distal end of a branch—that is, in the youngest hydranths; for

growth proceeds by the elongation of a branch, new polyps

coming into being next to the terminal polyp, w hich is the oldest

and the largest on the branch.

Pictet’s careful comparative study of colonies of typical

P. cavolinii from Naples, and of typical P. australis from

Amboyna, makes clear that the latter form falls within the

range of variation observed in the former species. But so

definitely limited is the basal ringing of the ramules from

which the hydranths arise, and so great a difference is there

* Wor synonymy, see Bedot, 1901, p. 459.

1910.] FROM MERGUI ARCHIPELAGO. 807

in the proportions of the European and the Indo-Pacific forms,

that I retain for the latter a distinct varietal name.

Locality. St. 29, reef off High Peaked Island, shore.

Although P. cavolintdi in its varying forms has been recorded

from Eastern Australia (Bale, 1884, 1893; Lendenfeid, 1885),

and from the East Indies (Pictet, 1893; Campenhausen, 1896 ;

Weltner, 1900), the only records for the Indian Ocean are those

of Warren (1906, 1907, and 1908, as Halocordyle cooperi, syn.

Pennaria australis, var. coopert) and that from Christmas Island

in the paper following.

II. CALYPTOBLASTEA.

Family HaLEeciIpDs&.

HALECIUM sIMPLEX Pictet, 1893. (Plate LX XVII. figs. 10 &

NS)

This very rare species is represented by closely woven colonies

eovering the surface of a Polyzoon which is clustered at the base

of colonies of Corydendrium sessile. ‘The presence of the gono-

some, so far undescribed, renders these specimens of particular

interest.

TropHosoME.—T'o the naked eye the minute colonies are

almost invisible, the hydranths being most easily discerned

they project from the surface on which the colonies grow,

in close groups, resembling clusters of the smaller species of the

entoproctan polyzoon, Pedicellina.

The stolon strands are of small diameter, but can often be

traced for a considerable distance. Their courses are com-

plicated, however, by the occurrence of offshoots, which, uniting

with other stolons, form a network which les closely upon the

substratum, or is interwoven with it.

Short, unbranched, unwrinkled hydrocauli arise from the stolon

at short distances from each other. Basally their diameter is.

small, but it gradually increases until, at the base of the hydro-

theca, the girth of the hydrocaulus is about equal to that of the

stolon. The hydrothec are very small. Their walls, which are

well developed, form an inward curve to the margin, so that,

instead of being flaring or trumpet-shaped, as in many species of

Haleciwm, they are rather saucer-shaped. Very close to the

margin occurs a ring of bright dots, chitinous prominences on

the inner surface of the: hydrotheca to which the hydranth was

attached. Here the internal prominences are more highly deve-

loped than in any other species I have examined, for they sometimes

rise into pronounced spines 7 p in length (Pl. LXXVII. fig. 11).

Occasionally two or even three hydrothecee occur in succession, a

new hydrocaulus arising from within an old hydrotheca.

The hydranth is of great size. When contracted it is about

twice as long as the hydrocaulus, but when extended it reaches a

length of one millimetre or even more. It consists of a long

8 a

808 MR. J, RITCHIE ON HYDROIDS [May 24,

cylindrical neck, the base of which is attached to the hydrotheca

at the chitinous projections already described. Distally this

swells out into a very definite bulb, which is surmounted by a

circle of from 18 to 20 tentacles. At the bases of the tentacles a

ring of very large bean-shaped nematocysts, 30 » long by 6 pu

broad, surrounds the hydranth. Similar cells occur in the

ceenosare of hydrocaulus and stolon.

GonosomE.— This does not appear to have been described. A

gonangium arises from the side of the hydrocaulus, about half-way

between the hydrotheca and the stolon. The female gonangium

is borne on a short stalk, and is roughly calceolate in shape,

resembling the female gonangium of H. beanii. except that here

the tubular orifice of the upper surface does not occur. All the

gonangia I have examined contain ova to the number of about

six. The male gonangium, therefore, remains unknown.

Dimensions :—

DUOLON) (ISIC AR ce sens sis's ais xa tne See 0:06—-0°07 mm.

fydrocanilig meneuibetes,.. 1-026 ss. chacee 0°18~0°28 mm.

Biydrouheea, Gepiny tes ecs.st. acne. nese sue 0:024-0:027 mm.

- diameter at margin ...... 0:08-0:10 mm.

ONAN CI CM EGN sec psn ssn ckanceees 0°42-0°50 mm.

: greatest diameter ......... 0:17-0:21 mm.

Hydranth, length extended ............ 1:05 mm,

Bs Py weecorbracted «4. .2.5..2e 0-78 mm.

Locality. Epizoic on a polyzoon associated with Corydendrium

sessile, from St. 35, between Warden Island, Howe Island, and

Lyall Island, 15 to 20 fathoms, rock and sand.

Hitherto recorded only from Amboyna, in the Moluccas

(Pictet, 1893, p. 22), this record adds the species to the fauna of

Indian seas.

HALECIUM TENELLUM Hincks (?), 1861.

The trophosomes are lax in habit, and in all other respects

agree with Hincks’s species ; but, in the absence of the gonosome,

confident identification is impossible, especially since the likeness

between the trophosomes of young examples of the Australian

HH, parvulum Bale and /, tenelluim appears to be almost perfect

(see Markt.-Turner., 1890, p. 218).

Dimensions :—

Sitomn, CAM etie re weempes aay eh os cas nan setens 0045 mm.

Ly droihecn mney Maer, cera res sa0sics 0:024-0:030 mm.

ms diameter at margin ...... 0:099-0°108 mm.

The dimensions of the Mergui specimens are considerably

smaller than those of British examples, ‘

Localities. Epizoic on Thyroscyphus vitiensis from Stations 15

and 16, Ravenshaw Island, Sir John Malcolm Island, and Alligator

Rock, 5 to 18 fathoms, rock and sand, or rock and mud.

Recorded with some doubt from Eastern Australia by Bale

(1884, p. 65) and Lendenfeld (1885, p. 405), this widely dis-

~~.

1910.] FROM MERGUI ARCHIPELAGO. 809

tributed species has been described from the Indian Ocean only

by Billard, from Macalonga, Mozambique, 22 metres (1907 0,

p. 338).

Family CAMPANULARIDS.

CAMPANULARIA CoRRUGATA Thornely.

Except at Station 1, where it was also associated with Sertu-

larella quadridens and Diphasia digitalis, this species always

occurred as an epizoon on Jdia pristis.

Localities. St. 1, east of Tavoy Island and Port Owen, 4 to 12

fathoms, sand and broken shells, and mud; short and stout,

strongly ribbed hydrothec on Diphasia digitalis, and long, cylin-

drical individuals on Jdia pristis. Stt. 15 and 16, Ravenshaw

Island, Sir John Malcolm Island, and Alligator Rock, 5 to 18

fathoms, rock and sand, or rock and mud; occasionally. St. 22,

Hastings Harbour, shore to 20 fathoms, rock and sand; fairly

common. St. 23, Five Islands, 8 to 12 fathoms, rock and sand,,

and mud; not common, St. 25, Gregory Group and Crichton

Island, 4 to 14 fathoms, stones and broken shells and rock: rare

eylindrical hydrothece with short stalks. St.35, between Warden

Island, Howe Island; and Lyall Island, 15 to 20 fathoms, rock

and sand; rare.

CAMPANULARIA RARIDENTATA Alder, 1857.

From British examples the majority of the specimens here

referred to C. raridentata differ in lacking the swelling which

intervenes between the base of the stalk and the stolon, in this

respect resembling examples recorded from areas so wide apart as

Calbuco, South America (Hartlaub, 1905, p. 567), St. Malo, France

(Markt.-Turner., 1890, p. 205), and the Indian Ocean (Hincks,

1889, p. 133, cf. pl. xii. fig. 5). From seven to eleven rings occur

at the base of the stem, and from three to five underneath the

hydrotheca, and sometimes a few annulations mark also the

middle portion of the stem. The margin of the hydrotheca

appears to be divided into from five to seven large teeth, the

extreme tenuity and transparency of which render them difficult

to observe. The hydranth bears twelve to thirteen tentacles,

No gonosome was observed.

The dimensions of the Mergui specimens are much less than

those of St. Malo specimens recorded by Marktanner-Turner-

etscher (1890, p. 205), and approach most closely those of a single

example from Saint-Vaast described by Billard (1907 @, p. 173),

with which they also agree in possessing few hydrothecal teeth.

Dimensions, in mm. :—

Simals St. 23:

Stem, lengtitsseeeeesttea ne cee cee eee 0°59 0:35-0:45

yo) GAME REee hose latoel eee: JOS In 0-07 0°04—0-06

ibiydrothecay depthitie? 2h Af: 2. ak 0°50 0:29-0:41

s diameter at mouth ...... Oki SOds=Osi5

810 MR. J. RITCHIE ON HYDROIDS [May 24,

Localities. St.1, east of Tavoy Island and Port Owen, 4-12

fathoms, sand anit broken shells, and mud; rare. St. 25, Five

Islands, 8 to 12 fathoms, rock and sand, and mud; rare. St. 35,

between Warden Island, Howe Island, and Lyall Island, 15 to 20

fathoms, rock and sand ; rare.

Except from St. 23, where this species also occurs on Sertwarella

quadridens, specimens are confined to /dia pristis.

From each of Stations 1 and 28 comes a solitary simply-stalked

hydrotheca with cone-shaped basal portion, sur mounted by almost

cylindric: al walis terminating in seven or eight sharp teeth. The

structure of the basal portion resembles that of the hydrotheea of

Obelia andersoni Hincks, but I regard the present specimens as

abnormally developed examples of C. raridentata.

HEBELLA CALCARATA A. Agassiz, 1865.

One of the most widely distributed of the species in the collee-

tion, occurring at eight of the fourteen stations from which

Hy droids were attained: It shows considerable diversity of

form, being usually represented by the long smooth hy drothece of

the typical form, but frequently assuming he shape of the variety

recorded by Marktanner (1890) as Lafoéu contorta. This epizoie

species, like several others, shows a remarkable constancy of

preference for /dia pristis, seeing that at all stations, with the

exception of Station 9, where it was creeping on Thyr oscyphus

vitiensis, it occurred on that species, and on it alone.

Localities. St. 1, east of Tavoy Island and Port Owen, 4 to 12

fathoms, sand and broken shells, and mud; not common, often

var. contorta. St. 9, between Bentinck Island and Courts Island,

12 to 26 fathoms, sand and shells. St. 14, Bushby Island

pearling-ground, shore to 21 fathoms, sand and mud; rare.

Stt. 15 and 16, Ravenebare Island, Sir Tole Maleolm Island, and

Alligator Rock, 5 to 18 fathoms, rock and sand, or rock and mud ;

occasionally present, often assuming the shape of var. contorta.

St. 23, Five Islands, 8 to 12 fathoms, rock and sand, and mud;

very common, mostly var. contorta. St. 25, Gregory Group and

Crichton Island, 4 to 14 fathoms, stones and broken shells, and

rock ; very rare. Moskos Telands, 3 to 26 fathoms, rock and

sand, or rock and mud; common.

FEBELLA CRATEROIDES Ritchie, 1909 4.

The hydrothece are smaller than those of the type specimens

from the Andaman Islands. The hydranths are decayed, and no

gonosome was observed.

Dimensions :—

Hy drothecagdepum mater ene... in -ss- 0:20 mm.

" diameter at mouth......... 0-15-0'17 min.

Localities. Stt. 15 and 16, Ravenshaw Island, Sir John Maleolm

Island, and Alligator Rock, 5 to 18 fathoms, rock and sand, or

vock and mud; rare, on Lytocarpus pheniceus.

1910.] FROM MERGUI ARCHIPELAGO. 811

THYROSCYPHUS REGULARIS Jiderholm, 1896. (Plate LX XVII.

eS) a

(7. equalis Warren, 1908.)

TropHosoMe. The specimens are somewhat larger than those

originally described, sometimes almost 20 em. in length, while the

branches may reach a length of 45 mm., in place of the maximum

of 18 mm. given by Jiiderholhn.

The species is readily distinguished from 7’. vitiensis by the

presence of a very distinct operculum, accompanied by deep bays

round the margin; further, the bright rim runs in this case

much nearer to the edge, and is not so pronounced, while the faint

ringing at the internodes is more marked. The joints, varying

in number, which are said to occur on the peduncle, are only

occasionally present (though sometimes up to three or four were

observed), and even then they are not essential to the specific

structure, for they are due to the destruction of the hydrotheca,

and to the subsequent regeneration of a new hydrotheca from

within the remains of the old one, the apparent joints being the

reliquiz of former calycles. No specific value can attach to

these accidental “joints,” the like of which I have already

observed in Thyroscyphus simplex = tridentatus (1909, p. 75,

fig. 1b) and in Lytoscyphus marginatus; but in 7’. ramosus the

semblance of a twist occurs in the peduncle.

The minute structure of the ccenosare is obscured by the

presence of opaque granules which occur in great abundance. In

these specimens the hydranth is attached, not to a continuous

ridge, as in the specimens from Natal, which Warren has de-

seribed as 7”. equatis (and which I am unable to distinguish from

T. regularis), but to a row of small chitinous prominences which

runs round the mside of the hydrotheca in an exceedingly

sinuous curve, having two maxima, one on the adcauline, the

other on the abcauline surface, and two lateral minima.

GonosomE. Gonangia, which have not previously been described,

were found on a single colony from Station 25 (Pl. LXXVII.

fig. 7). The ceenosare has disappeared, but the perisare is in the

form of a long cylinder, larger even than that of 7’. torresii (see

Jaderholm, 1903, p. 2738). Very indefinite and irregular wrinkles

appear on the perisare. The insignificant stalk of the gonangium

arises from the projection of the ‘internode upon which a hydro-

theea is perched

Dimensions :—-

Branch, dramietetamaas:. he -c neces ae 0°36 mm.

Distance between hydrothece ......... 1°13 to 1:3 mm.

Hl ydrothe@ea ed Speyer. co cecscts ame ee 0-96 mm.

- bre path Sie TaaVOKvITAY eae 0°63 mm.

Peduncle, diameters 5.02. sues escivses 0-20 mim.

Gonangium, cnet ve con One Cae aces 3 mm.

3 [email protected] : Sic eer. 1 mm.

Localities. A large bunch of colonies from St. 22, Hastings

812 MR. J. RITCHIE ON WYDROIDS [May 24,

Harbour, shore to 20 fathoms, rock and sand. St. 23, Five

Islands, 8 to 12 fathoms, rock and sand, and mud; two colonies.

St. 25, between Warden Island, Howe Island, and Lyall Island,

15 to 20) fathoms, rock and candi: many colonies,

This species has already been recorded from the China Sea

(Jiiderholm, 1896, p. 9), and from Bird Island, Algoa Bay, Cape

Colony (Warren, 1908, p. 344, as 7’. equalis).

THYROSCYPHUS VITIENSIS Marktanner-Turneretscher, 1890.

A widely distributed species, occurring at eight of the fourteen

stations from which Hydroids were obtained. In this, as in

7. reqularis, the hydranth is attached toa row of minute chitinous

prominences on the inside of the lower portion of the hydrotheca.

These are arranged as in the previous species, but they are more

strongly developed, and are placed on a slightly raised portion

of the hydrothecal wall.

Localities. St. 1, east of Tavoy Island and Port Owen, 4 to 12

fathoms, sand and broken shells, and mud; common, St. 3,

French Bay, King Island, and south end of tron Island, 3 to 8

fathoms, mud and rock, or sand; common. St. 9, between

Bentinck Island and Courts Island, 12 to 26 fathoms, sand and

shells; rare. St. 14, Bushby Island pearling-ground, shore to

21 fathoms, sand and mud; rare. Stt. 15 and 16, Ravenshaw

Island, Sir John Malcolm Island, and Alligator Rock, 5 to 18

fathoms, rock and sand, or rock and mud; rare. St. 19, Paye

Island and Pink Island, 7 to 9 fathoms, rock and sand; rare.

St. 25, Gregory Group and Crichton Island, 4 to 14 fathoms,

stones and broken shells, and rock; fairly common. St. 35,

between Warden Island, Howe Island, and Lyall Island, 15 to

20 fathoms, roek and sand ; rare.

Family CAMPANULINIDS.

OPERCULARELLA LACERTA Johnston, 1847.

Identification depends upon trophosome characters alone, the

gonosome being absent. I can detect nothing, however, which

could distinguish these specimens from North Sea examples of

the above species. It occurs in its simplest form, a creeping

stolon, sending up here and there short ringed stalks, on each of

which is poised an exceedingly hyaline hydrotheca I

Dimensions :—

Hydrotheea, depth *.................. 0:21 mm.

i, diameter ............+0. 0:08-0:09 mm.

Peduncle, diameter ooo. .....ccseee ees 0:04 mm,

Locality. Moskos Islands, 8 to 26 fathoms, rock and sand, or

rock and mud; rare, growing on a stem of Plumularia setacea.

Although the range of the species is a wide one, for it has

been recorded from the North of Europe, from both sides of the

* From top of operculum.

TSLOM FROM MERGUI ARCHIPELAGO, 813

North Atlantic Ocean, and from the Western Pacifie (China and

Kastern Australia), it has not hitherto been found in the Indian

Ocean,

CALYCELLA OLIGIsTA*, sp.n. (Plate LR XVI. figs. 3 & 4.)

TropHosomE. The colony is epizoic, and consists of a creeping,

delicate stolon which lies in close contact with the specimen upon

which it occurs. From this stolon hydrothec spring at irregular

intervals.

The hydrothece are exceedingly minute, delicate, hyaline, and

transparent, without a trace of the smoky tint which charac-

terises C. syringa. In shape they are tubular and cylindrical, of

approximately the same diameter for the greater portion of their

length, although the proximal half is usually slightly bulged, and

tapers, more or less rapidly, to join the stalk. The margin of the

hydrotheca, which is very slightly everted, is definite, and is of

considerable strength, for it is seldom that it is found in a

collapsed condition. It is divided into shallow crenulations,

similar to those in C. syringa, each of which subtends the base of

an opercular valve. Jn no case was a duplication of the margin

noticed. The operculum is distinct from the hydrothecal wall,

and is composed of about twelve to fourteen exceedingly delicate

valves, difficult to distingnish. The cavity of the hydrotheea is

separated from that on the stalk by a distinct but delicate

diaphragm. The stalk is usually very short, so that the hydro-

theca almost rests on the stolon, but in one case (Pl. LX XVI.

fig. 3) a well developed stalk was observed. This stalk was marked

by a few irregular indentations, but in other cases the stalks were

smooth, and ringing of a regular nature never occurred.

The hydr anth is large eompared with the size of the hydrotheca.

It is attached by a basal disk to the hydrothecal wall immediately

above the diaphragm, and is continued upwards as a cylindrical

body with conical hypostome, surrounded by a whorl of about

seven to ten tentacles. The ectoderm is thick, and the tentacles,

in contraction, are dumpy and stout.

The gonosome was not observed.

Dimensions :—

Stolon, diameter ......... WEES Se ea eee yaaa oO) fs

iydrctheca, ;depuimeensse ss .csaa.285. hn eT 120-138 p.

a greatest diameter ............... 45-66 p.

Localities. St. 23, Five Islands, 8 to 12 fathoms, rock and sand,

and mud; very rare, on J/dia pristis. St. 35, batw een W arden

Island, Howe Island, and Lyall Island, 15 to 20 fathoms, rock

and sand; very rare, on /dia pristis and Thyroscyphus vitiensis.

Systematic position.—This species differs remarkably in size,

colour, texture, and in the development of the peduncle and its

ringing from C. syringa, although there is a similarity between

* 6Xiy.oTos, smallest.

Proc. Zoou. Soc.—1910, No. LIII. 53

814 MR. J. RITCHIE ON HYDROIDS [May 24,

its hydrothece and some of the varieties of the latter species

(see Broch, 1909 6, text-fig. 22). It approaches most closely

C'. nuttingi Hargitt (1909, p. 378), from which it differs in being

smaller in size, in being cylindrical instead of tapering from

hydrothecal margin to base, and in lacking distinctly annulated

peduncles, with “annulations occasionally extending some distance

(rarely over the entire body) on the theeal walls.”

I regard the unnamed specimen, mentioned and figured by

Miss Thornely (1908, p. 83, pl. ix. fig. 5), from Khor Shinab, 10

to 12 fathoms, in the Soudanese Red Sea, as belongmg to this

species,

CusprpELta costata Hincks, 1868. (Plate LX XVII. fig, 8.)

To the short and incomplete description given by Hincks the

following observations may be added. ‘The hydrotheea is eylindri-

cal for the greater part of its length, but near the base it gradually

tapers downwards. Frequently the basal portion of the hydro-

theca lies along the object upon which the epizoon is growing,

the distal por tion bending upwards at an angle, slightly after the

manner of Lafoéa ser rata. Within the bent } portion the retracted

polyp is generally found. Similar bent hydrothece have been

described by Dr. Billard from La Hougue (1904, p. 165).

The lines of growth held to be characteristic of the species,

and clearly marked m our specimens, are not lines of growth in

the ordinary sense, marking the place where new growth has

commenced, They indicate, instead, the margins of earlier hydro-

thece, the opercular flaps of which occasionally remain attached

and give an appearance of longitudinal fluting to the succeeding

portion of the hydrotheca, as is suggested in the terminal seg-

ment in Hincks’s figures (1868, pl. xl. figs. 5, 5@). The secondary

hydrotheca (produced probably on the regeneration of an entirely

new hydranth) lies within the primary, and for some distance

their walls, though in close contact, remain distinct. The tertiary,

when such oceurs, lies within the secondary, and so on. Thus

the terminal segment. is, as Hincks observes, “ of thinner material

than the rest,” for its walls are those of a single hydrothec: a,

while proximal to the uppermost segment the walls, owing to the

telescoping arrangement, are consider vably strengthened. It is

noteworthy that the proximal segment is usually of distinctly

less calibre than its successors. The operculum of the hydrotheca

is composed of about 14 or 15 flaps.

The hydranth is strongly retractile, retirmg on contraction to

the proximal third of the hydrotheca. It appears to have about

six to eight tentacles, which are closely set with whorls of large

oval nematocysts, the whorls beng 7:5 apart. The average

size of the nematocysts im these whorls is 5-2 » long by 2 pw

broad, but at the tip of the tentacle larger examples oceur, 10°5

long by 5 » broad.

No gonosome was observed,

1910. ] FROM MERGUI ARCHIPELAGO, 815

Dimensions :—

Hydrotheca, lempibye. sees. -% 124 ---<2--- + 0-7-1712 mm.

BS CIA TINELOTRR a tyadase so 2kibiee oh 0-10-0714 mm.

Length of flaps of operculum ............ 0-11-0°183 mm.

Localities. St. 1, east of Tavoy Island and Port Owen, 4 to 12

fathoms, sand and broken shells, and mud; rare hydrothece on

Thyroscyphus vitiensis. St. 3, French Bay, King Island, and

south end of Iron Island, 3 to 8 fathoms, mud and rock, or sand ;

very rare, on U'hyroscyphus vitiensis.

Family LAro# rip &.

LAFOEA SERRATA Clarke, 1879.

A widely distributed, but moderately rare species. Repeated

regeneration of the hydranth had furnished one hydrotheca (from

St. 22) with the exceptionally large number of eleven successive

margins.

Localities. St. 1, east of Tavoy Island and Port Owen, 4 to 12

fathoms, sand and broken shells, and mud; fairly common, on

Idia pristis. St. 9, between Bentinck Island and Courts Island,

12 to 26 fathoms, sand and shell; very rare, on Thyroscyphus

vitiensis. Stt. 15 and 16, Ravenshaw Island, Sir John Malcolm

Island, and Alligator Rock, 5 to 18 fathoms, rock and sand, or

rock and mud; rare, on /dia pristis. St. 22, Hastings Harbour,

shore to 20 fathoms, rock and sand; fairly common, on 7hyro-

scyphus vitiensis and [dia pristis. St. 23, Five Islands, 8 to 12

fathoms, rock and sand, and mud; rare, on Hudendriwm attenua-

tum? St. 25, Gregory Group and Crichten Island, 4. to 14

fathoms, stones and broken shells, and rock; rave, on /dia pristis.

St. 35, between Warden Island, Howe Island, and Lyall Island,

15 to 20 fathoms, rock and sand; rare, on /dia pristis.

Laro&a venusta Allman (?), 1877. (Plate LX XVI. figs. 5 & 6.)

Various Hydroids have associated with them a small form

which I refer, with a query, to this species.

The trophosome consists of a creeping hydrorhiza from which

spring small cylindrical hydrothece, set on stout but variable

stalks, and zoned by numerous, well-defined, and regular corru-

gations. These decrease in distinctness towards the base of the

hydrotheca. The margin of the hydrotheca is everted, and the

stalk bears no regular rings. A marked and distinetive character

exists in the colour of the perisare which possesses a brownish-

smoke tint, slightly variable in density. An exceedingly delicate

membrane forms the floor of the hydrotheca.

The hydranth is cylindrical and minute, furnished with a

hemispherical hypostome, surrounded by tentacles varying in

number from nine to eleven. The tentacles and body of the

hydranth have a granular appearance owing to the presence

of scattered nematocysts, which do not seem, even in the tentacles,

5a"

816 MR. J. RITCHIE ON HYDROIDS [May 24,

to be arranged in whorls, and of large, deeply stainable cel!s in

the body-wall.

The habit of this epizoon is peculiar, for, in addition to the

ordinary external meandering, the ccenosare may live within the

perisarcal tube of the hydroid upon which it grows. Thus in

several cases, the hydrothece of this species project from within

the hydrothece of Jdia pristis and Sertularella quadridens. In

this respect much resemblance is shown to the habit of Lafoéa

dispolians, Warren (1909, p. 105), the wanderings of which

within the perisare of its host, Sertularia bidens Bale, have been

closely traced by Dr. Warren. In the present case. the material

being in a poor state of preservation, I have been unable to trace

the course of the parasite throughout.

Dimensions :—

Hydrotheca, length..................... 0-53-0-°84 mm.

as greatest diameter ...... 0:21-0°-27 mm.

Pedunele, GiaMeber .. .....62... cence 0:075—-0°081 mm.

Localities. St. 1, east of Tavoy Island and Port Owen, 4 to 12

fathoms, sand and broken shells, and mud; solitary hydrothecze

on Idia pristis and Sertularella quadridens. St. 14, Bushby

Island pearling- -ground, shore to 21 fathoms, sand and mud; rare,

on Idia pristis. Stt. 15 and 16, Ravenshaw Island, Sir John

Malcolm Island, and Alligator Rock, 5 to 18 fathoms, rock and

sand, or rock and mud; very rare, on /dia pristis. St. 23, Five

Islands, 8 to 12 fathorne. rock and sand, and mud; very rare, on

Sertularella quadridens.

Hitherto this species has been found only on the western

sea-board of the North Atlantic Ocean, and growing upon only

one host, Lytoscyphus marginatus Allman: Loggerhead Key,

9 fathoms (Allman, 1877), 10 miles north of “Zoblos Island

(Clarke, 1879), Anguilla, Antilles, 100-150 fathoms (Jaderholm,

1903), off Bermuda, 30 fathoms (Ritchie, 1909 @), and Prof. 8. F.

Clarke, in litteris 1909, mentions its occurrence at West Florida,

20 fathoms, again on Lytoscyphus marginatus.

Systematic position.——The general appearance of the specimens

described as Lafoéa venusta resembles that of a small parallel-

sided form of the exceedingly variable Campanularia corrugata

Thornely. As the gonosomes of both species are unknown, and

even the hydranth of the former is undescribed, the difficulties of

identification are thus greatly increased. The present specimens

are, however, specifically distinct from Campanularia corrugata,

on account of the much smaller size of their hydrothece, which

are also more regularly ringed, and are always cylindrical in

shape. The hydranths also differ in shape and in structure,

Those of the present specimens are more slender and possess 9 to

ll, in place of 22 tentacles; they have a more hemispherical

hypostome, and lack the peculiar development of those endo-

dermal cells, which, in C. corrugata, project from the base of the

tentacles into the gastric cavity, almost meeting there below the

1910. ] FROM MERGUI ARCHIPELAGO, 817

hypostome, and reminding one of the structure of Bonneviella

grandis, described by Broch (1909 a, p. 195). I have not noticed

that in Miss Thornely’s species the perisare is tinged with brown.

To Lafoéa venusta I have, with a little doupe referred my

specimens because of the shape, the small size, and the corrugated

walls of the hydrothece.

Family SERTULARID &.

SERTULARELLA CYLINDRICA Bale, 1888. Var. rusiLLA, nov.

(Plate LX XVII. fig. 9.)

Simple stems spring from a creeping stolon, and these are

divided into internodes of exceeding variable length by slightly

slanting nodes. At the distal end of each internode a hydrotheca

is borne.

The hydrothece are alternate, and have very thin, collapsable

walls, in strong contrast with the thick internodal perisare. In

comparison. with the stem upon which they are carried, the

hydrothece are very large. In shape they are cylindrical, the

cylinder expanding slightly at the mouth and forming an everted

margin, entire, and destitute of operculum. For about a third of

their length they are adnate to the internode, but beyond this

their free portion curves gracefully away from the stem, so that

the aperture faces upwards and outwards, the plane in which it

lies meeting the stem at an angle of about 45°, A slight break

in the even abcauline profile of the hydrotheca opposes the point

where the adcauline side becomes free from the stem. Often

the bottom of the hydrotheca is rounded. Stolons are occasionally

developed from the stems, and one was observed springing from

the inside of a hydr otheca,

Gonosome unknown.

Dimensions :—

Var. pusilla. Typical form.*

Internodes, length ............... 0°27-0°36 mm. | very variable.

33 diameter tecageseb ii: 0:06-0:07 mm. | up to 0°39 mm.

Hydrotheca, depth ............... 0-29-0°37 mm. | 0-80—0°84 mm.

mt diameter atmouth 0:10-0:11 mm. | 0°32 mm.

Localities. St. 32, south-west of Domel Island, 26 to 29 fathoms,

sand and mud; rare, on Lytocarpus pheniceus. Moskos Islands,

3 to 26 fathoms, rock and sand, or rock and mud; a single stem

amongst the rhizoidal tubes of /dia pristis.

The species has hitherto been found only on the east coast of

Australia, at Port Jackson (Bale, 1888, p. 765).

Systematic position.—W hile | do not find in this form characters

sufficient to separate it specifically from C. cylindrica, it can

readily be distinguished from the Australian form, examples of

which I have recently had an opportunity of examining, chiefly

* Specimens collected off New South Wales by the ‘Thetis,’ 1898, entrusted to

me for examination.

818 MR. J. RITCHIE ON HYDROLDS [May 24,

by its dwarfish size—the diameter of the stem is only +, the length

of the hydrothece Jess than 3, their diameter 3 those of

typical Australian specimens (ef. table of dimensions given above).

Var. pusilla is also to be distinguished by the slightly more

flaring margin of its hydrothece, and by its hyaline perisare, for

that of the robust typical variety is tinged with brown. It is

possible that the discovery of their gonosomes may separate those

two forms specifically.

Professor C. Hartlaub, in his fine monograph of the genus

Sertularella (1900, p. 65), regards S. cylindrica, somewhat doubt-

fully, as a synonym of S. integra Allman, described from New

Zealand specimens. The delicate texture of the hydrothecz is so

different, however, from that of the type of Sertularella which

bears intrathecal marginal teeth, that, apart from other obvious

differences of form, sculpture, and proportion, I regard the

presence of intrathecal teeth in Allman’s species as suflicient,

in this case, to indicate specific difference.

SERTULARELLA POLYZONIAS Linnzeus, 1758.

Var. cornuta Ritchie (?), 1909 6.

A single fragmentary colony, 34 mm. high, represents this

species. The hydrothece agree in structure with typical speci-

mens, but a single internode occasionally bears more than one

hydrotheca. The habit of the colony, too, is much more definite

than that of the lax variety (gracilis) of British seas, for from a

definite, thick stem spring alternate branches almost at right

angles. In this respect the specimen approaches var. robusta

Kirchenpauer (1884, p. 38), from Cape of Good Hope, and var.

cornuta, from the neighbourhood of the Andaman Islands. The

absence of the gonosome precludes certain decision as to which

variety it belongs, but the trophosome agrees in detail with that

of var. cornuta,

Dimensions :—

Stem, diameter ..... eee es Shi cee 0:63-0°70 mm.

Hydrotheca, length ot adnate portion . 0°32-0°45 mm.

a free 3 0:28-0°39 mm.

, greatest diameter ........ . 0°24-0°25 mm.

i diameter at mouth......... 0-17-0'18 mm.

Locality, St. 23, Five Islands, 8 to 12 fathoms, rock and sand,

and mud; rare.

SERTULARELLA QUADRIDENS Bale, 1884. (Plate LXXVII.

figs. 12 a, 12 6.)

The transverse chitinous projections from the bases of the

hydrothece are mostly absent, and where they do occur are much

less distinetly marked than in Bale’s figure (pl. vil. fig. 5) of a

Holborn Island specimen, The hydrothece are strictly ‘alternate,

but the internodes are very irregular in length, from eleven

to thirty-six hydrothecee occurring between successive nodes.

1910.] FROM MERGUL ARCHIPELAGO, 819

Considerable variation occurs also in the proximity of the hydro-

thee of the same lateral series, for while a marked gap separates

some—usually on the distal portions of branches—in many cases

the upper portion of one hydrotheca is pressed against the base of

the next (cf. Pl. LX XVII. figs. 12 a, 126), An abnormality in

the formation of the hydrothec is werth neting. Occasionally,

probably after some accident to the branch, the hydrothece first

formed thereafter, while normal so far as the adnate portion is

concerned, have an exceptionally long portion free (0°77 mm., as

contrasted with the average 0°l mm.), this part being tubular,

recurved, and having a rather indefinite margin, Its successor

is less, though still abnormally large, and the diminution con-

tinues until the normal size is reached after five or six hydro-

theca-pairs (text-fig, 79).

Text-fig. 79.

Abnormal hydrothece of Sevtularella quadridens, X 20.

The pinne, each of which arises beneath a hydrotheca, are

regularly alternate, three hydrothece being interposed between

two pinne on the same side. In one case a branch, of second

degree, was observed to spring from the lumen of a hydrotheca,

The structures of the soft parts of the species, to which I have

seen no reference, show points of interest. The ceenosare of the

stem is arranged in a varying number of longitudinal strands,

usually three or four, which are connected irregularly by anasto-

mosing canals, and which, with their connections, line the perisare.

In the branches the structure is less complicated. There, two

large longitudinal ccenosare tubes are to be distinguished, one

running along each side of the branch, and threading its way

820 MR. J. RITCHIE ON HYDROIDS [May 24,

between the hydrothece. These are connected at the base of

each hydrotheca by a strong bridge of coenosare, wpon the middle

of which the base of the hydranth rests. ‘These structures

closely resemble those which Nutting has observed in Selaginopsis

ornata (1904, p. 7), allowing for the differences in symmetry due

to the presence of four series of hydrothecze in that species, in

place of two in this. Although each tube possesses a thick

definite ectoderm, I have been unable to distinguish the perisare

which, Nutting supposes, surrounds each ccenosarcal tube.

The structure of the polyp is like that of Thuiaria robusta, as

figured by Nutting (1904, p. 11, fig. 15). It has a large blind

sac, and clumps of large endodermal cells project into the

hydranth cavity except from the abcauline wall of the blind-sae,

where the development of the endoderm is insignificant. The

protractor band is attached, not to the abcauline wall of the

hydrotheca, but to varying points of the lateral wall in the

neighbourhood of the margin of the hydrotheca.

The gonosome was not observed.

Dimensions :—

Stem, diameter ...... Ee eae 0:56-0°77 mm,

Stem internode, length ...,,............. 2 mm.

Hydrotheca, length adnate...,.....,..... 0:46-0°49 mm.

a 43 DAC Re eet 8 0:07—0'14 mm,

* diameter at mouth ...... 0:21-0:22 mm.

Localities, St. 1, east of Tavoy Island and Port Owen, 4 to 12

fathoms, sand and broken shells, and mud; several fragments.

St. 23, Five Islands, 8 to 12 fathoms, rock and sand. and mud;

one colony. St. 25, Gregory Group and Crichton Island, 4 to 14

fathoms, stones and broken shells, and rock ; one colony.

Previously known only from Eastern and Northern Australia ;

Port Curtis and Holborn Island (Bale, 1884), Peel Island,

Moreton Bay (Bale, 1888), Flinders Passage, Torres Strait

(Allman, 1888, as Zhuiaria vincta*), Thursday Island (Weltner,

1900).

Ipra pristis Lamouroux, 1816,

One of the commonest of the species in the collection, found at

eight of the fourteen stations at which Hydroids were obtained,

There is much variability in the length of the free portion of the

hydrotheca, the sloping opereulum appearing, in some cases, to

project directly from the surface of the stem, while in others a

large part of the hydrotheca stands out at right angles to the

stem. Occasionally, too, the adcauline operculum is lacking in

definiteness, the upper wall of the hydvotheca simply meeting

the lower as an indefinite flap. Many of the branches from

Station 35 end in stolons, indistinguishable from those which

project from the hydrorhizal tubes.

* See Billard, 1908, p. 2 of separate copy.

1910.] FROM MERGUI ARCHIPELAGO, 821

Localities. St. 1, east of Tavoy Island and Port Owen, 4 to 12

fathoms, sand and broken shells, and mud; common, St, 14,

Bushby Island pearling-ground, shore to 21 fathoms, sand and

mud; common. Stt. 15 and 16, Ravenshaw Island, Sir John

Malcolm Island, and Alligator Rock, 5 to 18 fathoms, rock and

sand, or rock and mud; common. St. 22, Hastings Harbour,

shore to 20 fathoms, rock and sand; common, with gonangia,

some colonies growing on a sponge. St. 23, Five Islands, 8 to

12 fathoms, rock and sand, and mud; fairly common. St. 25,

Gregory Group and Crichton Island, 4 to 14 fathoms, stones and

broken shells and rock; common. St. 35, between Warden

Island, Howe Island, and Lyall Island, 15 to 20 fathoms, rock

and sand; one colony. Moskos Islands, 3 to 26 fathoms, rock

and sand, or rock and mud; common, with gonangia.

DrpHasta DiGiTaALis Busk, 1852.

In these specimens it is clearly seen that the two so-called

opercular muscles are attached, not to the valves of the operculum,

but to the lateral walls of the hydrotheca near the margin, and

are, in function, protractor muscles (see Nutting, 1904, p. 13,

fig. 17). The hydranth possesses about twenty tentacles.

Locality. St. 1, east of Tavoy Island and Port Owen, 4 to 12

fathoms, sand and broken shells, and mud ; several small colonies,

on the bare axis of an Aleyonarian, and on /dia pristis.

Recorded from the Western Indian Ocean— Maldive Islands—

by Borradaile (1905, p. 842); this is the first record from the

Eastern Indian Ocean.

SERTULARIA TURBINATA Lamouroux, 1816.

=. loculosa Busk 1852 *.)

Several small, unbranched colonies of this species occur upon

Thyroscyphus vitiensis. They are pale in colour, in this, as well

as in the shortness of the internodes, agreeing with the specimens

described from Paumben, India, by Jiderholm (1903). But they

differ in the reduction of the lateral teeth, which are occasionally

so indistinct that the aperture appeurs to be almost round.

Besides an indistinct tooth on each flank, the hydrotheca is

surmounted by a small third tooth, from the summit of which a

membranaceous edge sometimes runs to the lateral teeth. Not-

withstanding difficulties of observation, I feel assured, after

examining many hydrothece, that the operculum is formed of a

solitary flap, hinged on the distal edge of a slight thickening

which occurs on the. abcauline margin of the hydrotheca.

Although membranes unite the superior with the lateral teeth,

these do not hinge inwards, and can scarcely, therefore, be

accounted part of the operculum. They are the less necessary

since the abcauline flap is of diameter sufficient completely to

close the aperture of the hydrotheca.

* Fide Billard (1909, p. 322), who has examined the type specimen of Lamouroux,

822 MR. J, RITCHIE ON HYDROIDS [May 24,

In the structure of the operculum S. turbinata appears to

differ from S. versluysi Nutting (1904)—regarded by Billard

(1908) as a synonym—for in the latter, as Congdon states (1907,

p. 482), and as careful examination of the lateral aspect of speci-

mens from Cape Verde Islands, in my collection, shows, the

operculum is furnished with a large abcauline and two smaller

latero-adcauline valves.

The protractor of the hydranth, which possesses only a small

blind-sae, is attached to the hydrothecal wall at the intrathecal

ridge.

Localities. Stt. 15 and 16, Ravenshaw Island, Sir John Malcolm

Island, and Alligator Rock, 5 to 18 fathoms, rock and sand, or

rock and mud; not common.

Family PLUMULARID A,

PLUMULARIA SETACEA Linneus, 1758 (¢).

A single immature colony, 15 mm. high, identical in all respects

with North Sea specimens. I have w en the designation with

a mark of interrogation on account of the impossibility of dis-

tinguishing the trophosome of this small variety from that of

P, strictocarpa Pictet (1893), from Amboyna.

Locality. Moskos Islands, 3 to 26 fathoms, rock and sand, or

rock and mud,

PLUMULARIA Sp.

An unidentifiable fragment, the structures of the stem and

hydroclades of which resemble those of P. setacea, but the frag-

ment shows traces of branching, and the hydrotheca of having

a slightly concave contour.

Locality, St. 32, Hastings Harbour, shore to 20 fathoms, rock

and sand.

ANTENELLA SECUNDARIA Gmelin, 1788-98.

The nematocysts of the nematophores measure 12 y by 4 p,

agreeing with Warren’s measurements of those in A, natalensis,

1.2 ~ my

which I regard as a synonym, There ave about 18 tentacles.

Locality. St. 1, east of Tavoy Island and Port Owen, 4 to 12

fathoms, sand and broken shells, and mud; not common, on

Idia pristis.

LyvrocaRPUS PENNARIUS Linneus, 1758,

A widely distributed species, represented by several fine

colonies, one of which reached a height of 95 em, Apart from

the prominent anterior tooth, that on the margin of the hydro-

theca second from the stem is distinctly the largest. It forms

the highest point on the outwardly curved margin of the hydro-

theca, ‘There is, however, some variation in the prominence of

1910.] FROM MERGUI ARCHIPELAGO, 823

the teeth in general. The margin of the mesial sarcotheca is

slightly sinuated.

The attachment of the hydroclades to stem and branches is

weak, for, whenever an attempt is made to clear the specimen

of flesh, by immersion in a solution of sodium hydrate, the hydro-

clades invariably fall off.

The soft parts of the specimens from Station 9 are well pre-

served, as the preservation of Plumularians in general collections

goes. The tentacles are about sixteen in number. The whole of

the inside of the hydrotheca, practically to the margin, is lined

with a thin layer of ectoderm, with which the hydranth is in

connection by means of exceedingly numerous strands which,

proceeding from its ectodermal layer, give the appearance of

filling the space between hydrotheca and hydranth with a delicate

open meshwork. An opening leads from the hydrotheca cavity

to that of the mesial sarcotheca, immediately proximal to the

point where the sarcotheca joins the hydrotheca, and through

this passes a strand of ccenosare, connecting sarcostyle and

hydranth.

A hydrotheca was observed, abnormal in lacking a mesial

sarcotheca.

Localities. Specimens from Stt. 9, 18,19, 25 bore phylactocarps.

St. 9, between Bentinck Island and Courts Island, 12 to 26

fathoms, sand and shells; one colony. St. 18, west and south-

west of Paye Island, 10 to 21 fathoms, sand, shells and rock ;

a large colony. St. 19, Paye Island and Pink Island, 7 to 9

fathoms, rock and sand; fragment. St. 25, Gregory Group and

Crichton Island, 4 to 14 fathoms, stones and broken shells, and

rock; three colonies. Moskos Islands, 3 to 26 fathoms, rock

and sand, or rock and mud; fragment.

LyTocARPUS PH@NICEUS Busk, 1852.

The hydrothecz of these specimens approach most closely those

of Bale’s fig. 2, pl. xv., 1884, although the chitinous septa and

walls are not so strongly developed in my specimens, and the

small lobe at the back of the hydrotheca is not produced into a

tooth. ‘There are two sarcothece at the base of each hydroclade

—one proximal, the other lateral—and, in addition, a sarcostyle

issues through a mere perforation with raised lips, on the anterior

of the stem-process on which the hydroclade is borne.

Localities. Stt. 15 and 16, Ravenshaw Island, Sir John Mal-

colm Island, and Alligator Rock, 5 to 18 fathoms, rock and sand,

or rock and mud; three colonies. St. 32, south-west of Domel

Island, 26 to 29 fathoms. sand and mud; one colony. St. 35,

between Warden Island, Howe Island, and Lyall Island, 15 to

20 fathoms, rock and sand; one colony. Moskos Islands, 3 to

26 fathoms, rock and sand, or rock and mud; one colony.

824 MR. J. RITCHIE ON HYDROIDS [May 24,

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”

ht)

”

EXPLANATION OF THE PLATES

PratEe LXXVI.

. Corydendrium sessile, sp.n. Complete colony. About nat. size.

S x Portion of branch with hydranths.

. Calycella oligista, sp.n. Hydrotheca and hydranth. X 230.

25.

3 9 39 99 39

Lafoéa venusta Allman (?). Hydrotheca and hydranth. X 60.

>b) » 3) >) 39

BV G19

PuatE LXXVILI.

Fig. 7. Thyroseyphus regularis Jaderholm. Portion of branch with hydrotheca

and gonangium. X 265.

8. Cuspidella costata Hincks. Hydrotheca, showing character of annular

markings. X 70.

9. Sertularella cylindrica, var. pusilla, nov. Portion of stem, with hydro-

thece. X 46.

10. Halecium simplex Pictet. Trophosome and gonosome. X 70.

11. » » Hydrotheca, showing internal chitinous pro-

jections. X 310.

12. Sertularella quadridens Bale. Showing variation in remoteness of

hydrothecw. (a) Proximal portion of branch. (4) Distal portion of

same branch, Each X 19,

826 MR. J. RITCHIE ON HYDROIDS [May 24,

4. Hydroids from Christmas Island, Indian Ocean, collected

by C. W. Andrews, D.Sc., F.R.S., F.Z.S., in 1908. By

James Rircniz, M.A., B.Sec., Natural History Depart-

ment, The Royal Scottish Museum*

{Received April 28, 1910. }

(Text-figures 80 & 81).

The present paper gives account of a small collection of

Hydroids which formed part of the Marine Invertebrate

collections obtained by Dr. Andrews during the autumn of 1908.

The smallness of the collection, however, is hardly to be taken

as an indication of the poverty of the Hydroid fauna of the

Island, for Dr. Andrews found that, at virtually all points, shore-

collecting was dangerous or impossible. The present specimens,

therefore, were obtained in a limited area: in Flying Fish Cove,

either on the shore, or on a reef laid bare at low tide.

The collection is of interest as being a first sample of the

Hydroid fauna of this isolated island ; and in spite of its

limitation, for it contains representatives of only thirteen species,

it adds four of these to the known fauna of the Indian Ocean,

The most remarkable of these additions are hydranths which

exactly correspond with those of /thizogeton nudum, described by

Broch, in 1909, from Spitsbergen.

I have to thank Mr. R. Kirkpatrick, of the British Museum,

for giving me the opportunity of examining the collection.

List of Species.

Gymnoblastea.

Family Ciayip2.

Rhizogeton nudum Broch (2)

Family EupENDRID&.

Hudendrium parvum Warren (2)

Family PENNARID®,

Pennaria disticha Goldfuss, var. australis Bale.

Family TuBuLarip&.

T'ubularia crocea Agassiz.

Calyptoblastea.

Family CAMPANULARID®,

Campanularia corrugata Thornely.

Clytia noliformis M:Crady (?)

Clytia serrulata Bale,

* Communicated by R. Kirkpatrick, F.Z.S,

1910.} FROM CHRISTMAS ISLAND. 827

Family SerruLarip&.

Sertularella polyzonias Linn.

Thuiaria tubuliformis Marktanner.

Family PLUMULARID

Plumularia buskii Bale.

Plumularia setacea Linn. (¢)

Plumularia sp. (megalocephala Allman, aff.).

Halicornaria hians Busk, var. lawa, nov.

Novres ON THE SPECIMENS.

GYMNOBLASTEA.

Family CLAVID&.

Ruizoceron NupuM Broch (?)

1909. Rhizogeton nudum Broch, Hydroiden, in Fauna Arctica,

Jena, vol. v. p. 137, fig. 1.

On the surface of a fragment of millepore there occurred a

delicate stolon from which solitary hydranths sprung at intervals.

The stolon is branched, forming a loose, irregular network upon

the substratum, and it is protected by a very thin layer of peri-

sarc, No perisare occurs on the hydranths, even at the base.

The hydranths are elongated, subcylindrical or spindle-shaped

according to the state of contraction. They are armed with

long filiform tentacles varying in number from seventeen to

twenty-four or twenty-five, and decreasing gradually in size, from

the longest which, in extension, surround the hypostome, to

short, stumpy individuals at the base. The tentacles are

scattered over all but the lower fourth of the hydranth; but

except for four oral tentacles which lie almost in one whorl, no

definite arrangement could be traced.

The soft parts are well preserved, and in the hydranth a thin

ectoderm with flattened cells could be distinguished, lined by

endoderm with elongated cells, which project for a short distance

into the digestive cavity, and within which, in some hydranths,

lie clusters of minute, deep-brown, almost opaque spherules,

probably refuse products of digestion. The internal portions of

the tentacles are filled by a septate skeletal endoderm. The

growing point of the stolon, in the one instance observed, was

formed of a thick pad of ectodermal cells, which gradually dwindled

into the insignificant layer which lines normal portions of the

stolon.

The gonosome was not observed.

Dimensions :—

Stolon, diameter.................. 0-060 mm.

Hydranth, length ............... 0-77 | 0:92 | to 2 mm,

», greatest diameter 0°17 | 0°21 | mm.

828 MR. J. RITCHIE ON HYDROIDS | May 24,

Text-fig. 80.

Rhizogeton nudum (?), X 65.

corp., dark corpuscles in endoderm of hydranth; ect., ectoderm ; end., endoderm ;

Op. growing point of stolon; p., perisare ; sé., stolon.

Locality. Flying-Fish Cove.

Owing to the absence of the gonosome, indentification with

R. nudum must remain a matter of doubt. The trophosomes, how-

ever, agree in Shape, in character, and in the number and position

of the tentacles; and Broch’s figures represent hydranths which

agree in size with those from Christmas Island, although he states

that his hydranths may attain a length of 4mm. The original

specimens of #. nudum were obtained by the ‘ Helgoland’

Expedition, in Bismark Sound, east of West-Spitsbergen, at a

depth of 35 metres.

Family EUuDENDRID4.

EvupeNDRIuM PARVUM Warren (°)

1908. Hudendrium parvum Warren, Ann, Natal Government

Mus. vol. i. pt. 3, p. 272, text-fig. 1, pl. xlv. figs. 1-4.

A single fragment of creeping stolon, from which spring two

1910. ] FROM CHRISTMAS ISLAND. 829

simple stems, and one, showing traces of a branch, occurred on the

surface of a fragment of millepore. The gonosome is absent, and

the characters of the hydranths have all but disappeared, but the

perisare of the trophosome shows the structures described by

Warren. The stolon measures 102”, and the hydrocauli from

8lu to lly, in diameter, but the unbranched hydrocauli are

very short, less than 2 mm. in height.

Locality. Flying-Fish Cove.

Family PENNARIDA,

PENNARIA DisticHA Goldfuss (= P. cavolinii Ehrenb.), var. aus-

TRALIS Bale.

*1820. Pennaria disticha Goldfuss, Handb. der Zool., Nurnberg,

p. 89.

1884. Pennaria australis Bale, Cat. Australian Hydroid Zooph.,

Sydney, p. 45. :

Several well-preserved colonies of this variety, distinguished

from typical P. disticha by the restriction of the annulation of the

peduncles to six, or fewer, rings at the base, and by the small size

of the colonies, were obtained. No gonosome is present, and

the specimens are overgrown by seaweed and diatoms.

Locality. Flying-Fish Cove,

Family TUBULARID &

TUBULARIA CROCEA Agassiz.

1862. Parypha crocea Agassiz Contr, Nat. Hist. U.S., vol. iv.

p- 249.

A considerable number of specimens of a small 7ubularia, with

stems up to 25 mm. high, show the following significant

characters. The stems are white, becoming faintly tinged with

yellow at the base. They are mostly unbranched, or have a few

straggling offshoots at tha base, and they are marked here and there

by groups of faint annulations. The hydranth is translucent, set

above a marked dilatation, the lower part of which is protected by

thin, wrinkled perisarc, while on the upper portion the ectoderm

is markedly thickened, appearing as if it hung downwards from

the base of the hydranth in a short flap. In this respect, and in

the separation of the hydranth cavity from that of the dilation,

the structures are similar to those of 7. bethe Warren, but the

ectoderm of the present specimens is also much thickened at the

neck beneath the dilation, and the endoderm lining the dilation

does not exhibit the development of elongated cells which Warren

describes?.

In these specimens the oral tentacles number about 15 to 17,

and the proximal tentacles about 18 to 20, set in single whorls.

* For synonymy see Bedot, Rev. Suisse de Zool., vol. ix. 1901, p. 459,

+ Warren, Ann. Natal Government Mus., vol. i. 1908, p. 282.

Proc. Zoor. Soc.—1910, No. LIV.

830 MR, J. RITCHIE ON HYDROIDS | May 24,

The blastostyles are short, generally erect, although some tend to

droop through the row of tentacles, and the +y are closely set with

male gonophor es, chavacterised by their elliptical shape, and by the

replacement of tentacles rudiments by a single terminal wart,

Unfortunately, female gonophores have not ioen observed. The

characters, however, point to identity with 7’. erocea, the male

gonophores of which, according to Allman, are ‘ destitute of

[tentaculiform] processes,” although Torrey’s observations show

that the presence or absence of tentacular processes is vaviable*,

Dimensions :—

SUGIN, UAMLOUGDS tr Reem Oey anc <s nesisins pisenneete 0-3] mm.

Hydranth, height (from ‘base of dilation)... about 3 mm.

* diameter at level of proxim: al

WOMOBOlGS dees ves. .s olde lee sitata about 2 mm.

Locality. Reef, Flying-Fish Cove.

7’. crocea Nas heen found on the Atlantic (Agassiz) and Pacific

(Torrey) coasts of North America, and in Plymouth Harbour on a

ship from Peru (Browne). Specimens were obtained growing

amongst barnacles on the screw-casing of the German Antarctic

Expedition ship ‘Gauss,’ while it was in the South Atlantic, and

Vanhéffen is of opinion that the specimens settled on the ship

during its passage through the south of the Indian Ocean*,

in which, until the present record, the species had not been

observed.

CALYPTOBLASTEA.

Family CAMPANULARIDA.

CAMPANULARIA CoRRUGATA Thornely.

1904. Campanularia corrugata Thornely, in Pleninag, Ceylon

Pearl Fisheries, Part 11. p. 114, pl. 1., fig. 2, Suppl.

Rep. viii. Hydroida.

A couple of hydrothece were observed on Plumularia buskii

Bale.

Locality. Flying-Fish Cove.

CiyriA Notirormis M‘Cracy (2).

1876. Campanularia noliformis M*Crady, Proc. Elliott Soe.

vol. i. p. 194.

A few hydrotheci, referred to this species, were growing upon

a fragment of sponge, and on Pennaria disticha, var. australis.

The hydranths bear twenty tentacles. Without the presence of

the gonangia the identification of this species is uncertain, owing

to the resemblance of the trophosome to that of Clytia johnstoni,

* Torvey, Univ. California Pub., Zool., vol. i. p. 44.

+ Vanhitfen, Deutsche Siidpolar-Exp, 1901-1903, vol. xi. Zool. iii, Berlin 1909:

die Hydroiden, p, 279.

1910.] FROM CHRISTMAS ISLAND. 831

but I have been guided by the very small dimensions of these

specimens, which are even less than those of specimens recorded

by Pictet from Amboyna*, and by Vanhoffen from the South

Atlantict, and approach most closely in size the original speci-

mens of Campanulari Z Calkins, regarded by Vanhoffe

: Jampanutaria atlenudta Calkins, regarded by Vanhotten

(I. ¢.) as identical with Clytia noliformis.

Dimensions :—

otallen athe ate seies fe eds cks ss 1:5-2 5 mm.

Stem, diameter ......... Pas sdesies beet sdton 0:054—0:066 mim.

El ydrotheca, Cepehy cares toi ice ons ae 0:32-0'45 mm.

= diameter at mouth ...... 0°21-0°28 mm.

Locality. Flying-Fish Cove,

This species has not hitherto been recorded from the Indian

Ocean.

CLYTIA SERRULATA Bale.

1888. Campanularia ? serrulata Bale, Proc. Linn. Soc. N.S

Wales (2), vol. iil. p. 757, pl. xii. fig. 4.

A single poor colony, growing along with Plumularia setacea

Linn., the str uctures and dimensions of which agree with those

given ‘py authors.

Locality. Flying-Fish Cove.

This species has not hitherto been recorded from the Jndian

Ocean, although Vanhoffen has recently described specimens from

Observatory B: ay, Kergueleng.

Family SERTULARID&.

SERTULARELLA POLYZONIAS Linn,

1758. Sertularia polyzonias Linneeus, Syst. Nat., ed. x. p. 813.

A solitary colony was found on the stem of Pennaria disticha, var.

australis. Although its structures appear to be indistin cuishable

from those of J. polyzonias, this specimen is exceedingly minute,

only 3 mm. high, reminding one of the small New Zeal: and

variety figured by Hilgendorf $; and the dimensions of the hydro-

thec are also very much smaller than those of typical specimens.

The hydranth has about twelve tentacles.

Dimensions :—

Internode, lemethe# ¥ oe. thet decd owt 0:28-0°35 mm

Pe diameter ......... SR MCOCE a 0-08 mim.

Hydrotihecamlenetbi 7. §o.220sci.4e. ones 0:28-0°31 mm

js Venu arees ao. fil e.2k. 8: 0°16-0°18 mm.

- diameter at mouth ...... 0-13 mm.

Locality. Flying-Fish Cove.

* Pictet; Rey. Suisse de Zool., vol. i. 1893, p. 31.

if Vanhoffen, op. cit. p. 299.

IGA anhétffen, op. cit. p. 802.

§ Hilgendorf, Trans. New Zeal. Inst. vol. xxx. 1898, pl. xx. fig. 2a,

832 MR, J. RITCHIE ON 1YDROIDS [May 24,

'THUIARIA TUBULIFORMIS Marktanner.

1890. Dynamena tubuliformis Marktanner-Turneretscher, Ann.

k.k. Hofmus. Wien, vol. v. p. 238, pl. iv. fig. 10.

Only a few colonies of this species, the largest 15 mm. high,

were found. The structures are typical, and agree more closely

with Marktanner’s and with Nutting’s* figures than with those

of Natal specimens by Warren‘, in which the hydrothece are

shown to be less crowded, and are more seldom arranged in groups

of several pairs. Our specimens show all the peculiarities of

arrangement of hydrothece on the stem internodes which Mark-

tanner cites. Most commonly, as in Nutting’s specimens, each

stem-internode bears ‘a branch and two hydrothecz on one side

and asingle hydrotheca on the other,” but occasionally, as Clarke

also has found, a hydrotheca-pair is added to this typical inter-

node, and occasionally a hydrotheca-pair is subtracted and

isolated on a separate internode, leaving one hydrotheca with

its associated branch-process alone on an internode.

Dimensions :—

Length, of Coloniyiyr sn. ...0.-+..-¥ iivwedenecinee, | Wp. tO) LD Tain

Typical stem internode, length ............ 1:06 mm.

Stem internode, diameter .................- 0'18-0°31 mm.

Branch, lemephretaag saan 0. <-s- 058 Seeks about 5 mm.

Hydrotheca, length of adnate portion...... 0-42 mm.

" ss free Sales Sian 0-11-0'15 min.

te GIR OLG ninco. o0 seeming ia aaa 014 mm.

Locality. Reef, Flying-Fish Cove. ’

This species has now been found in the Eastern Pacific Ocean

(Perico Island, Gulf of Panama—Clarke, 1907); in the Western

Atlantic Ocean (Bahia; Florida ; Bahama Banks—Nutting, 1904) ;

in the Eastern Atlantic Ocean (St. Thomas Island—Billard, 1907) ;

in the Red Sea (Jidda—Marktanner, 1890; Suez Bay—Thornely,

1908); from the West Indian Ocean (Natal—Warren, 1908); and

now from the East Indian Ocean. Pictet has recorded from

Amboyna, in the Hast Indies, specimens which he has named

Sertularia vege (Thompson), as a synonym of which he ranks

Dynamena tubuliformis Marktanner §. With Jiiderhotm ||, 1 am

of opinion that these specimens are not to be identified with

7’. vege. "They clearly belong to 7. tubuliformis.

Family PLUMULARIDAZ.

PLUMULARIA BUSsKIT Bale.

1884. Plumularia buskii Bale, Cat. Australian THydroid Zooph.,

Sydney, p. 124, pl. x. fig. 3, pl. xix. figs. 34, 35.

Several colonies occurred growing ona sponge. They are much

* Nutting, “ American Hydroids,” Spee. Bull. Smiths. Inst., Part 2, 1904, pl. xi.

figs. 1, 2, 3.

+ Warven, op. cit. p. 315, text-fig. 12.

+ Clarke, Mem. Mus. Comp. Zool. Harvard, vol. xxxv. 1907, p. 14.

Pictet, Rev. Suisse de Zool. vol. 1. 1893, p. 4.

Jiaderholm, Kungl. Svenska Vet.-Ak. Handi. vol. xly. 1909, p. 91.

p *

: ce

1910. ] FROM CHRISTMAS ISLAND 833

smaller than the two or three inch high specimens deseribed by

Bale, and even than Miss Thornely’s immature Ceylon examples ;

for the largest is under 15 mm, in height. On account of the

absence of intermediate athecate internodes from the hydrocaulus,

and of the presence of a minute median nematophore, with small

and delicate pedunculate sarcotheca, in the angle behind the

hydrotheea, the species can readily be distinguished from the very

similar P. cornucopie Hincks, and P, alternata Nuttmg. Hart-

laub* has described a specimen from Laysan, in which the minute

sarcotheca immediately behind the hydrotheca is assuredly absent,

and which, notwithstanding, he assigns to P. buskii; but I regard

the specific identity of the two forms as doubtful.

The following observations supplement Bale’s description, The

lower portion of the stem consists of a varying number of nodes

separated by transverse joints, the lower nodes being altogether

destitute of appendages, while those nearest the cladate portion

of the stem, from which they are separated by an oblique node,

bear a series of several sarcothece. The hydroclades spring alter-

nately from one side or the other of a hydrotheca. In rare cases,

opposite hydyoclades arise from the proximal cladate internode,

as in Miss Thornely’s specimens. The hydroclades rest on a very

insignificant process of the internode, which coincides with the

base of the lateral nematophore. A strong constriction separates

this stem-process from the hydroclade, although these specimens

do not show the excessive tendency to deciduousness of pinnz

which Bale noted.

At the origin of the hydroclades there are generally two very

short internodes (although rarely only one is present) without

appendages, and with transverse nodes; and these are succeeded

by a longer internode which bears, on its anterior surface, a single

pedunculate sarcotheca on a swelling near its proximal end, and is

limited by a proximal transverse node, and by a distal oblique

node which separates it from the first thecate internode. In these

specimens the hydroclade is very distinctly divided, by transverse

and oblique joints, into intermediate internodes furnished with a

single nematophore, and hydrothecate internodes; and although

Bale describes “a hydrotheca on each, except the first [inter node|

of each pinna,” his figure (pl. x. fig. 3) indicates the presence of

‘in obscure horizontal constriction immediately above the hydro-

theca. Only very exceptionally does the fusion of an intermediate

with the preceding hydrothecate internode oceur in these hydro-

clades, a phenomenon recorded by Billard in the lower portion of

the hydrocaulus of the closely related P. cornucopic f.

While emphasizing the fixedness of the median sarcothece in

his original description, Bale later regarded only the sarecotheca

beneath the hydrotheca as fixed. I am inclined to agree with

the earlier statement, as the large area of their attachment, and

Hartlaub, Zool. Jahrb. Syst., vol. xiv. 1901, p. 374, pl. xxi. figs. 22, 32, 36.

Billard, Exp. Se. ‘ Travailleur’ et ‘Talisman,’ vol. vii. 1907, p. 207.

Bale, Trans. Roy. Soc. Victoria, vol. xxii, 1886, p. 94.

t$+ %

834 MR. J. RITCHIE ON HYDROIDS [May 24,

the invariable position they assume, seem to show that there is

no movability in any of the median sarcothece.

The colonies are mature, male gonangia arising from the stem

at the proximo-lateral portion of a hydrotheca, on the side other

than that from which a hydroclade springs. They are eclub-

shaped in lateral aspect, resembling the gonangia of P. geminata

Allman, and are furnished with a stalk consisting of two short

internodes, each of which has a thick adcauline wall, from which,

in the second, a diminutive chitinous ledge projects inwards.

Although a nematophore usually occurs on each side of the

gonangium, occasionally only one side is furnished with an

individual,

Dimensions :—

Stem internode, length ..............0sceerescoee 0°35-0°43 mm.

5 dimater Bee urs gcc se Spats as mia 0-07-0711 mm.

Hy Gnoclade; lereu beet ce rire «css <= sone een up to 2 mm.

ry theeate internodes, length*...... 0-25-0°34 mim.

bs intermediate _,, er bli she . 018-021 mm.

Ey drotheca emilee starr: ssnch.--yneseueen sna 0:21-0°22 mm.

by diameter at mouth ............... 0-17-0°18 mm.

Gonangium (male), length..................2.+++ 0°36-0°42 mm.

a ee eatest diameter in lateral

BSIOCD oe ac scecuit armas 0:17-0°18 mm.

Locality. Growing on a sponge from the Reef, Flying- Fish

Cove.

The only previous Indian Ocean record is that of Miss Thornely,

from Gulf of Manaar.

PLUMULARIA SETACEA Linn. (¢).

1758. Sertularia setacea Linnzeus, Syst. Nat. p. 813.

Many small colonies apparently belonging to this widely-dis-

tributed species, and averaging 15 mm. in “height, were found

growing on a sponge. As it seems impossible to separate this

species from P. strictocarpa Pictet by characters of the trophosome

alone, and as the gonosome is not present on any of these speci-

mens, [ refer them ‘doubtfully, by courtesy, to the older, and cos-

mopolitan species, until the discovery of the gonothece shall fix

their identity. For guidance I append dimensions of parts.

Stem internode, length.................ce00e0 0 31-0°49 mm.

rs re dinmate EER ie 5 0°051—0°159 mm.

Hy droclade: GTA ESHD cs es fesse. heemetan meme 2-3 mm.

Hydrothe cate internode, length cit.esc0-aks 0°32-0'36 mm.

Intermediate “3 ial Clea 0-17—0°25 mm.

Hydrothechy-depihiie, f53 6... 000k Geeta 0-090 mm.

" diameter at mouth ............ 0:096-0°105 mm.

Locality. Reef and piles of the pier, Flying-Fish Cove.

* Measured from extreme points,

1910.] FROM CHRISUMAS ISLAND. 835

PLUMULARIA sp. (megalocephala Allman, aff.).

Growing on the base of a stem of Halicornaria hians, var. laxa,

were two simple stems of a Plwmularia which remains un-

identified. Most of the hydroclades have disappeared, but sufficient

remains to show that the trophosome is akin in structure and

relative proportions to that of P. megalocephala Allman, found on

the shores of the western Atlantic and eastern Pacific Oceans.

From this they appear to differ only in their simple habit and

small size, and in lacking that sarcotheca which occurs on the

stem internodes, immediately proximal to the hydroclade-bearing

process.

Dimensions :-—

tem, leno thie seater meses aaarsts co; cs,a cine 9 mm.

Stem internode, lemethy yas. ..1..-.:20. 50240 0°35-0°49 mm.

£ 53 GHAMELCD once r ee aes ceine ne 0:039-0:084 mm.

Hydrothecate internode, length ......... 0-31 mm.

2 . diameter ...... 0-019—-0-027 mm.

Basal athecate internode, length ......... 0-14-0-°19 mm.

ly drotheca, depunmmeece. .cc0s2..2--2<: 5005s 0-045 mm.

_ diameter at mouth ......... 0-048-0:051 mm.

Locality. Flying Fish Cove, 46 fathoms.

HIALICORNARIA HIANS Busk, var. LAXA, noy.

This is the species most abundantly represented in the col-

lection. The largest colony is 12 cm. in height, and the minute

characters of the hydrothece,agree closely with Bale’s * diagnosis,

except that the mesial sarcotheca, in overtopping the hydrotheca-

wall for some distance, approaches the condition of that described

by Stechowt. The ultimate portion lies, however, at a slight

angle to the course of the main portion of the sarcotheca.

There are differences in habit, however, which distinguish

this variety. The colonies are delicate and fine in appearance ;

and the stem is thin and shows no traces of nodal constrictions.

The hydroclades, which are of the unusual length of 22 mm.

(twice as long as those recorded by Stechow), are alternate, and

are wide apart, a space of 2 mm. separating successive individuals

on the same side of the stem. They are divided into long inter-

nodes, on the upper portion of which the hydrothece are set.

In this respect, and on account of the greater depth (as compared

with Bale’s and Stechow’s figures), and more erect posture of the

hydvothecz (the axes of which are inclined to that of the hydro-

clade at an angle of about 40°, instead of 55°), and of the greater

elevation of the intrathecal septum above the floor of the hydro-

theca, these specimens resemble var. profunda, from the

Andamans.

* Bale, Cat. Australian Hydroid Zooph., 1884, p. 179.

+ Stechow, Abh. K. Bayer. Akad. Wissens., Suppl.-vol. i. 3, 1909, p. 101, pl. vi.

o. 16.

t+ Ann. Mag. Nat. Hist. (8) vol. iii. 1909, p. 528.

836 ON HYDROIDS FROM CHRISTMAS ISLAND, [May 24,

Text-fig. 81.

Halicornaria hians, vav. lava.

Portion of hydroclade with hydrotheew, X 78.

Dimensions :—

Stem, greatest diameter...............:.eeeseee 0°45 mm,

Eydroclade; lam etait an. <a: .ssheneessana cere up to 22 mm,

Hydroclade internode, length .............4. 0'41-0°43 mm.

a = diameter: ).sasceeenst 0-10-0:15 mm,

Hiydrotinecs; Mepih waccu.ve. ..;. «uke supe Sik 0-19 mm,

diameter at mouth ............ 0-17 mim,

Locality. Flying-Fish Cove, 46 fathoms,

Re ern

ConTENts (continued),

Page

Miss Annie Porter and Dr. H. B. Fantham, F.Z.S. Ona new pain T. dendroceli,

sp. n., from Dendrocelwim lactewiti.s.ese se ceeecececces ete kOe eR Wiel swe sleds, oe 670

1, The Morphology and Life-History of Himeria (Coccidium) avium: a Sporozoén causing a

fatal disease among young Grouse. By H. B. Fanrnam, D.Se. Lond., B.A. Cantab.,

A.R.C.S., F.Z.8., Christ’s College, Cambridge, Assistant to the Quick Professor of Biology,

Cambridge, and Protozoologist to the Grouse Disease Inquiry. (Plates LV.—LVIIL.)..

2. Observations on the Parasitic Protozoa of the Red Grouse (Lagepus scoticus), with a

Note on the Grouse Fly. By H. B. Fanrmam, D.Sc. Lond., B.A.Cantab., A.R.C.S., F.Z.S.,

Christ’s College, Cambridge, Assistant to the Quick Professor of Biology, Cambridge,

and Protozoologist to the Grouse Disease Inquiry. (Plates LIX.-LXI.)............ 692

3. Experimental Studies on Avian Coccidiosis, especially in relation to young Grouse, Fowls,

and Pigeons. By H. B. Fanrmam, D.Se. Lond., B.A. Cantab., A.R.C.S., F.Z.S., Christ’s

College, Cambridge, Assistant to the Quick Professor of Biology in the University of

Cambridge; Protozoologist to the Grouse Disease Inquiry. (Plate LXITI.)

4. Observations on the Blood of Grouse. By H. B. Fantmam, D.Se. Lond., B.A. Cantab.,

A.R.C.S., F.Z.S., Christ’s College, Cambridge, Assistant to the Quick Professor of

Biology in the University; Protozoologist to the Grouse Disease Inquiry. (Plate

IPONGTATITE)) toy? tain, cbr el so) Soap ae RRP Tao: a sr scr sya ¥aachayalladore fates aia: ofa tsi Satiors 0’ isles Bdrm’ 722

2. Zoological Results of the Third Tanganyika Expedition, conducted by Dr. W. A.

Cunnington, F.Z.S., 1904-1905.—Report on the Ostracoda. By Prof. G. O. Sars,

Meiers CE lates LX 1 Vi USeMene an cheers cfdosad beoisie sce cinesers wvica ewe ealciee dee ns 732

6. On Tritylodon, and on the Relationships of the Multituberculata. By R. Broom, M.D.,

ID) ISH, OBI ASS SURIRRBEMPRAS cit cic 6.0) 00 SC oe AIG ERCCRMENO COIS CCRC CIE ISIPIERAICLIS EES SD Ei Are 760

May 24, 1910.

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

PUPEVE LALO) 6.5 °siuccai nt are cred Meee ira Y haven ata aan slehaiayate scans ok cciw es ceebe el diem cc 768

Mr. Ernest Gibson, F.Z.S. Exhibition of a skin of Felis geoffroyi 6... pee cece ee eeecee 769

1. Observations on the Anatomy and General Biology of some Members of the Larger

Cetacea. By D. G. Linu, B.A., Hutchinson Research Student of St. John’s College,

Gamonidpe. ; (Plate TAN maar ce ete slaves ore sin 'e > cielescledeitie ce ee ORR TE 769

- Zoological Results of the Third Tanganyika Expedition, conducted by Dr W. A.

Cunnington, F.Z.S., 1904-1905.— Report on the Rotifera. By Cuaries F. Rovsserer,

Benes... CE late: ER nas ames Lh coictay iene ih cualulcialsignts telat o'cek-ae s’aidieieacwiadacute 792

iS)

. The Marine Fauna of the Mergui Archipelago, Lower Burma, collected by Jas. J.

Simpson, M.A., B.Se., and R. N. Rudmose-Brown, D.Sc., University of Aberdeen,

February to May 1907.—The Hyprorps. By James Rivcure, M.A., B.Se., Natural

History Department, The Royal Scottish Museum. (Plates LKXVL. & LXXVII.).... 799

4. Hydroids from Christmas Island, Indian Ocean, collected by C. W. Andrews, D.Sc.,

F.RS., F.Z.S., in 1908. By James Rircure, M.A., B.Sc., Natural History Department,

ERE va Scottish: MyseMM wht ay ies 0s ales dsc eres deh Sah emeeenGks cc Sous 826

LIST OF PLATES.

1910, pp. 589-836.

Plate Page

XLVII

es Oandal fins of Nelegstomn’ 5..... 3.505. e0cenenee seen 590

LI. Tasmanian Frogs ....0+-..sseceeserseas ov.8 nisle's as oan 627

LIL. Photophores of Acanthephyra debilis ses cress eescerenee

iv Photophores of Acanthephyra and Sergestes ......+++++.++ 639

LV. Eimeria (Coecidium) avium. (Schizogony) ......+.ee-ee-

LVI. LKimeria ( Coccidium) avium. (Macrogamete formation). . 2 672

LVII. Himeria (Coccidiwm) avium. (Micr ie 5 Sporognay ls

LVIII. Kimeria (Coccidium) avium. (Sporogony)....-++++..4+++--

LUX. Lewcocytozodn lovatt oo... cc cecscecsyecceceecscceses -\

LX, 17-28. Leucocytozodn lovati. 29-40. Spirocheta lagopodis. |

41-49. Hamoproteus mansont .. ...ercerevcvecccscces 692

LXI. 48-50. Trichomonas eberthi. 51-57. Spirocheta ees)

58-65. Ameba lagopodis ..+...... sietete vere Sraaos = eves)

TUR. Atvian @Oceiasisaeiste + cic ic-viv's clele'r (ole olete aie es, ote crater ee eaters 708

LXIII. Blood cells of birds, chiefly of Grouse ..........-+-.+.. 722

LXV. 1-8. Paracypria declivis G.O. Sars. 9-10. P. complanata\

G. O. Sars. 11-12. P. conoidea G. O. Sars ........0s..

LXV. Paracypria obtusa G. O. Sars oo... ce eeeeceeeeescceceee

‘LXVI. 1-6. Paracupria deflexa G.O. Sars. 7-11. P. subangulata }

GESOMBAES GH eye ce «ein ine o 0: oo vale afore ints oft elele atene (ste sears

LXVII. 1-4. Paracypria reniformis G. O. Sars. 5-9. P. inmilis |

‘ G.O. Sars. 10-15. P. opaca &. QO) Sars.to 4s acsels Seem

LXVIIL. 1-10. Paracypria claviformis G.O. Sars. 11-17. P. flexuosa |

CROSS ANG ia ere varate oie 1s ein ele se cicietelshemvarera tote eaters ite emma

LXIX. 1-10. Cypridopsis obliquata G. O. Sars. 11-18. C. sinuata

GIO Sars ceteaisc. sie ois dc, 00, slecerelalejohanttehe tate ttate atehe tes ene

LXX. 1-3. Cypridopsis eunningtoni G.O. Sars. 4-6. C. pusilla

G. O. Sars. 7-9. C. monodonta G.O. Sars. 10-12. C.

SEN ALA Crew) SATS. « chclc o:8\ n'a 1010 aid cfolelete:ehela/aiehels tenia AS

LXXI. 1-3. Cypridopsis bidentata G. O. Sars, 4-6. C. tumidula

G. O. Sars. 7-9. C. congenera G.O. Sars. 10-12. C.

gibba G. O. Sars 16s. ce ence tesa es we sie on ee oe epi sien

LXXII. 1-3. Zonocypris costata Vavra. 4-7. Z. levis G. O. Sars.

8-10. Stenoeypris perarmata Brady. 11-14. Hotroceven

GiB AO MSAVG clave @ a: ss oins8la'e Rlele epalansre etolcteipe eae

LXXIII. 1-4. Llyocypris propinqua G. O. Sars. 5-7. LI. alta G. O. Sars,

8-14. Limnicythere obtusata G. O. Sate! 15-17. Para-

eypriaeurta. O. Sars... +20 sence wy ei8isis shilele saan )

LXXIV. Auditory apparatus of Balenoptera musculus ...... +0006 - 769

LXXV.” Notops WONG «since se seers gine eee sae See Sy hy

Ree | Hydroids from Mergui Archipelago ........+0..ee.seee 799

NOTICE.

The ‘ Proceedings’ for the year are issued in as aka paged consecutively,

so that the comps reference is now P. Z. S. 1910, p. .... The Distribution

is as follows :-—

Part 1 issued in March.

” 2 ” June.

eR eon September,

8 Sie Seer ae December,

* Proceedings,’ 1910, pp. 885-588, were published on August 23rd, 1910,

The Abstracts of the papers read at the Scientific Meetings in

April and May are contained in this Part.

PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCLETY

OF LONDON. ©

1910.

Pages 837-1038.

Part LV. conTaAINING PAPERS READ IN

JUNE.

DECEMBER 1910.

PRINTED FOR THE SOCIETY,

SOLD AT THEIR HOUSE IN REGENT’S PARK.

tpeONDON :

MESSRS. LONGMANS, GREEN, AND CoO.,

PATERNOSTER ROW,

[Price Twelve Shillings. }

LIST OF CONTENTS.

1910, pp. 837-1033.

June 14, 1910.

: Page

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

May 1910

Mr. R. I. Pocock, F.L.S., F.Z.8. Remarks upon, and exhibition of two drawings

illustrating, an instance of ant-mimicry by the Jarvx of a species of Mantis; with a

Note on the mimicry of the larva of the Ceylonese Leaf-Insect (Phyilium sp.?) -... 837

1. On the Specialised Cutaneous Glands of Ruminants. By R. I. Pocock, F.LS., F.Z.S.,

Superintendent of the Zoological Society’s Gardens ....--..eeeececererseeceeees 840

2. On a Wapiti and a Muntjac. By RB. LypexKar .......+<sis0seeseeus: = sees eaemene 987

3. On Three African Buffaloes. By R. Lypexxer

4, On a new Antelope and on the Spanish Chamois. By Prof. A. Casrura, C.M.Z.S..... 998

5. The Changes of Plumage in the Red Grouse (Lagopus scoticus Lath.) in Health and in

Disease. By Epwarp A. Witson, M.B., F.Z.S., Field-Observer to the Grouse

Disease Inquiry. (Plates LAX VITI-CI.) «0.252. cca es ote sn ae osecineleh eeennnen 1000

Titlepage cas cs as seme cals Aisle phat ae v's, o voy os oc ee nls cuin winnie nie ¢\nit# aie i

Last of Council and OMcars aempeents tits cc cise a's s coeds pie lolelememeneaiareigs » SU ciate ote li

List’ of Contents. oo i.'s 0.0 cutee ety dc eas vv nnivc cus ota maven c/n vail) on) + 5 Xie ili

Alphabetical List of Contributors 05.0 0.s esc ee ee ree een uent seen en eus¥ cu 6m pniaeieehaints vii

List of Plates: ooci%. uae ve = simeieelnrstaniaie telarp)s i's 0%: nloiecle Wote ate Cunt: stein cla ean xili

List of Toxt-figures ...... 2. es eseeueey [PME ty sere cevigutnosuvknusles «8 SME : xy

New Generic Term ....0 2 .esseeveuns \ sivmiedivesw a) cade eats ry xix

Lnd6x “.\.<'s cau qwiaiee e's balsa da ancan

1910. ] ON A NEW INSTANCE OF AN't-MIMICRY, 837

June 14, 1910.

Dr. 8. F. Harmer, M.A., 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 May 1910 :—

The registered additions to the Society’s Menagerie during the

month of May were 130 in number. Of these 80 were acquired

by presentation, 13 by purchase, 23 were received on deposit,

3 in exchange, and 11 were born in the Gardens.

The total number of departures during the month, by death

and by removals, was 156.

Amongst the additions special attention may be called to the

following :—

One Leopard Cat (felis bengalensis) and one Masked Palm-

Civet (Paradoxurus larvatus), from Szechuen, China, presented

by W. Thurlow-Lay, Esq., on May 23rd.

Two Hunting-dogs (Lycaon pictus), from Nairobi, presented by

H. H. Heatley, Esq., on May 23rd.

One Caracal (felis caracal) 3, one Chapman’s Zebra (Hquus

chapmanni) 3, and two Selous’s Marsh- Buck (Limnotragus

selousi), from Lake Ngami, purchased on May 7th.

One Sabre-horned Or yx On, yx algazel), from Northern Nigeria,

presented by Sir Hesketh Bell, on May 8th.

Two Ludwig’s Bustards (Otis dudwigi) and two Vigors’s Bustards

(Otis vigorsi), new to the Collection, from 8. Africa, presented by

Sir Charles B. Elliott, K.C.M.G., on May 14th.

Two Mantchurian Cranes (Grws japonensis), from N. China,

purchased on May 25rd.

Mr. R. I. Pocock, F.LS8., F.Z.8., Superintendent of the Gardens,

exhibited two drawings (text-fig. 82) illustrating an instance of

ant-mimicry by the larvee of a species of Mantis, and remarked :—

“In May of this year Mr. W. H. Boyle sent to the Society’s

Gardens, from Axim on the Gold Coast, the nest of a species of

Mantis. Thenest was about the size of a walnut but less regularly

shaped, and pale emerald-green in colour, with a glistening sugary

appearance. The day after it was placed in the Insect House, the

young insects emerged. They were 4 mm. or less in length, and

when crawling about the case looked exactly like a crow d of black

ants, their rapid darts and pauses recalling irresistibly the busy

method of progression so characteristic of these Hymenoptera.

When at rest they no longer resembled the models they imitated

when-in motion, but assumed the attitude of adult Mantises

watching for prey, raising the fore part of the body and head,

folding up their fore legs, and every now and then swaying gently

from side to side as if rocked by the wind. While thus em-

ployed, they were seen to be procryptically coloured. The long

Proc. Zoot. Soc.—1910, No. LV. 55

38 MR. R. I, POCOCK ON A [June 14,

filiform antennz were too delicate to be seen ; the large head was

mottled pale drab and brown; the long pronotum, which was

somewhat abruptly constricted behind the head and gradually

narrowed posteriorly, was uniformly black except for a pale

whitish spot upon its hinder end; the mesonotal, metanotal, and

abdominal terga were, like the head, pale drab relieved by sym-

metrically arranged brown spots. ‘The effect of the contrast

between the prothorax and the head in front of it and of the

tergal plates behind it was disruptive, the colour-continuity of the

upper surface being destroyed. The fore legs, folded beneath the

prothorax, were indistinguishable, and the remaining two pairs

Text-fig. 82.

A. Larva of West African Mantis in resting attitude.

B. The same when crawling, with the fore legs extended and abdomen

turned up.

of legs which supported the body were rendered invisible by their

greenish coloration matching the green of the leaves. These legs,

however, were not of the same tint throughout ; the base of the

femur was paler than the distal end; the tibia was marked with

a median and a distal dark ring, and the end of the tarsus was

darker than its proximal portion. The optical effect of this

alternate banding with dark and light of the distal half of the

legs was discontinuity of surface, the light areas being obliterated

against a light background and the dark areas against a dark

1910. ] NEW INSTANCE OF ANT-MIMICRY, 839

background. The whole of the underside of the body was black ;

the legs of the anterior pair were also mostly black, except for

pale annulations on the tibia and tarsus. Two factors were in

the main concerned in effecting the likeness to an ant, namely,

the blackness of the underside of the abdomen, and the habit of

the Mantis, when in motion, of curling the posterior half of the

abdomen up like a Scorpion’s tail. It resulted from this action

that the abdomen was in appearance reduced to about half its

real length, its shape being altered to an elongated oval, and

since its anal end was brought as far forwards as the posterior

extremity of the prothorax, completely covering the metanotal

and mesonotal terga, the black of its underside formed with the

black of the prothorax a continuous field only broken by the

white spot on the prothorax, which, being situated at the nar-

rowest part of the body, exactly simulated an ant’s waist. The

long and powerful fore legs were extended forwards in crawling,

their annulated distal ends projecting some distance in front of

the head on each side, so as to resemble the waving antennz of

an ant. ;

“The above described coloration persisted until the Mantises had

attained a length of about 7mm. At the next moult a remarkable

change took place, the prothorax losing entirely its black colour,

and with its loss the resemblance to the ants disappeared.

“* An apparently similar case of ant-mimicry by the larva of an

Indian species of Mantis has been described by de Nicéville in a

letter to Prof. Poulton (see Shelford, P. Z. 8. 1902, vol. i. pt. 2,

p. 232). The Mantises were said to be remarkably like a small

black ant; the deceptive resemblance was so close that careful

scrutiny was necessary to distinguish the true character of the

insects. Although de Nicéville gave no particulars as to the way

in which the mimicry was achieved, he noticed that the young

Mantises were incessantly moving, just like the ants they mimicked

but quite unlike the adults of their own kind. Shelford also

recorded an instance observed by himself at Kuchong of mimicry

between a Harpagid Mantid (Hymenopus bicornis) and a young

larva of a distasteful Reduviid bug (Lulyes amena).”

[Additional note, added June 15th, 1910, on the mimicry of

the larva of the Ceylonese Leaf-Insect (Phyllium sp.?).—Dr. A.

Willey, F.R.S., on his return from Ceylon, brought a number of

eggs of this insect as a present to the Society. The young upon

hatching were seen to differ remarkably in colour from the adults.

The head, thorax, and anterior part of the abdomen were crimson,

the posterior part of the abdomen being much darker and browner

in hue and the legs black. The crimson and black coloration

made these insects extremely conspicuous on green leaves, and

their general appearance was remarkably ‘bug’-like. Their

scheme of coloration fitted unmistakably into that depicted and

described by Shelford (P. Z. S. 1902, vol. ii. pt. 2, pl. xxiii.

figs. 1,2, 3) as illustrated by two species of bugs (Hetatops rubmiaceus

DD5*

840 MR. R, I. POCOCK ON THE [June 14,

and Serinetha abdominalis), both probably distasteful, and a moth

(Phaula limbata), all of which mimic a distasteful Lycid beetle

(Lycostomus gestroi). The resemblance was so close as to leave

no room for doubt that similar if not identical species of Lycide

and Hemiptera were mimicked in Ceylon by the larvee of the

Phylliwn. Yn the matter of coloration the larvee of this Ceylon

species of Phyllium differed markedly from those of the Seychelles

species, the only other Leaf-Insect that had been exhibited in the

Gardens. In the Seychelles species the young upon hatching

were always green like their parents and procryptically coloured. |

The following papers were read :—

1. On the Specialised Cutaneous Glands of Ruminants.

By R. 1. Pocock, F.L.S., F.Z.8., Superintendent of the

Zoological Society's Gardens.

{Received June 14, 1910.)

(‘Text-figures 83-142.)

Part I.—Inrropuction.

In the ‘ Proceedings’ of our Society for 1836 there is much and

varied information, some of it true and some of it false, about

the specialised cutaneous glands of Ruminant Ungulates. On

pp- 37-38 will be found a table compiled by Owen * showing the

presence or absence of preorbital and inguinal glands in some sixty

species of Antelopes. Bennett contributed a note on pp. 34-36

on the preorbital gland of Antilope cervicapra; and this was supple-

mented by remarks from Ogilby on the same gland in this species

and in an unidentified species of Gazelle. Bennett also communi-

cated (p. 39) a letter from Hodgson on the inguinal glands of

Pantholops and on the preorbital glands of Cervus aristotelis and

of Capricornis thar. The subject was further discussed by Ogilby

(pp. 131-139) in a paper dealing with the preorbital, inguinal,

and pedal glands, and other external features in genera of the

families Camelide, Giraffidee, Cervidee, Antilocapride, and Bovide.

Finally (on pp. 66-68), Gray described the tarsal and metatarsal

tufts of various species of Cervicee.

Eleven years later Hodgson (J. A. S. Bengal, xvi. pt. 11. pp. 685—

711, 1847) published a long paper on the same subject, based

mainly, but not solely, upon Indian genera of Cervide and

Jovide, without apparently being aware of the work already

done by Owen and Ogilby; and in 1868 Owen reprinted, with

slight modifications, his original paper, but without incorporating

* Apparently largely from data supplied by Hamilton Smith in vol. vii. of

Griflith’s ‘Animal Kingdom,’ 1827.

a eee

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 841

in it the facts established by Ogilby and Hodgson (‘ Anatomy of

Vertebrates,’ iii. pp. 632-638, 1868).

These papers contain by no means the first records of the

occurrence of specialised integumentary glands in Ruminants ;

but those of Owen, Ogilby, and Hodgson are apparently the

earliest which attempt to deal comprehensively with the subject

from the systematic standpoint ; they are also important from

the conclusion to be inferred from internal evidence, that they

ave the main source whence English writers of text-books and

monographs have borrowed most of the statements regarding these

glands. This is a point, however, on which it is impossible to

speak with certainty, because of the reticence of the writers

regarding their authority for the information they publish. Even

so careful and conscientious a zoologist as Blanford * did not say

whether his statements concerning the glands of Indian Rumi-

nants were taken from Ogilby and Hodgson or were based upon

his own observations upon new material which he assuredly had

the opportunity of collecting in India. The information he gave,

however, tallies so closely with that of Hodgson that it is im-

possible to doubt his great indebtedness to that author.

I think it probable that Messrs. Sclater & Thomas 7 and Mr.

Lydekker£ depended mostly upon the publications of Ogilby,

Hodgson, Blanford, Brooke, and Gray, and upon a few special

memoirs by authors like Flower, Garrod,and Forbes, supplemented,

no doubt in some cases, by their own observations upon the

glands as exhibited by dried or mounted skins. It does not appear,

however, that their works contain many new contributions to the

subject. The same may be said of Mr. W. L. Sclater’s treatise on

the Bovidee of 8. Africa $, except that the alleged details respect-

_ing the preorbital glands of the Tragelaphinz seem to be new, the

general statement as to their presence and size having been taken,

I infer, from Flower and Lydekker’s volume of Mammalia, a

standard source of information on this and other subjects since

its publication in 1891.

In the above-quoted works will be found many discrepancies

and contradictions with regard to the occurrence in particular

genera and species of the glands forming the subject-matter of the

present paper. These contradictions are difficult to understand,

unless the errors be attributable to the examination of defective

material. Unfortunately, most authors, like Owen, Ogilby, and

Hodgson, are silent as to the nature of the material upon which

their observations were made. This omission is important,

because inguinal and preorbital glands, obvious enough on fresh

skins, may be cut away or otherwise obliterated in the process of

+ ‘The Book of Antelopes,’ vols. i. to vi. 1894-1900: R. H. Porter, London.

{ ‘The Deer of all Lands,’ 1898; ‘ Wild Oxen, Sheep, and Goats of all Lands,’

1898; ‘Great and Small Game of Europe, Western and Northern Asia, and

America,’ 1901; ‘ The Game Animals of India, 1907; ‘The Game Animals of

Africa, 1908 : Rowland Ward, London.

§ ‘The Fauna of South Africa: Mammalia,’ i. pp. 123-258, 1900.

842 MR. R. I, POCOCK ON THE [June 14,

preservation or mounting ; and inferences as to the absence or

size of preorbital glands based upon the depressions on the

lacrymal bone may be quite false.

On account of their conspicuousness, more information about

the preorbital glands has been accumulated than about the inguinal

or pedal glands. Although statements as to the presence or

absence of inguinal glands are not infrequent, very little has been

recorded as to their number, depth, and nearness to the mammee.

The pedal glands have been described as large or small, deep or

shallow, present or absent, but no correct conception of their

variability in shape, size, and position is conveyed by published

descriptions. The available information, indeed, on such points is

so meagre as compared with the number of species in which they

have been recorded, that it has not been possible for systematists

to use these glands on a large scale for taxonomic purposes.

Apart from the, comparatively speaking, comprehensive works

above quoted, there are many isolated memoirs on the glands of

particular species. These are quoted in their appropriate places

in the text that follows. There is also a considerable literature

dealing with the physiology and histology of the glands. The two

papers that I have especially consulted are the following :—An

important memoir by Max Weber* upon the structure of the

preorbital glands and the nature and use of their secretions m

several ‘¢ Antelopes,” with which is incorporated a list of the

principal memoirs dealing with the subject of the cephalic glands

in this group; and a special treatise by Max Tempel 7, which has

for its subject-matter the pedal glands mostly of European species

of Artiodactyla. This treatise gives a brief summary of the

observations of many authors upon the specialised cutaneous glands

of this group, as wellas of other Mammalia. Max Tempel addsa

tolerably complete bibliography of the subject, and describes fully

the microscopical structure of the pedal glands of the twelve

species examined by him.

In the hopes of adjusting some of the discrepancies above

alluded to, of filling certain obvious gaps in our knowledge, and

of discovering possibly at the same time new and useful characters

for distinguishing genera and classifying higher groups, | began

about three years ago, with the sympathetic concurrence of Mr.

F. E. Beddard, F.R.S., the Society’s Prosector, to dissect, draw,

and describe the cutaneous glands of Ruminants that died in the

Gardens. But so many important species were unrepresented in

our collection, and were likely to remain so, that I soon perceived

the necessity of procuring material elsewhere if the work was to

* Aych. mikr. Anat. xxxi. pp. 500-540, 1888.

+ ‘Vergl. Anat.-physiol. Untersuch. iiber die Driisen der zwischen Klauenhaut

der Paarzeher,’ Inaug.-Diss., Leipzig, 1896 (pp. 1-67, 12 pls.). I was unable to see a

copy of this paper until my own observations upon the same animals were completed.

Except in one or two matters of detail, due perhaps to difference of preservation of

material, my results were completely in accord with this author’s. I am indebted

to Mr. B. B. Woodward for the information that a probable abridgment of Max

Tempel’s paper was published in Arch. Thierheilk. xxiii. pp. 1-48.

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 843

be made comprehensive in its scope. I therefore applied to

Rowland Ward, F.Z.S., and to Mr. Edward Gerrard for

skins, feet, or masks of specimens that might be discarded in their

stores as useless duplicates. To the generous response my appeal

elicited, I owe the opportunity of examining these glands in many

species I could not otherwise have seen; and I gladly take this

opportunity of expressing my indebtedness to Mr. Rowland Ward,

to his manager, Mr. J. B. Burlace, and to Mr. Gerrard for the kind

help they afforded me. I also applied to Mr. Herbert Bolton,

F.R.S.E., F.Z.8S., the Curator of the Bristol Museum, for the

use of discarded stuffed specimens, and with the sanction of the

Committee of the Museum he kindly sent me from time to time

material which has been of great use for my work. Finally, I have

to thank Mr. Oldfield Thomas, F.R.S., for allowing me free access

to the collection of skins under his charge at the Natural History

Museum, when I wished to examine representatives of species I

had no chance of seeing elsewhere.

But apart from the verification of previously observed facts,

very little satisfactory work can be done upon dried skins if left

intact. It is true that the absence of glands can in many cases

be inferred by superficial scrutiny ; buttheir structure, when they

are present, cannot be ascertained without maceration and dissec-

tion. It was here that the use of the material lent or given to

me by Messrs. Rowland Ward and Gerrard came in. I found that

softening the feet or the preorbital glands, when preserved, and

cutting them witha razor or sharp knife, revealed surprisingly

well the main features of the glands, especially those of the feet,

due allowance being made for shrinkage when, as was too

commonly the case, the back of the pasterns had been opened and

the connective tissue that normally holds the gland in place had

been cut away.

In no case did I attempt to ascertain the microscopical structure

of the glands themselves, this aspect of the subject having been

already worked out by Max Weber, Tempel, and other authors.

On the fresh specimens I generally made a practice of recording

and describing the scent of the secretions—that is to say, of com-

paring the smell to that of other and familiar objects whenever

the one reminded me of the other. In some cases this was a

simple task. There was, for instance, no mistaking the resemblance

between the scent of the secretion of the inguinal glands of Ovis

vignet and of Gazella subguttwrosa to the urine of Mus niusculus,

nor of that of the pedal glands of Zama to the same substance.

But in other cases no such comparison was possible, owing to the

fact that the scents were swi generis and were therefore not com-

parable to any substance known to me.

T noted incidentally i in most cases the number of mamme, but

I was disappointed in the hope of getting useful taxonomic features

from these organs. The typical number in the Ruminants is two

pairs, although the occasional presence of three pairs in domestic

cattle suggests that six may have been the original total. Two

844 MR. R. I. POCOCK ON THE (June 14,

pairs, however, seem to be invariable in the true Cervide ; never-

theless, J/oschus, sometimes at all events, has only a single pair,

and two pairs ave found in the majority of the Bovide. In the

Caprine and Antilopine, however, only a single pair is, as a rule,

found, and these represent the posterior pair of the quadrimaim-

mate genera; but in Sheep, Goats, and Gazelles the mamme of

the anterior pair ave so commonly retained in a yestigial state

that it is only possible to affirm of these genera that they usually

have but a single pair of mammie,

I found, on the contrary, as [ anticipated, that the glands, and

especially the pedal glands, sometimes supplied valuable con-

firmatory evidence of kinship between genera already affiliated for

other reasons, and sometimes showed greater divergence between

genera than was previously suspected to exist. But neither the

presence nor the absence of the glands could be considered as

furnishing in itself evidence of kinship, without other characters

being taken into account, because it was clear that the glands had

been independently suppressed in genera belonging to different

groups.

7 While w orking at the pedal glands I was greatly impressed by

the difference in the structure of the feet with respect to the en-

croachment of the “ web,” or junctional integument, of the digits

over the interdigital area, and to the manner in which it is folded.

In the case of dried skins it is generally impossible to substantiate

the presence of glandular layers ; but the interdigital clefts which

result from the folding of the integument. of ‘the pastern can

always be detected whether they be. specially glandular or not.

It must be remembered, however, that, str ictly speaking, glands

are never absent, since the skin is furnished with sebaceous

and sweat glands which form the basis of the specialised glands

here described. The statement that glands are absent merely

means that macroscopic inspection fails to reveal evidence of a

specialised area of glandular activity or of a particular receptacle

for the storage of secretions.

The terminology of the glands and of other features employed

in this paper is as follows :—

Foot: the terminal portion of the leg between the carpo-meta-

carpal or tarso-metatarsal jomt and the apex of the hoof.

Fetlock: the carpo-metacarpal or tavso-metatarsal joint.

Pastern : the area between the fetlock and the hoofs.

Hoof: the terminal portion of the foot, consisting of the pai

of distal phalanges, each of which is covered with naked skin which

is softer close to the pastern than distally, where it is protected in

front and laterally by the wail; the supporting area of the hoof

is composed of the sole in front and the heed behind ; the heel-tie

is the integument which joins the heels together ; and the inter-

ungual integument is the skin that extends forwards from the

heels to the front of the pastern above the hoof.

Pedal glands ave the larger or smaller pocket-like or flask-like

invaginations of the skin which open upon the front of the

1910. | CUTANEOUS SCENT-GLANDS OF RUMINANTS. 845

pasterns or between the base of the hoofs (= postdigital, inter-

digital, and interungulate glands of Owen ; fosse interdigitales of

Ogilby ; interdigital pits of Hodgson ; foot-glands or hoof-glands

of other English authors).

Carpal or - knee glands are the glands commonly represented by

a pad of long hair just below the knee or carpus in several

Bovide.

Larsal glands are the glands represented by a tuft or pad of

hair on the inner side of the tarsus of some telemetacarpal Cervidee

(=the calcic tuft or gland of Hodgson, at least in part).

Metatarsal glands ave the glands represented by a pad or tuft

of hair on the outer side of the metatarsus of some Cervide and of

pyceros amongst the Bovide (=the calcic tuft or gland of

Hodgson, at least in part).

Preorbital glands ave the glands in front of the eyes (= sub-

orbital and maxillary pits of Owen ; sinus lacrymales and glandule

maxillares of Ogilby ; eye-pits of Hodgson ; antorbital glands of

other English writers).

Inguinal glands are the saccular glands of the groin (=inguinal

pits of Owen ; folliculi inguinales of Ogilby ; inguinal elands and

pores of Hodgson ; ; groin- glands of other authors).

In addition to these there are glands of less common occurrence,

such as the post-cornual glands of Rupicapra, the occipital glands

of Camelus, the caudal glands of Capra, the preputial gland of

Moschus, and the wiaguicular glands ov glands in the false-hoofs of

Tetraceros.

Part I].—On THE STRUCTURE OF THE GLANDS AND THE SYSTEMATIC

RESULTS TO BE DERIVED FROM THEM.

Family BOVIDA.

The subjoined dichotomous analytical key, based mainly on

the structure of the feet and upon the specialised cutaneous

glands, shows the subfamilies into which the Bovide may be

provisionally divided on the evidence supplied by these characters.

Thave preserved all the subfamilies already established by previous

authors, and have added a few new ones for the reception of

isolated genera which I found could not be included in the divisions

to which they had been heretofore assigned without frustrating

the attempt to define the groups in question.

The erection of new subfamilies for the accommodation of these

aberrant genera appeared to me to be the most expedient course

to follow under the circumstances, although some authors may

perhaps demu to the separation thus entailed of Pantholops

from the Antilopine and of Oreotragus from the Neotraginz.

Furthermore, I am inclined to think that additional groups

of subfamily rank will probably be established in the future.

Goselaphus and Tetracerus, for instance, might perhaps be

846 MR. R. I, POCOCK ON THE [June 14,

wisely and advantageously separated from the African Tragela-

phines. Possibly a special subfamily should be erected for

Ammodorcas, which at present I am unable to classify; and a

good case could be made out for splitting up the Rupicaprine,

which no one can claim to be a homogeneous unit. But our

knowledge of the mutual affinities of the various sections of the

Bovide is so surprisingly incomplete that no attempt at a final

classification of them can at present be attempted.

Owing to the impossibility of arranging the subfamilies

according to their probable relationships in a linear series, it

comes about that in the annexed table divisions of this rank are

juxtaposed which have obviously no near relationship to each

other. Oreotragus, for instance, is probably, as is usually supposed,

allied to the Neotraginze and Pantholops to the Saiginze, Antilo-

pine, or Caprinze. Yet these two genera fall, by the differential

characters made use of, under the heading ve alongside the

Mpycerine, the Orygine, and others with which they have

certainly, as I at present believe, no close connection. That is

one of the obvious defects of such tables as the subjoined, which

are commonly much more useful than scientific.

IT have made no attempt to include Ovibos in this key, Lénn-

berg’s recent paper on this form making one conclusion abundantly

clear, namely, that the genus must for the present be left by itself

in the subfamily Ovibovine.

a. On the front and usually also on the hind leg a large inter-

digital cleft, forming a gland the orifice of which is a long

slit with its upper margin sharply defined by the abrupt

descent of the integument between the digits; or else the

orifice of the cleft is shorter and the gland more or less

tubular, extending up between the bones of the pastern, its

posterior wall being formed by a close fold of integument

forming the heel-tie inferiorly, so that the orifice of the

gland opens downwards between the hoofs.

6. Preorbital gland nearly halfway between the eye and the

nose, not valvular or invaginated, but marked externally

by a bare streak of skin studded serially with pores; web

of pastern reaching only to the heels, closely folded, the

gland tubular with the orifice Opening freely into the

space between the hoofs .......................... CEPHALOPHIN® (p. 867).

L’. Preorbital gland occasionally aborted, when present usually

a valvular inv agination, never *marked externally by a

streak of integument with serially arranged pores.

c. Glands equally or approximately equally developed on

both front and hind feet.

d. Mute large, naked above as far back as the posterior

angle of the nostrils .......................... NEOTRAGINE * (p. 879).

d’, Muftle small, hairy above almost or quite to the

anterior angle of the nostrils.

e. Pedal glands more or less tubular; the integument

of the back of the pastern closely folded at the heels.

Mapnoquin& (p. 876).

* Based upon observation of Ourebia and Raphicerus, the other genera being

unknown to me.

1910.] CUTANEOUS SCENT-GLANDS OF RUMINAN'tS. 847

e’. Pedal glands a deep cleft with a long aperture on

the front of the pastern, the web extending forwards

as aclose fold of integument from the heels towards

the front of the hoof and separating the cavity of

the gland from the space between the hoofs.

ANTILOPIN® (p. 887).

ce’, Pedal glands aborted or nearly so on the hind feet.

Bouparin (p. 900).

a’. Never a deep interdigital cleft opening by a long orifice, with

sharply defined upper margin, on the front of the pastern;

pedal glands when present and well developed somewhat

flask-shaped with a constricted orifice opening on the front

of the pastern above the hoofs (exc. Rupicapra), never

opening deep in the interdigital cleft above the heels.

Jf. A depression on the lower half of the pastern in front caused

by the gradual downward and backward descent of the

skin of the middle line towards the heels, where it meets

the posterior skin to form the heel-tie, allowing the

hoofs to be widely or moderately widely separated

apically ; only rarely (in Rupicapra) does the closely

folded integument extend forwards from the heels and

shut off the interdigital cleft from the space between

the hoofs; pedal gland, when specialised, folded upon

itself and opening by a small orifice on the front of

the pastern high above the hoofs.

RuPrIcaPRiIn» * (p. 848) and CaprRIn» (p. 859).

f’. At most a shallow depression on the front of the pastern ;

the hoofs joined together almost throughout their basal

extent by an interungual band of integument extending

forwards from the heels, where it meets at an obtuse or

right angle the skin forming the posterior wall of the

pastern; it similarly meets that of the anterior wall of

the pastern, which forms a continuous surface in the

middle line, except where its continuity is sometimes

broken for a short distance by the orifice of the pedal gland,

which, when present, opens close to the hoofs; thus,

except for,the glandular infolding, the anterior and pos-

terior walls of the pastern are widely separated from each

other in the middle line.

g. Hoofs truncated apically, the heels raised high above the

ground during progression; (interungual integument

hairy throughout ; a valvular preorbital gland; mufHe

large; no pedal glands)........................... OREOTRAGIN# (p. 885).

g'. Hoots pointed, the entire sole and heel in contact with

the ground during progression.

h. Nasal cavities considerably inflated; skin of the heel-

tie posteriorly folded upon itself in correlation with

the backward extension of the enlarged heels; (no

preorbital or pedal glands; a pair of very large

inguinal glands with orifice remote from the

MAMIM)) hee sexes ceheaage: eet ste. -es --4y | LANDHOLOPINE (p. 898).

h’. Nasal cavities not specially inflated ; skin of heel-tie

not folded, heels moderate in size.

i. A pair of large metatarsal glands on the hind legs;

no false hoofs; interungual integument continu-

ously hairy ; [no preorbital or pedal glands}.

JEPYCERIN”E (p. 918).

* This section should, perhaps, be divided into two or more subfanpilies, none of

the genera being closely related to one another. Saiga should possibly come under

the heading f as the type of a special group Saigine ; but the exact structure of

the feet and of their glands is unknown to me.

848 MR. R. 1, POCOCK ON THE (June 14,

i’. No metatarsal glands; false hoofs present; inter-

ungual integument naked.

k. Pedal glands present on all four feet and consisting

of a thick-walled, elongated sac gradually lessen-

ing in calibre towards the orifice, which is situated

close to the summit of the interungual web: or

. else the gland is small and opens behind a conical

excrescence developed from the summit of the

interungual web as in Addaw* ..... ORYGIN (p. 907).

k’. Pedal glands absent, except in Pe/ea, where they

consist of a thin-walled, compressed, roundish

sac with a short and narrow neck, the orifice of

which opens some little distance above the summit

of the interungual web .................. CERVICAPRIN® (p. 911),

TRAGELAPHIN® (p. 921), Bovry& + (p. 932).

Subfamily RupircarRin &,

Genus Rupicarra Blainv.

Ruricapra RupricaPRA Linn. (The Chamois.)

(Text-fig. 83.)

According to Owen the Chamois has no preorbital glands, but

possesses inguinal glands and a gland behind the base of each ear.

Mr. Lydekker, on the contrary, states that a small preorbital gland

is present $; and even so recently as 1904 Max Weber (‘ Die

Siiugethiere,’ p. 675) asserts that the male possesses postauricular

glands. The pedal glands were described by Max Tempel.

Living examples of Tyrolean Chamois in the Zoological Gardens

show no trace either of the preorbital gland mentioned by

Mr. Lydekker or of the post-auditory gland mentioned by Owen

and other writers ; nor could T detect a trace of them in the only

fresh specimens I have been able to examine, namely two newly

born kids, or upon dried skins in the British Museum.

The feet of these immature specimens were more like those of

Gazelles than of any Antelopes, Sheep, or Goats that I have seen.

When the hoofs were pulled apart a triangular interdigital

depression was shown. This, however, was both shorter and

much shallower than in Gazelles. It became gradually shallower

and narrower upwards to its apex, the deepest and broadest part

being at its lower end, where it was closed by the inter-

ungual web. The skin of this web was folded back in the same

way as in Gazelles, but much less deeply, so that the skin forming

the back of the glandular depression was separated some distance

from that of the back of the pastern. The walls of the depression

were somewhat scantily clothed with short hairs. Also, as in

Gazelles, the entire interungual web, extending from near the

* On account of the peculiar structure of the feet this genus should perhaps form

the type of a special subfamily, Addacine.

+ These groups are distinguishable by other characters than those supplied by the

feet.

t ‘Great and Small Game of Europe,’ p. 173, 1901. The authority for this

statement is not given.

1910. ] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 849

front of the proximal end of the hoof almost to the heels, was

clothed with long hairs, the heel-tie itself being naked.

The feet of these immature specimens agreed with those of an

adult Chamois described and figured by Max Tempel (op. cit.

p. 50, pl. xi.), except for the more scanty clothing of hairs on the

walls of the depression. In Max Tempel’s example the walls

of the depression were thickly clothed with long hairs loaded

basally with secretion, and the skin forming the posterior wall of

the depression was in contact with that of the back of the pastern

as in Gazelles.

Judging from the immature examples above mentioned and

from dried skins examined in the British Museum, Chamois have

four mamme, but no inguinal glands,

Text-fig. 83.

oe 5

OS he tate

Cotintey

Rupicapra rupicapra.

A. Head of male showing fully developed postcornual glandular swelling.

BR. The glandular swelling from above: a, anterior, 6, posterior end.

C. Summit of head of adult female drawn on the same day as A, showing

absence of swelling.

D. Foot of newly born kid showing depression on the front of the pastern.

That Chamois possess glands upon the head has been known for

many years. These glands have had a curious and confusing

history. They have been persistently described as ‘‘post-auditory ”

or “post-auricular,” whereas in reality they are immediately

850 MR. R. I. POCOCK ON THE [June 14,

behind the horns and well in advance of the base of the ears.

By no possibility can they be described with accuracy as behind

the ears. I think Hamilton Smith must have been the first to

start this error in 1827. In Griffith’s ‘Animal Kingdom,’ iv

p- 22, he stated that there was a gland behind the ear; but on

. 281 of this volume he described the same gland apparently a

occurring behind the horns. In 1836 Owen repeated the ee

that they are post-auditory, thus confirming my suspicion that he

borrowed many of the facts for his paper from Hamilton Smith.

In 1855 the glands were quite correctly figured and described by

Von Hessling as behind the horns (Zeitsch. f. w. Zool. vi. pp. 265—

271, pl. viii.) ; but in the illustration accompanying this memoir the

ears appear to have been added as an afterthought, for they are

drawn as projecting from the side of the head- skin much too far

forwards, the anterior edge of their bases being in a line with the

posterior base of the horns, so that part of the gland, at all events,

is behind the ears. The figure, however, is quite wrong in that

particular. As is well known, the horns of a Chamois rise verti-

cally nearly over the eyes and the ears are set far behind both. To

Owen’s mistake and Von Hessling’s inaccurate figure must probably

be attributed the statement made by Flower and Lydekker (‘ Mam-

malia,’ p. 12, 1891), by Max Weber (Die Siiug. p. 675, 1904), and

by Lydekker (‘ Field, 1909, p. 1100) that the glands in question

are post-auditory, the latter even speaking of them as “ occipital.”

Yet, curiously enough, Flower knew at one time that they were

behind the horns and in front of the ears, because there is a pre-

paration in the Museum of the College of Surgeons showing their

exact position ; ; and in the catalogue there isan entry in tks hand-

writing, giving an accurate description of the preparation with a

reference to von Hessling’s paper *. This specimen was received

from the Zoological Society in 1877, many years before the

repetition of Owen’s mistake by Flower in his classical work on

the Mammalia.

A further point of interest connected with this specimen is that

it was a female. The glands appear as a pair of suberescentic

slits, one behind each horn and about half an inch away from

it. This establishes the conclusion that the female possesses

the glands as well as the male, although they do not appear to

become enlarged at the breeding-season as in the latter. Their

appearance in the bucks was rec ently described by Mr. A. Buxton

(‘ Field,’ 1909, p. 1056), who found them in the autumn of the

year in various stages of growth, and said that they gave out a

strong goaty smell.

In the male T yrolean Chamois now living in the Gardens the

glands began to swell in the latter half of September when the

coat began to darken, were at their maximum through November

and the first half of December, then gradually dwindled away, and

by the end of the first week of January were represented by a flat

* IT am indebted to Mr. R. H. Burne, F'.Z.S., for this information.

1910. | CUTANEOUS SCENT-GLANDS OF RUMINANTS. 851

area of wrinkled skin buried in the hairs of the top of the head.

At no time did I see any discharge or secretion coming from them.

At their best they projected well above the longest hairs on the

parietal area of the head as a pair of rounded cushion-like excre-

scences. of purple skin studded with short hairs and folded into

deep intersecting grooves, recalling a pair of cerebral hemispheres

marked with a few sulci. Seen from above, the margin is

straightish in front up against the horn, lightly convex externally,

widely rounded posteriorly, and straight internally, where it is

closely in contact throughout its length with that of the opposite

side. The surface is convex from before backwards and from side

to side ; and from the antero-internal angle of each on the upper

side two deep sulci descend towards the external border, one close

behind the horns, the other backwards and outwards, giving off

two backwardly-directed short sulci in its course.

In a male eighteen months old the glandular area showed no

signs of swelling as late as November.

In one of the newly born kids I found these postcornual glands

represented by a pair of very shallow crescentic grooves entirely

concealed in the hairs of the top of the head.

Genus OREAMNOS Raf.

OREAMNOS MonTANUS Ord. (The Rocky Mountain Goat.)

(Text-fig. 84, A.)

The presence of glands behind the horns in this species has

been recorded by Mr. Hornaday*, Mr. Frederick Gillett +, and

by Mr. Madison Grant?. Mr. Hornaday described them as “a

peculiar bare patch of black oily skin, the size of a half dollar.”

To this Mr. Gillett added the important information that ‘“ under

the external skin these glands consist of a soft red tissue saturated

with a milky substance, like the udder of a cow. In the speci-

men at our {London Zoological] Gardens these glands are partially

covered up by long hair at the present time [February], but in

October and November they are more conspicuous. The older

the animal the more pronounced are the glands.” According to

Mr. Madison Grant the gland, which is as large as half an orange

and situated in a half-circle immediately behind each horn, ‘ is

sometimes so tough as to wear deeply into the base of the horn.”

This author cites the presence of this gland as the most striking

character of Oreamnos as compared with other genera of Rupi-

caprine. No doubt his failure to recognise it as the same gland

that is found in the Chamois (fupicapra) is to be attributed to

the erroneous description of the latter as ‘‘ post-auricular.”

My own notes upon an old male animal living in the Gardens

completely bear out what Mr. Gillett said as to the seasonal

development of the gland. In this respect it agreed closely with

+ P. Z.S. 1905, vol. 1. p. 56.

~ Ninth Annual Report of the New York Zool. Soc. p. 12 (eprint).

the homologous gland of the male Chamois; but the integument

covering the gland was much more horny and coarser in texture

than in the Chamois.

Text-fig. 84.

Situ im ees s ESE WS Torey

852 MR. R. I, POCOCK ON THE | June 14,

A. Median vertical section of front foot of Oreamnos montanus.

LB. The same of Budoreas tibetanus.

This Rocky Mountain Goat had no preorbital or inguinal

glands. The feet were constructed almost as in Goats (Capra) ;

that is to say, there was on the lower part of the pastern in front

a deep interdigital depression formed by the abrupt downward

and backward slope of the integument to meet that of the posterior

1910. ] CUTANEOUS SCENT-GLANDS OF RUMINANTS, 853

side of the pastern at an acute angle just above the heels, firmly

binding the latter together. As in wild and most tame Goats,

the walls of this depression were clothed with long hairs. On

the hind feet these hairs were clean; but on the fore feet they

were stained yellow and stuck together, showing dried scurfy

material at their roots and suggesting activity of the sebaceous

glands over a considerable part of the area.

The genus Oreamnos exhibits puzzling cross-affinities. With

the feet and beard of the Goats (Capra) are combined the short

cannon-bones of the Takin (Budorcas), the post-cornual glands of

the Chamois (Aupicapra), and the short, lightly curved, backwardly

inclined horns of Capricornis and Nemorhedus.

Genus NzmorHEDus H,. Sm,

N #®MORHEDUS GORAL Hardw.

(The Goral.)

According to Hodgson, WV. goral has no inguinal or preorbital

glands, but possesses feet-pits on all four feet. Owen is in agree-

ment with him as regards the absence of the glands in the groin

and on the face.

Ogilby also states that the preorbital and inguinal glands are

absent, and adds that the pedal glands are large and the mammze

four in number.

My own observations agree with these in every respect.

On a flat skin of WV. goral sent to me from Chamba by

Major Rodon, F.Z.8., I found the pedal glands present on all the

four feet. They presented a close resemblance both in position

and structure to those of Sheep (Ovis), opening by a small orifice

on the front of the pastern some distance above the hoofs. The

orifice led into a slightly dilating hair-lined sac running back-

wards to the posterior wall of the pastern parallel to the inter-

ungual web. Just below the orifice the integument descended

obliquely between the hoofs to the heels, where it formed a stout

horny tie. ‘This interungual web met the integument of the back

of the pastern on the heels almost at right angles. The hair on

the back of the pastern extended only for a short distance between

the heels ; but the interungual integument was covered with long

hairs, becoming progressively shorter towards the heel-tie, which

was itself smooth and hairless. The hoof itself was short and

compressed with the heels high, their web or tie being raised con-

siderably above the ground. Thus not only in the position and

structure of the pedal gland*, but also in the hairiness of the

interungual integument and the smoothness of the heel-tie, a very

close resemblance could be traced between the feet of Vamorhedus

and those of Ovis.

* T believe the pedal gland described above was incomplete, since the bones and

tissues of the pastern had been to a large extent cut away from behind and there

was a small tear in the skin of the bottom of the gland.

Proc. Zoot. Soc.—1910, No. LVI. 56

854 MR. R. I, POCOCK ON THE ‘June 14,

N £MORHEDUS RADDEANUS Heude. (Radde’s Goral.)

(Text-fig. 85, A.)

‘The entive skin of an adult example of this species * from

Lao-i-san, near Shensi in Central China, was kindly lent to me

by Mr. Edward Gerrard,

The structure of the foot was the same as in iV. goral, except

that the hoofs were rather longer and the pedal gland much

Text-fig. 85.

} Mt i ip iy

A y Y,

BoM aS K

WWW

HUE

=, TLE: ULES

A. Median vertical section of front foot of Nemorhedus raddeanus.

B The same of Capricornis crispus.

gl, pedal gland; o., its orifice.

(Dry specimens.)

longer and move capacious. Just within the orifice the duct was

directed for a very short distance upwards and backwards. It

then took a sharp bend downwards and forwards, running close

along the interungual web to the heel-tie. At this point it was

* Vide Pocock, P. Z.S. 1908, p. 196.

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 855

bent sharply backwards and upwards on itself, the terminal portion

which, roughly speaking, ran parallel with the integument of the

back of the pastern, being subequal in leneth to. the portion

running parallel to the inter ungual web. The gland was lined

with hairs and charged with secretion. Its structure was practi-

cally the same on both front and hind feet.

There were two pairs of teats, but no trace of inguinal glands

could be detected.

Genus CapricorNis Ogilby.

CaPRiIcoRNIS THAR Hodgs.

(The Nepalese Serow.)

According to Owen this species possesses preorbital glands, but

is without inguinal glands. Ogilby, however, asserts that it has

inguinal glands and pedal glands and four mamme.

In the example of the Darjiling race (C. thar jamrachi Poc.) *,

now living in the Gardens, a distinct but small orifice is observ-

able upon the pastern of the fore foot, and the white hair imme-

diately around it is at times stained yellow, proving that a gland

is present and active. This animal is a female nearly five years

old; and the preorbital gland is never prominent or turgid as

deseribed by Hodgson in the case of the male C. thar. It is

represented externally by a circular orifice, from which exude

drops of whitish fluid hardening into solid particles, recalling

both the appearance and scent of gum arabic. Lam unable to

state anything about the inguinal glands.

CAPRICORNIS (or CAPRICORNULUS) CRISPUS ‘Tei.

(‘The Japanese Serow.) (‘Text-fig. 85, B.)

Mr. Edward Gerrard kindly lent mea dried skin of this species.

The preorbital gland consisted of a simple hair-lined pouch

with a small subcireular orifice. The pedal glands were well

developed on all four feet, and resembled almost exactly in

structure those of Memorhedus raddeanus. The feet also were

constructed as in that species, though the hairy clothing was

longer. ‘There were four teats, but no inguinal glands,

IT am also indebted to Mr. E. Gerrard for the opportunity to

examine the flat skin of a female of C. argyrochetes from Szechuen.

On this I could find no pedal glands except a shallow depression

on one of the anterior feet, recalling the aborted pedal glands of

Capra, Psewdois, and Hemitragus. Moreover, the hair on the

front of the pastern did not extend so far downwards between

the hoofs as in C. crispus and Naemorhedus raddeanus. The state

of preservation of the feet on this skin does not justify the final

* Abstr. P. Z.S. no. 55, p. 12, 1908; P. Z.S. 1908, p. 183, fig. 34.

+ P.Z.S. 1836, p. 39.9 .

56%

856 MR. R. I. POCOCK ON THE [June 14,

conclusion that the glands are absent ; and until more is known

of the pedal glands of the other species referred to Capricornis,

no certain conclusions can be drawn regarding the taxonomic

value of the facts above recorded. If C. argyrochetes prove to be

without glands and C. thar to have them, there will be a strong

prima facie case for regarding these species as belonging to two

distinct genera. ©. thar is the type of Capricornis, and for

C. argyrochetes the name Nemotragus Heude will have to be

employed.

Finally, it may be added that Owen states that C. swmatrensis

possesses inguinal glands as well as the preorbital glands—a fact

which still further emphasises the importance of the examination

of fresh material of this little-known genus.

Genus Buporcas Hodgs.

BuboRCAS TAXICOLOR WHITEL,

(The Bhutan Takin.)

In the living example of this Takin in the Gardens there is no

trace of preorbital gland, and none of pedal glands on the fore

feet, as may be clearly seen when the animal rears herself up

with the hoofs spread against the bars.

Buporcas trserANus M.-Edw. (‘The Chinese Takin.)

(‘Text-fig. 84, B, p. 852.)

Mr. Gerrard kindly allowed me to cut the fore and hind feet of

a female example-of this species from Szechuen, There was no

trace of pedal glands on either the fore or the hind feet. On the

fore feet the integument on the anterior surface of the pastern

ended in the middle line below in a strong transverse web,

beneath which it passed between the hoofs nearly parallel with

the proximal edge of the inner surface of the nail, then turned

sharply at right angles, forming a strong heel-tie running parallel

with the sole of the hoof before turning up to be continued with

the skin of the back of the fetlock. Asin the Serow ( Capricornis

argyrochetes) practically the whole of the interungual integument

was naked, although anteriorly the hair on the front of the fet-

lock was continued a little lower, slightly below the transverse

web. The hoofs themselves were shaped very much as in Capri-

cornis. but the heel-tie was situated more forwards and nearer

the proximal edge of the inner lamina of the nail, making the

hoofs much less distensible than in that genus.

The structure of the hind foot was essentially the same as that

of the front foot, but there was no trace of the anterior integu-

mental web. ;

There was no trace of inguinal glands, but the four teats on

the dried skin were curiously invaginated, giving the appearance

of four small symmetrically arranged pits, which I at first mistook

for inguinal pouches.

1910. ] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 857

The genera of Rupicaprine: may be distinguished as follows by

their cutaneous glands :—

a. Mufile large, encircling the nostrils below and extending as far

as their posterior angle above; a well-developed pedal gland

like that of Ovis on all four feet.

b. Preorbital glands present: .......002..cccesssccesssenseeaeesteseossecee, Capricornis.

Grebreoupicalec lands) ab senimeeseeeeerrce aia s.deseas. sca Nemorhedus.

a. Muffle small; pedal glands, when present, otherwise formed.

e. A pair of glands behind the horns in both sexes ; interdigital

cleft deep.

d. Interdigital glandular cleft opening inferiorly between the

hoofs, the heel-tie Sim plemarereas en. . Oreamnos.

. Interdigital cleft shut below by a ances fold of PAteornent

extending forwards from the heels towards the proximal

margin of the nail in front ............... steers. Lepicapra.

ce’. No glands behind the horns; ‘vier areal cleft shallow Be. fe Budorecas.

Subfamily OVIBOVIN&.

Genus Ovreos Blainv,

Ovisos moscHatus Linn. (The Musk-Ox.)

(Text-fig. 86.)

According to Ogilby this animal has fowr teats, but neither

preorbital nor inguinal glands. Of the pedal glands he said

nothing. Lonnberg (P. Z. 8. 1900, pp. 163-165) agrees as to the

number of teats and apparently as to the absence of inguinal

glands ; but he found in his specimen the preorbital gland well

developed, being 23 mm. long, 18 mm. broad, and 8 mm. deep.

The interior of the pit was lined with hair, but I cannot judge

from the description whether the orifice was valvular as in Ovis

or simple as in Capricornis (the Serow).

Lonnberg also states that he failed to find any pedal glands.

I was able to verify this fact on a front and a hind foot of

Musk-Ox which Mr. E. Gerrard kindly allowed me to cut. On

the fore foot the skin of the middle line of the pastern in front

ends inferiorly in a small web, beneath which the integument runs

backwards and downwards to the anterior portion of the heel,

then curves upwards and backwards towards the posterior portion

of the heel, where it is continuous with the skin of the posterior

surface of the pastern. This skin is covered with long hairs, which

pass for a short distance between the heels, but the gre eater part

of the integument which ties the heels together is quite naked.

Above the heel- tie, however, the inter ungual integument is

covered with long hairs, which largely overlap the proximal inner

margin of the nail; and when the hoof is examined from below

these hairs appear as a long and thick tuft projecting forwards from

the anterior portion of the heel-tie and filling the greater part

of the interungual space. Long hairs arising from the bottom of

the pastern in front also overlap the proximal anterior portion

of the hoof. Asis well known, the hoofs are remarkably short

858 MR. R. 1. POCOCK ON THE [June 14,

and broad, the antero-external portion of the nail being enormously

developed as compared with the small internal portion, the soles

and heels being correspondingly large.

The hind foot is very like the fore toot, and has no trace of a

gland; nor on the specimen examined did 1 find any trace of the

small antero-superior web, and the integument of the front of

the pastern descended move abruptly between the hoofs down to

the heels, making a more marked depression between the hoofs

than was visible before the front hoof.

Text-fig. 86.

: “En hi

pp ME

Ovihos moschatus.

A. Median vertical section of front foot.

B. The same of hind foot.

C. Lower surface of hind hoof.

The systematic position of Ovibos must for the present be

left unsettled. According to Lonnberg, it should rank as a special

subfamily, for which the name Ovibovine is retained. Gray *,

it may be added, long ago placed the Takin (Budorcas) with Ovibos

in the family Ovibovide ; but the evidence that the two are really

related is not convincing, and cannot be settled until the anatomy

of Budoreas is known.

* Cat. Rum. Mamm. in the British Museum, p. 31, 1872.

1910.] CULANEOUS SCENT-GLANDS OF RUMINANTS. 859

Subfamily Carprin a,

Genus Ovis L.

(Sheep.)

Ogilby stated that in O. aries (Domesticated Sheep) the preorbital

glands are large but immovable, the pedal glands small and the

inguinal glands absent, adding that there is only a single pair of

mammee,

According to Hodgson, O. ammon, the Siberian Argali, and

O. ammon hodgsoni (=ammonoides), the Tibetan Argali, have

large but immovable preorbital glands, distinct inguinal glands,

and pedal glands on all four extremities, whereas O. musimon, the

Moutlon, for which on that account he created the genus Caprovis,

differs in having no pedal glands.

Owen (‘ Anatomy of Vertebrates,’ iii. p. 638, 1868) described and

figured the pedal gland presumably of a domesticated breed ; but

according to Isidore Geoffroy the glands are absent in some breeds ;

most modern text-books and systematic treatises state that the

preorbital, pedal, and inguinal glands are present, and that there

is a single pair of mamme, a statement with which my own

observations are entirely in accord except that there is occasionally

an additional pair of small mammee.

I have examined fresh specimens of two wild species, namely,

O. vignet and O. musimon, and a dried skin of O. canadensis.

Ovis vieNeI Blyth. (The Urial or Gad.)

(Text-figs. 87, 88.)

The preorbital gland consists of a hair-lined valvular pocket or

invagination closed by a flap of skin forming a movable upper lid.

The pocket in captive specimens is commonly blocked with a nearly

dry and waxy or quite dry secretion with no definite smell

but that of “sheep,” and the hairs around the orifice are generally

stuck together with the same substance. The dryness of the

secretion is probably pathological and due perhaps to insufficiency

of green or natural food.

The inguinal glands, two in number, consist of rather shallow

pouches, with the deep part underlying the teat, close to which

the wide suberescentic orifice opens. The secretion is yellowish

and waxy, and gives off a powerful odour, just like that of the

urine of the common House Mouse (Jus musculus). The area

of skin surrounding the teat and gland is naked. Normally

there is a single pair of teats. In one specimen, however, I

found an additional smaller pair lying in front of the pair

adjacent to the orifice of the gland.

The structure of the feet is very constant. When the hoofs are

pulled apart a triangular hairy depression is seen in the middle line

of the lower half of the pastern. Thisdepression is caused by the

860 MR. R. I. POCOCK ON THE [June 14,

Text-fig. 87.

a “)

TM. | \\\ (lw,

Ovis vignei.

A. Inguinal region of young male: m., mamma; gl., orifice of gland, the

size of the invagination shown by a dotted line; Se., scrotum.

B. The same of lamb three days old, showing the shallow glandular

depression (g/.).

C. Front foot of an adult showing the orifice of the pedal gland (o.).

Text-fig. 88.

os eS AWAY

£3 = N\\ \

Ovis vignei.

A. Median vertical section of foot of lamb three days old.

B. The same of adult.

gle, pedal gland; o., its orifice.

1910. } CUTANEOUS SCENL-GLANDS OF RUMINANTS, 861

somewhat abrupt descent of the integument just above the hoofs.

It descends straight down to the heels, which are united by a

strong tie running for a short distance parallel to the sole of the

hoof before turning upwards in continuation with the skin of the

back of the pastern. The heel-tie is the only portion of the

interungual integument which is hairless, the area above the tie

being covered with long hairs which project forwards, overlapping

the proximal margin of the nail in front.

The hoofs are short and pointed and the heels of the hoofs well

developed, being both high and long, so that the integumental tie

between them is raised considerably above the ground.

The pedal gland opens by a small cireular orifice near the

summit of the triangular depression on the front of the pastern,

some distance above the hoofs. The orifice leads into a’ narrow

duct which quickly but gradually expands into the gland. The

latter descends along the interungual integument towards the

heel, then turns sharply upwards and backwards upon itself for

a short distance along the integument of the back of the pastern.

The gland is lined with short hairs, and the secretion is a clear

semi-fluid substance with a pleasant scent like toffee (baked

butter and sugar), slightly infused with acetic acid.

Of this species [ have examined several specimens of various

ages from Persia. One example three days old had all the glands

developed, the pedal glands being as large relatively as in the

adult, the preorbital gland only a little smaller, but the inguinal

glands decidedly shallower. I have also noticed that the mguinal

pouches are sometimes quite shallow in Sheep of domesticated

breeds.

OvViIS MUSIMON Schr.

(The Sardinian Mouflon.)

This species has the glands of the same structure as in O. vignei,

with the preorbital gland smaller ; but in one specimen | noticed

an unexpected difference in the feet, the entire interungual area

from a point on a level with the front of the hoof downward

including the heel-tie being naked instead of hairy.

OvIs CANADENSIS Shaw.

(The Rocky Mountain Bighorn.)

In a dried skin of this species, for which I was indebted to

Mr. E. Gerrard, I found the preorbital gland, which Mr, Lydekker

has described as minute and almost rudimentary, to consist of a

small but very distinct valvular invagination, only differing in its

relatively smaller size from that of other species of Sheep. ‘The

presence of a pair of inguinal glands was unmistakably indicated

and there was a single pair of teats. Remains of the pedal glands

showed that these organs open as in ordinary Sheep, and the

interdigital area was clothed with hair down to the heel-tie.

862 MR, R. 1, POCOCK ON THE [June 14,

Domesticated Sheep.

Of Domesticated Sheep I have examined examples of many

breeds from different countries without finding any variations of

moment in the structure of the pedal, inguinal, or pr eorbital glands.

The rarest specimens were the following :—a lamb of three or four

months old from Fair Island between the Or kneys and Shetlands,

belonging to Mr. A. J. Sewell; a black lamb of about two months

old, probably belonging to a four-horned breed from Caithness in

Scotland ; an adult short-tailed ram of the breed from the island

of Soa in the Hebrides ; a Hunia or Fighting Ram from Nepal

and a four-horned breed from the same country, both short-tailed ;

two rains of the long-tailed maned breed belonging to the Haussa

tribes of N. Nigeria; and a fat-tailed South-African Sheep

belonging to Sir Claud Alexander *,

In three of these sheep, ni amely, in the Caithness and Soa

specimens and in the Hunia Ram, I noticed that the interungual

area was hairy, exactly as in O. vignei, and not naked as in the

example of O. musimon described above.

In no Sheep, domesticated or wild, have I found the pedal glands

absent ; and in every case where the scent of the secretion was

tested it had the sweet odour of toffee. The histology of the

pedal glands of Ovis aries has been fully deseribed by Max

Tempel.

Genus Ammorracus Blyth.

AMMOTRAGUS LERVIA Pall. (=Ovis TRAGELAPHUS).

(The Barbary Wild Sheep.)

| have had no opportunity of examining a dead specimen of

this species ; but careful inspection of a living male example gave

the following results.

There was no trace of preorbital gland. Inguinal glands also

were absent, and there was a single pair of teats. The feet were

constructed as in Sheep and Goats ; but there was no trace, either

upon front or hind feet, of any orifice leading into a special gland

such as is present in Sheep. In Sheep the orifice of this gland

may be seen at once by superficial scrutiny of the front of the

pastern, in spite of the hairy clothing. The most striking

peculiarity about the feet of this example of O. lervia was the

apparent smoothness of the interungual integument and the

* It is commonly stated in works on Natural History that domesticated breeds of

Sheep differ from all the wild species in having long tails. This is not true. Of the

above-mentioned Sheep the shortest tail, containing only 5 vertebrae with a short

terminal unsegmented portion, was found in the Soa breed. The Fighting or Hunia

Ram from Nepal had a tail 6} inches long, containing 10 vertebra. In the Caithness

breed the tail scarcely reached the hocks and contained 15 vertebrw. In the Haussa

Sheep, on the contrary, the tail nearly reached the fetlocks and was composed of

20 vertebra, as in a fat-tailed Asiatic Sheep mentioned by Pallas. It may be added

that Nepalese Sheep of the Hunia, four-horned, and one-horned races all have

short tails, the tail of an adult four-horned specimen measuring only 34 inches.

POLO CUTANEOUS SCENT-GLANDS OF RUMINANTS. 863

shortness of the hairs clothing the interdigital depression on the

front of the pastern.

In the absence of preorbital, of inguinal, and of specialised

pedal glands Ammotragus lervia agrees with Pseudois, Hemitragus,

and Capra, and differs from Ovis. The glands, in fact, completely

bear out the conclusion suggested by the structure of the skull

and certain external features described in text-books, that Ammo-

tragus cannot consistently be included in QOvis if Capra he

excluded therefrom.

I do not know who was the authority for the statement found

in Mr, Lydekker’s work ‘Wild Oxen, Sheep, and Goats,’ pp. 149

& 226, 1878, that this animal possesses pedal glands. The

invariable presence of these glands is cited as characteristic of

Ovis as contrasted with Capra; and the species in question is

included under Ovis, although subgeneric rank is assigned to it.

Genus PsEuDoIs Hodgson.

Psrupors NAyYAUR Hodgs. (The Bharal.)

(Text-fig. 89.)

According to Hodgson, O. nayaur has no preorbital glands, but

possesses small pedal glands on all four feet and two inguinal

glands, marked, however, by vague pores. It was principally on

Text-fig. 89.

Pseudois nayaur.

A. Median vertical section of front foot of female retaining the aborted gland (9/.).

B. The same of the male without trace of the gland.

the absence of the preorbital glands that Hodgson separated this

species from Ovis as the type of a distinct genus Pseudois.

Blanford repeats the information supplied by Hodgson, but

without quoting him as his authority.

864 MR, R. I, POCOCK ON THE [June 14,

[ have examined two fresh examples, male and female, of this

species, and can endorse what Hodgson says with regard to the

absence of the preorbital gland, though its position was marked

by a small patch of naked skin. I found, however, no trace of

inguinal glands, and Hodgson himself admits that the pores

(invaginations) are “ vague.” Perhaps the glands he found were

mammary glands. As for the pedal glands, I discovered on the

fore feet of one specimen (a female) a very small invagination

dipping into a subeutaneous glandular patch, exactly like the

gland described above as occasionally occurring in Goats. On the

hind feet a similar small pocket was present, but there was appa-

rently no differentiated gland. In a full-grown male there was no

trace of the gland or the invagination on any of the feet. Except

that the interdigital depression is perhaps a little shallower, the

feet do not differ structurally from those of Goats. The depression

is covered with long hairs, the tips of which overhang the margin

of the nail; only the bridge of skin holding the heels together is

naked, The practical suppression of all the glands found in typical

Sheep is,in my opinion, ample justification for regarding the Bharal

as the type of a genus distinct from Ovis, quite apart from other

structural differences such as those supplied by the skull. Whether

or not Pseudois can be distinguished from Capra, to which it is

linked by one of the races of C. cylindricornis, is another matter.

renus Capra Linn. (‘Text-fig. 90, B.)

All authors seem to agree that the preorbital and inguinal

glands are invariably absent in this genus; but there is no such

unaninity as regards the pedal glands. According to Ogilby they

are present ; whereas Hodgson says they are either altogether

wanting, or present only on the fore feet, and Isidore Geoffroy *

states that they are very rarely present. Hodgson records their

occurrence on the fore feet in specimens of three distinct breeds of

domesticated Indian Goats, namely the Chyapu, the Sinal, and the

Digi. He gives, moreover, an illustration of an extracted gland,

which shows it to be of the same general structure as in Sheep,

although relatively smaller t

Having examined the feet of many English Goats killed

in the Society’s Gardens for food, I can endorse the accuracy of

Hodgson’s statement that the glands are absent in the hind feet

and present or absent in the front feet. In the latter, however,

they are much more commonly absent than present, and I have

never found them so well developed as Hodgson represents $.

They are, when present, represented by a shallow pocket-like

invagination bearing a few short hairs, those that protrude from

the orifice being stuck together with secretion. This invagination

* Hist. Nat. gén. ili, p. 435.

+ J. A. 8. Bengal, xvi. pt. ii. pp. 1016-1026, 1847.

+ Max Tempel also only occasionally found an aborted gland in Dome sticated

Goats.

1910. | CUTANEOUS SCENT=GLANDS OF RUMINANTS. 865

occupies the same position as the orifice of the gland in Ovis; and

this fact, coupled with the close resemblance that Hodgson showed

to exist between the glands of Capra when well developed and those

of the former genus, suggests that the ancestors of existing Goats

had glands on the fore feet, and probably also on the hind feet,

like those of Sheep. The feet themselves also closely resemble

those of Sheep. There is a deep depression between the penul-

timate phalanges, formed by the abrupt downward slope of the

integument of the upper (anterior) side to meet that of the lower

side at an acute angle just above the heels, forming a strong some-

times horny union between them. The heels are thus incapable

of much distension, whereas the tips and proximal portion of the

upper side of the hoofs can be widely separated. The heels are

Text-fig. 90.

A. Median vertical section of front foot of Hemitragus jemlaicus.

B. The same of Capra hircus (var. dom.) retaining aborted pedal gland (g/.).

high and prominent behind. So far as the hairiness of the inter-

digital depression is concerned, great variation exists. In some

cases it is covered with longish hairs, in others it is quite naked,

an intermediate condition existing in others. Seeing how constant

a specific character the hairiness of this area is in wild species of

Ruminants, [ am at a loss to know what explanation to put upon

its variability in these domesticated animals, which are supposed

to be descended from C. egagrus.

Domesticated Goats have two shallow glandular pouches at the

base of the tail beneath, one on each side, close to the anus. I

have also found these in Capra cylindricornis and falconeri.

Of species usually referred to Capra 1 have examined the fol-

lowing specimens, in addition to the domesticated examples alluded

866 MR. R, 1, POCOCK ON THLE (June 14,

to above, and found no trace of preorbital, inguinal, or pedal

glands in any of them :—

C. egagrus trom Crete: one young male.

(", sithirica from Chamba: one young male.

C'. falconeri trom N.W. India: two adult males.

(. cylindricornis from the Caucasus: one young male and

one female kid that died at birth.

The structure of the feet resembled that of Domesticated Goats,

and the interdigital depression was in all cases hairy, but showed no

sign of secretion such as was observable in the example of Oreamnos

montanus described above.

Genus Hemirracus Hodgs.

HemrrraGus Jemiarcus Hodgs. (The 'Tahy.)

(Text-fig. 90, A, p. 865.)

According to Hodgson this species has no preorbital, inguinal,

or pedal glands. The specimens, three in number, which I have

examined bear out this statement, except that in one female

example [ discovered on the hind feet an abbreviated glandular

pocket like that found on the hind feet of a female Psewdois

nayaur. Sometimes, at least, there are four mamme.

Hodgson included in his genus /emitragus, HM. hylocrius

from the Nilgiri Hills; but I cannot find any evidence that

he examined a specimen of that species for the points mentioned

in his generic diagnosis. He separated Hemitragus from Capra

because of the larger size of the “mutte” or naked skin at the

end of the nose, the alleged presence of four teats, and the absence

of pedal glands, which he himself adinits may be altogether absent

in Capra. According to Blanford, however, //. hylocrius has but

a single pair of teats, thus agreeing with typical Goats.

Hodgson states that the male Tahr has a strong Caprine odour,

[t is true that in the winter and early spring the animal smells

strongly; but the odour is exactly like that of a Cormorant or

Frigate-Bird, and not like the odour of Capra c@gagrus ox

falconeri.

Judged by their glands, the genera of Caprine fall into two

categories, as follows :—

a. With well-developed preorbital, pedal, and inguinal glands, the

latter not always invaginated in domesticated breeds............... Ovis.

a’. Without preorbital and inguinal glands; pedal glands, when

present, represented by avery shallow depression, except in some

Indian Domesticated Goats, where glands resembling those of

Ovis ave found on the fore feet.

Ammotragus, Pseudois, Hemitragus, Capra.

1910. ] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 867

Subfamily CerpHALOPHIN &®.

(The Duikers.)

According to Owen the following species of this subfamily have

inguinal pits and preorbital glands lodged in a depression of the

maxilla :—Cephalophus silvicultrix, grimmi (=mergens, burchelli,

grimmi), monticola (perpusilla, pygmea), maxwell.

Two species were examined by Ogilby, namely, C. monticola,

cited as T'ragulus pygmeus, and C. grimmi, cited as Sylvicapra

mergens. According to this author C. monticola has neither

inguinal nor interdigital glands, whereas in C. grimmi there are

small interdigital glands and distinct inguinal glands, both species

having oblong maxillary [preorbital] glands lodged in depressions

of the maxilla. Ogilby also records the presence of two pairs of

mami in this genus *.

The preorbital gland is described in most recent systematic

works as consisting externally of a bare streak of skin studded

with a series of pores and situated some distance in front of the

eye of the corresponding side. Weber? describes in detail the

histological structure of the gland and the nature of its secretion

in C. pygmeus |[=monticola|, points out the difference it presents

in the male and the female, and gives a list of the species in which

he had observed its external form, together with much other

interesting matter. The pedal glands do not appear to have been

described.

Genus GuEVEt Gray £.

GUEVEI MAXWELLI H. Sm. (Maxwell’s Duiker.)

(Text-figs. 91, A, & 92.)

In a temale example from Portuguese Guinea, presented by

Dr. Ansorge, the external streak of the preorbital gland was

markedly bent downwards at its posterior end towards the anterior

corner of the eye, its lower margin being therefore concave. It

was studded with about 25 vertically elongate pores, forming a

single series, from each of which a few short hairs arose. With

pressure, liquid drops of secretion could be easily squeezed from

the pores. The gland itself underlying the bare streak did not

* The specific names perpusillus and pygmeus have been assigned to the so-called

Royal Antelope , Neotragus pygmeus; and Sclater and Thomas cite Tragulus

pygmeus of Ogilby in the synonymy of Neotragus pygmeus. This view is, |

think, incorrect, because Ogilby described the preorbital glands of Tragulus pugmeus

and of Sylvicapra mergens in exactly the same terms as “glandule maxillares

oblong ”; and since Sylvicapra mergens is admittedly a Cephalophine, one must

infer that Ogilby’s Tragulus pygmeus also belonged to that subfamily and not to the

Neotragine. That Owen’s Antilope perpusilla and pygmea weve also Cephalophiaes

is clearly indicated by his placing them with other species of Cephalophus ina group

characterized by “ maxillary ” as opposed to “ suborbital pits.” Moreover, he quotes

S. Africa as the locality of perpusilla. whereas Neotragus pygmeus comes from

Liberia. : in

+ Arch. mikr. Anat. xxxi. pp. 507-539, 1888.

t Reasons for the adoption of this genus are given below (pp. 874-876).

868 MR. R, I, POCOCK ON THE [June 14,

differ in structure and consistency from that of G. melanorheus and

Sylvicapra coronata described below. When alive the animal] had

the habit, not noticed in other Duikers, of rubbing the glandular

area against one’s hand when presented to her.

There were no inguinal glands, but two pairs of teats were

present.

The pedal glands were well developed. The skin of the back

of the pastern extended right down to the heels, forming a strong

union between them. It was then folded sharply back upon

itself, as far as the false hoofs, to form the posterior wall of the

gland. At its deepest or upper part the gland formed a laterally

Text-fig. 91.

A. Head of female of Guevei maxrwelli, showing the preorbital gland (g/.).

B. Longitudinal section of the preorbital gland of Sylvicapra coronata.

C. Transverse section of the same.

compressed subcireular sac which communicated with the tri-

angular interdigital depression, seen when the hoofs were pulled

apart, by means of a narrow tube with a gradually expanding

subeircular orifice. This interdigital depression was formed by

the folding back of the skin of the anterior side of the pastern

some distance above the proximal margin of the hoof, the infolded

portion forming the anterior wall of the gland and constituting a

somewhat deep anterior interdigital web which did not project

nearly so far inferiorly as the posterior shallower but stronger

1910. | CUTANEOUS SCENT-GLANDS OF RUMINANTS. 869

web tying the heels together. The walls of the interdigital

depression were thickly covered with long hairs projecting down-

wards over the proximal margin of the nail, those on the anterior

side of the posterior web being especially long, so as to conduct

the secretion downwards between the heels of the hoofs. The

dilated cavity of the glands was beset with short scattered hairs

which gradually lengthened towards the orifice of the gland. The

secretion itself was white and sticky, with a repulsive odour of

dog’s feces mixed with acetic acid.

Text-fig, 92.

Guevei maxvwelli.

A. Median vertical section of hind foot: o., orifice of pedal gland,

B. Front foot dissected to show the gland entire.

(Fresh specimens.)

In another female specimen from Sekondi also referred to

G. maxwelli, the preorbital streak, although lightly concave down-

wards, was not so abruptly angled at its posterior end as in the

one described above. The streak was 18 mm. long, 2 mm. wide,

and had a row of about 20 vertically elongated pores, from each

of which a few small bristles arose. Drops of fluid could be

squeezed in the same way from these orifices. There were no

inguinal glands, and the pedal glands resembled those of the first-

described specimen.

The absence of the inguinal glands in these two specimens of

G, maxwelli suggests that Owen was probably wrong in assigning

these organs to the species, for it does not seem probable that

Proc. Zoou. Soc.—1910, No. LVI. 57

870 MR. R. I, POCOCK ON THE [June 14,

such a character is subject to individual variation within specific

limits. 1 suspect, moreover, that Ogilby was correct, and, by

implication, that Owen was in error, in stating that inguinal

glands are absent in G. monticola, since the latter seems to be

nearly allied to G. maawelli. But I cannot doubt that Ogilby

was wrong in saying that pedal glands are absent in G’. monticola.

GUEVEI MQUATORIALIS Matsch. (Text-fig. 93, A, B.)

The cut foot of a dried skin given to me by Dr. Christy, who

shot the animal in the Chagwe forest, Uganda, had a gland of

nearly the same structure as in G'. melanorheus. The foot of this

specimen differs in colour from that of G. maxwelli, being brown

all over and showing no white patch just below the false hoofs

on the back of the pastern and no white on its front close to the

triangular interdigital depression.

The similarity in the structure of their pedal glands favours

the view that G. melanorheus and G. equatorialis may be sub-

specifically and not specifically distinct.

GUEVEL MELANORHEUS Gray. (Text-fig. 93, C.)

A single young specimen from Angola presented by Mr. H. F.

‘Varian,

As in G. maxwelli, the preorbital streak was lightly concave

below, but its curvature was less abrupt and situated farther

forwards than in Dr. Ansorge’s example from Portuguese Guinea.

The streak was 22 mm. long and 1 mm. wide, studded with a

single series of circular pores, from each of which a single short

hair arose. The gland itself, when extracted, measured about

27 mm. long, 10 mm. wide, and 5 mm. in vertical depth. In

section, both longitudinal and transverse, it was pale externally,

with its central third forming a dark core. Under slight pressure

drops of clear fluid, like liquid gum tragacanth, could be squeezed

from the pores. When rubbed between the fingers it became, as

it dried, quite sticky, giving out a faintly aromatic odour like that

of black-currant tea. Under hard pressure pale whitish-grey

maggot-like threads could be expressed from the pores.

Inguinal glands were absent as in G. maawelli.

The feet were constructed as in the last-named species, with

the marked difference that the pedal gland consisted of a simple

narrow cylindrical tube not extending so far as the false hoofs

and showing no sign of expanding between the bones into an

enlarged laterally compressed dilatation.

Although superficially G. maawelli and G. melanorheus are so

much alike that it has been suggested that they be regarded

merely as subspecies, it appears to me that the difference in the

structure of their pedal glands entitles the two to specific separation,

at all events until further examination has shown that this

character has not the value here claimed for it.

1910.| CUTANEOUS SCENT-GLANDS OF RUMINANTS. 871

Text-fig. 93.

A. Median vertical section of hind foot of Guevei equa

B. Do. of front foot of the same.

C. Do. of hind foot of G. melanorheus.

(All dried specimens.)

Genus CrePpHALopHuus H. Sm.

CEPHALOPHUS DORSALIS Gray. (The Bay Duiker.)

A single example from Sekondi presented by Capt. CO. H.

Armitage.

The preorbital streak was straight, 22 mm. long, 1 mm. wide,

and studded with a single row of 14 pores, from which grey-

coloured fluid could be pressed.

Inguinal glands were well developed and widely separated by

a space of 42 mm. from the two mamme of each side. There was

only a single pair of these glands. Each consisted of a long sac

very gradually expanding internally from the orifice, the walls of

57*

872 MR. R. I, POCOCK ON THE [June 14,

the bottom third of the pocket being corrugated and ridged. The

secretion was yellow and sticky. ‘The structure of the feet was

almost the same as in the species already described, except that

the fold of integument forming the web at the heels was enor-

mously thickened and horny, forming a very strong union between

the two hoofs behind. Moreover, this interdigital depression was

only scantily clothed with shortish hairs, the tips of which

scarcely overlapped the margin of the nail of the hoofs, except

Text-fig. 94.

hi.

A. Median vertical section of foot (? front or hind) of Sylvicapra qrimmi, subsp. ?

B. The same of hind foot of Cephalophus nigrifrons. h.t., heel-tie.

(Both dried specimens.)

quite in front. The pedal gland itself was not dissected out, but

its circular orifice was apparent. Apart from other features,

C. dorsalis differs from the species of the maxwelli group in

possessing inguinal glands; and in this connection it may be

noted that Owen records the presence of inguinal glands in

(. silvicultrix, one of the larger species of the genus usually

considered to be related to (’. dorsalis.

(EPHALOPHUS NIGRIFRONS Gray. (The Black-fronted Duiker.)

(Text-fig. 94, B.)

A female example of this species, kindly lent to me by

Mr. Gerrard, agreed with C. dorsalis in possessing a pair

1910. | CUTANEOUS SCENT-GLANDS OF RUMINAN'S. 873

of inguinal glands consisting of deep subcylindrical pouches,

with the orifiees remote from the mamme. ‘The interdigital

cleft was scantily hairy and the heel-tie was very thick as in

C. dorsalis ; and the pedal gland on all four feet consisted of a long

expanded sac with a small orifice, as in G. maaielli and in

S. coronatus.

CEPHALOPHUS SILVICULTRIX.

(The Yellow-backed Duiker.)

In two dried and flat skins from the Cameroons, kindly given

to me by Rowland Ward, Ltd., I found the feet and pedal glands

constructed as in C. nigrifrons. About the inguinal glands on

these skins I can affirm nothing. Owen, however, says they were

present in the species.

Genus Syiyrcapra Ogilby.

SyLvrcaPRA coronata Gray, 1842.

subsp. CAMPBELLI® Gray (2) 1843 *.

Of specimens referred to this species, | have examined several

from various parts of Nigeria, including two young males from

Sokoto, presented by Major Searight, and one from Ibadan in

the Yoruba country, presented by Mr. 'T. Christ.

In all of them the preorbital streak was lightly concave

upwards, thus differing in shape from that of the Duikers of the

maawwelli group, and also, though in a lesser degree, from that of

C. dorsalis. The streak was studded with a series of 15 or more

pores, from which fluid secretion with a pleasant aromatic odour

could be squeezed. Under continued pressure black coiling worm-

like threads of a waxy consistency exuded.

A pair of inguinal glands consisting of wide but (in the young)

shallow pits were present. They were much shallower than in C. dor-

salis; but their orifices, as in that species, were situated a long way

from the two mamme on each side, lying in fact just on the femoral

side of the fold or crease between the inner surface of the thigh

and the abdomen; and, as in ©. dorsalis, the whole inguinal

and mammary area of the abdomen was naked as far out as

the glandular orifices. The secretion of the inguinal glands

smelt like sour milk in two examples, but had an objectionable

feecal odour in another.

The feet were constructed as in other species of this group,

* According to Gray (Cat. Rum. B. M. p. 23, 1872), C. campbellie was in 1872

represented in the British Museum by a single skin from West Africa, which he

marked interrogatively as having been obtained at Sierra Leone. The skin agrees

tolerably closely with the Nigerian examples that I have seen. I think, therefore,

that Gray was probably correct in assigning his type to West Africa; and that it

was an error to suppose that the name campbellie belongs to a Natal form of

Sylvicapra (Lydekker, ‘Game Animals of Africa, p. 142, 1908). S. coronata Gray,

from Senegambia, appears to me to be a paler form of the same species.

874 MR, R. I. POCOCK ON THE [June 14,

but the walls of the triangular interdigital depression leading

to the gland were very sparsely clothed ‘with quite short hairs,

much fewer and shorter than in C. dorsalis. Only at the

inferior margin of the side walls of the depression were there

long hairs overlapping the proximal edge of the nail of the

hoof. The gland itself consisted of a large laterally compressed

sac, very much like that of G. maxwelli, but with the narrowed

portion much shorter and wider, and with no long hairs

jutting from the orifice, the walls of the sac being clothed with

very short sparsely set hairs, like those of the “triangular de-

pression. As in @. mawwelli, the glandular tissue could be traced

not only over the walls of the gland itself but from the commence-

ment of the anterior web of the hoofs to that of the posterior web.

There was, however, no secretion in the glands.

My observations on this species agree with those made on

S. grimmi by Ogilby, who stated that both inguinal glands and

pedal glands were present, the latter being small. Owen also

found inguinal glands in S. grimmi.

In an odd foot of a Sylvicapra closely related to the typical

S. grimmi, and given to me by Mr. Gerrard (text-fig. 94, A) I

found pedal glands resembling in all essential points those of the

legs of the examples identified above as S. coronata.

The featuresto be noticed in connection with the cutaneous glands

of the Cephalophine are the general constancy of the structure of

the preorbital gland and of the feet and the minor differences pre-

sented by each. The preorbital gland is always large, oval, tough

and india-rubbery in consistency, is situated farther forwards

than in other Antelopes, and has no valvular invagination of the

skin connected with it as in Gazelles, Sheep, Dik-Diks, and others.

Instead of this it communicates with the exterior by means of

a series of setiferous pores placed on a narrow strip of bare

integument. It differs, in fact, from that of all other Ruminants.

The structure of the feet, too, is very uniform. They do not,

however, differ materially from those of Jadoqua, as illustrated

by JW. phillipsii, nor from genera of Neotragine that I have

examined, namely Owrebia and Raphicerus. Only i in one little

point is there, within my experience, a constant difference.

Whereas in the Madoquine and Neotragine the web joining the

heels is covered continuously with long ha airs, in the Cephalophins

the back of this area of integument is always naked for a longer

or shorter distance.

Another peculiar feature of the Cephalophine is the wide

separation of the inguinal glands, when present, from the

mamme. In this particular, however, they resemble some

Tragelaphine and Pantholops.

In my opinion the absence of the inguinal glands * in the forms

* This difference, as well as others, between me/anorheus and dorsalis was noted

on the same material by Mr. F. E. Beddard (P. Z. S. 1909, i. p. 181).

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 875

above identified as maxwelli, equatorialis, and melanorheus, points

to their generic separation from those referred to dorsalis,

Text-fig. 95.

<1 ayy Mt, Ree

TAO HEREIN NSS)

LIN By) } VY \ns we

Zi il DN) ) Hae

ss UY) Wye e e 7

Sylvicapra coronata.

A. Inguinal region of male: p., penis; m., mamme ; gl., orifice of gland ;

se., scrotum.

B. Median vertical section of front foot.

€. The same of hind foot.

0., orifice of gland ; A.., heel-tie.

(All fresh specimens.)

876 MR, R. I, POCOCK ON THE | June 14,

nigrifrons, and coronata, of which abyssinica* is, 1 think, a sub-

species.

It is, moreover, well known that the group exemplified by

coronata differs from that exemplified by dorsalis in the

markedly more upright inclination of the ‘horns. If generic

importance be attached to that character, a proceeding desirable

on the grounds of the unwieldiness of the genus Cephalophus as

at present recognised, the genus can be split at all events into

three genera which may be distinguished as follows :—

. Inguinal glands absent ........ .. Guevei.

. Inguinal “glands present, consisting of a pair of deep pouches,

the orifices of which are remote from the mammr.

b. Horns inclined backwards in the plane of the eo ears

NOM cass .. Cephalophus.

. Horns inclined upw ards and backw ards, forming : an obtuse

angle with the plane of the face; ears long Seis scdseReces hee, GS LOC Open

The type of the genus Guevei, as selected by Messrs. Sclater

and ‘Thomas, is maaweili, with which equatorialis and melanorheus

must be associated. The type of Cephalophus is said to be silvi-

cultriv, @ species in which Owen affirms the existence of inguinal

glands. As defined above, the genus also contains dorsalis and

nigrifrons. Thetype of Sylvicapra is mergens, but grimmi appears

to be the oldest name given to the species.

For the characters of 7etraceros, usually referred to the Cepha-

lophinze, see p. 921.

Subfamily MapoQquiIn 4,

Genus Mavoqua Ogilby.

(Dik-diks. )

The available evidence suggests that the species of this genus

possess preorbital and pedal glands, but are without inguinal

glands. Im the case of J/, saltiana, both Owen and Ogilby are i in

agreement about the presence of preorbital and the “Abeenee of

inguinal glands, and Ogilby adds that the pedal glands are also

present.

MApoqua puiiiipsit Thomas. (Lort Phillips’s Dik-dik.)

(Text-fig. 96.)

Of this species I have examined, in the fresh state, one example

which agreed with the spec simens of M. saltiana seen by Owen and

Ogilby in having preorbital and pedal glands, but no inguinal

glands. Mr. Beddard also noted the absence of inguinal glands

(P. ZOOS. 1909s),

The preorbital glands were small. The orifice was a small

vertical crescentie slit, with its concavity looking backwards,

* There adopt, without prejudice, Mr. R.C. Wroughton’s opinion that S. abyssinica

differs from the southern form ©. grimmi in the shortness of its ears. It is quite

immaterial to the work in hand whether abyssinica be specifically or subspecifically

distinct trom grimmi, since the close relationship between the two is admitted by

everyone,

1910. ] CUTANEOUS SCENT-GLANDS OF RUMINANTS, 877

situated in the middle of a naked oval patch of skin just in front

of the orbit. The orifice led into a shallow integumental pocket,

dipping into the substance of the black underlying gland. Under

pressure thickish black secretion could be squeezed from the

orifice.

Text-fig. 96.

158 ta caep

anal tem —

Madoqua phillipsii.

A. Median vertical section of front foot.

B. The same of hind foot.

C. Preorbital gland with orifice distended.

D. Ditto ditto closed.

EK. Ditto in section.

The two halves of the feet were united in the middle

line by two “ webs,” an anterior and a posterior, the posterior

extending as far downwards as the heels and constituting the

posterior wall of the interdigital space, the anterior being some

little distance above the proximal margin of the hoofs, which

were thus capable of much greater distension anteriorly than

posteriorly. Beneath the anterior web, the integument receded

obliquely backwards and upwards, and gradually passed into the

anterior wall of a short subcylindrical tube running upwards

878 MR. R, I. POCOCK ON THE [June 14,

between the phalanges, the posterior wall of this tube being formed

by the posterior web of integument. In longitudinal section the

gland had the appearance of a narrow cylindrical tube opening by

a wide orifice into the interdigital space. The tube and this

space were alike lined with hair, which on the lower lip of the

orifice formed a longish tuft to guide the secretion down between

the heels of the hoofs. The posterior web was continuously hairy

all along its back and inferior edge.

Genus Dorcorraaus Noack.

DorcorraGus MEGALOTIS Menges. (The Beira.)

(Text-fig. 97.)

Of this rare and anomalous Antelope I have seen only the>

dried skins in the British Museum. ‘Through the kindness of

My. Oldfield- Thomas I was enabled to examine the pedal glands

of one of these; and their structure convinces me that Doreco-

tragus is allied to Madogua, but adapted to a mountainous

Text-fig. 97.

mn meme CES B

Dorcotragus megalotis.

A. Median vertical section of hind foot.

B. The same of front foot on a slightly larger scale.

gl., gland; h.t., heel-tie; h., heel.

(Dried specimens. )

habitat. The feet have acquired a superficial resemblance to

those of Goats and Sheep, owing to the shortness of the nail and

the great development of the sole and heel of the hoofs, the

latter forming large cushion-like pads. ‘The strong integumental

bridge joins the hoofs together just above the heels; and the

anterior edge of this is raised so as to form an upstanding lower

lip to the orifice of the gland. The latter isa depressed elongated

1910. } CUTANEOUS SCENT-GLANDS OF RUMINANTS. 879

poueh, recalling that of Madoqua phillipsti, except that the

orifice is much smaller in consequence of the ridge-like lower rim.

This rim seems to be developed in connection with the vertical

position of the feet when the animal stands, not only increasing the

size and strength of the interdigital union, but serving to check

the otherwise uninterrupted downward flow of the secretion from

the gland by forming a barrier, above which it accumulates. The

pocket of the gland is somewhat sparsely lined with short hairs ;

those on the upstanding lower rim of its orifice are longer, serving

to conduct the secretion outwards, asin Mahoqua. tis noticeable

that in Rhaphicerus campestris and Madoqua phillipsi the entire

interdigital integument is covered with hair, there bemg no naked

space behind the heel-tie such as is seen in the species of Cepha-

lophinee examined by me. The deep depression between the

penultimate phalanges of the foot into which the gland opens

resembles that of Sheep and Goats.

On the dry skin I can find no trace either of preorbital or

inguinal glands, and there appears to be a single pair of teats.

Messrs. Sclater and Thomas, though apparently not without

misgivings (‘ Book of Antelopes,’ ii. pp. 1, 2, 1897), placed this

Antelope in the subfamily Antilopine, ‘ near the Gazelles,” and

considered as “superficial” the “resemblance it bears to certain

members of the subfamily Neotragine.” It appears to me, how-

ever, that its affinities, as attested by the horns, skull, muzzle, and

coloration, not to mention the apparent absence of inguinal glands

and the structure of the pedal glands, are more Madoquine than

Gazelline.

Using the feet and preorbital gland as a basis, these two genera

may be distinguished as follows :—

a. Hoofs more or less elongate, with normal sole and heels; heel-

tie formed by a simple fold of integument; a preorbital gland

TORE. HI a CAB coo eadeco q0c. o60 oto ogu BeATBOUES 276 Bde De seCOOC DORE BOO OE REDO CEARCK

a’. Hoofs short, with pad-like soles and heels; heel-tie forming a

double fold, the anterior fold projecting asa ridge constituting

a kind of lower lip to the orifice of the pedal gland; no pre-

MANUS EHIGL Joc codecosomoaece Hegeeadbs Rab pac oeSecoTooDdeho Gap eBDOODODL OM Dorcotragus.

Madoqua.

Subfamily NEOTRAGIN 4.

Genus Rapuicerus H. Sm.

Rapwicervs campesrris Thunb. (= Pediotragus tragulus).

(The Steinbok.) (Text-fig. 98.)

The presence of preorbital and inguinal glands was recorded by

Owen; and according to Max Weber, who cites the species under

the names Pediotragus tragulus and rufescens, the preorbital

glands are present in both sexes.

Of this species I have examined two old stuffed examples,

which the Committee of the Bristol Museum and Art Gallery

kindly permitted Mr. Herbert Bolton, F.R.S.E., F.Z.8., the

Curator, to send to me.

880 MR. R. I. POCOCK ON THE | June 14,

Neither specimen showed any trace of inguinal glands. ‘This,

however, cannot be regarded as satisfactory evidence of their

absence, since the skins were mounted and dried.

Text-fig. 98.

ZB —

Sins _>—

yy SN

SM in, DING

My, a My repent SSW

AW ASS WAM 7, XQ

Wary r [{ TRY LUMI

Raphicerus campestris.

A. Median vertical section of hind foot: gZ., pedal gland.

b. ‘Che same of front foot.

C. Mufle ov rhinarium.

D. Ocular region of female showing orifice of preorbital gland.

E. Section of the same gland.

The preorbital glands were placed near the anterior edge of a

large avea of naked skin in front of the eye. The orifice formed

a small oblique sht, with a slight sigmoid curvature. This led

into a scantily hairy pocket, dipping into the gland, which, as in

Madoqua phillipsii, formed a small, nearly black mass. The upper

margin of the orifice formed a kind of lid over the pocket. The

gland was larger in the male than in the female.

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 881

The pedal glands were well developed, but different from those

of Madoqua. On both front and hind legs they consisted of large

wide-mouthed deep pouches passing up between the antepen-

ultimate phalanges of the feet, and opening some distance above

the proximal margin of the hoofs. The anterior wall of each was

formed by a closely folded web, and the posterior by an equally

closely folded posterior web; but neither of these webs extended

so far down the foot as in J/adoqua, the posterior web of the hind

foot not nearly reaching the heels, and extending only a little

farther than in the Tragulina, and reaching only a little farther

than the anterior web. This suggests that the heels, as well as

the rest of the hoofs, are capable of very wide separation. On

the front foot, on the contrary, the posterior web was longer, and

reached almost to the heels, extending considerably lower than

the anterior web. Its inferior extremity, moreover, was turned

slightly forwards for a short distance, making a distines ridge,

something like that of Dorcotragus. Although the walls of the

pocket were only scantily clothed with hair, the rest of the inter-

digital space was clothed with long hairs, and the edge of the

posterior web forming the posterior lip of the pocket was

furnished with long hairs projecting downwards, as in J/adoqua

and Dorcotragus. There were two pairs of teats.

RAPHICERUS sp.? or NESOTRAGUS sp. 2

Tam indebted to Mr. Gerrard for a front anda hind foot without

supplementary hoofs, and therefore referable probably either to

Raphicerus or Nesotragus. ‘These are worth describing, because

they are structurally nearly imtermediate between the feet of

Raphicerus campestris and of Ourebia nigricaudata.

In the hind foot the interdigital cleft was open in front

throughout nearly the whole of its length, being only slightly

overlapped above by a short angular fold of integument. Its

posterior wall descended with a gradual curvature downwards,

meeting the skin of the back of the pastern considerably nearer

the heels than in 2. campestris, and forming a strong thickened

tie, which was turned slightly forwards. ;

The interdigital cleft on the front foot hardly differed from

that of Our ebia nigricaudata, except that its upper extremity was

overlapped to a greater extent, although not so much as in

LR. campestris, by the antero-superior integumental web,

Since the hind foot was of a more rusty tawny hue than the

front foot, and measured half an inch more from the fetlock to

the tip of ‘the hoof, there can be very little doubt that it belonged

to a different species or subspecies, possibly to a different genus,

of Antelope. Considering the differences the two feet present

from those of R. campestris, the evidence is, I think, in favour of

their belonging to the genera Weotragus or Nesotragus, perhaps

one to moschatus and the other to livingstonianus,

882 MR. R. I, POCOCK ON THE [June 14,

Nototragus* melanotis Thunb. (the Grysbok) has, according to

Owen, preorbital and inguinal glands. Max Weber, however, says

that although the preorbital gland (‘ Thrinengrube ”) is present

in the male, in the female it is represented by a naked area of

skin. This statement must be taken with reserve, in my opinion,

since it does not appear to have been based upon fresh material

(see under Ourebia ourebi). I have not examined material of this

species.

Genus OvuresiA Laurill.

OvREBIA OUREBI Zimm., (=scoparia Schr.).

(The Cape Oribi.)

According to Owen, this species has large preorbital glands and

inguinal pits. Max Weber, on the contrary, states that, although

the male has a deep preorbital gland, this organ is apparently

represented in the female by a naked spot in front of the eyes.

This observation must, I think, have been made upon a mounted

specimen, for I cannot reconcile it with the fact that in the

female of O. nigricaudata described below, and in one of O. mon-

tana, now living in the Gardens, the preorbital gland is well

developed and valvular.

OUREBIA NIGRICAUDATA Brooke.

(The Black-tailed Oribi.) (Text-figs. 99, 100.)

I have seen two young examples, male and female, of this

species from Senegambia, presented by Sir George Denton. The

preorbital gland was marked externally by a longish slit running

downwards from a point close to the anterior angle of the eye.

This was capable of considerable distension, and led into a simple

pocket-like invagination dipping into a large, oval, black, glandular

mass. Under pressure, the secretion, black and adhesive like

lamp-black, could be squeezed from two areas marked by depres-

sions, one near the middle, the other upon the upper corner

of the gland, Each depression contained several pores, from

which long coiling threads of the substance exuded. The gland

and its orifice were relatively longer in the male than in the

female.

The so-called glandular patch beneath the ear was rather

scantily clothed with very short hairs. In section the skin

was seen to be markedly thinner than that of the surrounding

area, and showed no evidence of specially developed glands.

There were two pairs of teats.

The inguinal glands were represented in the female by a pair of

* This generic name was proposed by {Thomas and Schwann for the Grysbok

(Abstr. P. Z. S. No. 27, p. 10, 1906), because it differs from Rhaphicerus in

possessing supplementary hoofs. Mr. Lydekker’s statement, therefore, that the

genus was based upon RF. campestris (the Steinbok), which has no supplementary

hoofs, is erroneous (‘The Game Animals of Africa,’ p. 181, 1908).

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 883

shallow depressions, each marked bya tuft of hair arranged in a

whorl and situated in the ordinary position nearthe mamme. In

the male, however, they were far back alongside the scrotum, a very

unusual position. Each consisted ofa short sac, with tumid orifice,

whence a large tuft of hairs protruded.

Text-fig. 99

‘

Qe AN / ms 4, Ve SK

ASS SENS

son

DELILE

Ourebia nigricaudata §.

A. Head to show orifice of preorbital gland (g7.) and nearly naked patch

below ear.

B. Preorbital gland with orifice distended.

C. Section of preorbital gland.

D. Median vertical section of front foot. (N.B. The hairs within the cleft

are too long, coarse, and clearly defined.)

The tuft of hairs below the knees covered an area of thickened

There were no special indications of

integument as in Gazelles.

clandular activity, however, beyond a certain amount of powdery

seurf at the base of the long hairs.

MR. R, 1, POCOCK ON THE [June 14,

884

The pedal glands, present on all four feet, were very like those

s e ti

al glands

of Gazelles, except that the fold of skin forming the heel-tie

Text-fig. 100.

Vay

}

ay)

Ourebia nigricaudata 2.

A. Median vertical section of front foot

gl, inguinal

B. ‘The same of hind foot.

penis; m., mammie; se.,

C. Inguinal region: p.,

gland with tuft of hair protruding from the orifice.

scrotum ;

extended less to the front, so that the cavity of the interdigital

cleft was to a greater extent in communication with the space

1910.] CUTANEOUS SCENY-GLANDS OF RUMINANTS. 885

between the hoofs. This was more marked in the male than in

the female, and in the former sex the heel-tie was thinner and

less produced in the hind than in the front leg. The walls of

the glandular interdigital depression were scantily covered with

short hairs; but the heal- tie itself was thickly clothed with long

hairs, largely overlapping the base of the hoofs on their inner

aspect. The chief difference between the feet of this species and

those of Raphicerus campestris lies in the fact that the interdigital

cleft is not overlapped above by a long integumental fold, and

that the fold of skin along the back of the pastern extends right

down to the heels, tying them together, and extends further

forwards towards the anterior edge of alae hoofs. From this

arrangement it comes about that the hoofs of Ourebia are much

less distensible than in Raphicerus; they are also shorter: the

combination of these two features suggesting that Owrebia

habitually frequents harder ground.

The structure of the feet in Ouwrebia, correlated with that of

the preorbital glands and the presence of well-developed tufted

carpal or knee glands and of a single pair of inguinal glands,

is strongly suggestive of aftnity between that genus and Carella.

By the structure of the feet, Ourebia and Raphicerus may

be distinguished as follows :—

a. Oritice of glandular interdigital cleft much shorter than the

cleft itself, which is partially overlapped in front and above

by a longish integumental fold or web: posterior web of the

pastern not reaching the heels, which ean thus be widely

separated... sc pumasann COUS/GEE Geb Ano cuban eee eee Raphicerus.

. Orifice of elendulee interdigital clete approximately as Tene

as the cleft itself, which is not to any appreciable extent

overlapped above and in front by an integumental fold;

posterior web of the pastern reaching the heels, which are

thus coralie of being separated only to a hae! limited

extent . Boo TOs c0Or 36.000 006 cu cod EU BEBUDE Gin DUB CGO Rad BOO OOEODEE EE BEE same OLLIE

Subfamily OREOTRAGIN &.

Genus Orrorracus H. Sim.

OREOTRAGUS OREOTRAGUS Zimm. (The Klipspringer.)

(Text-fig. 101, A, B.)

According to Owen, this species has preorbital glands but no

inguinal pits. Hi. Smith, on the contrary, says ‘that inguinal

glands are present.

In a foot of this species, given to me by Rowland Ward, which

had the bones of the pastern cut away from behind, but with the

skin of the interungual area and of the front of the pastern

entire, I could find no trace of pedal gland. The skin on the

middle line of the pastern in front showed no sign of depr oe

Stopping short a little above the proximal margin of the hoot,

turned sharply backwards at right angles between the hoofs i

Proc. Zoou. Soc.—1910, No. LVIII. 58

886 MR. R. I, POCOCK ON THE [June 14,

form the interungual web, which in section was shown to be thick

and horny and to meet that of the back of the pastern just above

the heels.

Thus the foot of this genus differs essentially from that of the

genera of Neotragine above described, in that the integument of

the back of the pastern is not folded back upon itself to form the

posterior wall of an interdigital cleft or depression. But, as in

Text-fig. 101,

ly ff

——_

ALMA La (ih %,

bs fi /

gees .

4 Mee 4

Vij) [I IT papa HEY

y, WL gif TERRE

Pe aiuils

ses

NSS

<—S

A. Anterior view of pastern of Oreotragus oreotragus (? of front or hind foot)

with hoofs distended.

B. Median vertical section of the same foot.

C. The same of front foot of Pantholops hodgsoni.

the Neotragin, the interungual area is covered with long hairs

from the front of the pastern to the back, there being merely a

small triangular naked spot at the summit of the interungual

web. In spite of the thickness and strength of this web, the

hoofs can be widely separated and, as is well known, their points

are very broad and truncated, the animal resting upon these

truncated extremities instead of on the heels.

1910. } CUTANEOUS SCENT-GLANDS OF RUMINANTS. 887

Subfamily ANTILOPINA.

Genus GazeLiA Licht. (Text-figs. 102, 103, 104, A-C.)

According to Owen, the following species of Gazella have large

preorbital glands and inguinal pits:—G. dorcas, subgutturosa,

bennett, arabica, semmeringit, mhor, ruficollis, rufifrons (he-

vella)*. With these he contrasts colus and gutiurosa as having

small preorbital glands.

Ogilby states that this genus has distinct and movable preorbital

glands, very large pedal and inguinal glands, and one pair of

mamme,

Hodgson examined G. dorcas and G. bennettii. He correctly

says of the former that it possesses distinct preorbital glands,

large pedal glands on all four feet, and large and distinct inguinal

glands. Of G. bennettiz, on the contrary, he asserts that the pre-

orbital glands are absent, and he consequently erected the genus

Tragops for its reception. Blanford and others, however, say

that these glands are present in 4. bennetiii, an observation quite

in accord with my own,

Of the genus Gazella I have examined one or more specimens

of the following species :—

G. bennettii, Several examples of both sexes, mostly from

Persia.

G. subgutturosa. One male and one female from Meshed.

G. marica. ‘Two males from Central Arabia.

G. muscatensis. Several specimens from South Arabia.

G. dorcas. An adult pair from Egypt.

G. pelzelni (2). One quite young female from Somaliland.

G. cuviert. One old female from Morocco.

G. rujifrons. One young male from Senegal.

G. semmeringi. One skin of an adult male lent by Mr. E.

Gerrard.

So constant in structure and occurrence are all the glands that

a detailed description of them, as exhibited by each species, is

unnecessary.

The preorbital gland is valvular. When closed it has the

form of a vertical slit of black nearly hairless skin extending

downwards from a point a little in front of the anterior

corner of the eye. When opened to its fullest extent, it appears

as a subcircular patch of skin, scantily beset with short hairs, and

marked with a shallow central depression, which is, in reality, the

bottom of the invaginated pocket. The gland itself, when

extracted from the preorbital pit of the skull, is seen to consist of

a black integumental thickening formed round the invagination.

In the case of G. dorcas I found that the inner surface of the

pocket had about seven large holes, from which, under pressure,

* T infer that rufifrons is the species cited by Owen under the name “ kevelle,’’

because he gives Senegal as its locality.

58*

888 MR, R. I, POCOCK ON THE [June 14,

secretion could be squeezed first in the form of black waxy threads,

then as fluid drops, the waxy consistency being due, I expect, to

the pores being blocked and inoperative for some little time before

Text-fig. 102.

~SSS WON (

~ \

aS = SS SY) {

~~. —= SSS S (

oe 2 al

Bz Ae sa

eZ Se “ WN

=. a © 9 > //,

fF! EE ao. F UG, i

= ZF = ; Al

— PLLE= —S, nee

= EES PO ev. i

Nee

bf fA

A. Section of preorbital gland of Gazella dorcas: o., orifice of gland

m., utuscle,

B. Superficial view of the same gland flattened. 5

C, Superficial view of the flattened preorbital gland of G. euviert 2.

D. The same gland in section.

death. Fluid drops could also be squeezed from the hair-pores

studding the surface of the integument. The secretion contained

ra ae

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS, 889

in the pocket itself is black and sticky. It may here be men-

tioned that Ogilby long ago (P. Z. 8. 1836, p. 38) described the

preorbital gland of a Gazelle as being furnished with six excretory

ducts placed nearly in a circle and with one central duct, from

the orifices of which there issued strings of a dense ceruminous

matter.

The inguinal glands are always two in number, consisting of

deeper or shallower pouches opening by larger or smaller orifices

situated just externally to the mamme. The pouches dip into

a mass of vascular tissue just above the mammez. They are

usually about one inch deep and about as wide inside as deep or a

little wider, the integument surrounding the orifice and the mamma

being naked. Ina male G. swhguttwrosa the orifice was nearly as

large in diameter as the scrotum ; in G. dorcas and G. muscatensis

it was considerably smaller and the lumen of the pouch was cor-

respondingly smaller in the two latter than in the former. The

largest gland of all was found in the female of G. ewvieri, in which

it was two inches deep and about twice as deep as wide, and as

wide at the orifice as within.

The walls of the pocket were, in all cases, sparsely covered with

short hairs, and the secretion was waxy and drier at the orifice

than at the bottom of the pouch. It has a very strong odour.

In G. bennettit and subgutturosa it is yellow in colour and smells

exactly like the urine of the common House Mouse (ws musculus) ;

in G. cuviert the colour is greenish yellow and the smell a mixture

of mouse urine and flour-paste.

In a female of G. dorcas the inguinal gland secretion was pale

green and resembled in scent a mixture of flour-paste and cheese ;

the pedal glands had a strong and decided, but peculiar and

indescribable, smell; the secretion of the knee glands had a

repulsive odour of dogs’ dung, while that of the preorbital glands,

although faint and quite indescribable, was not unpleasant. This

specimen furnished an admirable instance of the fact below alluded

to, that scents of totally different character may emanate from the

glands of one and the same individual. This was further borne

out by a specimen of G. muscatensis, in which the green secretion

of the inguinal glands smelt exactly like cream-cheese, that of the

knee gland like a Domesticated Sheep, while the pedal glands had

a faint and indescribable, but different odour from either.

It may be noted in passing that the smell of the inguinal

secretion in specimens of G. bennettii and G. subguiturosa from

Persia is the same as that of Ovis vignei from the same country.

Although there is typically only a single pair of mamme in

Gazelles, now and again there are two pairs, indicating, no doubt,

the descent of these animals from quadrimammate ancestors.

Similar atavism is sometimes seen in Sheep. When there are two

pairs in Gazelles, the inguinal glands are connected with the pos-

terior pair, showing that the anterior pair are supernumerary.

The pedal glands are well and equally developed on all four feet.

When the hoofs are together, the orifice has the form of a narrow

890 MR, R. I. POCOOK ON THE [June 14,

slit dividing the hairs on the front of the pastern. When the hoofs

are forcibly distended, it appears as a triangular cleft, pointed

Text-fig. 103.

7 BF

)

ant

- Superficial view of knee gland of Gazella dorcas.

- Longitudinal section of the same.

C. Transverse section of inguinal gland of G. euviert 2.

D. Inguinal region of @, subgutturosa : p., penis; m., mamma; se., scrotum;

gl., orifice of inguinal gland, its area beneath the skin shown by dotted

line.

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 891

above and wide below, above the base of the hoofs. The summit

of the cleft is arched over to a small extent by a slight web of

integument, and the bottom of it is closed by a strong and deep

web which passes forwards from the heels of the hoofs to a point

nearly on a level with their proximal edge in front. This web

firmly ties the hoofs together, so that, in a living animal standing

naturally, they never splay apart like those of Sheep and Goats,

although when it rears up against anything the front hoofs often

spread toa considerable extent. The skin forming the interungual

web is folded sharply upon itself in front and passes back to the

heels; its inner layer thence passes upwards only a little in front

of the skin of the back of the pastern, then curves forwards about

on a level with the joint between the penultimate and ante-

penultimate phalanges to meet the skin of the front of the

pastern in the small web described above, thus forming the

bottom, back, and top of the long interdigital cleft, the ovitice

of which is almost as long as the cleft itself. The walls of the

cleft are only sparsely covered with short hairs; and I have

never detected any signs of secretion either on them or on the

longer hairs fringing the edges of the orifice or on the tufts

of still longer hairs overhanging the proximal portion of the

hoofs in front.

The interungual web is continuously hairy from front to back,

though the hairs often become somewhat thinner towards the

heels. The latter are well developed, and the hoofs are never

long, though in these two particulars they are not specialised to

the same extent asin Goats and Sheep, which stand rather more

upright than Gazelles.

Dr. Murie long ago pointed out that the knee-tutts of Saiga are

glandular. My. Lydekker made the same suggestion with regard

to Gazelles (‘ Field, March 14, 1908). This suggestion proves

to be correct. The hairs are directed downwards, those on each

side of the middle line usually crossmg at the tips, somewhat

as in the metatarsal gland of the Impala (Apyceros melampus).

The underlying skin is much thicker than that of the vest of the

leg; but the thickening, accompanied by increase in size of

the sebaceous glands, is not abrupt but gradual, thinning away

into the normal skin all round the circumference of the area.

The hairs covering the latter hold a quantity of scurfy secretion.

As a general rule, the knee-glands are covered with long hairs

forming a continuous pad and not showing, when undisturbed,

any central longitudinal line; but in three examples of G. dama

rujicollis, now living in the Gardens, the hairs are quite short and

their inwardly directed tips do not meet in the middle line, so

that the median gutter is plainly visible.

According to Hodgson the Tibetan species of Gazelle known as

picticaudata has no preorbital or inguinal glands and has small

pedal glands on all four feet. If his account be trustworthy, the

species differs from all the true Gazelles I have examined in the

absence of the preorbital and inguinal glands, not to mention the

892 MR. R. I. POCOCK ON THE [June 14,

seeiningly smaller size of the pedal glands. Hodgson erected the

genus Procapra tor piecticaudata ; and, in view of the constancy in

occurrence and development of the cutaneous glands in the species

of Gazella I have examined, I am disposed to think that Procapra

should be admitted as a valid genus, comprising probably, in

addition to picticaudata, guiturosa and przewalskii. 1 do not

know whether these three species are in actual agreement as

regards cutaneous glands; but they at least differ from all the

southern species of Gazelles in the shortness of the tail.

Text-fig. 104.

A. Hind foot of Gazella subgutturosa showing slit-like orifice of pedal

gland (0.).

L. Median vertical section of front foot of G. bennettii: gl., glandular clett.

C. Foot of G. euvieri dissected to show the gland in situ.

D. Median vertical section of foot of Antidorcas marsupialis.

The validity of Procapra was (liscussed by Messrs. Sclater and

Thomas (* Book of Antelopes,’ iii. p. 73), and the claims of the

genus to recognition were dismissed on the grounds that three of

the characters in which the above-mentioned species agree (namely,

1910. ] CUTANEOUS SCENT=GLANDS OF RUMINANTS. 893

the absence of knee-tufts, of preorbital glands, and of horns in

the female) are not absolutely distinctive of the group; G. sub-

gutturosa, tor instance, has the female hornless and ‘ several

species besides [the three in question] are without knee-brushes,

while the antorbital fosse in others are so shallow as to be prac-

tically non-existent.” In accordance, however, with the modern

more restricted application of generic names, it may be doubted

whether these reasons will be regarded as invalidating Procapra

from recognition, especially as the shortness of the tail seems to

be an absolutely distinctive feature of the three above-mentioned

species, as compared with Gazelles proper.

Pallas, according to Gray *, says that the male of G. guttwrosa

has a follicle, like the musk- bag of Moschus, in front of the pre-

puce. Still more inter esting is the recor d by Hodgson of the

presence of a large sinus behind the horns in picticaudata. This

probably represents the posteornual gland of Rupicapra. Indeed,

four of the characters cited by Hodgson as diagnostic of Procapra,

namely, the absence of preorbital g glands and “of inguinal glands,

the presence of small pedal glands, as compared with those of

Gazella, on all four feet, and of a posteornual sinus, apply equally

well to "Rupicapr a; and it may be thatin P. picticaudata we have

the link between Rupicapra and Gazella. Possibly also the organi-

sation of Procapra may throw some light upon the true position

of Saiga and Pantholops.

Genus AnTIDORGAS Sund.

ANTIDORCAS MARSUPIALIS Zimm. (=euchore).

(The Springbuck.) (Text-fig. 104, D.)

According to Owen, this species, which he appears to quote

twice, under the names Antilope euchore and pygarga, has large

preorbital glands and inguinal glands.

No information as to the glands is contained either in Sclater

and Thomas’s ‘ Book of Antelopes; or in W. L. Sclater’s volume on

‘The Mammals of $. Africa,’ or in Rowland Ward’s ‘ Great and

Small Game of Africa.’

I have examined three examples of this species, two males and

a female. The preorbital glands resemble those of the Gazelles

described above, except that the gland itself is pale yellow instead

of black in colour. When everted and squeezed, the gland yields

a clear fluid.

In spite of what Owen says, there is no trace of inguinal

glands.

Pedal glands are present on all four feet and so closely resemble

those of Gazelles as to need no redescription.

There are no knee glands, the knee-tufts being absent, as has

already been recorded by previous describers.

* Cat. Rum. Mamm. in B. M. p. 37, 1872.

894 MR, R. I, POCOCK ON THE | June 14,

To compensate, as it were, for the absence of inguinal and knee

glands, the Springbuck has an immense dorsal gland, the skin

underlying the well-known white dorsal shield “having highly

developed sebaceous glands secreting a yellow sticky substance

which stains the bases. of the white hairs yellow or forms a scurfy

kind of powder about their roots. No such substance is found

upon the hairs of the rest of the back ; and the skin of the area

in question has a markedly fatty appearance when stripped, its

colour differentiating it at once from that of the shoulders and

sides of the body. Mr. F. E. Beddard, F.R.S., long ago noticed

the glandular nature of this patch and drew my attention to it.

Genus ANTILOPE Pall.

ANTILOPE CERVICAPRA Pall. (The Indian Blackbuck).

(Text-fig. 105, A-C.)

The large size of the preorbital gland in this species has

attracted the attention of zoologists from early times. The

presence of inguinal glands was recorded by Owen in 1836 and

independently in the same year by Ogilby, who stated at the

same time that there are large pedal glands and only one pair

of mamme. These observations were repeated by Hodgson and

have been copied by recent systematic authors.

Through Mr. Herbert Bolton, F.R.S.E., F.Z.S., Iam indebted

to the Committee of the Bristol Museum for the stuffed skin of a

male of this species, and Mr. Gerrard kindly supplied me with

another.

Mr. Gerrard’s specimen had the preorbital glands mtact and

by cutting them I was able to see that they differed in no essen-

tial par ticulars from those of Gazelles, except that they were very

much larger, the infolding forming a long deep hair-lined space,

and the elandular portion being very thick and pale in colour,

The structure and general features of this organ have already been

described by Bennett, Owen, and others.

The orifices of the i inguinal glands were clearly shown opening

close to the mamme, of which there was a single pair,

The pedal glands were present in all four feet. and were very

like those of ‘Gazelles. The walls of the interdigital cleft were

furnished with longer hairs; the interungual integument was

much thicker and more horny, forming a stronger tie between

the hoofs; its anterior extremity was slightly recurved and

furnished with a somewhat radiating tuft of hairs, the longest

of which overhung the proximal edge of the hoof in front, while

between this tuft and the heel-tie the skin was naked. There

was no naked horny pad on the knees: but the knee gland was

developed as in Gazelles, the roots of the long hairs being packed

with scurf-like secretion.

The number, position, and structure of the cutaneous glands

show close relationship between the genera Antilope and Carella,

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 895

the only difference I can find between the two being the thickness,

smoothness, and curvature of the interungual integument.

Text-fig. 105.

7, y a Se Oe, — h J

YY EMP,

i ——

A. Transverse section of preorbital gland of Antilope cervicapra ¢ :

a, skin extending towards the eye; 6, towards the nose.

B. Vertical or longitudinal section of the same gland: a, upper, 6, lower

extremity.

C. Median vertical section of front foot of Antilope cervicapra.

D. The same of hind foot of Lithocranius walleri. (The fold in the integu-

ment of the wall of the gland probably absent in fresh specimen.)

~ In a living male specimen of this species the secretion of the

inguinal glands smells exactly like flour-paste and might be

described as starchy. The secretion of the preorbital gland is

peculiar and indescribable. I cannot agree, however,with Bennett

that the odour is urinous*.

* Tt is not, I believe, generally known to naturalists and sportsmen, though the

fact is perhaps familiar to some who have paid attention to animals in captivity,

that the adult male of this species undergoes a marked seasonal change of colour.

During the summer months, that is to say, from about May to August, the colour,

except of the face, is like that of females and young males, namely tawny. In the

latter part of August or thereabouts, the coat begins to darken and gradually but

rapidly assumes the dark hue whence the trivial name of the species “ Blackbuck ”

has been derived. At or about the beginning of May the opposite change rapidly

896 MR, R. I. POCOCK ON THE [June 14,

Genus LrrnocraNnius Kohl,

LirHoCRANIUS WALLERL Brooke.

(The Gerenook or Waller's Gazelle.) (Text-fig. 105, D, p. 895.)

No fresh material of this species has been available for

examination.

Elliot deseribes the preorbital gland as a ‘‘ conspicuous promi-

nence with a central aperture, from which exudes a black secretion

that stains everything it touches in the same way that ink does”*.

Judging from this description the gland is better developed than

in Gazelles, but the colour of the secretion appears to be the same.

On a fore and a hind foot kindly lent to me by Mr. Edward

Gerrard I have found well-developed pedal glands, each con-

sisting of a large saccular invagination on the front of the

pastern. The orifice is as large as in Gazelles, and extends up the

pastern as far as the joint between the penultimate and the ante-

penultimate phalanges. ‘The inferior edge of the orifice is formed

as in Gazelles by the interungual web, which, however, differs from

that of Gazelles in passing only halfway along the space between

the heels and the anterior proximal margin of the hoof. It is

also much thicker and more horny than in Gazelles ; but, as in the

latter, is folded upon itself to form in the same way the back and

upper wall of the glandular interdigital cleft. In the specimen

examined, this wall was thrown into folds, suggesting a sae too

voluminous for the space containing it. The w valls of the gland

were clothed with longish hairs, all projecting forwards or down-

wards towards the margins of the orifice, a downwardly directed

fringe of hairs clothing the edge of the iInterungual w eb.

The entire pastern is Eibehed’ with very short ieee. set hair, with

a small black tuft over the proximal end of the hoof in front ; ; a

felting of thick close-set hair clothes the integument on the inner

surface of the hoof above the proximal margin of the nail and of

the heels.

The hoofs are longer than in Gazelles, and capable of greater

distension, on account of the web being shorter and_ situated

somewhat higher above the hoofs.

Although the pedal glands of this genus are more like those

takes place, the first sign of it beg a gradual lightening along the middle of the

back. In August, when the blackening sets in, it appears first most pronouncedly

npon the neck, legs, and fore quarters, and gr: idually spreads backwards.

For three consecutive years I have observed this in a specimen in our Gardens. I

do not know what theory Indian sportsmen hold as to the disappearance of “ Black-

bucks” from the herds during the months they are in “ eclipse ” pelage, to borrow an

ornithological expression. But, since most of them, I imagine, believe all tawny

individuals to be does or young bucks, they must suppose, I conceive, that the adult

bucks have separated themselves from their companions, an inference clearly not

justified by the facts.

* ‘Great and Small Game of Africa,’ p. 371, 1899 (Rowland Ward).

1910. ] CUYANEOUS SCENT-GLANDS OF RUMINANTS. 897

of the Gazelles above described than they are like the glands of

any other Antelope, they present well-marked differences from

them.

There was a well-developed knee-pad of black and reddish hairs

much Jonger than those of the surrounding skin and stuck

together with plentiful dry secretion.

On skins of this genus in the British Museum I found two pairs

of teats, but no evidence of inguinal glands.

As attested by other characters, this genus is an aberrant type

most nearly allied to Gazella; but in the structure of the feet it

seems to be less closely related to Gazella than is Antilope or

Antidorcas. 1 do not feel sure, however, that the shape of the

upper wall of the interdigitai cleft in the example described and

figured was not due to mutilation, since the skin of the pastern had

been cut away at the back, perhaps thus loosening the connective

tissue which normally holds the invaginated sac in place. It is

very desirable that fresh examples of “this species be examined,

Excluding Procapra, which I have not seen, the genera here

referred to the Antilopinz may be distinguished as follows by the

structure of the feet and by their cutaneous glands :—

a. An immense dorsal gland extending from the middle of the

back to the rump, feet, and preorbital gland ee Leeeataly

as in Gazella, but no inguinal GALT OOKS “Sog:coecebonvenqoposeerncce LIMA OV REND.

a’. No dorsal gland.

b. Feet with the folded interungual web extending only a

little more than halfway between the heels and the

proximal edge of the hoof in front ; no inguinal glands ;

two pairs of mamme. ; ss... Lithocranius.

. The folded interungual integ ument extending "forwards

nearly to the proximal margin of the hoofin front 3 a pair

of inguinal glands; normally a single pair of mami.

ce. Interungual integument thinner, hairy, the interdigital

cleft practically naked . . Gazella.

ce’, Interungual web smooth; ‘interdigital cleft more hairy. Antilope.

In addition to the four genera just described, the Antilopine,

according to accepted views, comprises the genera Dorcotragus,

Ammodorcas, Pantholops, Saiga, and Lpyceros. 1 do not, how-

ever, think these can be included in the subfamily without

unwarrantably upsetting its homogeneity. Kinship is possible

between Ammodorcas and -Zpyceros on the one hand and between

Saiga and Pantholops on the other, but it does not appear to me

that the two latter have any close relationship with the two former

it is indeed very far from obvious why the genera should he

associated in the pairs suggested. Pending the ‘demonstration of

relationship such as would justify such a course, | propose to relegate

Pantholops, Saiga, and Aipyceros each to its own subfamily and

to leave Ammodorcas for the present Eougleced- In the following

pages Ammodorcas will be found near the Cervicaprinze (p. 917)

It may be added that these four genera differ widely in the

structure of the feet from those here restricted to the Antilopine.

898 MR. R. I. POCOCK ON THE [June 14,

Subfamily SaAr@in #&

Genus Saiea Gray.

Saga TARTARICA Linn. (‘The Saiga.)

According to Dr. Murie (P. Z. 8. 1870, p. 500) this animal has a

pair of small preorbital glandular pits; a pair of carpal or knee

glands; a pair of shallow inguinal pouches in one specimen

examined, but not in another, a younger example, although the

secretory area was very evident; and pedal glands in all four

feet. These are described as “about 13 inch in depth and

opening by a narrow constricted aperture” at the “front and

upper part ” of the foot. Although on p. 502 of this memoir

Dr. Murie remarks that these glands “hardly form satisfactory

erounds for basing affinity upon,” it is nevertheless quite

clear from his description of the pedal glands that they are

much more like those of a Sheep than of a Gazelle or Black-

buck. The description is not sufficiently detailed to permit

the conclusion that the pedal glands exactly resemble those of

Ovis; but it is quite certain that with their narrow constricted

orifice they bear no similarity to the long interdigital cleft of

Gazella or Antilope. Their structure, in fact, bears out Dr. Murie’s

opinion that the Saiga may be regar ded as an Antelopine Sheep.

In any case, it seems evident that the genus is an isolated one.

T think, therefore, that it should form, provisionally at all events,

the type of a special subfamily.

Subfamily PANTHOLOPINA,

Genus PanrnoLors Hodgn.

PantruoLors HopGsoNt Abel. (The Chiru.)

(Text-fig. 101, C, p. 886.)

In 1836 Hodgson (P. Z.S. 1836, p. 39) described the inguinal

glands as “ exceedingly large sacs which hang by a long narrow

neck from the groin. He re-aftirmed the | same fact in other

words in his paper in 1847, where he adds that preorbital glands

are absent, but the pedal glands are present and large on all four

feet. Blanford and Lydekker repeat his observations, without

alluding to the pendent, inguinal glands, while the latter supple-

ments his account with Kinloch’s observation that these glands

penetrate a considerable distance into the body.

Hodgson’s statement as to the pendulous nature of the inguinal

glands is puzzling. It is hardly likely that they can be volun-

tarily eversible and retractile, and I can only suppose that he saw

them hanging as loose sacs on a stripped skin,

The only material of this species I have been able to examine

fully was a mounted head and two feet, fore and hind, of a male

specimen, for which my best thanks are due to Mr. Edward

—

1910. ] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 899

Gerrard, and a single foot shown by its colour to be that of a

female, which was kindly given to me by Messrs. Rowland Ward,

Ltd.

The head enabled me to confirm Hodgson’s statement as to

the absence of the preorbital gland, and the feet to refute his

statement as to the presence of the pedal glands—they are

absent. The integument along the middle line of the pastern

forms a very shallow depression. Near the proximal end of the

hoof in front it turns backwards at a rounded angle and runs

almost parallel to the inner proximal margin of the nail to the

heels, joining them together throughout their length. The

posterior portion of the heels is well developed and the skin

which unites them behind is folded sharply upon itself for a

short distance, then turns up to be continuous with the skin

of the back of the pastern. The back of the pastern and the

fold in question are thickly hairy, and the interungual integu-

ment or web is covered with long hairs as far down as the point

where the heel-tie begins. These hairs are especially long at the

base of the hoof in front, and overhang the anterior edge of

the nail nearly to its tip. Thus the integumental web which

ties the hoofs together is as deep and strong as in Cattle (Bos)

or Elands (Yaurotragus), and the feet are quite different from

those of typical Gazelles, to which Pantholops is usually held to

be closely related ; and the hoofs clearly have nothing like the

power of extension seen in the feet of Sheep and Goats.

According to Hodgson, there is only a single pair of mamme,

and I only succeeded in finding one pair in a dried skin of a male

in the British Museum. I noticed in this skin that the orifice

of the inguinal gland on each side les some four or five inches

away from the teat. On one side the gland was loose or pendent,

as Hodgson described it; but on the other it was adherent and

directed inwards from the orifice as a slightly expanding tube,

its inner extremity lying close to the teat. This, no doubt, is its

natural position, Judging from the direction of the gland in other

Bovidee that possess it.

In 1872 Gray recognised the peculiarities of this genus by

placing it in a special family, Pantholopide.

Saiga is commonly held to be related to Pantholops, perhaps

correctly ; but the relationship is not close. Apart from the

well-known differences between them in the structure of the

skull and nose, they may be distinguished as follows:

a. Pedal glands, apparently ovine, on all four feet; preorbital

elands present ; inguinal glands small and close to the mamme,

of which there are two pairs sbeobdoccoseoc cco gngncobanepoucodea | ISUIAI IE

a’. No trace of pedal glands; no preorbital glands; inguinal

elands very large and deep, with orifice remote from the

mamme, of which there is a single pair ........................... Pantholops.

In the characters here used, however, the two genera do not

differ from each other much more than Ovis differs from Pseudois

or Capra.

900 MR. R. I, POCOCK ON THE | June 14,

Subfamily BuBALIN &.

(Hartebeests and Gnus.)

Of this group Owen records the presence of preorbital glands

and the absence of inguinal glands in the following species :—

Damaliscus lunatus, Bubalis boselaphus (= Antilope bubalis),

B. caama, Gorgon taurinus, and Connochetes gnu.

Ogilby, who examined Bubalis boselaphus, which he quotes as

2B. mauretanicus, confirms Owen’s statement regarding the above-

given particulars, adding that the pedal glands are large and that

there is a single pair of mammez.

Recent systematic writers agree with Ogilby as to the number

of mamme; but in spite of his use of the epithets “ exigui,

distineti” in describing the preorbital glands, these are quite

commonly, but erroneously, stated to be sinall.

Of Hartebeests belonging to the genus Lubalis I have seen

no complete material, and have been compelled to rely upon

soaked feet and masks usually with the skin only of the preorbital

gland retained. Of the White-tailed Gnu (Connochetes gnu) I

have seen two freshly dead specimens. I have, however, seen

living examples of the two species of Gnu (G@. tawinus, C. gnu

and of two species of Damaliscus (D. pygarga and D. albifrons,

the Bontebok and Blesbok respectively).

Genus BuBauis Frisch.

Bupauis CAAMA F. Cuy. (‘Text-fig. 106.)

LT am indebted to Mr. E. Gerrard for a mask, containing

entire preorbital glands, and front and hind feet of this species.

The preorbital glands were covered with hairs which in the

individual examined had a thickened appearance, owing to the

coating of brown earthy-looking secretion which covered them

individually. The centre of the area was marked by a distinct

orifice leading into a narrow duct about half an inch long and

lined with short hair. This duct penetrated about halfway into

the large gland which was about as large as a walnut. :

Pedal glands were present only on the fore feet, or if present on

the hind feet they attained only a comparatively slight degree of

development. On the fore feet they very closely resembled those

of Gazelles, consisting of a very large interdigital cavity opening

by a long orifice on the anterior aspect of the pastern and

extending upwards from the proximal margin of the hoof in

front. As in Gazelles the two hoofs were united by a deep

interungual web, which extended from the heels to the proximal

margin of the hoof in front, then folded sharply over and was

continued backwards along its course nearly as far up as the

false hoofs, where it turned obliquely forwards and downwards,

ineeting the skin of the front of the pastern at an acute angle to

1910. | CUTANEOUS SCENT-GLANDS OF RUMINANTS. 901

form the anterior web joining the two halves of the pastern

together in front. Although essentially of the same nature as in

Gazelles this gland differed in one or two points. For instance,

the height or length of the cavity along the back of the pastern

was considerably greater than the length of the orifice, owing to

the cavity being continued farther up the pastern than the edge

of the anterior upper web. In the second place, the walls of the

cavity were clothed with long blackish hairs which projected from

the orifice, forming only a small tuft over the proximal margin

Text-fig. 106.

\ UNIS \Y

= ‘)\

Bubalis caana.

A. Vertical median section of preorbital gland.

Bb. The same of front foot.

C. The same of hind foot.

of the hoof. Lastly, the interungual web was quite naked back to

the heels, and instead of meeting the skin of the back of the

pastern at right angles met it at an obtuse angle, the difference

of inclination being obviously correlated with the less upright

position of the foot when standing, the long hoofs of the

Hartebeest being much less vertical than the short hoofs of

the Gazelles.

In the hind foot there was only a shallow depression just above

Proc. Zoou. Soc.—1910, No. LIX. 59

902 MR, R. I. POCOCK ON THE June 14

the hoofs, the skin of the interungual web being folded back for

only a short distance, then inclining obliquely upwards, forming

the back wall of the depr ession in question, This depression on

the hind foot might be compared to the depression on the pastern

of the Chamois, except that its walls were hairy instead of nearly

smooth.

Of the genus Bubalis I have also examined the face-skin and

part of the gland of . cokei supplied to me by Rowland

Ward, Ltd.. and by Mr. E. Gerrard. In this species the pre-

orbital gland has essentially the same structure as in 4. cawma.

The following species also have the gland clothed externally

with a mat or tuft of hairs, namely, L. major, tora, swaynei,

lelwel, and jacksoni. In Bb. lichtensteini, on the contrary, the

glandular area is naked as in Damaliscus, the orifice in the dry

skin being a small slit-like aperture leading into a duct. The

hind foot of B. lichtensteini is like that of B. caama, having only

a shallow depression on the front of the pastern. For the

material of the species, enabling me to establish these data, I

am indebted to Mr. E. Gerrard.

Genus Damatiscus Scl. & Thos.

DAMALISCUS KORRIGUM Ogilby, Subspe. JIMELA Matisch,

(Text-fig. 107.)

I am indebted to Mr. E. Gerrard for the opportunity of

examining a complete skin of this Hartebeest.

The preorbital gland itself was cut away, but the orifice was

represented by a sant elongated aperture in the middle of a bare

patch of skin in front of the eye.

The pedal glands, present only on the front feet, resembled

those of Bubalis caama in a general way, with the exception that

the interdigital pit was much less deep, owing to there being a

Houatnalala space between the integument of the back of the

pastern and that forming the posterior wall of the cleft; more-

over, the skin of the front of the pastern, forming the upper edge

of the orifice, was not folded backwards and upwards upon. itself,

but descended almost at right angles to form the short upper wall

of the interdigital cleft. A further difference was seen in the

presence of a Trebtical fascia joining the skin of the back of the

pastern to that of the interdigital cleft just above the heels. The

walls of the cleft were Bicrhed with long rusty-red hairs, the ends

of which projected from the orifice, overhanging the proximal

end of the hoof in front.

The hind foot was like that of 4. caama, except that the

depression on the front of the pastern just above the hoof was

shallower. A connecting fascia, corresponding to that of the hind

foot, was also present.

The area of the preorbital gland on a face-skin of D. korrigum

tiang, for which I am indebted to Rowland Ward, Ltd., resembled

that of D. k. jimela in all respects.

]

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 903

The area in question is also bare in D, hunteri, albifrons, and

pygargus. In an adult male of D. albifrons now living in the

Gardens, the gland is usually active and tumid, secreting a

substance which passes on to the face, staining the edges of the

white blaze yellow. The orifice of the gland is circular and lies

in the centre of the smooth glandular swelling. The gland itself,

which I saw in a dead Blesbok many years ago, was, I clearly

recollect, about as large as a filbert nut, with a short duct leading

Text-fig. 107.

WSS

Se << LZ: — SS

WR L777 SSS ~

AR ~~ 32 Y L; y ;

Ain : ZZ =

: We Lae ~

Ay Sse

Wh = Ss

A W

OEE

Damaliseus korrigum jimela.

A. Median vertical section of hind foot.

B. The same of front foot.

f., fascia.

into it, exactly as in B. cawma. In two young Bonteboks now

living in the Gardens, the gland is much less prominent than in

the Blesbok, possibly because the specimens are not yet adult.

The above-described specimen of D. korrigum jimela had no

inguinal glands ; but the hair on the skin a short distance away

from the one teat on each side was arranged in a whorl.

59*

904 MR, R. I. POCOCK ON THE [June 14,

Genus Connocu#tes Licht.

Connocu&res GNu Zimm. (The White-tailed Gnu.)

(Text-fig. 108.)

Of this species I have examined two specimens, an immature

male and an adult female. In both there were no inguinal

glands and a single pair of teats.

Text-fig. 108,

: Zr See

LS Yj35

Connochetes gnu.

A. Median vertical section of front foot.

B. The same of hind foot.

C. Section of preorbital gland.

D. Superficial view of preorbital gland (g/.).

1910. | CUTANEOUS SCENT-GLANDS OF RUMINANTS. 905

skin, sparsely pitted with setal pores, whence hairs arose. In the

young male the gland was as large as a hazel-nut, but markedly

longer than deep. In section it was pink in colour and resembled

a cyst enveloped in the thickened skin. It was not lodged in

any depression of the skull, but merely rested on the bone of the

face. The hairs which appeared upon its surface were seen to be

rooted deep in the gland and to reach the surface through the

pores above described. No secretion could be squeezed from the

gland. This was perhaps due to the fact that the animal was

immature and had been ailing for some time before death. It

seems probable that the secretion natur ally exudes through the

setal pores. Nevertheless, 1 have never seen the preor bital gland

of a living Gnu, either of this species or of G. taurinus, ‘show

any sign of secretion or even of moisture. I did not cut the

gland of the adult female, since the specimen was required for

preservation.

Pedal glands, as in Lubalis, are well developed only on the

front feet and appear to be absent on the hind feet. The

feet and the glands have essentially the same structure as in

Bubalis caama. In the adult female, however, the cavity of

the gland differed in some interesting particulars from that of

B. caama above described. The posterior wall of the cavity

was not folded so closely upon the skin of the interungual web

and of the back of the pastern, but ran obliquely upwards and

backwards, then turned forwards at a right angle, then down-

wards and forwards at a right angle closely in contact with the

skin of the front of the pastern, which it joined at the upper end

of the orifice at a very acute angle, forming a close fold. The

anterior upper web thus constituted extended nearly as far

downwards as the edge of the interungual web, leaving a much

shorter orifice for the gland than in Bubalis caama. In the

latter the gland was rhomboidal in section, whereas in C. gnu t

was suboblong. It was thickly clothed with long hairs, pro-

jecting like a fringe or tuft from the orifice. The roots of all

the hairs were covered and stuck together with blackish secretion

smelling exactly like sour milk or cream-cheese. In the young

male the cav ity of the gland was nearly intermediate in shape

between that of the adult and that of B. cauma, the slit-like

orifice being much longer than in the adult owing to the absence

of the anterior wall, the anterior web or fold stopping short

much higher up the pastern above the hoofs. In section the

upper portion of the cavity resembled that of B. caama, while its

lower portion was like that of the adult C. gnu.

On the hind feet there was only a very shallow hair-lined

depression, recalling that of Bubalis caama.

The front and hind feet of another adult exainple, kindly given

to me by Rowland Ward, exactly resembled the one above

described.

The preorbital gland consisted of a subcireular patch of black

906 MR. R. I, POCOCK ON THE [June 14,

Genus Goreon Gray.

GorGon TAURINUS Burch.

(The Brindled Gnu or Blue Wildebeest.)

Living examples of this species have the preorbital gland like

that, of Connochetes gnu; and in a dried skin, lent to me by

Mr. E. Gerrard, I found the front and hind feet constructed

exactly as in that species. One pair of teats only is visible in a

pregnant female now living in the Gardens.

Gray long ago placed the White-tailed and the Brindled Gnus

in distinct genera; and it was evidently the opinion of

Messrs. Sclater and Thomas that there was a great deal to be

said in favour of this view, although they did not give it

practical expression in their ‘Book of Antelopes.’ Judged by

modern standards it appears to me that the well-known dif-

ferences between the two are quite worthy of generic recognition.

There can be no evasion of this conclusion if specifie value be

given, as is commonly done, to the differences between the

Brindled and White-bearded Gnus. The White-tailed Gnu is a

very differently built animal from the two just mentioned. The

back is not sloped as in the rest of the Bubaline, but horizontal,

and the hind-quarters are markedly elevated on account of the

prominence of the summits of the iliac bones. Other external

features separating this species from the Brindled Gnu and from

its ally, the White-bearded Gnu (@. albo-jubatus), are well known,

and are in my opinion equivalent to those separating Bubalis

and Damaliscus.

Using the preorbital glands as a basis, the genera of Bubalinz

may be distinguished as follows :—

a. Preorbital gland marked with a central orifice leading into a

hair-lined tube, which dips into the substance of the gland.

b. Surface of the preorbital gland naked or nearly so.

Damaliscus and Bubalis lichtensteini.

b’. Surface of the preorbital gland thickly and uniformly covered

Wed DL OFT DMI MRE Rose Live wc ces hss cho has een ee ne iene

a. Preorbital gland without any central orifice and tube, its free

surface consisting of an area of skin sparsely covered with

VSATS sch ach v MUREMES CEST «pyres hsss.srson0e>suae cus neencaas gu COMMOCAQLES UUM TONE

Bubalis.

Since Lichtenstein’s Hartebeest, hitherto referred to the genus

Bubalis, is known to depart from the other species of that genus

in the structure of its horns and also in the nakedness of the

preorbital gland, it should perhaps be referred to a distinct genus.

In diagnosing the genera in this table, no use has been made

of the differences recorded in the structure of the pedal glands

of Connochetes gnu, Bubalis caama, and Damaliscus korrigum,

striking though they be, because they have only been noted in a

single species of each of the genera.

The structure of the feet and the presence of pedal glands

only on those of the anterior pair bear out the evidence from

other sources that the Gnus and Hartebeests form a natura

1910.| CULANEOUS SCENT-GLANDS OF RUMINANTS. 907

group. Moreover, it seems to me that the size and structure of

the preorbital gland and of the glands in the front feet, coupled

with the hairiness of the rhinarium and the presence of a single

pair of mamme, suggest tolerably close kinship between Dama-

liscus, the least-specialised member of the Bubalinz, and the

Antilopine. At all events, | think it may be claimed with some

assurance that the relationship indicated is closer than that

between pyceros and the Antilopine.

Subfamily ORYGIN «A.

I prefer to use the term Orygine for the group called Hippo-

tragine by most modern authors, because the generic nainme

Hippotragus will soon, probably, no longer be admitted as a valid

generic designation, being antedated both by Ozanna and

Hgoceros.

Genus Oryx Blainv.

According to Owen the following species of this genus have

neither preorbital nor inguinal glands: O. gazella(= A.oryx) and

O. algazel (= A. leucoryx).

Ogilby also says that these two species have neither preorbital

nor inguinal glands, but states that the pedal glands are large

and that there are two pairs of mamme.

Oryx BEISA Riipp. (The Beisa.)

(Text-fig. 109, C.)

In the dried skin of a young male example of this species, for

which I am indebted to Mr. Gerrard, I could find no trace of

preorbital glands nor of inguinal glands, but two pairs of mamme

were present.

Pedal glands, approximately similar in structure, were present

on all four feet. They opened by an elongated, somewhat lanceo-

late interdigital cleft on the front of the pastern just above the

hoofs. This cleft led into a shallow depression, which at its

deepest part, close to the interungual web, passed into the

principal part of the gland, which consisted of a moderate-sized

sac extending for a short distance upwards and backwards almost

as far as the posterior integument of the pastern. Describing

the gland another way, it might be said to consist of a deep but

rather short interdigital depression subdivided into an anterior

and posterior part by a ridge of integument projecting from its

upper wall, thus constricting its lumen.

The floor and posterior wall of this depression were formed by

the closely infolded layer of the interungual web. Its opposite

walls were formed by the integument of the front of the pastern,

which sloped obliquely downwards and forwards to the aforesaid

ridge, then obliquely upwards and backwards. ‘The entire inter-

digital depression on both sides of the constriction was clothed

with longish white hairs clogged with secretion. Inferiorly, just

below the orifice, the hairs were longer and overlapped the

908 MR. R. I. POCOCK ON THE [June 14,

proximal edge of the nail of the hoof. The interungual web was

thick and horny and, as in the Antilopinz and Bubaline, ex-

tended from the heels to a point near the anterior margin of the

hoof before being closely folded upon itself; and, as in the

Bubaline, Antilope, Hippotragus, and the Tragelaphine, it was

quite smooth.

Text-fig. 109.

a :

i >

ats

A. Median vertical section of front foot of Addawx naso-maculatus: e., conical

excrescence with the depression behind it.

3. Anterior view of the same foot ; e., conical exerescence.

C. Median vertical section of hind foot of Oryx Leisa.

The hoofs were pointed, compressed above, with broad flat heels

and flat soles. The only difference noted between the glands of

the front and hind feet was the greater development of the

constricting ridge in those of the hind feet.

The interdigital glands of Oryx beisa with their longer cleft-

like orifice occupy a nearly intermediate position between those

of Gazelles and those of /Tippotraqus.

OryxX LEUCORYX Pall,

(The White Oryx = O. beatrix of most authors.)

In an old dried skin of a male example from the shores of the

1910. | CUTANEOUS SCENT-GLANDS OF RUMINANTS. 909

Persian Gulf, kindly lent to me by Mr. Gerrard, the pedal glands,

though imperfect, had apparently the same structure as in

O. beisa and there was no trace of inguinal glands.

OryX ALGAZEL Oken.

(Lhe Soudanese Oryx = O. leucoryx of most authors.)

Both Owen and Ogilby state that this species has no pre-

orbital gland. I presume this observation was based upon an

examination of dried skins or of skulls. It is not, however, true ;

for in the example now living in the Gardens there is quite

commonly a small patch of secretion sticking the hairs together

about an inch in front of the eye. This proves the existence of

a specialised glandular area beneath the hairs; but probably the

gland is not highly developed.

Genus Hipporraaus Sund.

According to Owen, the following species of this genus have

neither preorbital nor inguinal glands: 7. lewcophwa, equina

(=barbata).

The only species of this genus examined by Ogilby was

H. leucophwa, which he referred to the genus Oryx. He agreed

with Owen that this species has neither inguinal nor preorbital

glands, but adds that the pedal glands are large and that there

are two pairs of mamme. W. L. Sclater also states that im the

genus Hippotragus the preorbital gland is absent and that there

are two pairs of mamme.,

Hipporracus NicER Harris. (The Sable Antelope.)

(Text-fig. 110.)

Tn an adult female of this species which died in the Gardens I

found a preorbital gland underlying the white preorbital tuft of

hair. It was very little differentiated, however, consisting, like

the knee glands of Gazelles and some Sheep, of thickened vascular

integument, The bases of the white hairs were crowded with

pale yellow grains like scurf, smelling of a mixture of sour milk

and new-mown hay. .

There were no inguinal glands.

The pedal glands were well developed on all four feet and

opened to the exterior by a small subcircular orifice in the centre

of a naked patch of skin in a shallow depression close to the

proximal end of the hoofs. The gland was a large sac passing

obliquely upwards and backwards and reaching as far as the

posterior integument of the pastern, and with its lower wall closely

applied inferiorly to the interungual web which was sharply folded

back upon itself, the infolded portion constituting the wall of the

gland in question. The walls of the gland were covered with

. . 5 .

short hairs, becoming longer towards the orifice of the gland.

910 MR. R. I, POCOCK ON THE [June 14,

The interungual web of the hoof was smooth from the orifice

of the gland to the posterior area of the heels.

Text-fig. 110.

Hippotragus niger.

A. Median vertical section of front foot: gl., pedal gland; o., its orifice.

B. Anterior view of front foot: o., orifice of gland.

C. Longitudinal section of preorbital gland; a., ocular end of gland.

I was able to verify these facts upon a second specimen, a dry

skin kindly lent to me by Mr. Gerrard. This specimen had two

pairs of teats, but no trace of inguinal glands was discoverable.

HipporraGcus equinus Is. Geoff.

(The Roan Antelope.)

In a female specimen with two pairs of teats, but no inguinal

glands, for the examination of which I am indebted to

Mr. Gerrard, I found the pedal glands to be of substantially

the same structure as in H/. niger, except that there was a slight

constriction of the lumen of the gland, caused by a small in-

tegumental ridge rising from its upper wall and partly dividing

it into an outer and an inner chamber, as in Orya beisa.

Genus AppAx Raf.

AppAX NAsO-MACULATUS Licht. (The Addax.)

(Text-fig. 109, A, B, p. 908.)

According to Owen, this species has neither preorbital nor

inguinal glands.

[ have seen only the front ‘and hind feet of one dead example,

no other material being available.

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 911

The feet were very different from those of Oryx and Hippo-

tragus, being low, flat, long-heeled, and broad at the point in

adaptation to a soil of soft sand. In section, the integument

of the posterior surface of the pastern was seen to be folded just

above the heels like that of Pantholops. From that point it

passed forwards for some distance parallel with the sole of the

foot before turning abruptly upwards towards the front of

the pastern. Here it passed into a somewhat conical excrescence

or eminence, covered with short hair and defined behind and

laterally by a semicircular infolding or depression of the skin.

Behind this excrescence, the integument of the anterior surface

of the pastern was continued straight up this area without sign

of insinking and was covered with normal longish hairs. The

latter inferiorly encircled the excrescence, which jutted up like

an island from their midst. I believe the excrescence to be

glandular. The structure of the two feet was the same, except

that the excrescence was smaller on the foot which I judged to

belong to the hind leg.

Examination of the feet of a very young example now living in

the Gardens revealed no signs of the excrescence, but there was a

distinct orifice, representing, I suppose, the infolding of the skin

defining the excrescence in the adult. This orifice was in almost

the same position as the orifice of the gland in Oryx, and its

glandular nature was attested by the staining yellow of the white

hairs round it.

Addax has a marked patch of longer hairs below the eye

on each side of the face. This probably overlies a glandular

thickening of the skin, such as is present in Hippotragus. I have,

however, no direct evidence of the fact.

From the structure of the feet dddax would seem to be less

closely allied to Oryx and Hippotragus than these two are to one

another. But more and better material of the genus must be

examined before any satisfactory conclusion as to its true

systematic position can be arrived at.

Subfamily CERVICAPRINA.

Genus Prin Gray.

PELEA CAPREOLUS Bechst. (The Vaal Rhebok.)

(Text-fig. 111.)

According to Owen, this species possesses inguinal glands, but

has no preorbital glands.

The absence of preorbital glands has been aftirmed by sub-

sequent authors, and correctly so, judging from the one specimen

of this species I have examined, a female brought by Dr. Lyster

Jameson from 8. Africa, which lived two years in the Gardens.

This specimen, however, had no trace of inguinal glands. Pedal

glands were well developed on all four feet. They opened by a

912 MR. R. I. POCOCK ON THE ‘June l4,

small circular orifice on the front of the foot ina shallow depression

lined with long hairs. The orifice led into a short hair-lined

duct which expanded into a capacious but laterally compressed

thin-walled sac reaching almost to the posterior integument

of the fetlock. The sae was filled with semifluid, pale-coloured,

evil-smelling secretion, recalling exactly the odour of dog’s dung.

Close to the orifice in the depression arose a number of long

hairs overhanging the base of the hoofs in front, and these were

stuck together with the secretion, which could be easily squeezed

from the gland by pressure applied to the sides of the pastern.

Text-fig. 111.

Pelea capreolus te

A. Median vertical section of hind foot: g/., pedal gland; o., its orifice.

B & C. Lateral and dorsal views of the nose to show swollen rhinarium.

The structure of the feet was almost the same as in Cervicapra

redunca, the interungual integument being quite naked, meeting

that of the back of the pastern at a right angle and forming an

obtuse angle with the floor of the depression on the front of the

fetlock. Thus the integument or “web” joining the hoofs

1910. } CUTANEOUS SCENT-GLANDS OF RUMINANTS. 913

together extended from the heels almost up to the proximal end of

the anterior ridge of the nail. The apical angle of the nail was

also less acute than in Cervicapra redunca and Adenota kob, the

hoofs more approaching those of Goats and Sheep in shape—in

accordance, no doubt, with the mountain habitat of Pelea.

A well- known distinguishing feature of Pelea as compar ed

with Cervicapra and Robus is the enlar ged and swollen rhinariun,

which has a widely rounded posterior border extending bac k-

wards beyond the posterior notch of the nostril. This rhinarium

is glandular. In the living animal it became studded with drops

of moisture under the stimulus of excitement or pleasure, and

after death was easily made by pressure to yield the same

substance.

Like Cervicapra and Kobus, Pelea has two pairs of mamme.

Genus CervicAPRA Blainy.

Owen noticed the absence of preorbital and the presence of

inguinal glands in three species of this genus, namely, in

OF redunca, the Nagor, a Senegambian Species ; in C. arundinwn,

which he quoted as Antilope eleotr ‘agus; and in C. fulvorufula, if I

am right in supposing he meant this species by A. landiana, the

latter name being, I surmise, written in error for lalandia or

lalandir.

I have examined only fresh material of one species, namely the

following.

CERVICAPRA REDUNCA Pall. (The Nagor.)

(Text-fig. 112.)

Owen does not state how many inguinal glands he discovered

in the species he examined, but in two examples of C. redunea,

from Senegambia, presented by Sir George Denton, I found two

pairs of sacs. The anterior sac on each side was wide-mouthed

but shallow, the glandular area into which is dipped being only

slightly vascular. The posterior sac, lying rather nearer the

middle line and close to the scrotum, was, on the contrary,

13 inches deep, of even width throughout, the orifice being about

= an inch in diameter and not half the width of the orifice of the

anterior sac. Its glandular area was very thick and vascular.

Both pouches secreted a semifluid substance, turning yellow and

waxy at the orifice and giving off an odour of flour- -paste going

stale, which to me was most nauseous.

There were no preorbital glands.

In one specimen I found no trace of pedal glands; but in the

other there was a very small hair-lined pouch opening into the

shallow hair-lined depression on the front of the pastern just

above the angle formed by the junction of the integument of this

surface with the strip of naked skin extending from the front to

the heel of the hoofs and binding the two together. There

914 MR. R, I, POCOCK ON THE [June 14,

appeared to be no gland connected with this pocket, since there

was no secretion on the hairs projecting from its orifice; but

beneath the pocket between the phalanges there was an elongated

mass of vascular gelatinous tissue extending almost as far up as

the false hoofs, just such as is found in the hoofs of Oxen. The

fore and hind feet were alike, except that the anterior depression

was somewhat deeper on the hind feet, allowing a slightly wider

spread to the hoofs, and whereas the false hoofs were united by

thick horny material on the fore feet, those of the hind feet were

connected by naked skin.

Text-fig. 112.

Cervicapra redunca.

. Median vertical section of front foot taken a little to one side of the aborted

gland (gl.) ; gel., gelatinous interdigital mass.

The aborted gland of the same in section.

Median vertical section of hind foot : lettering as in A.

. Inguinal region showing four mamme (m.); scrotum (se.) and inguinal

lands, the anterior pair (g7.) wide and shallow, the posterior pair behind

them narrow and deep.

BE. Section of inguinal gland of posterior pair,

F. Side view of nose to show rhinarium.

dow >

In both specimens there were two pairs of mammz; but in

neither was there a bare patch of skin below the ear. The knees

had a bare patch of thickened horny skin, which was apparently,

however, not glandular. So far as the pedal glands are concerned,

the genus Cervicapra, as represented by redwnea, stands in the

same relation to Pelea capreolus as Capra stands to Ovis ; and

there cannot, I think, be much doubt that the ancestors of

existing species of Cervicapra are descended from Antelopes with

pedal glands like those of Pelea.

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 915

Genus ADENO?A Gray.

AveENotA KoB Erxl. (Buffon’s Cob.)

(Text-fig. 113.)

According to Owen, this species has inguinal pits but ‘no

preorbital glands,

Text-fig. 113.

. VA Yop: (\n

MAY LU iy

aD:

rary iw)

Me:

RATA

I4,

TEA “2

“aoe.

Adenota kob.

A. Longitudinal section of preorbital gland: a, upper or ocular end;

b. lower end.

B. Inguinal region: m., mamme; sc., scrotum; gl., orifice of gland, the

subcutaneous extension of which is shown by the dotted line. _

C. Median vertical section of front foot; gel., gelatinous interdigital mass.

I have seen two examples of this species, a subadult male from

Upper Nigeria presented by Lady Constance Stewart Richardson,

916 MR, R. J. POCOCK ON THE | June 14,

and an immature female from Sierra Leone presented by

My. ‘Tinling.

In the female there was a distinct preorbital gland, marked

externally by a tuft of hair darker in colour and a little longer

than that of the surrounding area. Beneath this the skin was

considerably thickened, the thickest part being situated above

towards the anterior angle of the eye and the thinnest part below,

where the thickening gradually merged with the normal skin

of the cheek. In its essential characters this gland resembled

that of the Sable Antelope above described (p. 910). The male

was not examined for this character.

Both specimens had two pairs of teats and a single pair of

inguinal glands representing the posterior pair found in Cervi-

capra redunce ; ; but in addition to these there was upon the right

side in the female a very minute pocket lying far out and in advance

of the mammz. There was no trace of Saha on the left side,

The orifice of the large gland was subcircular, about 3 an inch

in diameter in the male onl ; Of an inch in ithe fooralee the

pouch itself was capacious, widening considerably Ba ci the

orifice and extending for 1 inch in the female and 13 inches in

the male in the direction of the scrotum and mamme. The

yellow secretion had exactly the same nauseating smell as that of

Cervicapra redunca, namely of flour-paste or starch. There were

no pedal glands and no trace even of the duct. The structure of

the feet was the same as in Cervicapra redunca, with the same

elongated mass of vascular gelatinous tissue lying between the

penultimate phalanges along the inner surface of the skin of the

back of the pastern. The skin round the false hoofs was naked.

Genus Konus Smith.

IXOBUS ELLIPSIPRYMNUS Ogilby, Derassa Riipp., and MARLE Gray.

(Water Bucks.)

According to Owen, 4. ellipsiprymnus has neither preorbital

nor inguinal glands.

It is on account of the alleged absence of the inguinal glands

in this species and their presence in A. kob that I ‘keep the two

species under separate generic headings. Trouessart regards

them as representing distinct subgenera, presumably for other

reasons.

On the skin of the head of an example of A’. defassa, from the

White Nile, lent to me by Rowland Ward, Ltd., I could find no

trace of a preorbital eland: but I think a eland vesembling that

of Adenota hob will probably be found when fresh material is

examined, The fore and hind feet of A. marie (Mrs. Gray’s

Water Buck), given to me by Mr, Gerrard, showed no trace of

pedal glands, the feet in section resembling those of A. hob.

The genera of this subfamily, so far as my observations extend,

1910. ] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 917

may be distinguished as follows, using the glands as a basis for

their separation :—

a. Pedal glands present on all four feet and opening by a short

duct with a small circular orifice on the front of the pastern

just above the hoofs; summit of rhinarium swollen and

glandular, and extending back beyond the posterior edge of

pheinosirilsis| Mo ime mMAlANAS | fs ....4:..sccssenceecespaciessen Peleme

a’. Pedal glands represented at most by the duct of the glands

present in Pelea; summit of rhinarium normal, not ex-

tending behind the posterior notch of the nostrils.

6. With two pairs of inguinal glands ; tail bushy as in Pelea.,. Cervicapra.

b’. Inguinal glands absent or represented by a single pair of :

pouches ; tail tufted at the end.

c. A single pair of inguinal glands ...................-.0..e.»..... Adenota.

coe Nonneuinalielandspepeen eeeeriteeee-cesce ctor ircccsn ce), OU

In their volume on the Mammalia, Flower and Lydekker

classified the genera above referred to the Cervicaprine with the

Neotragine. The Cervicaprinz, however, differ markedly from the

Neotragine, so far as my observations extend, in the structure of

the feet, since they entirely lack the deep and long interdigital

cleft on the front of the pastern found in the Neotragine.

It seems to me that, in looking for the relationships of the

Cervicaprinz on the lines followed in this paper, the following facts

should not be ignored. With the exception of the Indian genera

of Tragelaphine, the preorbital gland is absent in that group;

and in the Cervicaprine it is at most represented by an integu-

mental thickening. In both groups, and in no other subfamilies

of Bovide, two pairs of inguinal pouches have been recorded.

More significant still is the fact that the structure of the feet,

whether pedal glands be present or absent, is the same in the

Cervicaprine as in the Tragelaphine. Corroborating these

characters are the presence of two pairs of mammeze and a mode-

rate or large rhinarium, to which may be added the invariable

absence of horns in the females of Cervicaprine and in most

genera of Tragelaphine. I do not, however, attach much im-

portance to this character, despite the value given to it by some

writers *.

At all events, it must, I think, be conceded that the Cervicaprinz

are more akin to the Tragelaphine than they are to the Antilopine,

Bubaline, Neotragine, Cephalophine, or Oryginz, although the

retention by Peleaw of pedal glands, somewhat recalling those of

the Orygine, is a fact full of interest.

The following genus is possibly related to the Cervicaprine

(see p. 897). ;

Genus AmMmoporcas Thos.

AMMODORCAS CLARKEI Thos.

(The Dibatag or Clarke’s Gazelle.)

The only materials of this genus and species I have seen are

* W. L. Sclater, for instance, who uses it as a basis for grouping the S. African

subfamilies of Bovide.

Proc. Zoou. Soc.—1910, No, LX. 60

918 MR. R. I, POCOCK ON THE [June 14,

some dried skins in the British Museum. I made, however, no

dissections. Superficially the feet showed no trace of pedal glands.

Certainly there were none such as exist in the Antilopine ; and I

judged that the feet resembled those of Zpyceros or Kobus. Also

there were two pairs of well-developed teats, and I could find no

evidence of the existence of inguinal glands, These features,

coupled with the long “ bovine” tail and the shape and direction

of the horns, which are more Cervicaprine than Gazelline, should

exclude the genus from the Antilopinse. On the other hand, the

preorbital glands appear to be like those of Gazella, and the

muffle is hairy as in that. genus.. The skull also is very Gazelline.

These cross-resemblances make Ammodorcas extremely diflicult

to classify. It has been described as intermediate between

Gazella and Lithocranius. This is true of the length of the neck,

the structure of the upper lip, and some characters connected

with the skull. On the other hand, in the structure of the feet,

horns, and tail, the genus is less Gazelline than Lithocranius : and

T am disposed to think that the degree of relationship to Gazella

that the two exhibit may be fairly gauged by the fact that the

type-species of Lithocranius was referred to Gazella by so high an

authority as Sir Victor Brooke, and was for a long time cited

under that genus; while the type-species of Ammodorcas was

originally assigned on equally good authority to the genus Cervi-

capra. From this it may be inferred that the kinship between

Lithocranius and Gazella is tolerably obvious and that between

Ammodorcas and Gazella by no means evident,

Subfamily A pycERIN&,

Genus AZpycEeros Sund,

Apyceros MELAMPUS Licht. (The Impala or Pallah.)

(Text-figs. 114, 115.)

According to Owen, this species has large preorbital glands and

inguinal glands as well. All recent authors, however, are in

agreement that there are no preorbital glands, and I could find no

trace of them on any of the dried skins in the collection of the

British Museum,

Tam indebted to Messrs. Rowland Ward, Ltd.,and to Mr. E.

Gerrard for the opportunity of examining and cutting open the

hind feet of two specimens, and of thus being able to establish

the absence of pedal glands on these limbs ; and, so far as I can

judge from looking at the fore feet of skins in the British Museum,

they are equally absent in the front legs, the feet of which seem

to be constructed exactly like those of the posterior pair except

for the absence of the two metatarsal glands, well known and

often recorded as present on each hind leg.

Of these glands W. L. Sclater says: ‘‘attached to the lower ends

of the cannon-bones of the hind legs isa brush of very dark brown

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 919

hairs surrounding the opening of a peculiar gland containing a

fatty secretion.” This is not quite an accurate description of the

glands, according to my observations. The long blackish hairs

marking each gland form a subelliptical elongated mat. The hairs

point downwards, but those on each side of the middle line of the

mat are inclined inwards, meeting at their tips and marking a

central streak. When these converging hairs are pulled on one

side it may be seen that they overlap an elongated area of naked

skin, broadest across the middle, pointed above and below, and

more than twice as long as wide. This naked area was covered,

in the specimen examined, with secretion and the skin composing

Text-fic. 114.

Aipyceros melampus.

Z ape vertical section of hind foot.

. Hind foot to show the mat of hairs (gl.) marking one of the metatars:

glands and the tuft (4) representing the false hoofs, Scere

it was shown in section to be exceedingly thin. The skin sur-

rounding it was, however, markedly thickened, especially exter-

nally, where it formed a distinct upstanding ridge. It was upon

this thickened skin that the long hairs forming the mat ae

and, since the bases of these hairs were clogged with secretion,

I infer that the thickened rim of skin is the secreting area of the

gland and that the depressed naked area constitutes the floor of a

kind of reservoir for the secretion, which is conducted downwards

by the downward-growing long hairs.

The area of the false hoofs, which are themselves absent, is

60*

| es

920 MR, R. I, POCOCK ON THE [June 14,

covered with a thick mat of longish mostly black hairs directed

backwards and upwards. The skin beneath this mat was thick

and horny, but I could not find any evidence either upon the hairs

or in the skin of special glandular development.

Text-fig. 115.

Mpyceros melampus.

A. One of the metatarsal glands with the hairs undisturbed.

B. The same with the hairs pulled aside to show the area of naked skin

beneath.

C. Longitudinal section of the gland: a, its upper, J, its lower end.

D. ‘[ransverse section of the same: e, its external, 7, its internal edge.

The feet themselves were constructed as in the Tragelaphingw and

most Cervicapriney, except that the whole interungual integument,

forming the web, was covered with short hairs from front to back.

1910. ] CUTANEOUS SCENT GLANDS OF RUMINANTS, 921°

Although Gray long ago made the genus Wpyceros the type of

a special family, it has of late years found a place amongst the

Antilopine. Apart, however, from the hairiness of the rhinarium

and of the interungual integument of the feet, it has very few

‘“« Antilopine ” features, in the strict sense of the word. This

latter feature, moreover, 1s as much Neotragine as Antilopine,

In other respects, the feet may be described as Tragelaphine or

Cervicaprine, except for the disappearance of the false hoofs and

the presence of the metatarsal glandular mats of hair. The horns

moreover, are as much Ceryicaprine as Antilopine in character,

and the absence of the preorbital gland is another feature ex-

cluding the genus from the Antilopine.

Hence, taking into consideration the sum total of its characters,

the peculiarities of Apyceros seem to me to be best expressed by

setting it in a subfamily apart from all others.

Subfamily TRAGELAPHINA,

Genus Terraceros Leach.

TEPTRACEROS QUADRICORNIS Blainv.

(Lhe Four-horned Antelope.)

(Text-figs. 116, A, B; 117, 118, 119, A, B, C.)

Hodgson described the preorbital glands of the genus J'etraceros

as linear and longitudinal, and stated that inguinal glands are

absent and pedal glands absent or present only on the hind feet.

Blanford repeated this account with the additional information

that the pedal glands are confined to the hind feet.

I have seen three examples of this species, a still-born young

one and an adult male and female. In all the preorbital gland

was well developed and valvular, consisting of a longitudinal

pocket, the upper rim of which was produced into a large over-

lapping flap, forming a longitudinal slit where its free margin

came into contact with the skin below it. When the flap was

lifted a blackish nearly naked area was exposed; the posterior or

ocular end of this area was the deepest portion of the invagina-

tion, which from that point forwards became gradually shallower.

The gland itself was thick and pale in colour. In the adult male

it was much larger than in the adult female, and in the summer

discharged a quantity of soft cheesy-yellow secretion. The odour

of this was not noted after death, the specimen having been pre-

served in alcohol; but in the fresh female specimen the drops of

clear fluid that were squeezed from the gland had a faint aromatic

scent.

There were two pairs of teats, but no trace of inguinal glands.

The feet were constructed as in the genus Zragelaphus, the

hoofs being short and but little distensible, owing to the extension

of the interungual integument from the heels forwards for some

distance before curving backwards to join that of the front of the

922 MR. R, I, POCOCK ON THE [June 14,

pastern, merely a shallow depression being formed, There was

no trace of any folding of the integument constituting an inter-

digital cleft or gland such as is seen in the Cephalophine, Anti-

lopin, and Neotragine, The interungual skin, moreover, was

Text-fig. 116.

< oer ke,

SA we

; NAL Ge

(iG PON ANE) ae

1 «ww { /)\ AWS oe Mtl “és Za

ie a OE

gE guia is

= \ Le dee

an be

19a pe te a ae

lig See

Eas,

LY)

SQN gS 4

A, Head of Tetraceros quadricornis @ : orifice of gland (gl.); area of gland

represented by dotted line.

B. Transverse section through the gland; 0, its orifice; 7, flap of skin

constituting its upper lid.

C. Inguinal region of Tragelaphus scriptus scriptus $: m., Mammex; se.,

scrotum ; gl., orifice of gland; the subcutaneous extension of the gland

shown by a dotted line.

naked. In the male there was a small accumulation of material

on the hairs in the shallow depression of the front of the pastern

just above the smooth area of the interungual integument ; but

there was no evidence that this was an odorous secretion. In the

adult female, a much older animal, however, there was on all four

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 923

feet a large black horny excrescence occupying the same position

and large enough to be visible without separation of the hoofs.

The skin from which this excrescence arose was thrown into a

multitude of columnar folds. Since I have seen no such excre-

scence on the feet of any other specimen of this Antelope, I must

regard its development in this example as abnormal, perhaps

pathological.

Text-fig. 117.

OA LE LL:

Ke ree 7?

Tetraceros quadricornis.

A. Head of newly born young: gl., preorbital gland.

B. Head of female: gl., preorbital gland.

C. Section of the preorbital gland of the same: /7J., flap of skin constituting

its upper lid. :

D. Preorbital gland of the same with the lid (//.) raised.

One of the most interesting features connected with this genus

is the presence of a pair of well-developed glands between the

‘false hoofs of the hind legs. In both the male and the female

the false hoofs were large and the skin between them was black,

tumid, horny, but soft and naked, except for a longish fringe along

924 MR. R, I, POCOCK ON THE [June 14,

the middle line. When squeezed a yellowish buttery substance

with a strong sour odour exuded from a pair of smallish apertures,

one on each side close to the middle line, so that the hairs became

clogged with it. This aperture led into a voluminous sac, very

scantily clothed with short pale hairs, the two sacs being separated

by a narrow partition and occupying otherwise the whole area

between the two false hoofs. Each was filled with the odorous sub-

stance above-mentioned, which, in the male at least, presented a

concentrically stratified arrangement and was greenish in colour.

Text-fig. 118.

Tetraceros quadricornis §.

A. Jafesior view of hind foot ; fA., false hoof with its inner half and the adjacent

skin cut away to show the glandular mass (s.).

B. Front view of the same foot showing the large horny accretion (e.).

C. The same in median vertical section.

This is the only Antelope known to me which possesses such

glands. Whether these sacs are the glands that Hodgson men-

tioned as sometimes present in the hind feet or not I cannot say.

In any case, it is quite evident that they have no sort of con-

nection, except a functional one, with the interdigital glands of

other Bovidee.

1910.] CUTANEOUS SCEN'T-GLANDS OF RUMINANTS. ~ 925

Text-fig. 119.

»\)

Za,

A. Median vertical section of hind foot of Tetraceros quadricornis g: fF,

tendon; hk, small horny accretion at anterior end of interungual web ;

J, fringe of hair between false hoofs; s, secretion within the sac shown

by cutting away the partition between the two glands.

B. Transverse section.of the pastern of 7. qguadricornis g, through the false

hoofs ; ¢, tendon; ¢, glandular sae with arrow marking the oritice; a, false

hoof; f, fringe of hair between the glandular areas of the false hoofs, with

the partition above it; s, secretion within the other sac, the orifice of which

is not seen.

C. Inferior view of hind foot of 2. quadricornis g: a, false hoof; b, horny

but soft integument forming lower wall of gland; 0, orifice of gland.

_D. Transverse section through the region of the false hoofs of Zvagelaphus

scriptus g, for comparison with B: ¢, tendon; a, false hoof.

926 MR. R. I. POCOCK ON THE [June 14,

Tetraceros is usually regarded as belonging to the Cephalophine.

I do not think, however, that the evidence for this relationship

is strong. T'etraceros and Cephalophus differ widely in the struc-

ture of “the feet and of the preorbital glands; but Zetraceros

approaches Boselaphus in these particulars. The presence of

four mamme, the absence of inguinal glands, and the size of

the muflle may also be cited as “corroborative evidence of this

relationship. Most suggestive, too, is the presence of white patches

on the pasterns, and sometimes at all events of two white specks

on the cheeks in 7'etraceros. When I noticed these marks in the

living animal, before a dead one was available for examination,

I felt sure that the affinities of the genus would prove to be with

the Tragelaphines, a group in which these markings are extra-

ordinarily persistent. I subsequently found that Sir Harry

Johnston* had noticed the same colour-features in 7'etraceros,

and pointed out the resemblance the genus presents to the Trage-

laphines in those particulars. It is true that the horns of Vetra-

ceros have no anterior crest or keel; nevertheless, they are more

like the horns of Boselaphus than the latter are like the spirally

twisted horns of the African Tragelaphines.

Genus BoseLapuus Blainy.

BosELAPHUS TRAGOCAMELUS Pall. (The Nylghaie.)

(Text-figs. 120, 121.)

According to Owen, this species has inguinal glands ; Ogilby,

on the contrary, says they are absent. The two authors agree,

however, as to the presence of preorbital glands; and Ogilby adds

that there are distinct pedal or interdigital glands. Blanford

supports Ogilby’s statements.

Turner described f the preorbital gland as a slight pit imme-

diately in front of the orbit with a small longitudinal fold of skin

in front of it. In the middle of this fold (or flap) of skin “ there

is a small round pore, through which exudes a yellowish secretion

furnished by a gland placed just underneath. The gland itself

is slightly larger than a hazel-nut and is laid upon the surface of

the bone without any fossa to receive it.”

In an adult female example kindly sent to me by the Duke of

Bedford, K.G., I found the preorbital gland marked externally

by a short longitudinal slit some distance in front of the eye, the

area round the slit and between it and the corner of the eye being

practically naked. When opened, this slit was seen to be the

orifice of a shallow elliptical depression containing some dried

waxy secretion. Near the centre of the depression, but varying

in position on the two sides, was a conspicuous circular hair-follicle

with a hair protruding from it. ‘There were also one or two

+ P.Z.S. 1851, p. 116.

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 927

additional but smaller hair-follicles, also varying in position.

From these follicles yellow waxy secretion could be squeezed ;

but I could detect no special odour about the secretion. As

described by Turner, the underlying glandular mass was of large

size and elongated, but heart-shaped in transverse section. Its

colour was yellowish.

There were two pairs of teats, but no trace of inguinal glands

could be detected.

Text-fig. 120.

os ea ~

‘Al (UUW uuree : os

= |

DD yyWVZy ic) dyynny es”

ISS aE

ACS! ANS

él. iS

Boselaphus tragocamelus ° .

A. Portion of the face showing the eye and the orifice of the preorbital

gland (g/.).

B. Preorbital gland with the lid raised showing two large hair follicles.

C. Section of the gland passing through the larger hair follicle showing the

protruding bristle.

D. Rhinarium.

The feet were constructed exactly as in other Tragelaphine,

the interungual integument being smooth from the heels for-

wards towards the proximal edge of the nail in front, then

inclined obliquely upwards to meet that of the anterior surface

of the pastern, leaving a small smooth triangular area at the

bottom of the pastern in front just above the hoofs. Between

928 MR. R. I, POCOCK ON THE [June 14,

the phalanges of the pastern was the gelatinous mass of tissue

found in Vaurotragus, Bos, and Kobus. In connection with the

relationship here claimed between Boselaphus and Tetraceros, it

was especially interesting to note that the skin between the false

hoofs of the hind feet was white, thick, and highly glandular,

the hairs being packed with secretion, smelling like the fat of

tallow candles.

Text-fig. 121.

ty bm)

Boselaphus tragocamelus 9.

A. Median vertical section of hind foot: gel., gelatinous mass between the

phalanges.

B, The same of the front foot.

This specimen proves that Owen’s statement about the inguinal

glands and Ogilby’s statement, apparently copied by Blanford,

about the pedal glands are alike erroneous. It further shows

that the prevalent view as to the kinship between Boselaphus and

the African Tragelaphines is correct, and affords no support

whatever to the suggestion found in Max Weber’s ‘ Siiugethiere,’

pp. 675-676, that Boselaphus probably belongs to the Bubaline

and not to the Tragelaphine.

1910. ] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 929

Genus TRAGELAPHUS Blainy,

TRAGELAPHUS SCRIPTUS and its subspecies.

(Text-figs. 116, C, p. 922, & 122, C, D.)

Owen records the absence of preorbital and the presence of

inguinal glands in specimens of the genus he identified as Antilope

scripta and A. sylvatica. W. L. Sclater, however, says that the

preorbital gland is “present, but small and with a very small

aperture,” in the genus 7'ragelaphus, with which Limnotragus was

included.

Text-fig. 122.

SAO

A, Anterior aspect of foot of young specimen of Taurotragus oryx.

B. Median vertical section of foot (? front or hind) of adult female of Zauio-

tragus ory« (dried).

C, Median vertical section of front foot of Tragelaphus scriptus ? roualeyni:

gel., gelatinous mass between the phalanges (dried).

D. Inguinal region of female foetal example of Tragelaphus scriptus (subsp. ?)

from Unyoro; gl., the two inguinal pouches; m., mamme,

The only entire specimen of this genus that I have examined

was a female fcetal example, of which J do not know the subspecies,

930 MR. R. I, POCOCK ON THE [June 14,

from Unyoro in Uganda, which was preserved in spirit and kindly

given to me by Dr. Cuthbert Christy, F.Z.8. It had no preorbital

glands, but two pairs of well-developed inguinal glands resembling

those of Cervicapra redunca, the pockets of the anterior pair being

shallow but wide-mouthed, and those of the posterior pair deeper

with circular orifice somewhat less in diameter than the diameter

of the pocket itself. Between the orifices of the inguinal pockets

were the two pairs of teats. The early development of the in-

guinal glands is a fact of considerable interest.

There was no trace of pedal g elands and only a very shallow and

short depression on the anterior surface of the fetlock close to the

base of the hoofs. The interungual area was naked as in Tawro-

tragus, and the web was deep, extending from near the proximal

end of the front of the hoof to the heels.

I have also examined the head, feet, and skin of the inguinal

region of a young male of 7. scriptus scriptus from French

Guinea, presented to the Society by Mr. Fenwick Owen. This

material was preserved in alcohol. The feet were constructed as

above described, and there was no preorbital gland. There were

two pairs of teats, but only a single pair of inguinal glands, each

consisting of a pouch about an inch deep opening by an. orifice

some distance away from, and in front of, the mamme of the

anterior pair (text-fig. 116, C, p. 922). I think it probable that

these glands correspond, to the deep posterior pair, and not to

the shallow anterior pair observed in the feetal example above

described.

L also received from Mr. E. Gerrard the dried front and hind

feet of a Bushbuck, probably belonging to 7’. scriptus roualeyni,

which also had no trace of pedal glands; but im which I was able

to discover the shrivelled remains of the vascular gelatinous

(fatty) mass lying along the back of the pastern between the true

and false hoofs, which has been already described in Cervicapra

redunca and Adenota kob.

TRAGELAPHUS ANGASII Angas.

(The Inyala,)

Of this species I received from Messrs. Rowland Ward, Ltd.,

a single dried foot of a male, which differed in no important

respects from the foot of 7. scriptus, except that the naked

shallow depression at the base of the hoofs on the front of the

pastern was obliterated and the hoofs themselves were shorter

with higher heels, suggesting the habit of walking on firmer

ground,

Genus Limnorraaus Sel. & Poe.

LIMNOTRAGUS GRATUS Sel.

As stated above, W. L. Sclater says the preorbital gland is

present in the southern race of ZL. gratus, namely, L. gratus

selousi Roths., the true Sitatunga.

fou] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 931

IT have examined no dead examples of this genus in a fresh

state, but examples of the typical Z. gratus and of L. gratus

seloust living in the Gardens show no trace of a preorbital gland,

nor of pedal , glands, the widely splayed hoofs making observation

of the absence of the pedal glands comparatively easy. The feet

appear to resemble those of Z’ragelaphus, apart from the length of

the hoofs and the nakedness of the back of the pasterns. Max

Weber (Die Siiug. p. 35, fig. 31, 1904) has figured the inguinal

region of a male ‘anid female On 3G gratus, showing the presence

of four mamme and of a single pair of small, almost aborted .

inguinal sacs, with orifices remote from and well in advance of

the anterior mamme. These obviously represent the larger

inguinal sacs seen in the specimen of 7ragelaphus scriptus scriptus

above described. It may further be noted that the glands as

figured by Weber closely resemble in size and position the single

minute inguinal sac seen on one side in the young example

of Adenota kob (p. 916): both are suggestive of a larger sac in

process of suppression.

Genus SrrRepsiceRos H. Sm,

STREPSICEROS STREPSICEROS Pall.

(The Greater Kudu.)

According to Owen, this species has inguinal pits but no pre-

orbital gland. Ogilby agrees with Owen as regards the absence

of preorbital glands and the presence of inguinal glands, and adds

that pedal glands are also absent. He also records the presence

of four mamme. W. L. Sclater, however, says that the preorbital

gland and its orifice are small. I have examined no example of this

species, but the available data suggest that Strepsiceros resembles

Tragelaphus or Limnotragus in the matter of cutaneous glands,

as it is well known to do in most other structural characters,

apart from the slight difference exhibited in the structure of

the horns. It is impossible, however, to guess whether this genus

has ‘inguinal glands like those of the foetal T’. scriptus (subsp. 2)

or the young L. s. scriptus, described above, or like those of Limmno-

tragus gratus figured by Max Weber. I suspect, however, that they

will prove to resemble in size and number those of 7. s. scriptus.

Genus Boocercus, Thos.

Boocercus EURYCEROS Ogilby.

(The Bongo.)

In an old skin of this species which Mr. Gerrard allowed me to

examine, I found the feet to be constructed as in 7'wurotragus, there

being no trace of glands and the interungual web being naked

throughout its extent. I could detect no trace of inguinal glands,

but found two pairs of teats.

The apparent absence of inguinal glands in this genus and in

932 MR. R. I, POCOCK ON THE [June 14,

Taurotragus and their presence in 7'ragelaphus and, as is alleged,

in Strepsiceros are quite in accordance with the’view that Boo-

cercus and Taurotragus are more nearly allied to one another

than either is to Strepsiceros or T'ragelaphus, a view supported by

the terminally tufted tail and the presence of horns in the female

in the two former genera,

Genus Tavrorracus Wagn.

TAUROTRAGUS ORYX Pall. (The Eland.)

(Text-fig. 122, A, B, p. 929.)

This species, according to Owen, who quotes it as oreas and

canna, has neither preorbital nor inguinal pits. W. L. Sclater,

however, says that the preorbital gland is small, ‘‘ with a very

small circular opening in front of the eye in a narrow bare tri-

angular space.” In two young male examples examined by me,

I could find no trace of preorbital gland or of inguinal or pedal

glands. The pedal glands were also entirely absent on a foot of

an adult kindly supplied to me by Messrs, Rowland Ward, Ltd.

The feet are constructed as in the genus Bos, the entire inter-

ungual area being naked, the integument that joins the hoofs

together forming a deep “ web” extending forwards from the heels

where it is thickest, to the anterior portion of the pastern only a

short distance above the proximal margin of the front of the

hoof, so that the hoofs are susceptible of but little separation.

By their glands and other external features the genera of this

section may be grouped and contrasted as follows :—

a. A well-developed preorbital gland opening by a longitudinal

valvular slit; horns short, not spirally twisted, present

only in the male (tail not terminally tufted; no inguinal

glands).

b, Preorbital gland relatively large; horn not crested; a pouch-

like gland in each false hoof of the hind leg, &e. ......... Tetraceros.

b'. Preorbital gland relatively small; horns crested basally ;

skin between posterior false hoofs highly glandular ...... Boselaphus.

a’. No preorbital gland; horns longish, spirally twisted.

c. Inguinal glands present ; tail hairy throughout ; no horns

in female .........s00ccecseeeeeeeeee Lragelaphus, Limnotragus, Strepsiceros.

ce’, No inguinal glands; tail terminally tufted ; horns present

ATU OLMIRGRON Mees Mere ene rans ducyessi'vce sus ccuineeaecnee Boocercus, Taurotragus.

Subfamily Bovin #.

tenus Bos Linn. (‘l'ext-figs. 123, 124.)

It is agreed on all hands that preorbital, inguinal, and pedal

glands are absent, and that two pairs of mamme are present in

all the species of Bovine, whether they be referred to the one

genus Bos or the several genera Los, Bibos, Poephagus, &e. In

these respects Cattle are indistinguishable from the large Trage-

laphine Antelopes Zauwrotragus and Loocercus and from the

larger forms of Cervicaprine like Kobus ellipsyprymnus, They

also resemble them in the structure of the feet, the interungual

‘an

‘

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 933

membrane being naked and continued straight backwards from a

point near the anterior proximal edge of the hoof to the heels,

forming a rectangular continuation on the middle line in front

and behind with the integument of the anterior and posterior

surfaces of the pastern respectively.

Text-fig. 123,

A. Median vertical section of foot of Bos sondaicus, three days old: gel.,

gelatinous mass between the phalanges.

BL. Anterior aspect of the same foot.

C. Median vertical section of foot of calf of Bos taurus; gel., gelatinous

mass between the phalanges.

T have verified this fact upon feet of specimens of the following

species :—

Bos taurus L. (Ox); fore and hind feet of a calf given to me

by Mr. R. E. Holding.

Bos indicus L. (Zebu); feet of adult male that died in the

Gardens.

Bos frontalis Lambert (Gayal); ditto.

Bos gaurus H. Sm. (Gaur); foot given to me by Mr. R. E.

He ding.

Proc. Zoou. Soc.—1910, No. LXI. 61

934 MR. R. I. POCOCK ON THE {June 14,.

Bos sondaicus Schleg. & Miill. (Banting); calf four days old

that died in the Gardens.

Bos grunniens (Yak); a female specimen.

Bos caffer Sparrm. (African Buffalo); dried foot lent to me by

Rowland Ward, Ltd.

Bos bubalus L. (Indian Buffalo) ; ditto.

Text-fig. 124.

A. Median vertical section of foot of Bos bubalus (dried).

B. The same of Bos caffer (dried).

In fresh specimens there is always a mass of vascular gelatinous.

tissue between the penultimate phalanges of the pastern, corre-

sponding to that which is found along the posterior inner surface

of the pastern in fresh specimens of Adenota hob, Cervicapra

redunca, Tawrotragus, Boselaphus, ete.

In the feet of the two Buffalos mentioned above I found the

interungual integument much thicker in B. caffer than in

B. bubalus, and the former had a deeper depression on the front

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 935

of the fetlock above the hoofs than the latter, the interungual

web not extending so far to the front and allowing the hoofs to

be rather more widely spread.

Family ANTILOCAPRIDA.

ANTILOCAPRA AMERICANA Ord. (‘The Prongbuck.)

(Text-fig. 125, A.)

In his paper upon the anatomy of this species (P. Z.S. 1870,

p. 340), Dr. Murie summarised his own observations and those

Text-fig. 125,

ULL

LET Z)

A. Median vertical section of front foot of prematurely born example of

Antilocapra americana; gl., pedal gland; o., its orifice.

B. Median vertical section of hind foot of Giraffa camelopardalis, three

weeks old; gel., gelatinous mass between the phalanges.

of previous authors on the cutaneous glands as follows:—(1) no

preorbital or inguinal glands; (2) a gland an inch and a half

61*

936 MR. R. I. POCOCK ON THE [June 14,

below the ear which exudes a yellow glutinous secretion with a

strong goaty odour; (3) an unpaired “dorsal gland six or eight

inches in front of the tail; (4) a gland on each side of the tail

above the ischiatic prominence ; (5) a gland on the posterior

aspect of each hock ; (6) interdigital elands on all four limbs.

My observations on this species were limited to a prematurely

born feetus. On this I could find no trace of the glands below the

ears or on the hocks or on the ischia, except for the crease of skin

at the root of the tail on each side. The dorsal gland was in a

rudimentary condition, the hairs on each side of the spine of the

lumbo-sacral area showing a black spot and being slightly stuck

together.

‘In view of the absence or rudimentary condition of the glands

mentioned, it was interesting to find the pedal glands highly

developed. They opened by a small circular orifice on the front

of the pastern some distance above the base of the hoofs. The

orifice led into a long and spacious gland directed at first down-

wards towards the hoofs, then shar ‘ply curved upwards along the

posterior wall of the pastern, occupying almost the entire space

between the phalanges of the feet, nearly as far up as their

articulation with the cannon-bone. These glands, present on all

four feet, were crammed full of white cheesy secretion. They

resembled on a very large scale the glands of Sheep more than

the glands of any of the Bovidee or Cervidee, although the orifice

was situated higher up the pastern than in Ovis, recalling in this

particular the orifice of such a Deer as Capreolus. It was notice-

able, too, that there was no depression on the front of the pastern

for the orifice to open upon. The hoofs were united by a very

deep and continuously hairy web, extending from the heels as

far forwards as the anterior proximal margin of the hoofs.

Family GIRAFFIDA.

GIRAFFA CAMELOPARDALIS Linn, (The Giraffe.)

(Text-fig. 125, B.)

According to Ogilby, the Giraffe has neither preorbital nor

inguinal glands, but has small interdigital fosse and four

mamme.

A young specimen, three weeks old, that died in the Zoological

Gardens, presented the characters mentioned by Ogilby except

that there was no trace of ‘ interdigital fosse,” the feet being

completely webbed, with low heels and no trace of a depression on

the front of the pastern. ‘The interungual integument, moreover,

was quite naked from front to back, as in Bovine and Tragelaphine

Ruminants, and the same gelatinous mass of tissue was present

between the penultimate phalanges of the pastern as is found in

the latter animals.

There was a thick pad of hairs over and below the knees ; but

| could detect no evidence of glandular activity in the underlying

skin, there being plenty of underwool but no secretion at the base

1910. ] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 937

of the long hairs of the pad. The same pad, I noticed, was well

developed in a feetal Giraffe taken from its mother that died some

four years ago in the Gardens (see F. E. Beddard, P. Z.S. 1906, ii.

p. 630, fig. 109).

The horns of the three weeks old specimen, which were rela-

tively as large as in the adult, had no connection, apart from

contact, with the skull, a fact already recorded by Sir Rav

Lankester (P. Z. 8. 1907, p. 101).

Family MOSCHIDAK.

Moscuvus moscurrerus Linn, (The Musk-Deer.)

(Text-fig. 126.)

The peculiar musk-gland of the male of this species has been

described by many authors, and I have nothing to add to what has

already been said of this organ*. Ogilby and Hodgson state that

the preorbital and pedal glands are absent. Flower tT confirms

this, saying ‘“ nor were there any interdigital glands on either feet,

the depressed space between the toes, where the glands usually

open, being covered with hair.” Brandt also described a gland on

the outer side of the thigh in the male; and Hodgson discovered

a gland opening on each side of the tail in that sex §. Neither the

femoral nor caudal glands were found by Flower in the female

example dissected by him.

I have only seen one example of this species, a male from

Chamba presented to the Society by Major G.S. Rodon, F.Z.S.; at

the time of its death I merely noted the structure of the feet and

the absence of the preorbital gland. On the front feet it was very

obvious that the long hairs of the interdigital depression were

stuck together with a brown sticky substance, having a decided

but indescribable smell and indicating considerable activity of the

sebaceous glands. On the hind feet there was no such secretion,

the hairs being clean and separable. The depression between the

phalanges of the feet in this species is deep, the integument of

the upper and under sides meeting at an acute angle to form a

web which extends nearly as far as the heels of the hoofs ; but on

the hind feet the webbing is wider and not so thick as on the front

feet, so that the hoofs of the former are capable of being rather

more widely distended. The entire interdigital and interungual

area is thickly covered with long hairs the tips of which overlap

the proximal portion of the hoofs to a considerable extent.

On the dried skin I find no trace of the femoral gland described

* The scent of the secretion of this gland when fresh is decidedly unpleasant, with

a peculiar pungent urinary odour. It bears no sort of resemblance that I can detect

to the prepared “musk” of commerce. This is one of the criticisms that may be made

on Mr. Beddard’s perhaps satirical suggestion (“Mammalia” in the Cambridge Natural

History, p. 13, 1902) that the musky scent of this Deer may be mimetic, in the

sense of frightening its enemies by suggesting the proximity of Crocodiles.

+ P. Z. S. 1875, p. 159 &e.

{ Bull. Sci. Acad. St. Pétersb. i. p. 174, 1836.

§ J. A. S. Bengal, x. p. 795, 1841.

938 MR, R. I. POCOCK ON THE [June 14,

by Brandt nor of the so-called calcic (? tarsal or met

mentioned by Hodgson. The tail, however,

is completely buried in the long hairs

atarsal) gland

is very peculiar. It

of the anal region and is for

Text-fig. 126.

Moschus moschiferus 2.

A. Lateral aspect of tail (dried),

B. Median vertical section of front foot (dried).

C. The same of hind foot (dried),

the most part naked,

Its tip, however, is furnished with a large

tuft and its naked up}

ver and lateral portions are overhung by long

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 939

hairs projecting backwards from its base on the upper side. Just

behind the anus on the lower side there is a mat of short hairs

which extends upwards laterally so as to cover the lower half of

the side of the tail. Just above this the lateral surface of the tail

is scooped out longitudinally, forming a wide gutter, which is filled

with yellow crumbly secretion, and this extends over the summit

of the tail and almost to its apex, both above and at the sides.

Before the skin became shrunken with drying, it is probable that

it was full and flaccid, and that the wide gutter above described

had the appearance of a narrow integumental slit such as Hodgson

figured.

From the great development of the caudal gland as well as of

the preputial gland in the male of the Musk-Deer, it may be

inferred that the females seek the males at the rutting season.

One other point may be noted: both Ogilby and Hodgson

declare that this Deer has two pairs of teats. Flower, on the

contrary, found only a single pair on the female he examined, and

I noted that the male above mentioned also had but one pair.

Family CERVIDA.

In describing the cutaneous scent-glands of the Deer I have

not taken into account the statements made by Fitzinger in his

diagnoses of the genera of this group (SB. Akad. Wien, Ixviil.

pp. 348-362, 1873), because many of them are contradicted by the

plainest facts. He says, for example, that Rangifer has no tarsal

gland and that Cervulus, Awis, Hyelaphus porcinus, Dorcelaphus

americanus, Mazama nemorivagus, and Dama have no pedal glands.

‘Since these glands are extremely well developed in all these Deer,

it is clear that no reliance is to be placed on this author’s state-

ments as to their absence in other species.

From the works of previous authors it is well known that one

ov more pairs of the following glands are present in all species of

this family :—

1. Pouch-like, shallower or deeper, preorbital glands, which are

apparently absent only in Capreolus,

2. Frontal glands, which are well developed in some species of

the Muntjac group and, according to Weber, in the Sambar.

3. Tarsal glands, consisting of a hair-covered patch of thickened

skin on the inner side of the tarsus or hock and especially

characteristic of the American Deer.

4, Metatarsal glands, which usually resemble the last struc-

turally, but are placed on the outer side of the metatarsus or

cannon-bone of the hind leg at a varying distance below the hock.

5, Pedal glands, which when well developed consist of a large

or small cleft or pouch on the front of the pastern usually of the

hind foot only, more rarely of the front as well.

Inguinal glands do not exist, and there are always two pairs of

teats.

940 MR. R, I. POCOCK ON THE [June 14,

Genus Cervus Linn.

Cervus (Cervus) MARAL Ogilby.

(The Caucasian Red Deer.) (‘Text-fig. 127.)

In an old Stag of this species that died in the Gardens the

orifice of the preorbital gland formed a subvertical §-shaped slit

at right angles to the long axis of the eye-aperture. The gland

itself was a simple but deep infolding of enormously thickened

skin, the pocket being scantily lined with short hair and filled

with a yellow waxy secretion.

The metatarsal gland was represented by a thick mat of hair

paler in colour than that of the rest of the leg. The area beneath

this mat was nowhere naked, as it is in Awis axis.

Text-fig. 127.

Cervus (Cervus) maral.

A. Transverse section through preorbital gland.

B. Longitudinal section through the same.

C. Median vertical section of front foot.

D. The same of hind foot.

On the front hoof the integument of the anterior surface of

the pastern formed a rather shallow depression uniformly clothed

with hair almost to the bottom. The interungual integument

was very thick and horny throughout its extent, forming a

strong but elastic union between the hoofs. Along the back of

the pastern up to the false hoofs the skin was also moderately

thickened ; but I could detect no evidence of any special glandular

activity in the skin of the front feet.

On the hind feet the interdigital depression on the anterior side

of the pastern was much deeper, the integument being much more

steeply inclined backwards and downwards towards the heels, the

1910. | CUTANEOUS SCENT-GLANDS OF RUMINANTS. 941

hoofs being joined together by a moderately deep and strong

smooth tie just above the heels. The integument of the deeper

portion of the interdigital depression was almost naked, being

merely sparsely clothed with short hair, and instead of forming

a continuous and level surface was sinuous or slightly folded, so

that the depression was somewhat deeper above and below than

in the middle. In section it was speckled with a layer of small

dark spherical glands, which extended not only over the skin of

the depression itself but round the interungual tie and some little

distance above the heels on the back of the pastern. These dark

specks, clearly visible to the naked eye, sharply define the position

of the skin with active glands from the rest.

The main differences, then, between the front and hind feet are

the presence of a deeper nearly naked interdigital depression,

of a shorter and weaker interungual web, and of more highly

developed sebaceous glands in the latter than in the former.

Cervus (Cervus) ELAPHUS Linn. (The Red Deer.)

(Text-fig. 128, A.)

According to Ogilby, C. elaphus has distinct and movable pre-

orbital glands and large pedal glands. Hodgson described the

Text-fig. 128.

GFE

ee ey eaeeL hi ea a

LE ecerst crys a

UE EOE

A, Median vertical section of hind foot of Cervus (Cervus) elaphus.

B. The same of Cervus (Rucervus) duvauceli.

942 MR. R, I. POCOCK ON THE [June 14,

preorbital glands as moderate in size and mobility and S-shaped,

the pedal glands as large but doubtfully present on all four feet ;

and added that the metatarsal gland is posterior and external. I

am indebted to Sir Claud Alexander, Bt., F.Z.S., for the front and

hind feet and the mask of a freshly killed Stag from Scotland.

The preorbital gland resembled that of C’. maral, except that

the orifice, when closed, formed a long slit, lightly concave

upwards and inclined obliquely at a very obtuse angle to the long

axis of the aperture of the eye.

The feet also were very like those of the Caucasian species,

except that the horny heels of the front hoofs were united below

the strong cutaneous heel-tie and a larger area of the deep portion

of the interdigital depression was very scantily clothed with short

hair. On the hind foot the depression, as in C. maral, was con-

siderably deeper and was divided by a transverse bridge into a

smaller wpper portion and a larger lower portion, both better

marked than in that species.

Cervus (Cervus) CANADENSIS Erxl.

(The Canadian Wapiti.)

The glands of this species resemble in all essential respects

those of C. elaphus and C. maral.

In an old male that died in the Gardens the orifice of the

preorbital gland was inclined at an obtuse angle to the axis of

the ocular aperture; the pocket was deep with nearly naked walls ;

the skin of the bottom at its deepest part just in front of the

orbit was very thin, the thickest part being close to the orifice,

especially on its anterior side. The secretion, which was very

dry, smelt like green grass just beginning to decay.

The metatarsal gland consisted of thickened skin completely

covered with hairs, which were white at the base and clogged with

secretion smelling like a pen of Domestic Sheep.

The feet in section were very like those of C. elaphus and

C. maral, except that the depression on the front of the pasterns

was shallower and the nail of the hoofs shorter. The heels of

the front hoofs were separated as in C. maral, not united as in

the example of C. elaphus examined.

According to Hodgson, the Hangul or Cashmere Stag (Cervus

cashnirianus), which he quotes as C. wallichii, and the Shou

Cervus affinis) agree with C. elaphus in their cutaneous glands.

Cervus (Pseupaxis) sikA Temm. (Japanese Deer.)

(Text-fig. 129.)

Two young male examples, about eighteen months old, agreed

closely in a general way with C. maral and C. elaphus, except that

the preorbital glands were much less developed, the infolding being

comparatively quite shallow. The metatarsal gland was marked

by a large elliptical white patch, consisting for the most part of

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 943

hairs with white tips. The bases of the hairs were covered with

adhesive secretion and the underlying skin was markedly thick

and glandular. In both the front and the hind feet the anterior

interdigital depression was naked and nearly hairless in its deeper

portion, but in the hind foot it was not subdivided into an upper

and a lower portion, which are so well marked in C. elaphus, the

floor of the depression sloping gradually and evenly downwards

and backwards from the tarso-metatarsal joint as in the front leg,

and the interdigital web was developed to about the same extent

in the two feet. In section the skin of this area showed no

especial signs of glandular activity.

Text-fig. 129.

"SS

Cervus (Pseudaxis) sika.

A. Median vertical section of front foot. | B. Median vertieal section of hind foot.

Although, on the evidence supplied by the antlers, Mr. Garrod

believed Pseudaxis to be nearly related to Dama, the structure of

the feet affords no support to this view.

Pseudaxis sika veadily interbreeds with Cervus elaphus.

Cervus (RucERVUS) DUVAUCELI Cuv.

(The Swamp-Deer or Barasingha.) (Text-fig. 128, B, p. 941.)

Hodgson, who refers this species to two genera under the names

Rucervus elaphoides vel duvauceli and Procervus dimorphe, says

that the preorbital glands are moderate, and moderately movable

in the former, medial and vertical in the latter, that the pedal

944 MR. R. I, POCOCK ON THE [June 14,

glands are absent, and that the presence of metatarsal glands is

doubtful.

In a young example, two months old, the preorbital gland was

scarcely developed, and consisted of little more than a shallow

naked channel running obliquely downwards and forwards from

the corner of the eye, There was no trace of metatarsal gland ;

nor can I detect this gland in living examples of the species.

The feet were very aimalae to those of the previously deseribed

species. On the front leg the skin of the anterior surface of the

pastern descended somewhat abruptly downwards and backwards

just above the hoofs, meeting at an acute angle the portion form-

ing the heel-tie which extended for a longer distance in a direction

parallel to the sole of the foot than in the other species. There

was no naked area along the middle line of the front of the

pastern; on the hind foot, however, this naked area was well

marked all the way up the front of the pastern, which was but

little depressed and showed no indication of being deeper above

and below than in the middle. The imterungu: a integument or

web formed an obtuse angle with that of the front of the pastern

and a right angle with that of its posterior surface, the heel-

tie being “long as in the front foot. On the posterior side of the

pastern ‘the hairs did not extend between the heels, as they did

tor a short distance in the case of the previously described species.

It is important to note the very slight differentiation of the

preorbital gl: and in this young specimen, and also its smaller

size in the eighteen months old examples of C. sika as com-

pared with that of the adult. From these facts it may be

inferred that the gland in question develops with age in these

Cervide. They are quite large, however, in newly born Muntjacs

(Cervulus).

Cervus (Panouta) evpr Guthr. (The Thameng or Thamin.)

(‘Text-fig. 130.)

In a very old male of this species the preorbital gland was

well developed, the invagination being about one inch deep at

its deepest part above, the direction of the long orifice, when

closed, being intermediate beween that of C. maral and C. elaphus.

The metatarsal gland was marked by a patch of hairs paler than

those of the rest “of the leg, with their bases clogged with black

secretion, the underlying skin being thick and markedly ¢ elandular

to the naked eye.

There was scarcely any depression on the front of the pastern

of either the front or the hind foot. On the front foot the

very shallow depression was hairy throughout as in C. duvaueeli,

without the naked area seen in C. maral, elaphus, and sika; and

even on the hind leg this naked area was of very small extent.

Since the skin of this area did not slope downwards and backwards.

between the hoofs in front, it resulted that the interungual tie or

web was of much greater extent than in the other species, the feet,,

1910.| CUTANEOUS SCENT-GLANDS OF RUMINANTS. 945

in section, being constructed very much as in the Tragelaphine

Bovide ; but, as in the other species of Cervidee hitherto described,

the interungual web was quite naked. The portion of this web

which formed the heel-tie was wider than in the other Deer,

giving greater distensibility to the hoofs, and the skin surrounding

the false hoofs was naked, although the back of the pastern was

normally hairy between the false hoofs and the heels *.

Text-fig. 130.

Cervus (Panolia) eldi.

A. Median vertical section of front foot. | B. Median vertical section of hind foot.

Genus Exaryurus M.-Edw.

ELAPHURUS DAVIDIANUS M.-Edw.

(The Milou or Pere David’s Deer.)

According to Mr. Lydekker (‘ Deer of all Lands,’ pp. 233-237) the

preorbital gland is large, the metatarsal gland covered with hair

and situated in the upper third of the cannon-bone, and the foot-

glands and tarsal glands absent in this species.

Examination of a dried skin in the British Museum has

enabled me to verify these statements, and to add to the informa-

tion supplied by Mr. Lydekker the fact that the interungual

integument or web is naked as in C. elaphus, maral, sika, eldi,

* In one race of this species the whole of the back of the pastern is naked as in

Limnotragus.

946 MR. R. I. POCOCK ON THE [June 14,

and duvauceli. Apart from the greater length of the false hoofs,

the feet of Llaphurus seem to differ from those of the species.

just cited in having the integumental groove on their upper or

anterior surfaces covered with short hair like that of the rest

of the foot, instead of being naked at the bottom.

Opinions of authors as to the systematic position of this Deer

are divided in accordance with the value attached to par-

ticular characters. Those, like Gordon Cameron and Lydekker,

who rely mainly upon the antlers, place HLlaphurus with the

American Deer; Brooke, on the contrary, depending largely upon

the structure of the lateral metacarpals, maintained that Hla-

phurus was an aberrant type of the group to which C. elaphus,

dama, wnicolor, and others belong.

The structure of the interdigital area of the pastern—that is to

say, the absence of the glandular pouch and depression on the

hind leg, and the smoothness of the interungual integument—and

the high position of the metatarsal gland completely bear out

Sir Victor Brooke’s opinion as to the affiliation of Hlaphurus with

the Old World Deer, especially the Oriental types (ef. infra,

pp. 969-970).

In opposition to Mr. Cameron’s view that HLlaphurus is not

related to the Old World Deer, the Marquis of Hamilton (‘ Field,”

July 1910, p. 199) pointed out that hybrids between Hlaphurus

and Cervus have been bred in Paris and that one of these hybrids

formed fertile unions with both parent stocks at Woburn. This.

is an important piece of evidence in favour of relationship

between the two genera.

Genus Rusa Ham. Smith.

Rusa untcotor Bechst. (The Sambar.)

(Text-fig. 131.)

According to Ogilby, this species has distinct and movable

preorbital glands and large pedal glands, the same epithets being

applied to them as to those of C’. elaphus, suggesting identity of

development and structure. Hodgson supplements this deserip-

tion by saying that the preorbital glands are very large and

completely reversile, that the pedal glands are large and present

on all four feet, and that the metatarsal gland is posterior and

external. According to Blanford, however, Hodgson says the

interdigital glands are wanting. I have seen one old male

example of this species.

The preorbital gland was a large and deep, naked invagination

half filled with a quantity of waxy secretion with a strong but

indescribable odour. Theintegument at the bottom of the pouch

was thin and apparently not glandular; but round the orifice,

especially anteriorly, if was much thickened, and it was here

apparently that the secreting area was concentrated,

The supraorbital gland, mentioned by Weber as present in

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 947

Rusa equina, I did not find; and no living Sambar that I have

seen has any gland on the forehead comparable in development

to that of the Muntjac (Cervulus).

The metatarsal gland was represented by a tuft of hair, which, as

in Llaphurus and Cervus, completely covered the underlying skin.

No evidence, however, of the glandular activity of the skin was

supplied by secretion at the base of the hairs.

Text-fig. 131.

Z Kus

sealers aus

CRS

Rusa unicolor.

A. Median vertical section of front foot. | B. Median vertical section of hind foot.

Neither on the front nor the hind feet was there a trace of

interdigital depression such as is seen in the Axis and Hog

Deer. The front of the pastern in both feet was only shallowly

depressed and was covered with hair as in Llaphurus, and the

interungual web was quite naked as in the latter and in the

typical Old World Elaphine Deer,

The difference in the structure of the hind feet between the

Sambar and the Axis justifies the generic separation of the two;

with this is correlated the nakedness of the metatarsal gland in

948 MR. R. I, POCOCK ON THE [June 14,

the latter animal and its hairy clothing in the former. On the

other hand, although the Sambar resembles the Bérasingha

(C. (Rucervus) duvauceli), the Thamin (C. (Panolia) eldi), and the

Japanese Deer (C. (Pseudawis) sika) in the structure of the feet,

it may be generically distinguished from these as well as from the

typical Elaphine Stags by the longer bushier tail and the extension

of the muffle as a distinct broadish rim of moist hairless skin

beneath the nostril.

According to the Marquis of Hamilton this species interbreeds

with Cervus elaphus (¢ Field,’ July 1910, p. 199).

Genus Axis Ham. Smith.

Axis (Axis) Axis Erxl. (The Chital or Spotted Deer.)

(Text-fig. 132, A—E.)

Hodgson says that the preorbital glands are large and very

movable, the pedal glands large and present only in the hind

feet, and that the metatarsal gland is posterior and external.

This is perfectly correct, according to my observations.

In an old male that died in the Gardens, the infolding of the

preorbital gland was decidedly shallower relatively to the size of the

animal than in (. maral, or C. elaphus, or C. eldi, and the line

of the orifice, when closed, was nearly parallel to that of the long

axis of the aperture of the eye and lightly concave upwards. The

pocket was lined with hair of the same colour as that of the face.

By squeezing, the gland could be made to yield drops of pale

moisture with a faint but decided musky smell.

The metatarsal gland was represented by a horny, hairless,

blackish oval patch of skin, exactly like the wart of a Zebra or

Wild Donkey and surrounded by a narrow rim of pale naked

skin. This patch was entirely concealed by the long hairs growing

immediately round it, recalling in this respect the metatarsal gland

of the Impala (Apyceros).

On the front feet there was only a very shallow depression on

the anterior surface of the pastern; but on the hind feet the

pedal gland was well developed and represented by a long and

deep depression starting at the top of the pastern and extending

almost to the hoofs. At its deepest part the wall of the depression

almost reached the skin of the posterior surface of the pastern.

From this point it gradually shallowed above and below. Its upper

edge reached the summit of the pastern, but its lower edge was

sunk some distance below and above the proximal portion of the

hoof, where it was continued into the interungual tie, which

extended backwards from near the middle of the hoof to the

heels, equalling in depth the corresponding web or tie of Cervus

maral and elaphus. As in those species this tie was naked, the

hairs beginning just between the heels and being continued up

the back of the pastern, From the above-given description it

may be inferred that the skin forming the lower rim of the inter-

1910. ] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 949

digital depression is separated by a considerable space from the

skin of the inferior part of the back of the pastern above the

heels and that the interdigital depression, instead of communicating

freely with the space between the hoofs, is to a great extent shut

Text-fig. 132.

zy)

Mii

WY SES, Ni

ea \ S \

Vy \ i ~ A i]

TAG | GON Sa" :

Set, i Ae) oo ate

Hy Hy) | A wan Miz, wot OZ w

J “th APSR) WE NAYS

HNN —— AMES

Ns fas

is a | \

4 (eres \ B

j ————S—

SSS

S25

EOIN

See ae

UW ;:44

. Transverse section of preorbital gland of Asis (Aais) axis.

. Part of the face of the same showing eye and gland (g/.).

. Anterior aspect of hind foot of same: gl, slit-like orifice of gland.

. Median vertical section of the hind foot of the same.

: aMetatarsal gland of the same with the hairs pulled aside to show naked

skin.

. Median vertical section of hind foot of Avis (Hyelaphus) porcinus.

my woaenS

off from it by the deep interungual tie. The walls of the

depression were scantily clothed with quite short hairs stuck

Proc. Zoox, Soc.—1910, No, LXIT, 62

950 MR. R. I. POCOCK ON THE [June 14,

together and stained with brownish secretion, and the layer of

secreting cells was restricted to the walls of the interdigital

depression and were much more highly developed than in

Cervus maral.

Axis (Hyertarnus) vorcinus Zimm. (The Hog-Deer.)

(Text-fig. 132, F.)

According to Hodgson, the Hog-Deer agrees with the Axis

in the development and structure of its cutaneous glands, both

species being referred to the genus Avis.

In a general way, this was true of the one example, an adult

female, which I examined in a fresh state. The invagination of

the preorbital gland was, however, shallower than in the male

Axis. The pedal gland differed, too, in one interesting particular, —

Its walls were covered with a growth of long hairs projecting

forwards from the orifice of the depression exactly as in the

Bubaline Antelopes. These hairs were clogged basally and

stained yellow by a waxy secretion with a pungent but in-

describable smell. Moreover, the interungual skin of the hind

foot was hairy; of the front foot it was naked, recalling what

oceurs in the Muntjac. The naked metatarsal patch was over-

lapped by the long marginal hairs as in 4. awis.

I have seen hybrids between the Chital and the Hog-Deer in

the possession of Sir Claud Alexander, by whom they were

bred.

Genus Dama Frisch.

Dama pvAMA Linn. (The Fallow-Deer.)

(Text-fig. 133.)

The structure of the feet and of the pedal glands of this Deer

have been described accurately by Max Tempel. The material I

have examined were a fawn about 24 hours old received from

Mr. R. E. Holding, some hind feet purchased ina shop in London,

and preparations of the feet in the Museum of the Royal College

of Surgeons.

Tn the fawn the preorbital gland consisted of a small but very

distinct pit of naked skin. This pit was considerably deeper

relatively than in the much older young example of Cervus

(Rucervus) duvauceli, but was shallower than in the fawn of

Cervulus muntjac.

The area of the metatarsal gland was quite covered with long

white close-set hair; but the gland showed no signs of activity.

The cleft of the pedal gland of the hind foot was as long as

in Axis avis or A, porcinus; but the cleft itself was decidedly

deeper, the skin of the posterior wall being almost in contact with

that of the back wall of the pastern. As in A. axis the deepest part

of the depression was at a point nearly midway between the heels

and the false hoofs; above the deepest part the skin sloped

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 951

somewhat steeply upwards to the joint of the pastern, but below

it the skin rose to only a very slight extent before passing into the

heel-tie, with which it formed a close fold. Thus the interdigital

cleft communicated quite freely with the space between the hoofs

and was not shut off therefrom by any extension of the integument

to form an interungual tie or web. The hoofs were united only

at the heels; but by way of compensation for this shallow

junctional area the heel-tie was very thick and horny. It was

naked behind. The whole of the interdigital cleft was thickly

Text-fig. 133.

< E ZU LEY fy y

= BLLALAE_

BZN

———

SS is Se

ae ////,

i YGYYLLI

Dama dama.

A. Median vertical section of front foot, from specimen in the Museum of the Royal

College of Surgeons.

B. Median vertical section of hind foot.

clothed with hairs, growing for the most part downwards and

grey or white in colour, but stained a deep yellow and stuck

together basally with secretion. Inferiorly these hairs spread

over the lower rim of the orifice formed by the closely folded skin

above the heel-tie. The hair along the back of the pastern spread

for a short distance between the heels, and the secreting cells of

the gland were restricted to the walls of the interdigital cleft.

In the Museum of the Royal College of Surgeons there are

preparations of the feet of this Deer. The hind foot agrees with

62*

952 MR. R. I, POCOCK ON THE [June 14,

the one above described, and the front foot shows a very deep

cleft similar to that on the hind foot but shallower, the web of

the heel-tie being much less closely folded.

Although, in the opinion of some authors, Dama is merely a sub-

genus of Cervus, the structure of the feet shows that it must rank

as quite a distinct genus from all Elaphine Deer to which it is

commonly supposed to be nearly allied. The only Deer, so far as my

knowledge extends, which have the hind feet so deeply cleft are the

Muntjacs (Cervulus) and the Chinese Water-Deer (/ydropotes).

IT am not aware that the Fallow-Deer ever interbreeds with the

Red Deer.

Genus CErvutus Blainv.

CervuLus MuNtsAc Zimm. (The Common Indian Muntjac.)

(Text-fig. 134.)

According to Ogilby, this species has very large movable pre-

orbital glands, two large movable supraorbital glands at the base

of the horns, and large pedal glands. Hodgson confirms what

Ogilby says about the preorbital and supraorbital glands, and adds

that the pedal glands are confined to the hind feet and that the

metatarsal gland is absent.

My own observations upon a specimen six days old that

died in the Gardens, agreed precisely with those of Hodgson.

The frontal gland was represented by two anteriorly converging

strips of skin covered with very short hair, and shallowly grooved

or rather longitudinally depressed along the middle line. This

area of skin was in no sense thickened, and in section showed

no macroscopic indication of being specially glandular, This

was probably attributable to the immaturity of the specimen,

for in living adult examples of this species now living in the

Gardens this gland appears as a long crease with thickened

upstanding edges, and, be it noted, it is much better developed

in the male than in the female.

The preorbital gland when closed has the form in the adult of a

crescentic slit with the concavity directed upwards. ‘The posterior

extremity of the slit, however, curves back towards the shallow

groove running forwards from ‘the eye, and its opposite extremity

curves slightly the other way, giving in reality a slightly sigmoid

curve to the slit. The two ends of the slit are deeper than its

median portion, and when the gland was everted in the young

individual its area was seen to be subcircular and marked with

an anterior and posterior groove representing the deeper parts of

the slit.

There was no trace of tarsal or metatarsal glands or tufts of

hair; and the pedal glands were present only on the hind feet,

the front of the paste’ rm bearing a deep triangular cleft when the

hoofs were separated. The skin of the back of the pastern

reached the heels, and was folded somewhat closely back upon

~1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS, 953

itself, though not quite so closely as in Dama. The anterior skin

of this fold or web formed the posterior wall of the interdigital

Text-fig. 134.

a =

: cs

J AN KRKOSSL

cue Nl MSs

Tos ” :

—— EE

GUC G Ge

eS So

S11 iq

y ; —

if <= —_

. yy 5 { \ Ss

SS =

SSS Wy tip) ff =

SL ees ltt ; E

— “

LS SSS

47)

Cervulus muntjac.

A. Median vertical section of hind foot of specimen six days old.

B. Anterior aspect of hind foot of the same.

C. Median vertical section of front foot of the same.

D, E, F. Preorbital gland, in three stages of distension, of the same.

G. Frontal glandular areas of the same.

954 MR. R, I. POCOCK ON THE [June 14,

cleft, the upper portion of which was depressed to form a shallow

pocket with a narrow elongated orifice, whence long hairs,

stuck together with secretion, projected. This pocket was the

glandular area of the cleft. The sides of the cleft were covered

with hair, as also was the interdigital web.

On the front foot the skin of the anterior surface of the pastern

formed a very slight and gradual slope between the digits until

it reached a point on a level with the heels, whence it receded

somewhat abruptly, forming a moderately deep interungual web,

which met the skin of the back of the pastern at right angles,

constituting the heel-tie. In this foot the heel-tie and interungual

web were smooth and not hairy as in the hind foot.

In a young male of about a year old examined since the one

just mentioned was described, I found the glands similarly but

better developed. For instance, the dark glandular layer of the

pedal glands was much thicker, and the white hairs of the

depression were stained a rich grass-green colour by the secretion,

the smell of which might be described as “ musteline,” being

decidedly unpleasant and reminding me of the scent of a Skunk

(Mephitis). When the frontal gland was excised and squeezed,

minute drops of clear fluid appeared on the skin. This had

quite a pleasant odour, very like that of a living Civet (Viverra

civetta) or of the waxy substance cobblers call ‘“ heel-ball.”

CERVULUS REEVESI Ogilby. (Reeves’ Muntjac.)

(Text-fig. 135, E.)

I had an opportunity of examining the glands on a fresh nearly

adult female specimen of this species sent to the Gardens by

Mr. W. Jamrach for determination. The frontal glands were not

nearly so well developed as in C. muntjac: in the latter the

nearly naked areas of skin converge and meet on the summit

of the nose; but in the specimen of C. reevesi examined they

were considerably shorter, subparallel, and separated throughout

their extent by a normally hairy area of skin.

The preorbital glands were of substantially the same structure

in the two. The secretion was white and adhesive, with a strong

smell of cream-cheese.

The interdigital cleft of the hind foot had no differentiated

glandular pocket in its upper half, but was evenly deep throughout

its length, very closely resembling that of Dama dama, the inter-

digital web consisting of a close fold of skin running along the

back of the lower half of the pastern and tying the heels

together where the fold began. his heel-tie and the walls of

the cleft were covered with hairs, and long hairs projected forwards

and downwards from its deeper portion, exactly as in the Hog-

Deer (Axis porcinus).

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 955

Genus Erarnopus M.-Edw.

ELAPHODUS CEPHALOPHUS Or MICHIANUS.

(The Chinese Tufted Muntjac.) (Text-fig. 135, A—D.)

In his memoir on the anatomy of 2. michianus, Garrod does

not mention the pedal glands. He records, however, the absence

Text-fig. 155,

ee an JN

iS xf ait Z| \

rs a

ee Por,

SS Saal

SAS Av SSS S

1 WAS

A. Orifice of preorbital gland and part of the eye of Llaphodus michianus,

B. Median vertical section of hind foot of the same.

C. Do. of front foot of the same.

D. Rhinarium.

E. Median vertical section of hind foot of Cervulus reevesi.

of frontal glands and the presence of preorbital glands in both

sexes *,

* P. Z. S. 1876, p. 757.

956 MR. R, I, POCOCK ON THE [June 14,

T am indebted to Mr. Edward Gerrard for the opportunity of

examining a dry skin of a female of one of these Deer, probably

referable to the Ningpo subspecies /. cephalophus michianus.

There was no trace of frontal gland.

The preorbital glands differed in no respects from those of

Cervulus, consisting of a subcircular area of short-haired integu-

ment with a subcrescentic crease deeper at the two ends than in

the centre. The pedal gland of the hind foot resembled that of

C. reevesi, except that the cleft was relatively longer. On the

front foot there was no gland and only a shallow depression, the

interungual web being moderately deep, owing to the anterior

and posterior skins of the pastern being considerably separated in

the middle line. The heel-tie and interungual web were quite

naked. The false hoofs were of comparatively large size and

the hoofs were shorter than in C. reevesi.

There were two pairs of teats as in other Deer. The absence

of the frontal glands and the presence of well-developed false

hoofs show that Llaphodus is a less specialised type than Cervulus

muntjac or reevesi. False hoofs, however, are by no means always

absent in specimens in the Gardens referred to C. muntjac.

Genus Hyproporrs Swinh. (= Hyprenarnus Lydd.).

HyDROPOTES INERMIS Swinh.

(The Chinese Water-Deer.)

I have seen no example of Hydropotes (= Hydrelaphus Lydd.),

but, according to Garrod (P. Z. 8. 1877, p. 780), the preorbital

gland was present, but quite small, in a newly-born specimen of

H. inermis, and the metatarsal gland was absent. Of the pedal

glands this author wrote :—‘‘ In the fore-limb the interdigital

skin is inflected but slightly, and there is no special gland

differentiated, although the surface of the skin is apparently

studded with minute gland-openings. In the hind-limb the inter-

digital skin forms a deep pocket which almost completely separates

the toes, except that they are joined by a thin transverse skin-

fold along their posterior edges. The included skin is studded

with small glands.” I infer from this description that the

posterior pedal gland resembles that of Cervulus or Dama and is

quite unlike the corresponding gland of Capreolus, the genus

near which Sir Victor Brooke placed Hydropotes on account of

the structure of its lateral metacarpals and vomer. Since W. A.

Forbes came to the same conclusion touching the affinities of these

two genera judging from their visceral anatomy (P. Z. 8. 1882,

p. 637), there is no reason to doubt the correctness of Sir V.

Brooke’s view. All the more interesting, therefore, is it to find

the difference in the pedal glands above alluded to; because

Hydropotes is the only member of the Telemetacarpal Deer yet

recorded as having the interdigital glandular cleft communicating

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 957

freely with the interungual space as in Dama and Cervulus, this

type of gland being, in my opinion, as is explained below, the most

primitive type in the Ruminants.

Genus CApreouus Frisch.

Capreotus cAPREOLUS Linn. (The Common Roe.)

(Text-fig. 136.)

According to Ogilby, this species has no preorbital glands, but

large interdigital glands ; and since he described the pedal glands

both of Cervus elaphus and of C. capreolus as “ fosse interdigitales

magne,” it might be inferred that they have the same structure

Text-fig. 136.

ZS >

ZZ f

ES Sen oa

ZA ZA DT

2 —— IL

LZ LM Yj

ey, :

o7. GYfe they

eee S,

Capreolus capreolus.

A. Median vertical section of hind foot. | B. Median vertical section of front foot.

in the two species. This inference, however, is as little in accord

with the facts as the statement that the pedal glands of Capreolus

are very like those of Hydropotes*. Max Tempel accurately

figured and described the glandular structures both on the front

and hind feet of Capreolus.

The only material of this species which I have been able

to examine was some limbs of freshly killed specimens kindly

* Lydekker, ‘ Deer of all Lands,’ p. 223.

958 MR. R. I. POCOCK ON THE [June 14,

procured for me by Dr. Plimmer, F.R.S., Pathologist to the Society,

and by Mr. R. EK. Holding, and a face-skin for which I was also

indebted to the latter gentleman. Mr. Lydekker describes the

face-gland of Capreolus as “almost obsolete.” I could find no

trace of it on the face-skin that I examined.

The metatarsal gland was represented by a thick pad of hairs,

the bases of which exhibited blackish secretion derived from the

thickened skin beneath. I could detect no distinct smell in

connection with this gland,

The pedal gland of the hind foot was represented externally by

a smallish orifice opening on the front of the pastern high above

the hoofs. This constricted orifice led into a capacious sae ex-

tending as far back as the posterior skin of the pastern and also

to a certain distance upwards, its upper extremity reaching as far

as the false hoofs. The walls of the sac were scantily clothed —

with long pale hairs all stuck together with thick yellow secretion

smelling like cheese. The dark brown gelatinous-looking glan-

dular layer enveloped not only the sac itself but the skin of the

front of the pastern below it and the skin of its posterior side up

as far as the false hoofs. The hair between the hoofs was all

matted with secretion. Below the orifice of the glandular pouch

the skin of the middle line of the pastern was only slightly de-

pressed ; the anterior and posterior walls of the interungual web

were moderately widely separated and in no sense closely folded

as in Cervulus or Dama. This web was, however, uniformly and

thickly covered, both above, below, and between the heels, with

long hairs.

On the front leg the glandular sac was represented by a very

shallow depression, but the skin of the pastern below this down

to the hoofs and of the back of the pastern from the heels up to

the false hoofs was lined with a dark secreting layer. This.

layer did not, however, extend up the interungual web, which

was deeper than that of the hind foot but similarly hairy. The

hairs both on the front and the back of the pastern were stuck

together basally with blackish sticky secretion smelling of mouse

urine.

Genus ALcE Frisch.

Ace ALCES Linn. (The Moose.)

(Text-fig. 137.)

According to Ogilby, this species has large preorbital and pedal

glands. Subsequent authors have also recorded the presence of

preorbital, tarsal, and metatarsal glands, the latter, according to

Nitsche (Zool, Anz. xiv. p. 181, 1891), being represented by a

naked area of skin overlapped by a marginal fringe. Max

Tempel accurately described the structure of the feet and the

microscopic structure of the glandular pocket of the posterior foot.

I have nothing new to add on these points, the only material

of the genus I have been able to procure for examination being

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 959

two feet which, from certain differences in their structure, I judge

to be those of front and hind legs. The former I received from

Mr. R. E. Holding, the latter from Messrs. Rowland Ward, Ltd.

On the one judged to be the hind foot the interdigital pocket

found in Capreolus and Rangifer was represented by a distinct but

shallow depression, with hairs paler than those of the surrounding

skin projecting from it. A little below this the integument dipped

obliquely backwards and downwards towards the heels to meet

that of the posterior side of the pastern, which extended for some

distance between the heels but was not markedly thickened to form

a strong heel-tie. The entire interungual area and heel-tie were

covered with long hairs, there being only a small less thickly

haired area a little below the glandular depression.

Text-fig. 137.

qeeeelog

Alce alces.

A. Median vertical section of front foot (dried).

B. The same of hind foot (dried).

On the front leg there was scarcely a trace of the glandular

pocket seen on the other, but the slightly depressed skin along

the middle line of the front of the pastern towards the hoofs was

scantily hairy, this area being of much greater extent than on

the foot previously described. Moreover, the interungual web

960 MR. R. I. POCOCK ON THE [June 14,

was much thicker and deeper, the two halves of the hoof of the

front foot being evidently much more strongly united than those

of the hind foot. As in the latter, the whole interungual tie was

thickly covered with long hairs, this hairy clothing constituting

a marked difference between the feet of the Moose and those of

the Red Deer group, and showing equally marked similarity

between the Moose, Roebuck, Reindeer, and typical American

Deer.

Genus RANGIFER Frisch.

RANGIFER TARANDUS Linn. (The Reindeer.)

(Text-fig. 138.)

According to Ogilby, this Deer has large preorbital and small

pedal glands. Max Tempel shows that there is a well-developed

glandular pocket on the hind foot, but none on the front foot.

My own observations are completely in accord with this.

The only material of this species 1 have been able to examine

was the fore and hind feet of the Caribou (2. tarandus caribou)

lent to me by Mr. Gerrard, and the fore foot of a second

example received from Messrs. Rowland Ward, Ltd.

The tarsal gland consisted of a large area of thickened skin

covered with a mat of long coarse white hairs without underwool,

the bases of the hairs being clogged with colourless scurfy

secretion.

The pedal gland opened by a large orifice on the front of the

pastern some distance above the proximal edge of the hoof. The

orifice led into a deep subcylindrical pocket, the deeper end of

which extended obliquely backwards and upwards in the direction

of the false hoofs. The cavity of this sac was filled with long

hairs, all directed towards the orifice and stained yellow, and a

well-marked dark-coloured secreting layer enveloped the walls of

the sae within the pastern. ‘The interungual web was very short,

and was situated some distance from the heels towards the

proximal end of the hoof in front, forming in section a semi-

elliptical figure. Like the back and the front of the pastern, it

was thickly covered throughout with long hair.

The front foot was, in a general way, like the hind foot, but

had no trace of the glandular pocket, and the interungual web

was even less extensive and still further removed from the heels.

The resemblances between the glands of the feet of the Rein-

deer and those of the Roe (Capreolus) or Brocket (J/azama) are

unmistakable, and are quite in accord with Sir Victor Brooke’s

view of the relationship between these Deer.

Writing of the foot of the Reindeer, Flower and Lydekker

remark :—“ The cleft between the two main hoofs is very deep,

so that these hoofs can be spread out as the animal traverses the

snow-clad regions in which it dwells” (¢ Mammalia,’ p. 325, 1891).

This expression, although true in a sense, by no means conveys

an accurate impression of the cause of the potential expansion of

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 961

the hoofs of this species, nor does it suggest the difference between

the distensibility of the hoofs in the Reindeer and those of the

Red Deer for example.

Text-fig. 138.

SALA

G D 4, Ve tt

aa TELL =

Rangifer tarandus.

A. Median vertical section of front foot (dried).

B. The same of hind foot (dried).

The peculiarity of the foot of the Reindeer consists in this :—

The integumental web joining the two hoofs together passes

straight forwards some distance above the heels and terminates

behind and nearly on a level with the upper posterior margin of

the nail. From this arrangement it results that whereas the

superior and proximal portions of the hoof are capable of com-

paratively little distension, the heels can be widely separated,

giving to the hoof a wide plantar area which is increased by the

long false hoofs. It is only when the hoofs are thus distended

962 MR. R. I, POCOCK ON THE [June 14,

that the plantar surface, or sole, of each is applied flat to the

ground, on account of the much greater size of the external as

compared with the internal lamina of the nail.

Genus DorceLaruus Gloger.

DorcELAPHUS (or CARIACUS) AMERICANUS Erxl.

(The Virginian Deer.) (Text-fig. 140, C, p. 965.)

I have had no dead example of this species for examination,

but in the collection of the Royal College of Surgeons there are

mounted specimens of the fore and hind feet showing the pedal

glands. ‘The glands on the hind foot resemble in a general

way both in position and development those of the Roebuck

(Capreolus), except that the orifice is relatively larger and less

constricted. As in the other Telemetacarpal Deer that I have

seen, the interungual integument is covered with longish hairs,

continuous in front and behind with those of the anterior and,

posterior sides of the pastern respectively.

The front feet are similarly constructed, and have a large

pocket-like gland resembling that of the hind feet but smaller.

This is the only species of Deer known to me which has a large

pedal gland on the fore foot.

The presence of preorbital, tarsal, and metatarsal glands in this

and allied species has been recorded by other writers, and they

are all apparent upon a living specimen in the Gardens, from

Venezuela, which I identify as D. americanus savannarum.

Genus Mazama Raf.

MazaMA NEMorIvAGuS F. Cuy. (‘The Wood-Brocket.)

(Text-fig. 139.)

I have examined one adult female of this species.

The preorbital gland was a small shallow infolding, linear when

closed and subelliptical when spread, resembling that of an

eighteen months old Cervus sika.

The hock- or tarsal gland was marked externally by a small

tuft of whitish or ashy-grey hairs arranged in a whorl and with

their bases encrusted with dark brown secretion, the underlying

skin being thick in section, with the roots of the hairs penetrating

deeply into it. There was no metatarsal gland.

The pedal gland was well developed upon the hind foot. The

front of the pastern was marked by a shallow triangular de-

pression, at the summit of which was the orifice of a cylindrical

sac resembling that of the Reindeer (/tangifer), the deep end of

the pocket reaching nearly to the false hoofs. The walls of the

pocket were sparsely lined with hairs covered with secretion

which stained black the longer hairs of the depression below the

orifice of the gland. The secreting layer of cells enveloped the

pocket and extended along the skin forming the posterior wall of

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 963

the depression down to the point where it passed into the inter-

ungual web. This was deep and projected straight backwards from

a point about on a level with the middle of the proximal edge of

the nail nearly to the posterior part of the heels, meeting at right

angles the skin of both the anterior and posterior walls of the

Text-fig. 139.

Mazama nemorivagus 2.

A. Median vertical section of front foot.

B. The same of hind foot.

C. Ocular region and preorbital gland.

D. Section of tarsal gland.

E. Rhinarium.

pastern which were widely separated. ‘The interungual web was

uniformly covered with longish hairs barely overlapping the

proximal edge of the nail. The hoofs were long and pointed.

The front hoof was constructed almost exactly like the hind,

964 MR. R. I. POCOCK ON THE [June 14,

except that the pocket was represented by a very shallow

depression, and the area of the pastern below this was scarcely

depressed. The secreting layer of cells lined the integument in

exactly the same way.

I believe this species to have been correctly determined ;

yet, according to Fitzinger, If. nemorivagus has no tarsal gland,

and was consequently made the type of that author’s genus

Doryceros.

MazaMA BRICENII Thos. (Briceni’s Brocket.)

(Text-fig. 140, A, B.)

A single adult female example from Venezuela, The preorbital

gland resembled that of J. nemorivagus. Superficial examina-

tion of the hock showed no trace of the tarsal gland, but when the

hairs over the appropriate spot were rubbed aside they were seen

to have darker bases than elsewhere on the leg and the under-

lying integument was thickened as in J/. nemorivagus although

to a much less degree. On a dried skin these glands would

probably have been wholly overlooked, since they were not marked

by any tuft of hairs thicker or coarser than those surrounding

the area.

The feet were rather differently constructed from those of

M. nemorivagus, the hoofs being very much shorter, the inter-

digital cleft on the front of the pastern deeper, and the inter-

ungual web shallower. On the hind foot the glandular pocket

was capacious and deep, extending backwards and upwards and

touching the skin of the posterior side of the pastern just below

the false hoofs. It was nearly elliptical in section, the lumen of

the sac being larger than the orifice. Just below the orifice the

integument formed an arched curve down to the strong heel-tie.

The walls of the pouch were thickly and uniformly covered with

long hairs projecting towards the orifice; and the walls of the

interdigital cleft and the heel-tie were also covered uniformly

with hairs. An olive-brown glandular secreting layer enveloped

the pocket and extended inferiorly nearly to the heel-tie. On the

front foot the pocket, although small as compared with that of

the hind foot, was better marked than in any species of Deer

I have examined with the exception of Dorcelaphus americanus,

consisting of a very definite hair-lined invagination. Below the

orifice the skin of the anterior surface of the pastern passed towards

the hoofs in a direction parallel to that of the posterior side of the

pastern, then descended obliquely downwards and backwards to

the heel-tie, which for a short distance was hairless, thus differing

from the corresponding area in the hind foot, and recalling the

similar difference between the two feet of the Muntijacs (Cervulus).

Since pedal glands essentially similar in structure and position

are found in such widely-divergent genera of Telemetacarpal

Deer as Rangifer, Capreolus, Dorcelaphus, and Mazama, it may be

inferred that they are present at least upon the hind feet in the

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 965

other Deer of this group, the genus Pudu alone constituting an

exception.

Text-fig. 140.

wah c

ae Hiss

| es

yD ILE

at :

é EN eA

A. Median vertical section of front foot of Mazama bricenii 9.

B. Do. of hind foot of the same.

C. Do. of hind foot of Dorcelaphus americanus from specimen in the

Museum of the Royal College of Surgeons.

I have had no opportunity of examining the feet or fresh skins

of any examples of the following types of Telemetacarpal Deer.

The principal external features of the glands will be found

described in Mr. Lydekker’s monograph.

Proc. Zoou. Soc.—1910, No. LXITI. 63

966 MR. R. I, POCOCK ON THE [June 14,

Genus Evuceryus Gray.

Evucervus HEMIONUS Raf.

(The Mule Deer.)

Nothing, so far as [ am aware, is known of the structure of the

feet ; but the preorbital, tarsal, and metatarsal glands are present,

the metatarsal gland being unique in its length, since it extends

at least halfway down the cannon-bone towards the fetlock. This

character I have used as a generic feature to distinguish Hucervus

from Dorcelaphus (p. 971).

Genus BLAsrocerus Sund.

BLASTOCERUS DicHOTOMUS II]. (=PALUDOsUS).

(The Marsh- Deer.)

The preorbital and tarsal glands are present and the metatarsal

gland absent, according to Mr. Lydekker. In this last-mentioned

character the Marsh-Deer resembles Mazama.

BLASTOCERUS BEZOARTICUS Linn. (=CAMPESTRIS).

(The Pampas Deer.)

This species appears to resemble the foregoing in the nature

of its glands,

The difference in the antlers suggests that this Deer should

rank as a genus (or subgenus) distinct from B. dichotomus.

Genus HirrocaMetus Leuck.

HiprocaMetus Bisutcus Mol.

(The Chilian Guemal.)

Under the name NXenelaphus, Mr. Lydekker admits this and

its ally H. antisiensis as a subgenus on account of the simply

forked antlers, the coarse brittle hair, and the absence of the

metatarsal gland. The latter, however, is also wanting in

Blastocerus and Mazama, leaving the subgeneric status dependent

on the character of the coat and antlers. This authoralso remarks

that the simple form of the antlers is seemingly due to degra-

dation and cannot therefore be regarded as an ancestral type,

because the absence of spots in the fawn and the suppression

of the metatarsal gland are obviously specialised features. Yet

there is no commoner morphological phenomenon than the

association of primitive with specialised features in a species.

Moreover, the antlers of Hippocamelus and Mazama may be due

to simplification from a much more complex type and yet be

“ancestral” in the sense that their characters result from the

retention to maturity of an early growth-stage.

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 967

Genus Pupv Gray. ,

Pupu pupu Mol, (The Chilian Pudu.)

I have had no opportunity of examining any material of this

genus beyond the dried skins in the collection of the British

Museum. According to Flower (P.Z.8. 1875, p. 160), the

preorbital gland in P. pudu “is a distinct involution of thickened

integument .... with an aperture half an inch in length,” and

the pedal glands are not represented by distinct pouches, “ but

the skin in the depression between the toes on the dorsal surface

of all the feet is bare and evidently has a free sebaceous secre-

tion,” representing, in the author’s opinion, “ the most rudimentary

or earliest stage of an interdigital gland” (see p. 978). So far as

it is possible to form an opinion from this description, it may be

inferred that the interdigital depression in this aberrant genus

closely, and it must be admitted surprisingly, resembles that of

such species as Cervus elaphus or C. eldi.

Lydekker says both tarsal and metatarsal glands are absent

(¢ Deer of all Lands,’ p. 307).

The Classitication of the Cervide.

From the account just given of the cutaneous glands of the

Deer, it may be inferred that I think Sir Victor Brooke’s classi-

fication of this family is correct, and Mr. Gordon Cameron’s

incorrect in the particulars in which it differs from Brooke’s.

It is important to refer to this matter, because in Mr. Lydekker’s

volume on Deer, to which recent authors not unnaturally refer

as a standard work, Brooke’s classification is set aside in favour

of Cameron’s, which gives primary importance to the antlers as a

basis for the major divisions of the Cervide *.

Briefly stated, Mr. Cameron classifies the Deer into four

groups :—(1) Rangifer, with antlers in both sexes; (2) Alce, with

antlers rising laterally on the sides of the skull; (3) mostly

American Deer of small or medium size, termed collectively

Cariacus, with which are associated Capreolus and Hlaphurus, a

section characterised by ‘‘forked” antlers without a brow-tine ;

(4) all the Deer of the Old World, except those mentioned above,

including Cervus and Cervulus, which are distinguished by antlers

consisting of a “beam,” usually itself branched with a basal

“ brow-tine.” These third and fourth sections are, however,

derivable from a common stock, from which Rangifer and Alce

are excluded as “strangers to one another and to all other groups

from the time when antlers first distinguished the family”

(The ‘ Field,’ 1892, p. 703).

Rangifer is considered to be a primitive type on account of the

occurrence of antlers in both sexes and their early appearance in

* The ‘ Field, 1892, pp. 625, 703, 741, 860. In the ‘ Field’ for 1910, pp. 154, 215

(July), this classification was upheld by Mr. Cameron in opposition to the views

adyocated in this paper and summarised in the ‘ Field,’ 1910, p. 97 (July).

63*

968 MR. R. I, POCOCK ON THE [June 14,

the young. This assumption I believe to be false. If true, we

should expect to find antlers of a primitive type in Rangifer.

This, however, is not claimed, Mr. Cameron describing those of

the Caribou ‘tas the most specialised of existing antlers.” Yet

their fundamental resemblance to the antlers of the Deer, in

which these weapons are well developed, is clear enough; and

this fact is difficult to explain in conjunction with Mr. Cameron’s

double claim that the antlers of Kangifer are highly specialised

and that the genus has followed a line of its own in evolution,

independent of all existing species. The characters of Rangifer

are, in my opinion, much more easily interpreted on the hypothesis

that the genus is a specialised representative of the Dorcelaphine

(Car iacine) Deer.

In the case of Alce, the lateral direction of the antlers is

admittedly a peculiarity ; but it is as obviously a derivative, not

a primitive, feature. As for the antlers themselves, they are, m

their simple form, clearly referable to the so-called “ forked type”

characteristic of Mr. Cameron’s third section of Deer ; and Alce,

like Rangifer, may be best interpreted as a specialised member of

that group.

Mr. Cameron also claims some support for his view of the great

antiquity of these two Deer on the ground of their wide range

‘in space; but although it may be true, very broadly speaking,

“that types which have a wide range in time have also a wide

range in space,” there are probably no species in the world to

which this remark applies with less force than it does to those

inhabiting subarctic latitudes. For it is known that in eom-

paratively recent geological times subarctic regions formed a

continuous tract of land with similar physical conditions, offering

no barrier to the eastward or westward migration of the indigenous

species adapted to the surroundings,

The antlers of Mr. Cameron’s third group (Cariacus, Capreolus,

Elaphurus) are derivable, using his own words, from alleys

consisting of a ‘ forked beam with equal or subequal prongs

the Neotropical Guemal (Hippocamelus) being cited as an illus:

tration; while the antlers of his fourth group (Cervus, Cervulus)

are derivable from an “unbranched beam with a true brow-

tine.”

The terminology here employed is apt to confuse the issue.

It suggests a fundamental difference between the two types of

antlers described, whereas, on the face of it, it appears that

Mr. Cameron is describing in different words structures which

are to all intents and purposes identical. An ‘ unbranched beam

with a brow-tine” and a “forked beam” are alike describable

as biramous antlers consisting of an anterior and a posterior

prong arising from a common base, the anterior branch of the

“forked baat? corresponding to the br ow-tine” and its posterior

branch to the so-called “ beam.” Until this suggested homology

he disproved, Mr. Cameron’s assumption of the fundamental

diversity of the two types of antlers cannot be regarded otherwise

1910.} CUTANEOUS SCENT-GLANDS OF RUMINANTS. 969

than as an interesting speculation ; and his conclusions are open

to the criticism of being based upon premises which cannot, on

the evidence, be accepted as firmly established.

Looking at the antlers of an adult Red Deer, Wapiti, Bara-

singha, Thameng, or Japanese Deer, it seems absurd to claim

that the “brow-tine” is equivalent to the so-called “ beam”

which constitutes the rest of the antler. It appears merely to

be the lowest of the series of branches which arise from the beam

and to be comparable to those that are above it. The accepted

terminology of the tines bears out this interpretation. | believe,

nevertheless, that the ‘“ brow-tine ” and the “ beam” are respec-

tively the anterior and posterior branches of an originally sub-

equally biramous or dichotomous antler, like that, for instance,

of the Guemal (/ippocamelus), and that their real equivalence in

the Deer above-named is obscured by the immense development

and complex armature of the posterior branch. It is this growth

of the posterior branch at the expense of the anterior which

characterises the antlers of most of the Deer of the Old World.

A marked approach to this inequality is seen in the Reindeer,

but in this genus the fundamental equivalence between the two

branches is more apparent. Limitations to the growth of the

anterior branch are imposed probably by necessities of feeding.

If the anterior branch projected beyond the muzzle, a Stag could

not feed with freedom off the ground; and even if it were pro-

longed obliquely upwards over the face, it would be a hindrance

to browsing. No such limitations exist in the case of the posterior

branch.

Support for the hypothesis that the antlers of the so-called

Klaphine group were originally biramous like those of the

Guemal (Hippocamelus), and that the biramous antler was pre-

ceded in evolutionary development by an unbranched antler

such as is retained in J/azama, is supplied by the annual growth

of an individual antler. The antler first appears as an undivided

bud corresponding, I conceive, to the unbranched antler of

Mazama. This bud then gives off an anterior and a posterior

branch. For a longer or a shorter time these grow with equal

rapidity, forming a biramous antler which in species with the

anterior branch inclined upwards, as in the Axis, Sambar, or

Japanese Deer, is shaped like a short-stalked Y, resembling, in

fact, the permanent antler of the adult Guemal (//ippocamelus).

These stages are not equally evident in all Stags. In Cervus

itself, for example, the early appearance of the “ bez ”-tine makes

the biramous appearance of very short duration; but in the

Thameng or Panolia Deer (C. eldi) the anterior and posterior

branches grow at the same rate until the anterior branch has

reached nearly its full size. Thereafter the posterior branch

rapidly surpasses it and, increasing in length, develops its supple-

mentary tines.

If this view of the fundamental resemblance underlying

the antlers of all Deer be correct, it minimises the difficulty of

970 MR. R. I, POCOCK ON THE [June 14,

classifying HLlaphurus with the Elaphine Deer, where Brooke,

in my opinion, correctly placed it. For, according to this reading

of the facts, the antlers of Llaphurus have not passed far beyond the

biramous stage, seen transitorily but quite clearly in the growing

antlers of such Deer as Rusa, C. (Panolia) eldi, and others. The

peculiarities of the antlers of Llaphurus consist in the lengthening

of the basal portion, the lengthening and bifurcation of the

anterior branch which concomitantly has a strong upward trend

so as to clear the face, and development of the posterior branch

into a long slender usually undivided tine.

I suggest this homology of the anterior and posterior branches

of the antlers of Elaphurus with the so-called “ brow-tine ” and

“beam” of the antlers of the Elaphine group, because this Stag has

evidently been the stumbling-block in the way of the acceptance

of Brooke’s classification both to Mr. Cameron and Mr. Lydekker.

The latter, indeed, writes strongly on the point. He says: “If

antlers count for anything in classification ... the genus

[ Elaphurus] has nothing to do with any of the living Old World

Deer with the exception of the Roes, while its alliance with the

American Deer seems close” (‘ Deer of all Lands,’ p. 234). Of

course, antlers count something in classification; but whether

they count as much as Mr, Cameron and Mr. Lydekker believe is

quite another matter. In my opinion they do not; and I think

the resemblance between Llaphurus and the American Deer, or

at all events some of them, consists in the fact that the antlers in

both have been specialised toa comparatively small extent beyond

the biramous stage, which was, I believe, common to all Deer

after the one- pronged or mazamine stage was passed. I do not

think it can be claimed that such a resemblance is strong evidence

of affinity.

Adopting, then, as a basis for the classification of the

Cervide the skeletal characters pointed out by Mr. Garrod and

Sir Victor Brooke, I give the subjoined analytical key of what

appear to me to be unquestionably valid genera tested by the feet

and glands alone. Of course, I speak on this point with reserve

as regards the American forms like Hippocamelus, Llastocerus, and

Jucervus, Which I have not seen, Very possibly there are more

genera to come into his section than IT have admitted. Of the

Deer of the Old World I am more confident. It must be remem-

bered, however, that the genera of this family have been established

for the most part upon characters other than those forming

the subject-matter of this paper, the branching and mode of

growth of the antlers being the principal one. i have not in all

cases made use of these characters because they are well known

and have been repeatedly described.

In the preceding pages I have, without prejudice, adopted such

names as //yelaphus, Rucervus, Panolia, and Pseudaxis as con-

noting groups of subgenerie rank. I have, however, no confidence

in the permanent admission of any of them as subgenera. In

the future they will probably be granted full generic status; but

this course or that of regarding them as subgenera or as synonyms

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 971

of Awis and Cervus will depend upon the views of each individual

taxonomist. The point I wish to emphasise is that the characters

on which they were established have not, in my opinion, so high

a systematic value as those to which full generic importance is

here attached. JI do net, for example, think that in the future

' Dama will be set down as a subgenus of Cervus, whatever course

may be adepted with regard to Rucervus, Panolia, or Pseudamis.

a. Distal portion of lateral metacarpals persistent, interungual

integument, where known, thickly hairy. (Telemetacarpalia.). CAPREOLINE

6. Naviculo-cubeid and external cuneiform benes of tarsus united ;

no pedal, tarsal or metatarsal glands ............0..ceeees ese eon eee Pudu.

6°. Naviculo-cuboid and external cuneiform bones of tarsus

separated ; pedal glands present at least on the hind feet

(2 in Rucervus, Blastocerus, Hippocamelus) ; tarsal or meta-

tarsal gland, sometimes both, present.

e. Vomer dividing the posterior narial aperture.

ad, Rhinarium large ; hoofs narrower, tied together posteriorly

or throughout their posterior half ; pasterns not depressed

behind.

e. Metatarsal gland very long... .:........001-- ees severe reese ss DUCE DUS.

Metatarsal : gland nor mal or absent.

“3 Metatarsal gland present; a large pocket-like pedal

gland on the front as well as on the hind foot ...... Dereelaphus.

J’. Metatarsal gland absent; at most a small glandular

pocket en the tront feet (? in Blastocerus and Hip-

pocamelus).

g. Preorbital elands moderately large; antlers branched.

Blastocerus, Hippocamelu

g’. Preorbital glands small; antlers unbranched ...... Mazawa.

. No rhinarium, muzzle hairy ; hoofs broad, widely sepa-

rableat the heels, owing to the anterior position of the

web ; pedal glands on the hind feet only, and consisting,

as in Mazama, Dercelaphus, and Capreclus, of a deep

pocket opening high up the Het ee ; tarsal and preorbital

glands large ...... BIL Rota e. pean oe Deane g els

ec’. Vomer not dividing the posterior nares.

A. Pedal gland either a deep and long interdigital cleft or

a large pouch with constricted orifice on the hind foot ;

rhinarium large ; no tarsal gland.

é. Pedal gland a large pouch, with constricted erifice

opening high up the pastern; preorbital gland absent;

metatarsal | gland present... Capreolus,.

. Pedal gland a deep interdigital. cleft, ‘the hoofs united

at the heels by a fold of integument ; preorbital

gland present ; metatarsal gland absent ....... Hydvropotes.

. Pedal glanda comparatively small inv agination situated

as in Caprealus and Rangifer ; rhinarium very small;

tarsal and preorbital glands present ; metatarsal eland

Small Gm AWSenbaerecc =! Magee cace tence canes gseaeciecle acer vegas Alce.

a’, Proximal portion of lateral metacarpals Bears or occasion-

ally wanting. (Plesienetacar palia.) .. : CERVIN&.

k. Tarsus with naviculo-cuboid and cuneiform bones united ; pedal

gland a deep clett on the hind feet; hoofs united only at the

heels ....... ; Cervulus, Elaphodus.

k’. Tarsus with ‘naviculo-cuboid and cuneiform bones separated.

1. Hoots of hind feet united only at the heels by a close fold of

skin; the gland a long and deep hair-lined interdigital

cleft as in Cervulus; a moderately deep cleft also on the

pastern of the front foot . . Dama.

. Hoofs of hind feet united almost. throughout their basal

depth by a deep interungual web ; no deep cleft on pastern

of front foot. ;

m. Pedal gland a Jong and moderately deep cleft on the

pastern of the hind feet; metatarsal gland naked, over-

Jappeds bye DIALOUALS aU Seteece mike) ecincechcuscedsavenssnterrnAsbees

972 MR. R, I, POCOCK ON THE [June 14

. No specialised gland or deep cleft on front or hind feet ;

metatarsal gland overgrown with hairs, occasionally

absent ; interungual integument naked.

n. Rhinarium extending some distance beneath the nostrils,

as in Avis; tail long.

o. Tail bushy; antlers with short base, short undivided

anterior branch and Jong, stout, two-pronged posterior

branch.. * Ses a uaa:

o’. Tail tufted; antlers with long base, ‘Jong, usually

two-pronged anterior branch, and long, slender,

usually undivided posterior branch ...... . Elaphurus.

. Rhinarium extending only a short distance below the

nostril; tail (except in Pseudawis) quite short ......... Cervus.

Family CAMELIDA.

(Text-fig. 142, A, p. 981.)

According to Ogilby, Camels and Llamas have neither preorbital,

inguinal, nor interdigital glands,

T have had no opportunity of examining fresh examples of

either species of Camelus, but, according to Max Tempel, C. drome-

darius has specialised glands in the interdigital cleft of both the

front and hind feet. In the foot of a Vicuna (Lama vieugna)

that died in the Gardens I found the pedal glands to be as well

developed as in some Cervidee, in the sense that the integument

of the interdigital cleft secreted a strong-smelling substance.

The hairs on the front of the pastern passed about halfway down

the walls of the cleft, leaving the deeper half quite naked. In

the specimen examined this naked area was covered with brown

sticky substance smelling exactly like the urine of JZws musculus.

The depth of this cleft is due to the fact that the integument of

the back of the pastern, which extends to a point halfway between

the heel and the nail of the hoof, is closely folded back upon

itself, forming a shallow but long interdigital web.

The peculiarity of the foot of the Llama as compared with that

of the Bovidee is the small size of the nail and the length of the

sole and of the heel as compared with it. The apex and lower

edges of the nail, however, extend for a short distance on to the

plantar aspect of the foot.

From the remark in Flower and Lydekker’s ‘ Mammalia’ (p. 295)

that there is “a broad cutaneous pad, forming the sole of the

foot, on which the animal rests in walking, aie of on the

hoofs,” one might be led to infer that the manner of walking in

the Camelide differs essentially from that of other Ruminant

Ungulates in that the latter walk upon the nails and the former

upon a cutaneous pad behind the nail. This is not the case. All

the ruminant Ungulates the feet of which I have examined,

with the single exception of the Klipspringer (Oreotragus), wi: alk

upon the cutaneous pad forming the sole and heel of the hoof,

and upon more or less of the inferior edge and apex of the nail in

front. The Camels form no exception to this rule, the only

difference being that the small nail does not invade the area of

the sole to anything like the same extent, and that the sole and

1910.| CUTANEOUS SCENT-GLANDS OF RUMINAN'S. 973

the heel are continued farther backwards. In other words, it is

only a difference in the degree of development of the parts

composing the distal portion of the foot.

As is well known, the plantar area of the hoof in late foetal

and many, perhaps all, newly born Cervide and Bovide is quite

soft right up to and in some cases beyond the apex of the nail.

This terminal portion soon shrivels and hardens, and becomes, in

most cases at all events, indistinguishable from the inferior

portions of the nail, especially externally ; but usually the line of

demarcation between the nail and the sole can be quite easily

detected, especially on the inner aspect of the hoof, showing that

the sole, although hard and horny, still persists and forms with

the heels the main area of support during progression. The one

exception to this rule known to me is the Klipspringer, which

appears to support itself entirely upon the truncated apex of the

nail of the hoof, with possibly a small piece of the hardened sole

wedged in the angle formed by the two sides of the nail.

The males both of Camelus dromedarius and of C. bactrianus

have a pair of well-developed glands on the summit of the neck

just behind the occiput. In the living animal these may be easily

felt as a pair of oval or roundish lumps lying close together. The

hair overlying them has no underwool. In a male of (C. drome-

darius I have seen black watery secretion running from these

glands and dropping off the hairs lke sweat. This was in the

month of June, when the glands of a male C. bactrianus were

quite inactive. In the latter animal the glands were active and

giving off a powerful repulsive odour much earlier in the year—

that is to say, in March. The smell of the secretion from the

glands in the Dromedary was equally repulsive, but reminded me

of no substance in particular. It stains the skin of the fingers in

much the same way as walnut-juice does. The position of the

glands in the Dromedary is easily seen by the post-occipital patch

of black hair. I have had no opportunity of observing the gland

in the female Dromedary ; but in the female Bactrian ‘Camel it is

much smaller than in the male, and never apparently noticeably

active as it is in the latter when rutting.

Function of the Glands.

I do not know who first propounded the theory that the use

of the cutaneous scent-glands in Ruminants is to keep the indi-

viduals of a species together. As early as 1836 Owen discussed

this view ; but, finding that the glands were present in some

species of Antelopes of solitar ‘y habits and absent in some gre-

garious forms, he dropped this hypothesis as untenable. He does

not appear, however, to have attached sufficient importance to

the mutual need of discovering each other on the part of the

sexes of solitary species at the breeding-season, nor to the

necessity of the mother and young being together, at all events

974 MR. R, I, POCOCK ON TIE [June 14,

periodically, in the course of the day. A hind, for example, which

leaves her newly dropped fawn in the jungle or bush while she is

away feeding, must have some means of retracing her steps to the

spot; and it may well be that she does so by means of the scent

her own foot-tracks have left.

This obvious criticism of Owen’s reasons for rejecting the

theory has no doubt suggested itself to others; for the view that

the secretion of the scent-glands serves to guide individuals of a

species, whether gregarious or not, to one another appears to be

commonly held, and was clearly expressed by Flower and Lydekker

in their volume on the ‘Mammalia’ (pp. 315-316), That the

secretion from the pedal glands must scent the ground over

which the animal passes seems clear, for the pouch is commonly

furnished with long hairs projecting from the orifice or the edge of

the orifice, and the: function of these hairs seems to be to guide the

secretion downwards between the hoofs. Similarly the substance

secreted by the tarsal, metatarsal, and carpal glands would touch

against higher herbage, and might leave a taint behind. But

the position of these glands suggests that their function is also

to indicate where the animal has been lying. In the habitual

resting attitude of Ruminants, the area below the knee (carpus)

on both the front legs is pressed to the ground, and in the case

of the Cervidie the metatarsal gland on the outside of one hind

leg and the tarsal gland on the inner side of the other in the

same way come simultaneously into contact with the soil whether

it be bare or overgrown with vegetation. Now the Deer which

possess these glands on the hind legs have no inguinal pits or

knee glands. Conversely the Bovide, which have no tarsal or

metatarsal glands, not uncommonly possess inguinal pits and

sometimes knee glands. The suggestion, tiererore e, is obvious

that the inguinal pits of Sheep and many Antelopes, so-called,

represent functionally the tarsal and metatarsal glands of the Deer

and scent the ground where the animal has been lying.

It is clear that the above proposed explanation of the use of

the inguinal and crural glands applies equally well to species

which inhabit jungle, plain, sandy desert, or barren mountain

heights. It is therefore more satisfactory than the view that the

scent of the secretion is only left behind on the vegetation by the

animal’s passage through it. Sheep, for example, which possess

inguinal glands, frequent hilly country, feeding on grass often not

high enough to reach the inguinal region. Except, therefore,

when these animals are in a state of re pose the secretion of these

glands can seldom be left behind adhering to natural objects.

Tt is also likely that one individual can Paltons another or detect

his whereabouts by the wafting of the scent through the air,

irrespective of contact with the ground. This wafting of scent

is probably the sole function of the caudal glands of Goats (Capra),

which can in no circumstances dispose of “their secretion by con-

tact with the soil, unless a certain amount of it falls to the ground

sticking to the feces, Be it noted, too, that the habit of turning

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 975

up the tail in flight, so common in Ruminants, would expose the

secretion of these glands to the air.

It has also been suggested, by Owen for example, that the use

of the secretion of the pedal glands is lubrication of the hoofs.

This is possible. The suggestion, however, does not exclude the

explanation advocated above; and it appears to me to be probable

that the function of the secretion of the pedal glands is the same

as that of the carpal, tarsal, and metatarsal glands, which cannot

be for lubrication. In this connection it is significant that

pyceros and Tetraceros, which have no pedal g glands, have very

specialised glands on the hind legs, occurring in the former above

and in the latter within the false hoofs, in situations, that is to

say, where the secretion must taint the ground where the animal

lies.

There is a good deal of evidence that the preorbital gland has a

sexual significance, at all events in part. Bennett, for example,

noticed that an immature example of Antilope cervicapra had the

lips of the sinus small and closely applied, so as to hide entirely

the internal lining of the sac. In a full-grown animal, on the

contrary, the sac, under the influence of excitement, was everted

so as to form a projection rather than a hollow. At such times

the animal delighted to rub the sac against any substance offered

to it, loading it with the secretion, which had a slightly urinous

odour. Ina castrated individual, believed to be about the same

‘age as the last, the gland, on the contrary, was only developed to

the same extent as in the young male and was never everted as in

the normal adult buck (P. Z.S. 1836, pp. 35-36). Hodgson also

believed that in the Serow (Capricornis thar) the gland was “ con-

nected with the generative organs”; for he noticed that, in the

spring especially, a thin viscid secretion was constantly poured out

from the sinus, and he suggested that this profuse secretion was a

means of relieving the animal during the time of extraordinary

excitement to w Bich it is liable in Aye breeding-season (P.Z.S.

1836, p. 39). Max Weber, moreover, has collected a good deal

of evidence showing that the preor bital gland is better developed

in the male than in the female of the many species of Antelope. I

have noticed this myself in the case of Raphicerus campestris and

Tetraceros quadricoriis. I have also observed the glands of the

latter species enlarge and discharge copiously in the summer, and

those of specimens of Gazella rwfifrons and Cephalophus dorsalis

to be markedly swollen at that season. I have also seen them

discharging actively in July in a female example of Sylvicapra

grimmt, and to be quite inactive in the same animal in August.

Moreover, in some Deer (Pseudaxis and Rucervus) I have noticed—

and this is possibly true of all Deer—that the preorbital gland is

quite small and shallow in the young as compared with the

adult, a fact warranting the conclusion that this gland attains its

greatest size and activity with sexual maturity.

But I cannot, on the evidence, bring my self to believe that the

significance of the preorbital gland is w vholly sexual, It is certain,

976 MR. R. I, POCOCK ON THE [June 14,

for example, that the gland is in some cases under the influence

of other emotions than that of sexual excitement. A male Black-

buck, for instance, advancing upon a man with hostile intent, turns

up his tail, and everts the gland to its fullest extent ; and I have

seen a Sambar stag with ears pricked, nostrils dilated, and gland

completely everted, gazing intently at a cat crossing the yard, his

whole attitude forcibly suggestive of curiosity and doubt with a

strong admixture of fear.

The habit of rubbing the secretion of the preorbital gland

against the bars of cages or other objects was recorded by Bennett,

of Antilope cervicapra, by Ogilby * of a species of Gazelle, and by

Mr. Beddard t+ of Oreotragus. Ogilby, indeed, speaks of this act

as frequent. In my experience it is uncommon. I have only

noticed such behaviour in one Antelope, an immature example of

Maxwell’s Duiker, which invariably rubbed its gland against my

proffered hand, exactly as a cat rubs its mouth and cheek. Con-

sidering the immaturity of the specimen in question, I cannot

believe een sexual excitement was in any sense the cause of the

action. It may therefore be the case, as Ogilby suggested, that

“rubbing the crumens against the shrubs. or stones of their

desert and mountain habitats ... may serve to direct [ Antelopes]

in their wanderings and migrations, when the storms and fogs

incident to such places obscure all visible landmarks.” As

guide-marks, therefore, appeal to the sense of sight and enable

the members of a species to follow one another and kee

together when vision is unobstructed, so, it is believed, do the

scent-glands described in this paper serve the same purpose by

appealing to the sense of smell when, for any reason, free vision

is prohibited.

It would be idle, however, to pretend that this hypothesis

gives a full explanation of the use of the glands. It fails, for

example, to account for the fact that in many, perhaps all, cases

each gland emits its own peculiar scent, as recorded, for instance,

of a Muntjae (Cervulus) and of a Gazelle (G. dorcas). That

each scent tells its own tale to other members of the species, it is

almost impossible to doubt; but what that tale may be, I am

wholly unable to surmise,

Origin and Evolution of the Glands.

The preorbital glands in all the Cervide, and in the majority of

Bovidee that possess them, consist of an integumental thickening

forming a larger or smaller invaginated sac, which serves to store

the secretion. But in a few of the Bov idee, like the Duikers

(Cephalophus), the Gnus (Connochetes and Gor gon), Buffon’s Kob

(Adenota kob), and the Sable Antelope (//ippotragus niger), there

is no invagination, the secretion making its way directly to the

exposed surface of the skin. The simplest type of preorbital

* P. Z.S. 1840, p. 9.

inc Mammalia,” The Cambridge Natural History, p. 13 (1902),

1910. | CUTANEOUS SCENT-GLANDS OF RUMINANTS. 977

gland known to me is found in Buffon’s Kob (Adenota kob) and

the Sable Antelope (/ippotragus niger). The skin of the

secreting area is merely thickened and retains its normal hairy

clothing. The hairs, however, are long, forming a tuft or mat

and are stuck together or hold the secretion like a scurfy powder,

It is a natural inference that all the preorbital glands, however

specialised, arose from simple integumental thickenings, with

enlarged sebaceous and sweat-glands, resembling in essence that

found in the Sable Antelope.

Exactly similar to this and equally simple in structure are the

carpal or knee-glands of some Antelopes, like Gazelles and their

allies, and of some Sheep, the tarsal or hock-glands of the Tele-

metacarpal Cervide, and also the metatarsal glands of Deer

belonging to both the divisions of this family. il some species

of Deer (: dwis avis and porcinus) the glandular area itself is naked

and is overlapped by a fringe of long hairs surrounding it.

A further stage in the evolution of this kind of leg-gland is

found in the Pallah (dpyceros), where the glandular ‘layer is

restricted to the periphery of the area, forming an elevated ridge

round it, the secretion accumulating on the smooth skin thus

circumscribed.

In connection with the inguinal glands it is noticeable that

they are intimately connected with the mamme. In the majority

of cases their orifices are situated close to those organs, and the

deepest part of the pocket actually overlies them; and when, as

in Pantholops *, the orifice is remote from the mamma on each

side, the invagination is elongated so as to extend inwards as far

as the area overlying the mamma. Again, two pairs of inguinal

pouches are found only i in species which have two pairs of functional

mamme (Zragelaphus, Redunca).

These facts suggest that the secretion found in the inguinal

sacs 1s derived actually from the mammary glands. This I

believe to be the case; and the view is corroborated to a certain

extent by the fact that the secretion in the sacs not uncommonly

has an unmistakable odour of cream-cheese, or sour milk. The

presence of inguinal glands in both sexes cannot be alleged

as an objection to this supposition; for mammary glands are

found in males as well as in females, although they are normally

inoperative in the former.

It seems to me that for the origin of the pedal glands we must

look to one of two processes : that is to say, they have either arisen

as invaginations of the interdigital integument of a more or less

fully webbed compact foot, such as is seen in Bos, Tuurotragus,

Pantholops, dipyceros, and others; or they are the remnants,

larger or smaller according to the genus, of a primitive inter-

digital space.

The former explanation appears at first sight to be the more

probable; and from the remark made by Flower as to the

* But not in immature examples of Sylvicapra.

978 MR. R. I, POCOCK ON THE [June 14,

seemingly primitive nature of this gland in Pudu, I infer that

this was the view he held. If this be so, the shallow or short

pocket seen in such genera, for example, as Alce, Redunca, and

Pseudois preceded, in an evolutionary sense, the well-developed

gland possessed respectively by such genera as Capreolus, Pelea,

and Ovis. Further, it will follow as a possibility, perhaps as a

probability, that the pedal glands have been developed inde-

pendently more than once in various groups of Bovide and

Cervide ; and that the genera within these families characterised

by pasterns without any ; interdigits il space have an earlier form of

foot than those in which such a cleft is found : that the hind foot

of Cervus, for instance, to take a concrete case, is more primitive

than that of Dama.

I believe this view to be erroneous; for everything we knowof the

evolution of the feet in Ruminantia points to the conclusion that

their structural peculiarities are due to descent from four-toed feet

with the phalanges separated to a greater or less extent. Gradually

the lateral digits became shortened, raised from the ground, and

practically functionless so far as affording support to the body was

concerned. The weight was borne by the two middle toes, which

became gradually strengthened for the purpose by the extension

of the integument between them. Now when Mammalia of other

groups develop an interdigital integument or “ web,” this web

always, I believe, extends along the posterior or inferior aspect of

the toes, leaving more or less of a cleft between them in front or

above. ‘This is well shown in such formsas Zudtra and the Canidee

and in various amphibious genera of Rodents and of other orders.

For these reasons I believe that the earliest type of Ruminant

foot is that in which the interdigital web consists of a fold of

integument extending along the back of the pastern only, leaving

a wide and deep depression between the bones above the hoofs in

front. But when two surfaces of skin are opposed to form a

cavity or crease, conditions favouring the activity of the skin-

glands, or at all events conditions preventing the evaporation of

integument: al secretions, are set up. Hence, in a foot with a

posterior web holding the digits together behind and a cleft

between them in front, we have conditions conducive to the

development of a scent-gland ; and I conceive that it was from a

foot of this structure, containing a potential pedal gland, that the

various types of feet of the Ruminantia, whether containing

glands ov not, have been derived, and that the glandular pockets,

whether large or small, are the remnants, specialised for the

secretion of scented material, of a primitive anterior interdigital

cleft.

This conception seems to me to supply the most satisfactory

explanation of the prevalence and diversity of the pedal glands of

the Ruminantia, all the glands being thus traceable to a common

origin.

Of existing forms of this group the most primitive type of foot

is found in the Tragulina, itself a primitive group combining

1910. ] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 979

characters of the Suid, the Camelide, and the Cervide, In

specimens of Zragulus stanleyanus (or an allied form) and of

7’. meminna, which lived in the Gardens, I find that the web on

the posterior side of the pastern consists of a fold of skin which

stops short at a point about halfway between the false hoofs and

the heels; the back of the pastern is open in its lower half, and

its front is marked with a long triangular depression, the apex of

which extends up to the level of the false hoofs.

Text-fig. 141,

A. Median vertical section of foot of Tragulus meminna.

J. Anterior aspect of foot of 7. stanleyanus.

C. Median vertical section of foot of Dicotyles.

D. The same of Phacocharus.

These feet are the only ones known to me amongst the

Ruminantia in which the web ceases behind far above the heels.

The inner walls of the two digits of the pastern are clothed with

hair, which is short and scanty in 7’. stanleyanus, longer and less

980 MR. R. I. POCOCK ON THE [June l4,

scanty in 7. meminna*; but I could detect no trace of any

secretion indicating special activity of the skin-glands of the

surfaces in question.

In the Camelide, as exemplified by Lama vicugna, the foot is

more specialised with respect to the extension of the web to a

point halfway between the heel and the nail of the hoof ; but the

anterior depression on the pastern is very deep and long and

glandular in its deeper portion, and the nail is small, (Text-fig.

142, A.)

Still more specialised is the foot in the Suide; for although

there is a deeper or shallower depression on the front of the

pastern, admitting of considerable distension of the hoof, the

anterior and posterior walls of the pastern are widely separated

and there is a strong deep interungual union extending forwards

from the heels to a greater or less extent according to the

species. In a Peecary (Dicotyles collaris) the heels were separable

and the interungual union shorter and the anterior depression

deeper than in a Wart-Hog (Phacochwrus), in which the depression

was quite shallow and the interungual union long in comparison

(text-fig. 142, B,D). In the foot of the Wart-Hog, moreover,

which was kindly given to me by Rowland Ward, there was an actual

horny union between the heels, which were thus quite inseparable.

In both these species the interungual area was quite smooth ;

but I detected no trace of specialised glands f.

Reverting once more to the Ruminantia, I find, as is not sur-

prising, that the foot of the Musk-Deer (JZoschus) comes nearest to

that of Zragulus. The pastern-web of J/oschus nearly reaches

the heels of the hoofs but not quite; the feet indeed, apart from

the wealth of hair that clothes them, are like those of 7'ragulus,

but very much shortened and having the glands of the depression

active.

It is only a short step from the foot of MJoschus to the hind

foot of Dama or Klaphodus, the chief difference being that in

these Deer the pastern-web extends right down to the heels, tying

* In 7. stanleyanus also the skin of the back of the pastern is coriaceous and

visible through the hairs, which are as scanty as on the inner surface of this part ;

whereas in 7’. meminna the skin is invisible on the back of the pastern when the

hairs are undisturbed. I have not investigated the submandibular glands of

Tragulus.

+ The secretion of the dorsal gland in both species of Peccary has a strong and

pungent odour, like concentrated human perspiration. It may not be without

significance that the scent of the South American Tree Poreupines (Coendu villosus

aud prehensilis) is exactly the same, as I have already recorded (P. Z. S. 1908, p. 946).

And since this scent has, I believe, a warning significance in the Porcupines, it may

be that we have here an instance of mimicry, Batesian or Miillerian, in smell. On

the other hand, the function of the gland in the Peccary may serve the purpose

merely of keeping the members of the herd together. Owen’s figure and description

of this dorsal gland in the Peecary are not very satisfactory. He describes and

figures the gland as opening by an orifice in the mid-line of the back, without

mentioning any nipple (‘Anatomy of Vertebrates,’ iii. p. 635, 1868). Possibly his

preparation was not made from fresh material, for in the specimens I have

exuinined the gland was marked externally by a conspicuous upstanding nipple.

This was traversed by a central duct dipping into the lobular fat-like tissue of the

gland. When the gland was squeezed, fluid secretion could be easily made to exude

from the orifice at the summit of the nipple, (Text-fig. 142, C.)

1910. ] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 981

them tightly together; and the upper portion of the*interdigital

cleft in front is deepened, showing the beginning of the differ-

entiation of the glandular pouch. Hardly more advanced is the

hind foot of the Muntjacs of the genus Cervulus, except that in

C. muntjac part of the glandular area is markedly more pocket-

like. A further stage is exemplified by the hind foot of the Hog-

Deer (A. porcinus) and of the Axis Deer (A. axis), where the web

Text-fig. 142.

A. Median vertical section of foot of Lama vicugna ; gl., glandular area.

B. Anterior aspect of foot of Dicotyles collaris.

C. Median vertical section of dorsal gland of the same.

D. Anterior aspect of foot of Phacocherus.

extends further forwards from the heels, holding the hoofs more

securely and tightly together, the interdigital cleft being shallower

owing to the greater space between the anterior and posterior

walls of the pastern-web. It is quite easy to imagine the steps

by which the comparatively narrow pocket-like gland of Mazama,

Cervulus, and Rangifer has been derived from a much longer and

Proc. Zoou. Soc.—1910, No. LXTIV. 64

982 MR, R. I, POCOCK ON THE [June 14,

shallower glandular depression such as is seen in A. axis and

A. porcinus; and also on the other side how by the gradual

shallowing of the depression in the feet of these last-named species

the condition of things found in the feet of the Red Deer (C.

elaphus) and its allies has been reached.

T do not by any means wish to suggest that the species named

stand in the exact line of descent to one another; but I do think

it probable that their pedal glands represent approximately the

stages through which the various types of pedal glands met with

in the Cervide have passed in their evolution.

Similar stages in the gradual obliteration of the interdigital space

accompanied by the evolution of differentiated pedal glands and

their final suppression may be traced in the Bovidee., In Madoqua,

Raphicerus, and Cephalophus, for instance, the digits of the pastern

are held together by a long and close fold of integument constituting

a web which extends down or nearly down to the heels; the anterior

skin of this fold forms the posterior wall of the glandular inter-

digital cleft, as in Cervulus and Dama. This cleft is at first open

below between the hoofs and remains so in Madoqua and Cepha-

lophus. But in both these genera, as well as in Raphicerus and

Dorcotragus, the orifice of the cleft is shortened toa greater or less

extent, according to the species, by the downward extension

towards the hoofs of the folded integument of the anterior

surface of the pastern, forming a longer or shorter anterior wall

to the enclosed glandular space. Another simple modification is

seen in the Gazelles, in which, while the orifice of the cleft retains

almost unchanged in its original length, its inferior portion is

closed and shut off from the interungual space by the anterior

extension of the posterior web from the heels to the front of the

hoof. Practically the same structural change has taken place in

the Oribi (Owrebia) and in the Hartebeests (Bubalis, Damaliscus),

except that in Bubalis the anterior web overlaps the cavity to a

considerable extent above and in front.

The adult White-tailed Gnu (Connochetes gnu) exhibits a

further stage in the evolution of the gland seen in the Hartebeest,

the orifice of the cleft being still more shortened and the posterior

integumental web not so closely folded. A further stage in the

closing up of the orifice of the cleft is exemplified by the feet of

Oryx beisa; and this is followed by the reduction of the orifice

toa small circular aperture as in Hippotragus, Pelea, Nemorhedus,

and Ovis; the final stage, sometimes certainly and always probably

of sporadic occurrence within the limits of a species, being the

retention of merely the orifice and part of the duct of the gland

as in Redunca, Capra, Hemitragus, and Pseudois.

If this be the correct view of the evolution of the feet, there must

be some reason or reasons for the disappearance of interdigital

clefts. It seems certain that large interdigital clefts must be a

source of weakness to the feet. Although a foot with its two

component digits united bya close fold of integument extending

from the heels up the back of the pastern is more suitable, on

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 983

account of its distensibility, for progression over soft and yielding

ground than a foot with its component digits united by skin

extending forwards from the heels to the front of the hoofs and

restricting their separation, it is obviously less well adapted for

Supporting heavy weight, for speedy propulsion over hard

unyielding ground, and for resisting the impact of a downward

leap on a mountain side, on account of the liability to rupture of

the integument under the strain put upon it.

Strengthening the feet, then, has probably been one of the

factors, perhaps the principal factor, that has operated in favour

of the reduction in size or total disappearance of the interdigital

clefts. This theory helps to an understanding of the fact that,

broadly speaking, large and heavily built Ruminants have small

interdigital clefts or none, and that large interdigital clefts are for

the most part found only in small animals. There are several

marked exceptions to this, however, and no doubt other factors

than the one suggested have influenced the formation of the feet,

such as the use or uselessness of pedal scent-glands. For example,

large Ruminants are much more easily kept in view by members

of their own species than small ones; or, if they live in thick bush,

are more easily followed by hearing as they crash away in a

‘State of panic through the vegetation. Of these and other matters,

however, we shall never have full knowledge until our information

is much more exact than it is at present with respect to the senses

and to the bionomics of the species concerned.

In the light of the above-made suggestions, the following con-

siderations are worth bearing in mind :—

1. Giraffes are large, heavily built animals living in open wood-

land country and gallop with speed over rough uneven ground.

The feet are compactly built and without interdigital clefts.

‘The great height of Giraffes gives them considerable range of vision,

enabling stragglers to keep with the rest of the herd. They can,

moreover, only with difficulty and delay reach the ground with

their nostrils. Hence specialised pedal scent-glands would be of

little use to them for tracking.

2. All the species of Sos are heavily built, mostly large animals

with thick legs. They live either in the open or in jungle or bush,

and can be followed either by sight or by the sound they make

passing through the foliage. The same applies to the Musk-Ox

and to Taurotragus.

3. The Orygine (Hippotragus, Oryx) are also Antelopes of large

or medium size and considerable weight. The interdigital cleft is

reduced to a gland witha small orifice, which is the only break in

the continuity of the integument of the anterior surface of the

pastern; and the orifice is smaller in the larger Hippotragus than

in the smaller Orya.

4. In the Bubaline, large or medium-sized Antelopes, although

the interdigital cleft is large and highly glandular on the front

feet, it is reduced to a shallow depression on the hind feet. It is

in the hind limbs that the galloping power principally lies; and

984 MR. R. I, POCOCK ON THE [June 14,

some of the Hartebeests, the Sassaby (Damaliscustlunatus), for

example, are extraordinarily fleet of foot.

5. In the Cervicaprine, the larger species of Kobus and Adenota

have no interdigital clefts or glands; only in the comparatively

small and lightly built genus Pelea are the clefts retained in the

form of saccular glands with a small orifice.

6. Allthe Duikers (Cephalophinz) are of small or very small

size and live in the forest on soft yielding ground or in fairly open

country overgrown with scrub. They have large interdigital

clefts and pedal glands, the scent of which is probably useful for

tracking individuals concealed by the bush, through which they

dive away when alarmed.

7. The Neotragine (Raphicerus, Madoqua, Ourebia) are also of

small size and lightly built, and mostly frequent country overgrown

with bush orsecrub. They have large interdigital clefts and glands,

by means of which one individual could silently follow by scent

another hidden from view. Dorcotragus, on the contrary, lives

on rocky hill-sides and has the hoofs shortened to lessen their

spread at the tips, the heel-tie thickened to hold them tightly

together, and the sole and heel modified to form a soft pad to

break the force of a leap and prevent slipping, the interdigital

cleft and gland being at the same time retained.

8. The Klipspringer (Oreotragus), commonly believed to be

related to the Neotragine, although of small size and light weight,

has lost the interdigital clefts. His habitat is rocky crags, where

he leaps surefootedly from point to point, the hoofs although

separable being firmly bound together by the thick interungual

integument.

9, All the Antilopine (Gazella, Antilope, Antidorcas) live in the

plains or more or less open country. They are, on the average, of

small size, although larger than the Neotragine, and of light build,

and depend for safety upon their great speed in the open.

Although the interdigital clefts are large, the hoofs are held

together by a fold of integument running forwards from the heels

and restricting their spread toa greater extent than in the Cepha-

lophine and, on the average, than in the Neotragine.

10. I do not know why Ammodorcas has lost and Lithocranius

has retained the interdigital clefts, unless the explanation may

be sought in the totally different modes of progression of the

two Antelopes, which more than one observer has recorded.

Lithocranius runs with a skulking action, ducking under the

bush, with the head stretched forwards; Ammodorcas, on the

contrary, moves with a bold stepping action, holding tail and

neck erect. This at least suggests that the Dibatag is more

easily kept in view by its companions than the Gerenuk, and has

therefore less need of pedal scent-glands to indicate the direction

taken.

11. In the Caprine the only wild forms which possess well-

developed pedal glands belong to the genus Ovis, the others (Ammo-

tragus, Pseudois, Hemitragus, and Capra) being without them.

The latter differ markedly from Sheep in superior activity and in

1910.] CUTANEOUS SCENT-GLANDS OF RUMINANTS. 985

their comparatively greater leaping powers, attributes which are

correlated with a difference of habitat, the Goats and their allies

habitually frequenting precipitous exposed rocky hill-sides or

mountain heights rather than elevated plateaus intersected by

ravines where Sheep are to be found. It seems to me probable

that the ground frequented by Sheep is better adapted by its

vegetation for holding the scent of the foot-glands and perhaps

affords more cover for concealment, requiring tracking by scent,

than the bare exposed situations to which the other genera are

adapted.

12. In the Rupicaprine well-developed foot-glands occur in

Rupicapra, Nemorhedus, and Capricornis. The last two never

seem to be found far from mountain forests, where individuals

may easily lose sight of each other and need scent-glands to keep

together. Although possessed of considerable activity and skill

in covering rough ground at speed, they have not the climbing

or leaping power of Goats. In this respect Chamois appear to

equal Goats, and they have thei feet strengthened by the for-

ward extension of the interungual web, while retaining pedal

glands which are useful perhaps in the mountain forests, where

at certain times of the year Chamois resort. Although not

apparently gifted with special activity or leaping power, Oreamnos

inhabits localities right away from forests and bare of all but

the scantiest vegetation. In this particular, as in the structure of

their feet, they closely resemble Goats. Somewhat of a mountain-

forest dweller like Vamorhedus and Capricornis, Budorcas is never-

theless a comparatively inactive animal of large size and heavy

build, requiring strong feet to carry his weight up and down the

mountain-side.

13. In the Cervide, also, small and medium-sized animals have

on the average larger interdigital clefts and more capacious glands

than large animals. Very deep and long clefts with highly

developed glands are found, for instance, only in small forms like

Hydropotes, Cervulus, and Hlaphodus and in the medium-sized

Dama. Shallower but equally long clefts occur in the small

Hyelaphus and the medium-sized Avis, the much larger Rusa,

which is allied to both these genera, being devoid of them, as also

are all the large Deer of the Old World. In the Telemetacarpal

Deer, setting aside Hydropotes, there is almost always, so far as is

known, a large pouch-like gland in the hind foot in genera

ranging in size from the small J/azama to the large Rangifer, two

exceptions being the diminutive Pudu, in which the glands are

aborted, and the giant Alce, in which the gland is considerably

reduced.

Very little is unfortunately known about the habitat of Pudu

beyond the fact that it has been recorded from Ecuador and

the Chilian Andes and appears to live normally at tolerably high

altitudes. From its compactly built feet, I infer that this little

Deer frequents hard or stony ground requiring firmly welded

hoofs. If this be so, we find a reason for the disappearance of

the glandular pouch in the feet found in its ally Mazama

986 ON THE CUTANEOUS SCENT-GLANDS OF RUMINANTS. [June 14,

if there is any truth in the supposition that such pouches are a

source of weakness.

In the case of Alce the nearly vestigial nature of the pedal

gland is accompanied by two features in this genus which have

an interesting bearing on the suggestions here put forward to

account for the retention or loss of inter digital clefts and glands,

In the first place, this Deer is of great stature and weight, the

latter requiring strong feet for support and the former making

concealment from sight, involving the necessity of tracking by

scent, less likely than in the case of smaller animals, In

the second place, length of limb is correlated with shortness of

neck, so that the Moose can only with difficulty reach the ground

with its nose. Scent-tracks left by the feet, therefore, would be

of comparatively little use in enabling one Moose to find another.

No Deer known to me have such strongly built completely

webbed feet as are found in the large Bovide like Vawrotragus

and Bos, or indeed as in some of the smaller members of that family,

like Zetracerus and Oreotragus. Generally speaking, there is a

ereater tendency towards plantigradism and there is nearly always

a more marked depression on the front of the pastern. This is to

be attributed, I suppose, to the circumstance that Deer are for the

most part denizens of woodland country, woods, jungle, or swamps,

where the soil is usually softish and yielding. This brings us to

the fact, otherwise perhaps seemingly opposed to the suggestion

here made as to the cause conducing to the suppression of the

pedal gland in the hind foot, that a large and heavily built animal

like the Reindeer none the less retains a well-developed glandular

pouch. The Reindeer, however, has a foot eminently adapted for

traversing soft ground, whether it be snow or swamp, and does not

require a foot of such strength as to call for the suppression of

the glandular pouch.

That the Cervide, like the Bovidz, originally possessed inter-

digital clefts and glands on the front as well as on the hind feet is

probable on general grounds, and also from the fact that a gland

is retained on the front foot in a state of greater or lesser

development in some American Deer, while a moderately deep

cleft is present on the front foot of Dama. The only suggestion I

can at present make to account for the greater compactness of the

fore feet as compared with the hind feet is the fact that all

Deer, I believe, both stags and hinds, commonly use their fore feet:

both in attack and defence, dealing blows with great precision,

swiftness, and strength. This I have myself witnessed on more

than one occasion. For such a purpose it seems obvious that

a strongly built foot is required.

These brief comments on the bionomics of Ruminants indicate

in a general way the lines along which research, likely to lead toa

better under standing of the structure of the feet and the reasons

for the retention or loss of the pedal glands, may probably, I

think, be pursued with profit

1910.] ON A WAPITI AND A MUNTJAC. 987

2. On a Wapiti and a Muntjac.

By R. LypreKxer *,

[Received May 4, 1910.]

(Text-figures 143 & 144.)

THe Wapiti (Cervus canadensis wardi, subsp. n.).

A few weeks ago the Rev. W. N. Fergusson, a missionary in

Sze-chuen, returned from China, bringing with him a small

collection of Mammals from that province and Tibet, among

them being a skin of Mlwropus and a pair of antlers of the

Sze-chuen Sambar (Cervis wnicolor dejeant). The collection also

included two shed antlers of an adult Wapiti (text-fig. 143), which

may or may not be a real pair, although from the fact that:

neither is quite symmetrical in form and size with its fellow,

while one is redder than the other, there seems considerable

probability that the two are not naturally associated. These

antlers, I was informed at the outset, came from Tibet, and,

owing to the fact that Wapiti and other antlers (although

generally, if not invariably, in the velvet) constitute an im-

portant article of export from the Altai and elsewhere to China,

IT was naturally suspicious— especially as Wapiti have been

hitherto unknown to exist on the southern side of the Gobi—

that Mr. Fergusson’s specimens might have been imported into

Tibet.

Enquiries were accordingly made from their owner as to the

history of the specimens, to which Mr. Fergusson replied as

follows :—

“ As you are no doubt aware, the native haunts of the Parti-

coloured Bear (Zlwropus) are in the dwarf bamboo and rhodo-

dendron forests so abundant in Sze-chuen at an elevation of from

9000 to 11,000 feet above sea-level. The Deer you identify as

Wapiti inhabit the region just above the tree-line; I have never

shot one myself in these regions, yet I have it on good authority

that they have been obtained. The specimen sent to you I

obtained from a native hunter within fifty miles of the place

where the Parti-coloured Bear was shot. The Deer, of course,

never enter the bamboo-thickets in which the Bear makes its

home, but graze on the grassy plains beyond.”

Although the Deer referred to by Mr. Fergusson may have

been the so-called White Deer or Sze-chuen Hangul described

by myself in the Society's ‘ Proceedings’ for 1909 (p. 588) as

Cervus cashmirianus macneillit, the late Mr. J. W. Brooke

referred to another and apparently larger Deer as inhabiting the

Sze-chuen frontier, and there accordingly seems to be a proba-

bility that the story told by the native hunter to Mr, Fergusson

* Published by permission of the Trustees of the British Museum.

+ Since this paper was written, Major McNeill has informed me that the specimen

was shot on the Tibetan side of the border.

988 MR. R, LYDEKKER ON [June 14,

may be true; the fact that the antlers are shed, and perhaps odd

ones, by no means necessarily discrediting the narrative.

Text-fig. 143.

Antlers of Tibetan Wapiti, Cervus canadensis wardi.

That a representative of the Wapiti should be found on the

mountains to the southward of the Gobi Desert is rendered

probable—or, at all events, not improbable—by the case of the

Argali Sheep (Ovis ammon), whose distribution (if poli and the

other allied forms be regarded as local races) surrounds the Gobi.

1910.] A WAPITI AND A MUNTJAC. 989

Here it is my pleasant duty to record an act of special liberality

on the part of Mr. Rowland Ward, F.Z.S., who, when their

interest was brought to his notice, purchased the two antlers and

presented them to the British Museum.

As is apparent from the text-figure, the antlers are unmistak-

ably those of a Wapiti, and are somewhat inferior in point of

size to the pair surmounting a specimen of the Tian Shan race

exhibited in the Natural History Museum. Satisfactorily to

distinguish some of the races of Wapiti by their antlers alone,

I find an exceedingly difficult, if not actually impossible, task.

Nevertheless, when the antlers forming the subject of this paper

are placed alongside Wapiti antlers from the Altai and Tian

Shan, it will be found that they differ in certain small details

from all; although such slight differences are almost impossible

to describe. One recognisable feature seems, however, to be the

narrowness of the terminal fork, as-compared with that of a

Tian Shan Wapiti. |

If I am right in regarding these antlers as indicating the

existence of a Tibetan Wapiti, it is a practical certainty that this

animal will represent a race by itself; and although I am not at

present in a position to define it, I venture to propose the name

of Cervus canadensis wardi for this presumed new race, in honour

of the donor of the type specimen.

THe Mountsac (Cervulus bridgemani*, Lydekker, Abstract

P, ZS. VOLO, p./ 38):

Passing on to the subject of the second part of this paper, I

have first of all to express my thanks to Lieut. the Hon. R. O. B.

Bridgeman, R.N., F.Z.8., for submitting to me a series of skins and

skulls of Muntjacs from the An-wei district of Western China,

obtained by himself during a shooting trip. Mr. Bridgeman is

of opinion that there are three distinct kinds of Muntjac in

An-wei, which he calls the red, the black, and the yellow or tawny ;

the last being the smallest.

Of these, the so-called red species is identified, and rightly, by

Mr. Bridgeman with Cervulus sclateri, which appears to be distin-

guished from C. lachrymans of Sze-chuen by the more marked

contrast between the yellow of the head and the rufous or olive

of the neck. Both are characterised by the absence of a dark

nuchal stripe; while in sclateri, at any rate, the young are

spotted, instead of uniformly coloured as in reevesi. The backs

of the ears are, I believe, yellow in both sexes. In An-wei

sclateri is the common species.

With regard to Mr. Bridgeman’s “yellow species,’ in which

the whole tone of the coat is yellowish olive, with yellow speckles

on the hair, I take this to be near akin to C. reevesi, which is,

however, described as being typically reddish chestnut speckled

* The complete account of this new species appears here; but the name anda

preliminary description were published in the ‘ Abstract,’ No. 86, 1910.

990 MR. R, LYDEKKER ON (June 14,

with yellowish grey. A mounted specimen in the Museum is

darker, with rufous speckles, which almost disappear on the hind-

quarters. All show a dark nape-stripe.

Mr. Bridgeman’s yellow Muntjac agrees with reevesi in the

relatively small degree of divergence of the antler-pedicles (text-

fig. 144, A), the great size of the lachrymal fossa, and the presence

of a distinct outer lateral prominence on the nasals where they

first come into contact with the maxillw.

Text-fig. 144,

A B

Skulls of Chinese Muntjaes.

A. Cervulus reevesi (?). B. Cervulus bridgemani.

In the male skin the back of the ears is covered with yellow

hairs, whereas in the female the hairs on this part are blackish.

No such difference appears to have been described between the

ears of the two sexes of reevesi; but this may possibly have been

1910.] A WAPITI AND A MUNTJAC. 991

overlooked. For the present, at any rate, I must leave this

yellow Muntjac without a definite name, suggesting, however,

that it is a local race of reevesi.

Of the black, or, as it might preferably be called, the brown

Muntjac, Mr. Bridgeman has sent home the skins and skulls of

several specimens of both sexes. The general type of colouring

of the fur of the upper parts is the same as in the yellow Muntjac,

but the tint is much darker. In the male the fur is blackish

brown, with the hairs on the middle line of the back and the

whole of the rump faintly ringed with yellow. In the female

the yellow rings on the hairs are more numerous and brighter in

colour, while the annulated area extends down to the flanks, so

that the whole body is gold-speckled. In both sexes there is a

dark nuchal stripe; and in the female, like that of reevesi, the

black frontal lines unite into a broad patch between the ears,

which is continued as the nuchal stripe. In the female the

ears are black externally, whereas in the males they are yellow,

although with the basal half black in one example.

The skull (text-fig. 144, B), while agreeing with that of the

yellow Muntjac in the great size of the lachrymal fossa, differs

in many instances, at any rate, by the greater divergence of the

antler-pedicles, and the invariable absence of any distinct lateral

projection on the nasals where they first come into contact with

the maxille.

This Muntjac appears to be a new form, which I propose to

regard as a species, with the name of C. bridgemani, although

it may be only a larger race of reevesi. The mounted female

presented to the Museum by Mr. Bridgeman is the type. The

height of this specimen is 19 inches, and when freshly killed its

weight was 29 lbs.

These brown Muntjacs live normally at a high elevation in the

Wei-Yas Shan Mountains of An-wei; and descend to the low

grounds only during spells of exceptional cold in mid-winter.

Information of their existence was given to Mr. Bridgeman by

Mr. Charles Maguire, a mining agent in An-wei; and the donor

suggested that the species should be named after that gentleman,

although this appeared to me, on the whole, inadvisable *.

These Muntjacs belong to the typical group of the genus, in

which the upper surface of the tail is chestnut; but in their

generally dark colour, and especially in the black ears of the female,

they form in some degree a transition to the plum-coloured

group, in which the upper surface of the tail is dark; the next

connecting species being C. few of Tenasserim, which lacks the

head-crest of the Chinese crinifrons and the allied genus

Hlaphodus.

It may be added that I have given preliminary notices—without

specific names—of both the Wapiti and the Muntjac in the

‘Field’ newspaper for April 1910.

* A male skin shot by Mr. Maguire was received at the British Museum after

this paper was read,

992 MR. R. LYDEKKER ON [June 14,

3. On Three African Buffaloes.

By R. LypexKer *.

[Received May 12, 1910.)

(Text-figures 145-148.)

Tn the present communication I take the opportunity of giving

figures of the heads or horns of three races of African Buffaloes,

two of which, in my opinion, indicate types distinct from any

of those described by Dr. P. Matschie in his paper on the local

forms of African Buffaloes published in the ‘ Sitzungsberichte

Ges. Naturfor.,’ Berlin, 1906.

1. Bos CAFFER THIERRYI Matschie.

As announced in the ‘ Daily Telegraph’ of December 30th,

1909, Dr. K. W. Kumm, during his journey through equatorial

Africa from Lake Chad to Wau, obtained evidence of the existence

in the Upper Shari Valley of what he at the time regarded as a

new Buffalo. This portion of the Shari Valley, it may be well to

mention, is situated in the French Congo, somewhat to the north-

ward, so far as I can determine from the narrative, of an east-

and-west line connecting Adamawa, in the German Cameruns,

with the British station of Wau, in the Bahr-el-Ghazal. The

single trophy of this Shari Buffalo brought home (and presented

to the British Museum) by Dr. Kumm consists of the frontlet and

horns of an adult, but not aged, bull (text-fig. 145). The animal

to which the specimen belonged appears to have been killed at a

comparatively recent date, but not, I should say, by a European ;

and the long, sharp tips of the horns afford decisive evidence as

to its relative age.

In the fact that they are situated throughout their length

almost in one plane, coupled with the great length of the slender,

cylindrical tips, which exceeds that of the basal portion, and the

right angle formed by the junction of these two portions with one

another, the Shari horns accord with the pair from the interior

of Togoland, German West Africa, described and figured by

Dr. Matschie on page 172, fig. 3, of the paper already cited, under

the name of Bubalus thierryi. The Shari horns are, however,

much deeper in the antero-posterior direction at their bases,

where they are more expanded and flattened, and also much more

closely approximated in the middle line than in the type of

thierryi. The latter is, however, a female, and this being so,

there seems no reason why the Shari horns should not pertain to

the same race. Accordingly, despite the long interval between

the localities where the two specimens were obtained, there seems

no possibility of separating the Shari horns from thierryi, and J

therefore associate them provisionally with that race,

* Published by permission of the Trustees of the British Museum.

1910.] THREE AFRICAN BUFFALOES. 993

The horns of the Lake Chad B. ¢. brachyceros are of a much

smaller and totally different type; while those of the Sene-

gambian B. c. planiceros, according to the specimen figured on

page 105 of my ‘ Wild Oxen, Sheep, and Goats,’ have, even when

unworn, much shorter tips.

Text-fig. 145.

Frontlet and horns of bull of Bos caffer thierryi.

From Dr. Kumm’s specimen in the British Museum.

The horns of Dr. Kumm’s specimen have a length of 317 inches

along the outer curve, with a basal span of 8 inches, and a tip to

tip interval of 264 inches.

2. Bos CAFFER SIMPSONI, subsp. n.

On page 156 of vol. exv. of the ‘ Field’ newspaper for 1910, I

gave a preliminary notice, accompanied by a figure, of certain

Buffalo-heads obtained by Mr. M. W. Hilton-Simpson on the left

bank of the Kwilu River in the Belgian (not, as I first thought,

the French) Congo. This river, which takes its rise in Portuguese

territory, runs nearly due north, and discharges into the Kwango,

a tributary of the Kasai, which, in its turn, forms the most

important affluent of the Congo. Of these Buffaloes, which were

found in herds of considerable size, the heads of two bulls and

two females were brought home by Mr. Hilton-Simpson, and of

these one female has been presented to the British Museum.

In the heavily fringed ears and the general form of the horns

these Buffaloes approximate to the well-known red Congo Buffalo

(B. c. nanus); but the horn-measurements of the largest bull

994 ; MR, R. LYDEKKER ON [June 14,

(text-fig. 146) exceed any recorded for the latter race, the right

horn measuring 253 inches along the outer curve and the left

one 243 inches; the basal girth of the former being 164 inches,

its maximum width 63 inches, and the expanse from tip to tip 132

inches. In the largest cow (text-fig. 147) the left horn measures

15 inches in length, with a girth iat 94 and an expanse of 82

inches. In profile the horns jalna upwards nearly in the plane

of the face.

Text-fig. 146.

Head of bull Bos caffer simpsoni. From Mr. Hilton-Simpson’s specimen.

The colour of both cows and bulls is a pure brown, and thus

much darker than the tawny red of nanus; the young only

agreeing approximately in hue with the latter, and the cows being

fully as dark as the bulls. Except in the bull whose head is

shown in text-fig. 146, the fringes of the ears are tinged with

tawny, with one white lock. In size, Mr. Simpson estimates that

the Kwilu Buffalo slightly exceeds the specimens of the red

Congo race mounted in the British Museum,

1910.] THREE AFRICAN BUFFALOES, 995

That the Kwilu Buffalo is distinct from B. ¢. nanus, whose

habitat includes Nigeria, Ashanti, and Sierra Leone, seems to be

clearly indicated by its darker colour and larger horns, the form

of which does not agree precisely with those of any example of the

latter that have come under my notice; and I cannot identify it

with any of the other races at present. Accordingly, I propose

to regard it as representing a distinct race, under the name of

Bos caffer simpsoni; taking as the type the head of a cow which

Mr. Hilton-Simpson has presented to the British Museum.

Text-fig. 147.

Head of cow Bos caffer simpsoni. From Mr. Hilton-Simpson’s specimen.

The Kwilu Buffalo evidently forms a connecting-link between

B. c. nanus and the short-horned blackish races of Bos caffer, as

exemplified by a pair mentioned by Dr. Graham Renshaw in the

Society's ‘ Proceedings’ for 1904, p. 130, as being then living in the

Antwerp Zoological Gardens. Those animals appear to have had

horns of the same general type as those of the Kwilu race, but

their general body- colour was much darker, being described as

dark blackish brow n; the ears show Smale heavy fringes. As

suggested by Dr. Renshaw, these Buffaloes may have been the

Senegambian B. c. planiceros.

996 MR. R. LYDEKKER ON [June 14,

3. Bos carrer corront (?) Lydekker.

So far as my information goes, the small-horned Bos caffer

brachyceros is at the present time known only by the two type

skulls brought home by Messrs. Denham and Clapperton after their

journey through the Lake Chad district, and named and described

by Dr. Gray in the ‘ Annals & Magazine of Natural History ’ for

1837, vol. i. p. 587. As to the precise locality where this so-called

Lake Chad Buffalo was obtained, there is no definite information,

the original description merely mentioning “ Central Africa”:

accordingly, the spot may well have been scores of miles distant

from the Lake. In ‘ Wild Oxen, Sheep, and Goats’ I have

tentatively regarded the two type skulls as respectively repre-

senting the bull and cow of the Lake Chad race; and I am now

more convinced than ever of the correctness of this interpretation.

A figure of the skull of the bull, which is fully adult, will be found

on page 114 of the volume cited

Text-fig. 148.

Bos caffer cottoni(?). Head of young male.

Krom the specimen in the British Museum.

This skull is characterised by the shortness and generally

smal] size of the horns, which are separated by a wide gap

in the middle line of the forehead, and show no marked

expansion or prominence at the base. Their basal portion, which

presents a nearly flat front surface, ascends upwards and outwards

1910.] THREE AFRICAN BUFFALOES. 997

at an angle of almost exactly 45° with the middle line of the skull

for some distance, after which the horns are curved inwards in a

regular sweep, so that the tips (which are quite unworn in the

specimens) are directed inwards in a nearly horizontal plane.

In my judgment, this type of horn is quite distinct from that of

both B. ce. nanus and B. c. planiceros.

Thus matters stood till 1907, when Prince EK. Demidoff pre-

sented to the British Museum the head of an immature bull

Buffalo belonging to one of the little-known reddish races, shot

by his brother, the late Prince Paul Demidoft, somewhere in the

interior of the French Congo. The immature condition of the

horns in this specimen (texb-fig. 148) is extremely unfortunate,

and led me at first to the conclusion that they indicated a short-

horned race, with a large interval on the forehead between the

horns of opposite sides, and a horn-curvature which would probably

grow into that of brachyceros; but a specimen from the same

district shown me by Mr. Rowland Ward proves this to be wrong.

In colour the abundant hair of Prince Demidoft’s specimen is de-

cidedly darker than that of the bull nanws from Ashanti exhibited

in the Museum; its general tint being tawny brown, tending to

blackish or black on the muzzle and chin. The most characteristic

feature of the specimen is, however, the colour of the excessively

abundant fringe of long hair on the margin of the ears. At the

base of the upper edge the colour of these hairs is light yellowish

chestnut; but on the remainder of the same margin, together

with most of the lower ones, the hairs are black, with two small

flecks of straw-colour near the middle of the lower border and a

larger patch near the base of the same. In these respects

the ear-fringes differ in colour from those of b.¢. nanus. The

specimen shown me by Mr. Ward is an older bull, in which

the colour of the hair is bright red, while the horns, although

smaller and more separated on the forehead, are of the type of

those of B. c. typicus. This head indicates a race allied to or

identical with my JB. c. cottoni from the Semliki (P. Z. 8. 1906,

p. 996).

Posrscrip?.—Since this paper was read I have received evidence

of the existence of another, and very well-defined, race of dwarf

Buffalo, inhabiting the Yala district of South Nigeria, and

characterised by the bulls being brownish black and the cows

dun or khaki-coloured. Heads of a bull and cow were brought

home by Mr. J. H. L. Thompson, of the Duke of Wellington’s

Regiment, and have been mounted at Mr. Rowland Ward's

establishment. Mr. Thompson informs me that the bulls of this

race stand from 3 ft. 6 in to 4 ft. at the withers, and that they

are short-legged and heavily built animals. In the adults of

both sexes the legs are light-coloured from the knees and hocks

to the hoofs. The calves are dark grey.

My informant also states that there occurs a larger race of

Proc. Zoou. Soc.—1910, No. LXV. 65

998 PROF, A, CABRERA ON A NEW ANTELOPE [June 14,

Buffaloes in the same district, whose colour is darkish rufous,

with the lower portions of the legs rather lighter than the body,

Although I am fully convinced of the distinctness of the small

Yala race, I do not propose to give it a name until a specimen

of the head of at least one of the sexes is available for the British

Museum. A fuller account will be found in the ‘ Field,’ vol. exv.

p. 1112, 1910,

4, On a new Antelope and on the Spanish Chamois.

By Prof. A, Caprera, C.M.Z.8.

[Received May 10, 1910.]

(Text-figure 149.)

Among a great number of hunting-trophies obtained last winter

by the Spanish sportsman, Mr. arcades de la Huerta, daring an

expedition in British East Africa, there are two Antelope heads

which seem to represent a new form of Damaliseus. They belong,

indeed, to animals similar in size and colour to D, jimela, but with

the middle part of the face not black, but pale whitish buff. That

they are not individual aberrations is demonstrated by the fact that

Mr. de la Huerta has met with two herds entirely composed of pale

blazed Topis—one on the Uasingishu plateau and another near

the Turkwell River. According to the natives, these antelopes

were also seen by the Roosevelt party, but no specimens were

obtained,

Without entering now into the question whether it is a true

species or only a local race of Damaliscus corrigum, I think this

form with whitish forehead needs a name, and I propose to call it

Damaliscus phalius*,

I take this opportunity to describe and name also the Chamois

from the Cantabrian Mountains, N. Spain, which is a very different

form from the Pyrenean one ( Rupicapra rupicapra pyrendica

Bonap.), smaller in size and redder in colour.

Following are the brief descriptions of these two new animals.

DAMALISCUS PHALIUS, Sp. 0.

Similar to D. jimela in size and in the colour of the body and

limbs, but with a pale whitish-buff blaze on the face, instead of

the black one common to the other forms of the corrigum group,

In old males, according to Mr, de la Huerta, the blaze becomes

pure white and almost as well defined as in D. albifrons and

D, pygargus.

Hab, The north-western part of British East Africa, east of

Mount Elgon.

* Phalios was, according to Dr. Ridgeway, the Greek name for a dark-coloured

hhorse with a white blaze from the top of the head down to the nose.

1910.} AND ON THE SPANISH CHAMOIS. 999

The type is the specimen from Uasingishu in the collection of

Mr. de la Huerta (text-fig. 149). It is an adult, but not old,

male, with horns 432 mm. in length along the front curve, The

Turkwell River specimen is a female.

Text-fig. 149.

Whitish-blazed Topi (Damaliscus phalius) from the Uasingishu plateau. Type.

RUPICAPRA RUPICAPRA PARVA, subsp. n.

Smaller than all the other forms of the species. General colour

brownish red, darker on the limbs, becoming dark brown on the

forearms, above the “knees,” and on the inner part of the hind

limbs. Sides of the head dirty greyish brown, leaving in the

middle of the face a broad space of yellowish buff. The throat

and under jaw are not whitish, as in &. r. pyrenaica, but only a

little paler than the body-colour.

Hab, Cantabrian Mountains, N. Spain.

The type is a female from Picos de Europa, Santander province,

in the Madrid Museum of Natural History. She is 570 mm. in

height to the shoulder, with horns 146 mm. in length on the front

curve. The hornsof a male skull in the same collection are stouter,

but not longer. A female from the Pyrenees, also in the Madrid

Museum, is 690 mm. in height to the shoulder, and has horns

232 mm. in length.

1000 DR. E. A, WILSON ON THE [June 14,

5. The Changes of Plumage in the Red Grouse (Lagopus

scoticus Lath.) in Health and in Disease. By Epwarp

A. Witson, M.B., F.Z.8., Field-Observer to the Grouse

Disease Inquiry.*

[Received June 14, 1910.]

(Plates LX XVITI.-CI.7 & Text-figure 150.)

Part I.

Plumage Changes in the Cock Grouse.

When a large number of skins of the cock Grouse are

arranged together, side by side, according to the month of the

year in which the birds were killed, it will be found that, even

taking into account the differences of well-marked local vari-

ations in plumage, the series can readily be divided into two very

distinct sets.

There is first a very marked uniformity in the plumage of the

cock birds killed from the middle of November to the end of

June; and likewise amongst those killed from the end of June

to the middle of November.

These two periods, November to June and June to November,

mark the two seasonal changes of plumage in the cock Grouse.

The first is a plumage worn throughout the winter, as well as

during the courting- and breeding-season of the spring.

The second is a plumage worn throughout the summer and

autumn.

It is necessary to lay stress upon this general broad division

of the cock Grouse’s plumage, and if a large number of skins

can be arranged as suggested, the exact time at which the

Grouse has definitely changed from the one plumage to the

other cannot possibly be overlooked. The birds obtained at

the end of May are definitely in the darker and redder winter-

plumage, and those procured at the end of June are definitely

in the paler and more buff-coloured summer-plumage ; those

killed at the beginning of October are still partly in the paler

summer-plumage, and by the end of November all are in the

darker winter-plumage.

It must, however, be added, that there is hardly a month in

the whole year, or a Grouse-skin in a collection of many hundreds

covering every month of the year,in which one plumage only can

be found unmixed with the other. This fact accounts largely for

the misunderstanding which at one time existed, but which has

now, we hope, been satisfactorily settled, in respect of the whole

* Owing to Dr. Wilson having left London on the Antarctic Expedition before

this Memoir was set up in type, he was unable to revise the proofs, but left full

power to the Editor and Mr. W. R. Ogilvie-Grant, who have made certain modi-

fications in the text.

+ For explanation of the Plates see p, 1032.

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1910.] PLUMAGE OF THE RED GROUSE. 1001

vexed question of moult and plumage-changes in the Red Grouse,

and their proper interpretation.

Without referring in detail to the points upon which

differences of opinion have before now arisen, it may be shown

that much misunderstanding upon such a difficult subject as this

is based upon a different rendering of facts into words, facts which

were recognized and perfectly well explained by Mr. Ogilvie-

Grantin 1893. Both he and Mr. Millais have made the subject of

plumage-changes in the game-birds, and especially in the Grouse,

a special study, and it must be admitted that there are very few

points upon which they have touched which seem to require

further explanation ; and still fewer points, if any, which can be

brought to light for the first time in connection with the plumage-

changes of the Red Grouse. A monograph on the Red Grouse, such

as the Report of the Grouse Disease Inquiry, would, however, be

obviously incomplete without an account of the plumage-changes of

the bird itself; and it so happens that during the past six years of

the Grouse Disease Inquiry’s existence the collection of some six

hundred Red Grouse skins, representing every age, phase and

change of plumage in that bird, has given a unique opportunity

for an independent revision of the work already done—an oppor-

tunity such as has never occurred before in the study of any

single species of British bird for observing the effect of disease

upon moult and feather-growth. So it happens that although

the work as it stands has been so nearly completed by the labours

of the two ornithologists already mentioned, there are still points

of interest to which attention may be drawn, especially in con-

nection with the marked effect which parasitism and other wasting

diseases have upon the moult and growth of feathers; and it

is to this influence of disease that attention will be particularly

drawn in the present paper.

It is almost incredible that a moult should be deferred from

one season to another or even to a third, and that the right

plumage should eventually be produced if the bird by means of good

food and good weather is at last enabled to recover its health and

grow any new feathers at all. It is interesting and to some

people, such as sportsmen and gamekeepers, even useful to know

that bare featherless legs and feet, which have so long been

considered a sure sign ‘of disease in the Red Grouse, may, in

certain months of the year, be an unavoidable accompaniment of

really good health, while thickly feathered legs in the same month

are a sure sign of deferred moult and of sickness. It is only when

the proper season for the moult of the leg and foot-feathering is

completely understood, that we can begin to understand the

reason for attaching an unfavourable prognosis to heavy leg-

feathering when the legs should have been featherless, and an

equally favourable prognosis to bare legs when the legs should

certainly have been bare (Pls. XCII. & XCIIT.).

To return, however, to the two plumages of the healthy cock

Grouse. They are distinguished by Mr. Ogilvie-Grant as the

1002 DR. E, A. WILSON ON THE [June 15,

autumn-plumage and the winter-summer-plumage, and he says

further that the cock “has no distinct summer-plumage.” It is

pertectly easy to see what is meant by this, and also by the

statement which follows, that the cock “yetains the winter-

plumage throughout the breeding-ses ison.’

Mr. Millais also, i in speaking of the cock Grouse, makes use of

the expression autumn-plumage which, he says, appears late in

June ; and he adds that the autumn-plumage together with the

“ Spring feathers” (or what Mr, Ogilvie-Grant considers the first

beginning of the autumn- _plumage on the Grouse’s neck) remain

till the main moult in August and September.

Mr. Millais also makes ‘the following statement, which appears

to be based on a misinterpretation. He says “as a matter of

fact the male Grouse sheds in September and August a plumage

which is a mixture of its Winter, Spring, and Eclipse feathers.”

These so-called “Spring” and “ Kclipse” feathers are no doubt,

as Mr, Ogilvie-Grant holds, the commencement of the plumage,

which is completed gradually during the summer months, and

which he has described as the autumn-plumage. It is naturally

a little misleading to find the autumn-plumage beginning to

appear in early summer, but so long as the term is understood

to mean the paler, more buff-coloured plumage with bolder bars

of black, which begins to appear first on the neck of the cock at

the end of May or ‘early in June and is eventually cast for the

winter-plumage in October, there need be no real misunder-

standing.

That feathers of the previous winter-plumage should be

mentioned in speaking of the moult of this autumn-plumage is

also quite intelligible, since the old winter-plumage of the breast

and abdomen is being quickly shed and replaced by a similar new

winter-plumage at the time when the autumn-plumage on the

rest of the body is being cast. There are in addition very fre-

quently a few feathers of the copper-red plumage on the chin

really belonging to and remaining over from the previous winter-

plum: age,

Instead of going into further details, however, with regard to

the two moults and plumages of the cock Grouse, it will be

simpler at this point to take its plumage-changes in detail, succes-

sively month by month, explaining as nearly as possible what can

be gathered from the examination of a series of skins such as has

been brought together by the Committee of Inquiry, including as

it does a great number of specimens in all stages of disease as well

as in health.

They illustrate every month of the year and most of the loeal

variations to be found in England, Scotland, and Ireland; and

there are a sufficient number of sick as well as healthy birds to

show the very great influence of disease in altering the capacity

of the individual for feather-growth. Unless this effect, which

results as a rule in the Red Grouse from excessive parasitism, is

fully recognized there will always be misunderstandings upon the

1910.] PLUMAGE OF THE RED GROUSE. 1003

moult of this bird, for almost every Grouse in the country is to

some extent infested with parasitic worms, and there are years

when irregularity of moult is the rule rather than the exception,

Moreover, it so happens that in autumn, when birds are being shot

in large numbers, the survivors of the worst two months of the

year for Grouse-disease mortality, that is, the survivors of May

and June, are all convalescing ; but they are convalescing with

their plumage-changes all retarded and put completely out of

order and routine. In this way it is possible in September to

kill two birds on the same day both of which have the chestnut-

coloured feathers of the winter-plumage on the chin and throat ;

but upon examination it may be seen that in one bird the

edges of these feathers are frayed and worn and the colour faded,

showing that they have survived from the previous winter-

plumage ; whereas in the other bird they are hardly free of the

scaly sheaths in which they grew, and are really precocious

feathers of the coming winter-plumage. This is only one of the

many traps which result from the deleterious influence which

disease exerts upon a bird’s capacity for feather-growth and

replacement and so upon the regularity of its moult.

There is another point to which attention must be drawn before

entering upon a systematic description of the monthly changes of

feather in the cock Grouse. It is as to whether the autumn-

plumage of the cock can be correctly described as an “ Eclipse ”

plumage, comparable as it obviously is in character with the

Spring breeding-plumage in the hen, but appearing just two

months later and after the breeding-season. In each sex the

general change from winter to summer may be described as a

change from a more richly pigmented, darker, black and chestnut,

or rufous-chestnut plumage with rather fine transverse black

markings, sometimes almost vermiculate in character, to a less-

richly pigmented, paler, buff or rufous-buff or tawny-buff plumage

with characteristically broad black bars and transverse markings.

To each sex, moreover, the characteristic buff and black broad-

banded summer-plumage is given its special appearance on the

dorsal aspect by the growth of feathers with large black centres

and a few buff or tawny-buff subterminal bars of considerable

width and a terminal border or spot of the palest buff, which is a

very conspicuous feature on the back of most hens, and often only

less conspicuous in the cock. In the cock, however, this plumage

appears just two months later, and is less beautifully developed

than in the hen.

There is without doubt a general broad resemblance first

between the cock and the hen Grouse when the former is in its

“winter-plumage” and the latter in its “ autumn-plumage ” ;

and, secondly, between the cock and the hen Grouse when the

former is in its “ autumn-plumage ” and the latter in its “ spring-

plumage.

The perplexing fact is that these general resemblances are not

synchronous in the two sexes, a peculiarity first discovered by

1004 DR, E, A. WILSON ON THE [June 14,

Mr. Ogilvie-Grant, for, as already pointed out, there is an interval

of two months between the moults of the cock and hen.

Again, it might reasonably be expected that, as the Ptarmigan

and the Scandins avian Willow-Grouse have not two plaumages in

the year, but three, some suggestion of the third plumage might

be forthcoming in the Red Grouse. But the Red Grouse has

only two moults. Mr, Ogilvie-Grant, however, explains the

position by saying that the buff and black plumage of the hen

answers to the spring-plumage of the hen Ptarmigan, while

the buff and black plumage of the cock Grouse answers to the

autumn-plumage of the cock Ptarmigan, ‘lhe grounds for this

position will be considered later in the light of the possible

effect which continued disease may have in permanently altering

the season of a moult.

Beginning now with the cock Red Grouse in January, and

taking its appearance from the ventral aspect first, the uniformity

of the series is a very conspicuous feature. Every healthy bird

is chestnut or rufous-chestnut and black, with fine, almost

vermiculate black cross-lines over it.

Even in the blackest birds the throat and fore-neck are always

of a rich copper-red colour with very little or no black edging at

the borders of the feathers, which are usually barred with black

only on the actual chin, Here there may be also more or less of

white tippings, even to the formation of two white moustachios

leading downwards from the gape, sometimes an inch in length,

This may be a feature either of the black type or of the red *

(Pls. LXXIX, & LXXX.). In some very red and black Red

Grouse the abdominal feathers are also freely and broadly tipped

with white; and this may sometimes be seen even on the feathers

of the upperparts (Pl. LXXXI.). The legs and feet are thickly

feathered, and are white, or white with brownish barring. The

claws are often in this month very long and strong.

Occasionally a pale bleached feather of the preceding “ autumn-

plumage “isto be found on the flanks, middle of the breast or neck,

and may be recognized by its frayed edges ; and occasionally (e. g.

No, 539), in a very backward bird, there may be many such worn

and faded feathers on the chest and flanks, but such a case is

invariably the result of sickness.

On the dorsal side there is again, broadly speaking, a general

uniformity of chestnut, bright or d: ark, or of blackish fe: vathers,

with fine black transverse ‘markings ; but in almost every bird

there may be found a considerable number of the old black-

centred “ autumn-plumage” feathers remaining, with their

frayed and faded edges of whitish-buff (Pl. LXXXII.), On

the lower back and rump the more worn and faded feathers

predominate,

The primary and secondary quills are all complete and are but a

* The whole paper deals with the Red Grouse (Lagopus scoticus Lath.). The

terms “ black Red Grouse ”’ and “ buff-spotted or white-spotted Red Grouse” must

not be confused with similar terms for other species of Grouse.

1910. ] PLUMAGE OF THE RED GROUSE. 1005

few months old, having been renewed between June and August

and the same may be said of the rectrices.

The following points in the cock Grouse of January are

characteristic :—

1. The rich copper-red, generally unbarred feathers of the

throat and fore-neck (Pl. XCIV. fig. 2).

2. The fine barring of the chestnut, dark rufous-chestnut, or

blackish-brown of the back, with the scattered black-centred

feathers of the last ‘ autumn-plumage.”

3. The thick white feathering of the feet and legs, which soon

becomes blackened and worn by the “ burrens” or ‘ colons,”

the charred stalks of old burnt heather.

4. The perfect flight-feathers of the wings and tail.

5. The enormous claws.

In February the cock Grouse is still in the darker winter-

plumage.

Young, sheathed and growing broad-barred feathers, the re-

mains of the “ winter-plumage,” may still occasionally be found

on the hind-neck, nape, and head in backward birds.

In March the cock Grouse normally shows no change; but

towards the end of the month in exceptional instances individual

birds may be found with a few precocious feathers of the autumn-

plumage making their appearance on the back of the head and

neck. These are very probably feathers irregularly acquired to

take the place of those which have been lost during encounters

with other males.

In April the cock Grouse still shows no change whatever.

In this month there are often greatly increased opportunities

for the addition of skins to a collection, because it happens to be a

month of very high mortality from “ disease.” The birds are found

and can be collected not only by the keepers who are out early inthe

month in search of fox-earths, and who are generally also burning

heather, but also, later on, by the shepherds, who are constantly

ranging the moor in the lambing time.

The result during the last five years has been a great accession

of Grouse-skins to the Inquiry’s collection, in March, with a very

large proportion of males badly diseased, and fewer birds in

perfect health. Therefore in the series of skins of cock-birds

representing April, the great majority are very backward.

Healthy birds have still the old rich red copper-coloured throat

of the winter-plumage, and fresh-looking “autumn” feathers

round the neck, upper-back and mantle, while the winter and

old autumn-plumage of the rump and back is bleached and faded.

The backward birds are easily picked out, as they have not yet

assumed their ‘ winter ” plumage, and are still mostly clad in old

worn autumn-plumage of the previous year.

It is important to note the extraordinary discrepancies which

so commonly occur in the plumage of the Red Grouse owing to

disease, whereby the deferred moult becomes in some years almost

1006 DR. E. A. WILSON ON THE [June 14,.

the rule, and the rule of health becomes almost the exception..

It is a very difficult matter, indeed, for anyone who has not

had the opportunity of examining an extensive series of Grouse-

skins, in disease as well as in health, and covering every month

of the year, to come to any true conclusion about the moult.

Diseased conditions often entirely mask the normal plumage-

changes from time to time, and it is far more important to

realise this than to examine thousands of more or less healthy

birds shot in the ordinary course of events in the shooting-

season. A study of abnormal plumage-changes in diseased Grouse

is essential if the discrepancies which arise in the moult of

what are often wrongly considered normal birds are ever to be-

explained. Once this point is grasped tne question becomes much

simpler ; and it is because the Grouse Disease Committee has had

such a unique opportunity for studying both sides of the question

that it has been deemed necessary to enter into these plumage-

changes at such length.

The legs and feet of a healthy bird in April should be getting

worn and thinly feathered. If a bird has newly and thickly-

feathered legs and feet, it means, almost certainly, that the-

“winter” plumage has been put on very late. The healthy

Grouse should now be moulting the feathers of the feet and legs,

so that bareness or lack of feathers becomes in them a sign of

health in April and thickly-feathered legs a sign of sickness ;.

this is the precise contrary of what has almost become proverbial

on the moor, that bare legs indicate disease ; though for the later’

autumn months the saying is quite true.

In May the preponderance of cock birds found dead, and there-

fore of skins of cock birds in the May collection showing belated

moult, is again a large one. The healthy cock is still in his-

much-worn winter-plumage, but on the head and neck some

feathers of the new autumn-plumage are beginning to make their

appearance (Pls. LXXXITI. & LXXXIYV.).

In June the disease mortality, due to Strongylosis, is, as a rule,

coming toanend. ‘This at least is true for the adults, but for

the young chicks June and July are often fatal months owing to.

Coccidiosis. Late in June the healthy cock Grouse can at last

be said to have changed into his complete ‘autumn-plumage.”

The winter-plumage persists only on the abdomen and lower

breast, on the actual chin which is blackish, with a few white

spots, and on the throat, where a few red feathers still remain.

The moulting of the quills and tail-feathers commences towards.

the end of the month The rump and back are now completely

covered with new black-centred feathers carrying broad-barred

buff and black bands and a few have a whitish terminal spot,

similar to that found in the female.

The head and neck, breast and throat, are now clothed with

broad-barred buff and black feathers, quite distinct from the more

chestnut and more finely black-marked plumage of the winter,

It is impossible, in seeing a series of the birds showing this dis-

1910.] PLUMAGE OF THE RED GROUSE. 1007

tinctive change, to avoid noticing how closely it approximates

to the nesting-plumage of the hen, and yet to think and

to speak of this “autumn” plumage as an “ Eclipse” plumage.

In every respect it closely resembles the breeding-plumage of the

hen, and the one prominent discrepancy is that it has arrived in

the cock just two months later than it is normally due in the

hen, far too late to be a breeding-plumage.

It appears almost as though the pathological postponement of

the moult, which is, after all, nothing but a sign and a symptom

of disease, has gradually developed into a normal habit in the

life of the bird ; and one is led to think that this habitual dis-

ability in the cock Grouse, which results from Strongylosis during

the nesting, courting, and breeding-season (a disability which

causes the death of about eight cocks to every hen in April and

in May), may have caused the alteration in the season of the

moult, simply because the wis vite of the cock bird, insufticient

as we now know it to be at the close of winter for the ordinary

calls of reproduction, would be still more disastrously insufficient

if preceded by an early moult.

At the present time the cock undoubtedly breeds in the winter-

plumage, without any further acquisition of new feathers, and,

as has recently been pointed out by Mr. Ogilvie-Grant, what

have been regarded as new ‘ spring-feathers” on the neck by

Mr. Millais are in fact the old autumn-feathers, which on that:

part of the body do not become worn and faded.

That any feather of the Grouse, either in the cock or in the

hen, was ever altered as to its pigment either in pattern, or in

tone, or in any other character, when once it had completed

growth and had been cut off from the circulation, is at present

an assumption which is not well supported by the physiology of

feather-growth.

Metchnikoff's observation upon the migration of leucocytes

into hair and their action in removing pigment cannot for one

moment be adduced as conclusive proof that the same thing may

happen in the case of a full-grown feather. While the circulation

is active in the feather-shaft, and for as long and in so far as it

continues, pigmentation may be altered, and so may growth;

but once allow, first that the circulation has ceased beyond the

entrance to the base of the shaft, and secondly that the feather,

although still attached to the epidermis, is cut off from the

circulation in the deeper living layer of the skin, then the

feather is no more likely or able to change the pigment which is

responsible for its pattern or its colour (except by external means

such as bleaching by sunlight) than if the same feather had

been plucked out and kept entirely separate from the bird.

Once the feather is full-grown and the circulation in it

stopped there is no reason to believe that any thing can alter

it save sunlight and water, and oil supplied as an external

unguent from the oil-gland. That appearances are most deceptive

in this respect must be allowed. Feathers may be collected from

1008 DR. E, A. WILSON ON THE [June 14,

the flanks of hen Grouse which show every possible graduation

between the perfect winter-plumage, almost vermiculate, flank-

feather and the broad-barred, breeding-season, flank-feather of

the summer hen. But it is very much more probable that the

growing period of these ambiguous or intermediate feathers is

one of great susceptibility to outside conditions, as we know

the case to be in respect of the metabolic processes which are

taking place within the hen-bird at the time. These are known

to be exceedingly sensitive. Pigment is indisputably a product

of tissue metabolism. It is often probably a mere waste product,

but it appears at times to serve a special function notwithstanding.

It is also certain that pigment is a production whose appearance,

or failure to appear, is open to considerable vicissitudes in

consequence of small recognized changes in physiological condition,

and many less easily recognized changes in the general metabolism

of the body.

In the hen Grouse during the breeding-season pigment

production is very actively at work, for we know that a very

large amount is being produced for excretion in the pigment-

glands of the lower part of the oviduct. This pigment, moreover,

is precisely of the shade and colour which is characteristic not of

the breeding- plumage, but of the winter dress of the hen and the

cock Red Grouse. It is deposited in abundance normally on every

egg, but it may also abnormally fail to be deposited or even

produced at all, not only for the eggs in the oviduct, but in the

circulating blood of the bird’s whole system. Thus the feathers

instead of becoming buff or brown, reddish or even black as they

proceed in growth, may be any intermediate paler shade of buff,

or even white, a character which is due generally to the complete

absence of all pigment granules. The place of the pigment

in such feathers is probably taken by shining air-globules, as

it is in the hair and feathers of the majority of white animals and

birds.

It is therefore easier to believe the suggestion that a sudden

check, either by a change of temperature, or by wet and cold, or

by want of sunshine, or by change in food, has for the time being

so far affected the tissue metabolism of the bird that a feather

which began to grow upon a circulation lacking pigment particles,

or the waste products which produce them, and which was

therefore originally planned for the paler plumage, may, by a

sudden increase in the metabolism of the bird and so in the out-

put of waste products to the blood, be completed as a feather of

the more deeply pigmented plumage, thus producing a feather

with the characters of both.

This is a plausible explanation, but is still open to some doubt,

for the difference between the broad-banded buff and black flank-

feather of the nesting hen, and the dark red-brown finely cross-

lined feather of the same bird in winter, is obviously greater as

regards pigment distribution than as regards the actual quantity

of pigment deposited in the feathers.

1910. ] PLUMAGE OF THE RED GROUSE. 1009

If there are, as has been held, distinct pigments, such, for

example, as buff, black, and orange-red, in the various colour-

tones of the Red Grouse, it becomes easier to see that the loss

of the red pigment which is utilized for the eggs, leaves the buff

and the black in greater quantity for the nesting-season plumage.

In the winter all three would once more be available.

The fat of the nesting hen is distinctly rich in colour, but in

no case that we have seen has it resembled the orange-coloured

fat which is often seen in overfed Pheasants, and quite commonly

in Gulls and Terns which have been feeding on red crustaceans.

In these birds the orange-red fat or oil, tints not merely the fat

beneath the skin, but the white feathers of the breast and body

which often pr esent a beautiful rosy flush.

The whole question of pigment production and pigment dis-

tribution, intimately connected as it is with the question of the

excretion of waste products and the deposition of fat both in

health and in disease, has not reached a stage which admits of

dogmatic statement upon the subject of pattern change in feathers

without moult.

One recognized method of changing a colour-pattern in feathers

without moult is to be seen in the male of the familiar House

Sparrow, which produces a handsome jet-black cravat in the

breeding-season, where before was a nondescript greyish throat ;

and this it does by the simple process of shedding the grey ends

of the feathers, leaving the blacker parts exposed. This method

is common among birds, but the Red Grouse has been credited

with changing in situ the colour and pattern of the flank-feathers.

Now, with still less reason as it seems, the cock bird has been

credited by Mr. Millais with achieving his summer or breeding-

plumage “ for the most part by repigmentation and pattern change

of most of the winter-feathers below the neck.”

This view cannot be upheld physiologically, and there is much

to uphold the contention that the feathers which are believed to

effect this change of pattern without moult, are actually new

erowing feathers. ‘This can readily be shown by the demonstration

of their unshed sheaths. The misleading birds are again in this

ease the cocks which have been too sick to shed the previous

“autumn-plumage,” and so are still struggling, and with

increasing success as the food improves, to produce a ‘ winter-

plumage,” which they should and would, in health, have achieved

in October,

That the cock bird should moult the feathers of the legs and

feet between March 30th and June 17th is no longer difficult

to understand when the prevalence of Strongylosis is fully

grasped. No bird is safe from the Nematode infestment, and one

is led to think that the majority of cock birds are so badly in-

fested that they are forced to defer the autumn-moult which should

precede that of the previous winter. It is therefore obvious that

between March and June there will be every stage of good or bad

leg- and foot-feathering between the newly acquired thick white

1010 DR, E. A. WILSON ON THE [June 14,

winter-stocking of the sick cock, and the naked featherless clean

moulted leg and foot of the really healthy male bird, in June,

In July, again, will be found the healthy cock bird beginning to

produce white feather-tips over the legs and feet.

In July the general appearance of the healthy cock is much

lighter in colour-tone and much more broken and mottled in

pattern-character than that of the same bird in the winter. The

claws are in many cases now ready to be shed, and the primaries,

secondaries and tail-feathers are in moult. Some six or eight

new clean grown primaries are often to be found in July, and

the long tail-coverts are broad-barred buff and black.

In August the cock Grouse has, of course, the appearance

of full summer- or autumn-plumage, but it requires very little

examination to see that he has already begun to put on feathers

of the winter-plumage.

He now rapidly sheds the old feathers of the last winter’s

plumage which remained throughout the summer upon his breast

and abdomen, and replaces them with the exceedingly handsome

narrow cross-barred red or brown or blackish feathers of the

coming winter-plumage.

There is no second moult or replacement of these feathers of

the breast and abdomen in the cock. Once in the year is enough

for this special area, and the feathers that carry through are

wholly of the winter-plumage. They are often broadly tipped

with white.

The chin-feathers which survived with those of the breast and

abdomen are now also replaced by new ones. It is noticeable

that in the Ptarmigan also it is the white feathering of the chin

and of the breast and belly, as wellas of the wings and tail, which

is changed once only in the year, exactly as with the winter-

plumage of the Grouse. It suggests that these two plumages are

analogous in each.

The plumage-changes in the Ptarmigan are, strange to say,

quite different to the changes in the Grouse. The Ptarmigan

has three distinct moults and plumages in the year. The Red

Grouse has but two.

In August, as has been said, the cock Red Grouse has begun to

put on his winter-plumage. The feathers of the breast and

abdomen are full of sheaths and sheath-scurf, the growth of these

feathers being very rapid and often scarcely noticeable.

On the rump, back, and to a less extent on the shoulders, new

rich red-brown feathers finely marked with black lines are showing

here and there.

Primaries, secondaries, tail-feathers and coverts are now re-

placed by new and blackish feathers with perfect and unbroken

outlines.

Even a few new rich copper-coloured feathers are appearing as

isolated touches of bright colour amongst the faded broad-barred

autumn-feathers of the upper breast.

The feet and legs are bare, save where new white feather-tips

1910.] PLUMAGE OF THE RED GROUSE. 1011

are just appearing through the skin, and the claws of all the

healthy birds are being shed.

In September the chin and throat of the cock Grouse are a

mixture of many pale autumn-feathers much worn and faded,

and a few new copper-red ones.

Most of the “ autumn-plumage ” feathers are now falling out,

more or less frayed and faded.

The breast and abdomen, wings and tail are now clothed with

altogether new feathers, while the head and neck, back, shoulders,

rump, and coverts of the tail are in a transition state, the

“autumn” feathers contrasting with the new rich chestnut and

darker brownish winter-feathers with their fine black transverse

markings.

The feathers of the legs and feet of healthy birds are rapidly

growing to form thick white stockings for the winter. Bare legs

in September are a sign of belated moult or, in other words, a sign

of sickness.

In October, for the first time since the preceding winter, the

red and black varieties of Red Grouse become once more con-

spicuously distinct. This result is due to the new growth of fully

pigmented feathers, either red or black, upon the under surface

of the body.

The upper neck is rapidly becoming copper-red. The chin and

throat still show a proportion of the faded buff “ autumn ”

feathers among the red, the former looking spotty and pale.

On the back the new chestnut and black feathers are rapidly

replacing the faded autumn-feathers.

Some perfectly healthy cocks still look as if in “autumn-

plumage,” while others, on the contrary, have nearly completed

their winter dress.

The legs and feet are thickly covered with white feathers, and

the nails are uniformly small, as the old claws have all been shed.

‘Their growth, however, is extremely rapid.

In November the cock Grouse drops most of the remaining

“autumn-plumage.” By the end of the month his moult is com-

plete, but on the neck and back a greater or lesser number of

these feathers are retained till the following summer.

The most striking characteristics of the winter-plumage are the

rich copper-coloured neck and throat, and, in the darker varieties

which are common in the Scottish Highlands, the contrasting

blackness of the upper breast and abdomen, often broadly flecked

with pure white tips.

Amongst the cocks there are several well-defined and easily

recognized varieties, which seem to have a certain regularity of

distribution geographically. | These will be found fully considered

below.

It must not, however, be forgotten that owing to innumerable

efforts, which have been more or less successfully made from

time to time, to transfer Red Grouse from one part of the country

to another, the distinction of local variations has become a thing

1012 DR. E. A. WILSON ON THE [June 14,

of the past and is now impossible except upon a very limited

scale.

The attempt, however, can be made, and the number of

specimens in the Committee’s Collection of Red Grouse skins

makes it possible to arrive at some conclusions.

Parr II.

Plumage Changes of the Hen Grouse.

The wo changes of plumage in the hen Grouse are completed,

as has already heer explained above, in the one case by the end

of April or the beginning of May, and in the other case by July

and August.

The actual feather-changes in both cock and hen are really

very comparable in character, notwithstanding the discrepancy as

to season ; and allowing for the difference of two months which

makes the moult in the two sexes asynchronous, they may be

described and explained in very much the same terms.

Mr. Ogilvie-Grant was the first to draw attention to the

exception: al want of agreement in the seasons chosen by the two

sexes of the Red Grouse for their moults, and as in the cock’s

plumage | he makes use of the terms “autumn” and ‘ winter—

summer” or “ winter” plumages, which have therefore been used

here, so in speaking of the hen’s plumages it will be well to adhere

aunt rly to the expressions used by him, and to eall them

“summer” and “ autumn—winter ” or autumn ” plumages.

Exception may be taken, and indeed has been taken, to these

names, as being inappropriate and inexact, but they are suffi-

ciently exact for all practical purposes, and so long as moults and

plumage- -changes are not completed in a week but ¢ are spread over

a period of several months, so long will there be some imexacti-

tude in the terminology of these moults and plumages if they are

named according to the months or seasons. It is immater ial so

long as the term is sufliciently defined, for it is obviously

impossible to use a term so exact as to require no definition,

The hen Grouse moults twice in the year, and wears her

“ summer-plumage ” as the breeding-dress from April to July,

and her “autumn” or “autumn to winter” plumage from

August to March.

These changes may be expressed in terms of comparison with

the cock, as a case of plumage change in which the hen has two

annual moults, exactly as has the cock, but both ioults occur two

months earlier in the hen than in the cock.

The hen’s “summer ” or breeding-plumage is a very beautiful

dvess, variable to a considerable extent it is true, but yet having

a general uniformity which be¢éomes the more obviousas a greater

series of skins in any particular phase of plumage is examined,

Opportunities for even seeing the hen Grouse, to say nothing

of obtaining her skin, in the full breeding- plumage are rare;

and thus it has fallen out that in even the large series of Grouse

1910.] | PLUMAGE OF THE RED.GROUSE. 1018

skins at South Kensington and at Cambridge, for example, this

phase is somewhat poorly represented,

The Grouse Inquiry Committee has been to some extent more

fortunate and has obtained a great many skins of hens in the

summer-plumage (see schedule of material, p. 1022), so that points

of resemblance can be noted at sight and individual variations per-

force take their proper and, as it turns out, rather less significant

places. It has been a marked feature in the whole collection

of 580 skins, that as the series grew and the general uniformity

became more marked, the individual variations of which one was

inclined to make much at first became gradually relegated to

their subordinate position in the background.

Uniformity, albeit with endless minor variations, is the rule in

the Grouse as it is in every other creature that leads an unpro-

tected existence under natural conditions. How long it will

continue in the protected, often over-protected, Grouse, remains

to be seen. It is possible that such variations as already occur

are to some extent a modern development, but on this point there

is at present insufficient evidence to amount to certainty.

Beginning once more with January, and considering the

changes of plumage from month to month in the hen Grouse, it

may be said that some hens when examined on the under side

are hardly distinguishable in this month by their plumage from

some cocks. On the dorsum it is different, and a healthy ben in

January is unmistakable owing to the terminal spots of buff which

appear almost invariably, though occasionally in limited numbers,

on the feathers of the back.

In some healthy hens the chin is sometimes still pale buff in

colour, owing to the persistence of summer-plumage feathers of

the preceding year. The throat and fore-neck, on the other

hand, are copper-red, but rarely so uniforinly red as in the cock

(Ee MIC TIN).

The copper-red feathers seem to begin on the fore-neck and

proceed towards the chin, so that the chin often remains buff and

black when the throat is alveady red.

Except in very backward birds, which have been sick, the old

and faded broad-barred feathers of the flanks are never found in

January. The legs and feet are white and thickly feathered and

the claws are long and str ong.

In February the bird is still in the same plumage as in

January. In a few forward birds the feathers of the summer-

dress are beginning to make their appearance on the back of the

neck about the middle of the month.

In March, the change from autumn-plumage to spring breeding-

plumage is, in healthy birds, now quite unmistakable, though

many birds are very backward owing to disease. All doubt as

to the sex, whether from above or below, is now removed except

in cases of disease.

The broad-barred buff and black feathers of the flanks are now

appearing and are most conspicuous and characteristic, while the

Proc. Zoou. Soc.—1910, No. LA VI. 66

1014 DR. E. A. WILSON ON THE [June 14,

whole of the lower breast and abdomen covered by the red-brown

or red-black finely barred feathers of September growth are still

in excellent condition and remain unchanged.

The chin, throat, neck, and upper breast are now mixed with

broad-barred black and yellow feathers in forward birds ; while in

backward birds the throat and fore-neck may still be clad in

copper-red feathers.

The legs and feet are already looking worn and less well-

feather ed, but the claws are long.

In April and in May, for the simple reason that many hen

Grouse died of Grouse Disease during the past six years of the

Inquiry’s work, the proportion of skins of backward hens is

large.

The birds thus picked up dead carry one immediately back

again to winter, for although they ought by this time to be putting

the finishing touches to their spring-plumage, they are, in fact,

but just succeeding in the belated effort to put on the autumn-

dress. They are thus a clear six months late, and afford the most

misleading seasonal characters imaginable. Their legs and feet,

instead of being worn and almost moulted clean, are at last, after

a winter spent with almost naked legs, well-clothed with thick

white feathers. The appearance of the legs therefore in the hens,

as in the cocks, is totally misleading to the Keeper or to the

Sportsman who considers bare unfeathered legs to be a sign of

Grouse Disease. It holds good for autumn only, and in spring

precisely the opposite is the case, for in April, May, and June

none but healthy birds have naked legs and feet.

The general character of advanced and healthy birds towards

the end of April and in May is that of a complete spring-plumage.

The whole of the upperparts ave broadly barred with buffand black,

and marked with conspicuous terminal whitish-buff spots or bars.

The underparts, again, are broadly barred with buff and black, from

the chin to the throat and neck, over the breast and down the

flanks, while the central lower breast and abdomen are still in

the autumn-plumage of the previous September (Pls. LX XX VI.—

LXXXVIII.).

White terminal spots may, of course, bepresent on the breast and

abdomen. These are a local or an individual character which

will be mentioned later in dealing with varieties of feather-

pattern and coloration.

The flank-feathers of the hen in the full spring-plumage

show much diversity of pattern. This same diversity even in

the same individual bird has led to the belief that the pattern

may be changed in an unmoulted feather from the autumn-

plumage ari angement of red-brown and reddish-black finely

barred with lines of black to a much bolder barring of buff and

black. It hay been surmised, from the examination of single

feathers, that) the change commences in the centre of the feather

on either side of the shaft, and gradually produces another

pattern of a totally different colour, But can this be possible

1910.] PLUMAGE OF THE RED GROUSE. 1015

in a feather which has long been fully grown, and which has

presumably been long cut off from any blood or lymph supply

and which is as dead as if it had been shed? (Pl. XC.)

It is almost certain that rearrangement of the pigment or of

the pattern in this way is quite out of the question, and the

reasons for so thinking have already been discussed.

The legs and feet of the hen Grouse in April and in May are

very poorly feathered and the claws are very long (Pls. XCII.,

XCIII.).

In June the legs and feet are almost bare, and the claws begin

to drop off. The precise date of this shedding of the claws is

again really a part of the moult, and is, in consequence, equally

dependent upon the health of the bird. Sick birds which have

survived the spring mortality are always late in the shedding of

their claws, and equally late in the changing of their feathers.

The claws are shed, both in health and in disease, but once a

year, and the casting is synchronous as a rule with the dis-

appearance of the autumn-dress. The figures (Pl. XCIII.) by

which this process is illustrated require but little explanation.

The whole of the year’s growth of horny black nail becomes loose

on the soft and growing vascular matrix, and when quite ready

to be cast can be easily pulled off like a little cap. The young

nail beneath is at first soft and pink and vascular and very short,

but soon hardens and deepens in colour, and in a month or two

has grown to be a useful nail of horn. The transverse or

circular groove which is left at the point of detachment of the

old nail is quite a useful indication of age in cases where there

is a doubt as to a bird being over twelve months old or of the

year. ‘The presence of the groove showing that the claws have

once at least been shed is conclusive proof that the bird is more

than twelve months old.

In June there is another characteristic appearance in the hens,

namely the bare patch of abdominal skin which results from the

shedding of the abdominal feathers, grown in the previous

September. The loss of these feathers leaves a naked patch of

skin on the abdomen of a hen that has been sitting, and this

patch remains naked for the next few months.

The general character of a June hen in health is that of the

completed summer-nestling plumage, broad-barred buff and black

over all the upper- and underparts, excepting the abdominal

area, the lower breast, wings and tail. But it looks already

somewhat faded and worn; and it is quite probable that in

acquiring so perfect a plumage for sitting unnoticed on a nest

built amongst the heather, the economic absence of the redder

pigment in the feathers is in part a result of the acknowledged

fact that for longer and more trying use and for wear and tear

in feathers, darker pigments are required, whereas for the short-

lived and less exacting requirements of the summer-plumage

in the Grouse from April to June the buff and black feathers,

with very much poorer wearing qualities, are found to be sufficient.

66*

1016 * DR. E. A. WILSON ON THE [| June 14,

The accompanying figures of a few worn-out and moulting feathers

taken from a hen in summer-plumage, show how distinctly

better the black pigmented parts of the feather stand wear and

tear than the yellow parts (Pl. XCI.). Certain pigments have a

value, therefore, of a very practical nature apart altogether from

the more wsthetic point of view of attractiveness or the often

rather hypothetical view of assimilation to surroundings for

purposes of safety or to assist in obtaining food. He would be

unwise, however, who denied that all “three views may be

reasonably held to account for the very beautiful nesting-

plumage of the hen Grouse.

It very occasionally happens that the hen Grouse, instead of

retaining the redder. plumage of the previous autumn’s growth

on the abdomen until it drops off during incubation, grows an

almost universal spring-plumage of buff and black broad-barred

feathers covering the lower breast and abdomen as well as the

remainder of the body from head to tail. A skin showing this

condition is preserved in the National Collection, and there is an

almost equally perfect specimen in the Grouse Disease Inquiry’s

Collection, No. 919.

The more usual procedure is that the abdominal patch of

autumnal plumage is lost during incubation and is then quickly

replaced by a renewal of the autumnal feathers when the spring-

plumage is also being shed. ‘There remains, however, in the

majority of birds a very quaint growth of belated spring-plumage,

consisting of buff and black- barred feathers in two lines down

each side of the centre of the naked patch, as though, for

some occult reason, the intention to ere “ spring-plumage ”

feathers upon this avea had never been altogether lost. ‘This

peculiar persistence of belated intention shows itself as a pateh

of yellow feathers made up of the two linear areas of feather-

growth in the midst of a much broader area of the autumn

red pigmented feathers which one would expect to find all over

the abdomen (Pl. LX XXITX.).

It is conceivable that a small persistent remnant such as this,

having no obvious connection with the surrounding plumage at

the time or with the habits of the bird, or with the seasons, may

yet have something to do with the third or lost ‘* Eclipse” plumage

which is still to be found in the grey plumage of the Ptarmigan,

but is almost completely lost in the case of the Red Grouse.

In July the summer-plumage of healthy hens is much worn out,

frayed at the edges and very definitely faded, and the feathers

are already dropping out. On the chin, throat, and fore-neck

new red feathers of the autumn-plumage looking rich and dark

are already making their appearance. The back is as it was, but

faded, and the flanks are still conspicuously broad-barred with

buff and black; but the abdominal hare patch is now growing

new autumn-plumage feathers with great rapidity from the

centre outwards.

The primaries and secondaries have now commenced to moult,

1910.] PLUMAGE OF THE RED GROUSE. 1017

There may be in July, in the hen, as many as six or eight old

primaries in each wing with frayed tips, still to be renewed.

Precocious young birds of the year can readily be distin-

guished in July from hens in moult, because in the former the

dark yved-brown black-lmed autumn-plumage is on the flanks,

while the broad-barred buff and black, and rather worn-out

chicken-feathers ave in the centre of the abdomen. In the

adult the distribution is reversed, The broad-barred buff and

black feathers of the spring-plumage are on the flanks and the

redder fine-barred autumn-plumage is appearing in the centre

(compare Pl. XC. figs. a, ¢, g, h, k, & n, and fig. @).

In skin No. 284 there seems to be an unusual compromise in

a very backward hen, owing to disease. The compromise is

between the broad-barred and the winter-plumage with its

very fine black cross-lines (Pl. XC. figs. e, f, J, & m).

The legs and feet in July are naked and the claws are very

small; but the feathers are already showing through as small

nite, points, not to be confused with broken shafts, which

occasionally result from wear and tear in woody heather.

The plumage of the hen Grouse in August is well known. It

has already been pointed out in another paragraph how, owing

to the sudden increase of observation and owing to the ‘sudden

arrival of opportunities for examining an enormous number

of birds over the whole country during this month, there

has sprung up an idea that disease amongst Grouse has a

recrudescence in the autumn. Eut this is not the case. There

are probably fewer diseased birds on the moor in August than

there are in July. In July, however, they are never seen, and

therefore not suspected, but in August they are almost invariably

shot and carefully picked out of every bag, and owing to the

general interest in the question of disease, are almost always

noticed, and in a large proportion of cases publicly notified.

Hence the idea that disease makes a new start in August and

September.

As a matter of fact, however, these wasted birds are almost

certainly conyalescent. ‘They have been diseased, and they are

still suffering from disease, but thanks, in the majority of cases,

to their sex (for the bulk of the sickly autumn birds are hens)

they have avoided actual death im the two highest mortality

months, April and May. Once tided over those fatal months

the food and general conditions of life improve, the weight of the

cock goes up and the balance is again in favour of recovery for

him; and although with the hen the exigencies of inc vubation

and the cares of the family continue to “militate against her

recovery until June and even to July, she then rapidly begins to

put on we ight, and im August and September has once more

everything in her favour towards complete recovery.

Many sick- looking “ piners” are shot upon the moors in

August, but it should be remembered that in that month they are

recovering from disease, not growing worse ; and in September

1018 DR. E. A. WILSON ON THE [June 14,

many that were not up to the average weight the month before

will now be practically normal and probably indistinguishable

from healthy birds, were it not that their serious indisposition

of the preceding months has put them behind their fellows in the

matter of feather-change.

In August, therefore, the Committee’s collection of skins

contains a large number of examples of hen-birds showing

deferred moult and belated growth of feather.

The normal healthy hen Grouse in August has already put off

most of the broad-barred spring-plumage feathers of her nesting

dress, and is very much like the cock-bird in appearance, with

the same dark red-brown vermiculate or fine-barred plumage

underneath, white-flecked or not, as the case may be, and above

with a mixture of feathers new and old.

The legs and feet of a forward hen are already showing quite

a fair growth of white feathers, and the nails have all been shed.

The claws are therefore short and rather soft and the transverse

sulcus or groove at the point of detachment is clearly marked.

In the wings there may still be a number of primaries to be

changed.

In the convalescent “piner,” on the other hand, the case is

often very different.

She has still a most deplorably bleached and weathered

breeding-plumage on her, with worn-out feathers, frayed or ragged,

often with saw-toothed edges , Showing the different effect of wear

and tear on the pale buff- pigmented and black-pigmented parts.

The bird in this belated plumage has quite naked legs and feet

and long unshed nails, or may at the most be just showing the

points of a new er owth of feathers thr ough the skin ; and in this

state she is conspicuously shabby and ill to ) look upon in comparison

with the splendid plumage recently acquired by her healthy

sisters, and by the now almost universally healthy cocks.

But the point above all others to be remembered in this

connection is that this hen is convalescent, and still has a couple

of months of good food and good weather, as a rule, in which to

complete her convalescence before the winter comes.

If the spring outbreak of disease has been severe—that is, if the

general weather conditions of the preceding winter and early

spring months have been such as to conduce to a heavy and

widespread infestment of the grouse with the microscopic larval

trichostrongylus—then both cocks and hens will be equally in-

fested. But the breeding-season and the concomitant needs of

the two sexes are, from April onwards, quite distinct.

The result of this is that there is often a large mortality of

cocks in April and in May, and a much less marked mortality of

hens, probably in the proportion of seven or eight cocks to one

hen, but definitely occurring in the same two months.

There is no great mortality from strongylosis in any other

mouths of the year. Yet, notwithstanding this, after May the

cocks are suddenly relieved and rapidly recover, so that by August

1910.] PLUMAGE OF THE RED GROUSE. 1019

there are almost no sick cocks; whereas, after May, the hens have

still two very trying months to face, and although, thanks to the

abundance of food, they probably most of them or nearly all of

them succeed in struggling through, yet by August they have only

just been relieved of their more pressing cares and disabilities, and

so in August a very great number of them are still found to be in

a very poor condition. The moment the disabilities are removed,

however, they begin to recover, and it is this point which has so

constantly been overlooked. Sick birds in August are convalescent,

and however many there may be, it is not a sign of a new out-

break of disease, but a sign that the past spring infection was a

heavy one, though less fatal than it might have been.

At the end of their own specially critical periods, the cocks

have at any rate June, July, and August in which to pull them-

selves together by means of good food assisted by good weather ;

whereas the hens, at the end of their own specially critical period,

have August and September. Hence the preponderance of sick-

looking hens when the shooting begins, and the widespread, but

erroneous, belief in a recrudescence of disease in autumn.

To return to the further consideration of the hen’s change of

plumage in September, her finest feature is now undoubtedly the

clean new growth of bright red, or dark red or even black and

white-flecked feathers of the breast and abdomen, with their

narrow but even blacker markings.

The whole of this tract has now been shed, but is so quickly

overgrown again that no bare skin is visible save in the middle

area of the abdomen quite low down, where, as has been already

pointed out, the new growth is of belated feathers coloured as

in the spring-plumage, and therefore quite different to the rest

around them.

There is still, as a rule, no access of new red feathers on the chin

or throat of the healthy September hen, or at the most but a

feather or two. But in the sick hen there is still often a sprinkling

of the old red feathers of the preceding autumn-plumage, very

faded, amongst the faded buff and black feathers of the belated

spring-plumage.

On the back of even forward hens there is still a mixture of old

and new plumage, and the scapulars are often faded to some-

thing like black and white, and are badly frayed at the ends.

The wings have now almost completed their moult, but there

may still be a primary or two to change, even in very forward

birds.

The legs and feet are rapidly becoming feathered for the winter,

though in backward birds which have been sick they are still

quite bare, and now, of course, this feature is quite truly to be

taken as a sign of sickness and disease, though in a convalescing

bird.

In October one may find a very backward bird with as many

as three worn-out primaries in either wing to change; but, as a

rule, the wing is perfect, the primaries and secondaries and their

1020 DR. E. A. WILSON ON THE June 14,’

coverts all completelyynew, and in the tail the rectrices are full-

grown.

The legs and feet are now also fully feathered, though the

thickness of the growth increases as the winter cold comes on.

On the back the bird now looks fresh and richly coloured, from,

head to tail, but a close search will always disclose » number of

spring-plumage feathers which have still to be thrown off.

Beneath: the rich red-copper colour is gradually replacing all

the previous buff on the chin and throat. ‘The change hangs fire

a little on the neck and uppev breast, but it is still progressing.

Whereas on the lower breast and belly the rich red or darker

winter-plumage with its beautiful fine blaek cross-lines and pure

white flecks is a very striking feature.

There are, in the Committee’s collection of skins, a number of

examples showing the result of disease in deferring the moult ;

many of these birds, even in October and November have made

no effort to get rid of the old, faded and completely worn-out

spring-plumage. ‘The majority of these birds have been so

diseased in spring that they have not bred at all. The ovaries

have throughout he season shown no development at all, and

there are no signs, even in the earlier months, of the shedding or

development of ova or of any increase in size of the oviduct.

They have been trué barren hens.

In some cases (e.:%. in No. -1247) there are feathers of three

separ ate plumages is November. ‘There are the faded spring-

plumage feathers of the current year, but mixed up with them

here and there are new feathers of fie autumn-plumage coming.

and here and there exceedingly old worn feathers of the antumn-

plumage of the year before.

No. 1225, an October hen, shows exceedingly well how the

bare broody patch of the abdomen grows delay ed broad-barred

buff and black feathers, instead of the: fine-barred dar)» autumn-

plumage feathers which surround the patch. “Uhese'broad-barred

feathers appear in two parallel rows, breaking through the skin

of the broody patch on either side of the median line; this

growth is also well shown in a specimen at the British Museum

of Natural History, which shows the point exceptionally well

(Pl TeX’):

In November the chief alteration is the completion of the

antumn-moult and the assumption of the autumn-plumage. The

feathers of the upperparts have black middles, and are barred

with rufous-chestnut and ornamented with the characteristic

white or buff-coloured terminal spot.

In December the hen is in full autumn-winter plumage.

On the legs and feet she is well and thickly feathered; and on

the under Bide, the chin and throat are dark red, as well as the

fere-neck, and marked with broader black bars than upou the lower

breast and abdomen, where the marking is of the finer type and

the colour distinetly of the redder and darker autumn-plumage.

1910.) PLUMAGE OF THE RED GROUSE, 1021

Parr IIT.

Local Variation in the Plumage of the Red Grouse.

The following notes are the outcome of an attempt to find

some broad differences between Grouse from the Highlands and

Lowlands, East Coast and West Coast, English, Welsh, and

Lrish moors,

It seemed possible that, with a large series of skins of a species

peculiar to the British I Isles and at the same time so variable, one

might discover points in the coloration of the plumage or in the

size of the birds which could be attributed to some of the varying

physical conditions under which they live,

The artificial transportation of Grouse from one county to

another, generally from the southern moors to the northern,

often far removed from one another and with very different food

and climate, has no doubt to some extent confused the issue.

But this is a difficulty which will increase rather than decrease,

and it is even possible that the purity of the British breed,

present the only species of bird peculiar tc our Islands, may

before long be entirely lost by the introduction of a foreign species

on the mistaken supposition that the latter is freer from the

parasite of Grouse Disease. This is not the case, and even were

it so the species has already been intre7uced here and there,

and there has been some interbreeding wit.. our own Red Grouse.

There was at any rate some reason for thinking that, unless the

opportunity for collecting a series of ynure-bred British Red Grouse

skins had been taken by the Committee, the same wide oppor-

tunity might not have occurred again before the introduction of

foreign species had become popula 12

The Grouse Inquiry’s colle tion contains 580 skins of the Red

Grouse ‘& eluding 540 adult birds of both sexes and 40 chieks and

pullets. Thesc, however, cannot be taken all together’ in one

series. It is essential, for purposes of comparison, that the male

birds in their two plumages should be taken separately in two

lots and the females in a similar manner. Therefore the skins

have to be divided as follows :—

No. of skins.

Male birds in winter-plumage jc pith Shs ud) 1241

Male birds in autumn-plimage SuanaEe Bs PAD

Female birds in autumn-plumage ... ... ... ... 108

Female birds in summer-plumage ....... ... ... 71

Immature birds of the first 6 months Remini. vs eee)

The largest series of skins is therefore that of the male birds in

winter- -pluma ge, and it so happens that this set, both as regards

sex and plumage, is best adapted by its general uniformity to give

some result when arranged map-wise over a large outline. of

Scotland and England,

1022 DR. E, A. WILSON ON THE [June 14,

The following is an analysis of the greater part of the collection

of skins :-—

rol Se.

—————_

0A OF

Jan.to May.|June to Dec. Novy. to Apr.| May to Oct.) Totals.

Gaythngss \.cccccscaes ee 22 3 5 3 33

Sutherland ........... 12 4 2 8 26

IROSS iat. hctes sees eee 6 16 3 4 29

INVEINOSS o..ccceceesse. 30 14 5 8 57

INAITT esr atree eee es «ve 0 i 0 0 1

Moray it..kee eeneees 11 0) 1 @) 12

BAM tie exer eters scale 5 0 0 0 6

NS Os ae Beg ae 3 3 0 3 9

Kincardine ........... 2 0 2 0 4

Monfair Uist 1 1 0 3 5

Perultestsccs seeks ane 17 16 5 13 51

Argyll and Mull...... 19+1 5 0 9 34

Dumbarton ........... 14 4 3 1 22

Stirling. Rates.k 1 3 0 4 8

UT e aes eaetr es suic seek ae 1 0 0 0) 1

Haddington ........... 3 0 i 1 5

Midlothian ............ 5 0 0 2 7

Danark ie sakes 1 1 0 1 3

IAW iets ones ecoseeiket i 2 2 5 10

Arran 4 4 1 1 10

Peebles Sa roe 2 1 0 0 3

Selkinkie esos eee 7 4 0 4 15

ROXDUNE Diss cesscesanni es 0 4 1 2 7

(Dumotiries soos 0 cess eos 8 A 4 2 18

Kirkeudbright......... 5 3 1 0 9

Wiiehown Tn ccsiee...: 0 1 0 3 4,

Northumberland...... 9 1 4 2 16

Cumberland _......... uf 0 0 4 5

Durham eae. eee 0 0 1 0 1

Westmorland ......... 8 1 2 1 12

Lancashire ............ 2 1 1 0 4

fos Slane a eenme ea ce 28 13 14 13 68

Derbyshire ......... 1 0) 0 1 2

Wales Ean a ae 10 0 3 0 13

TrGlandtvagacntoczsenuces 0 0 0 1 1

510

—

co)

[or]

v=)

Having thus sorted the skins into lots which are sufliciently

uniform to allow of comparison, and having arranged one of

these lots, the cocks in their winter-plumage, for instance,

according to the locality from which they were obtained, it

becomes possible to make the following general deductions :—

1, That the general uniformity is very much more marked than

might have been expected considering the character for

variability which has always been attributed to the bird ;

the variability is lost in the mass, though it is visible in

individuals.

2. That, allowing for a good many exceptions, there is certainly a

greater tendency to blackness in the birds of the northern

Highlands than in those of the south.

Or, one may say that in passing from the north of

1910. ] PLUMAGE OF THE RED GROUSE, 1023

Scotland southward and westward there is an increasing

tendency to the bright red and dark red types of Grouse,

which culminate in the very characteristically bright

red bird of Wales and of the Midlands of England, in

which the predominating colour of the feathers of the

breast and underparts generally is red with fine broken

black cross-lines, and these are sometimes almost absent.

3. This gradual change from north to south of black, or red-

and-black to dark-red cocks, and farther south to bright red

cocks, is accompanied (speaking very broadly, for there are

many exceptions) by a loss of the white terminal borders

which characterize the feathers of the abdomen.

There is no doubt that the blacker birds of the Highlands in

the north of Scotland are more frequently white-spotted beneath

than the birds obtained farther south.

Nevertheless the white spotting is not confined to the blacker

or to the darker birds, for it may be quite a conspicuous feature

in the bright red birds of Wales and England, though in the

lowlands and in the north of England, especially in Yorkshire, it

is a rare character, only exceptionally met with.

Mr. Ogilvie-Grant, in his ‘Handbook to the Game Birds,’

1896, says, ‘The ordinary varieties of the male may be divided

into three distinct types of plumage: a red form, a black form,

and a white-spotted form.”

The red form, he says, “is mostly to be found on the low

grounds of Ireland, the west coast of Scotland, and the Outer

Hebrides”; and this statement is borne out not only by the

Inquiry’s collection of Grouse-skins, but by the interesting collec-

tion, made by Mr. 'T. E. Buckley, now in the Cambridge Museum.

Similar birds have been obtained in some numbers from the

following areas :—Caithness, Sutherland, the Lewes, and Inver-

ness-shire. From Stirling, Selkirk, Northumberland, and Wicklow

only one or two have been examined, but in Wales the red type

is almost always met with. Welsh birds are often most typically

and uniformly very bright red. Dumfriesshire also undoubtedly

produces a large proportion of the same red type.

Bright red birds are not commonly characteristic of Ross-shire,

Stirlingshire, or Northumberland, notwithstanding the fact that

an occasional example of this type may be found there. Dum-

bartonshire, however, is said to produce more birds of a bright

red type than any other, so also is Argyllshire, and both these

counties fall in with Sutherlandshire as forming part of the west

coast of Scotland. i

The following specimens have been figured to illustrate the red

form of the cock Grouse :—

Plate LXXXII. Upper surface, red type in winter-plumage.

Male Grouse, No. 630. Wales, 18.iii.07.

Plate LXXX. Under surface, red type in winter-plumage.

Male Grouse, No. 407. Glendoe, Inverness-shire,

7.xii.06,

1024 DR, E. A, WILSON ON THE [June 14,

Plate LXXXIV. Under surface, red type, changing from

winter- to autumn-plumage.

Male Grouse, No. 915. Forfarshire, 4.vi.07.

Plate XCVII. Under surface, variety of red type in

autumn- ‘plumage,

Male Grouse. Loch Sween, 19.vi1i.92. C.C.S. Parsons.

Brit. Museum (Nat. Hist.), 92.7.13.1.

The second or black form of cock Grouse is, according to Mr.

Ogilvie-Grant, rarely met with, most of the black buds being

mixed with the red, or the white-spotted forms,

In the Committee’s Collection there are a tew very good ex-

amples of the really black type, and they come from the following

areas :—Caithness, Sutherland, Perthshire, Dumbartonshire, and

Yorkshire.

More or less typical examples have also been obtained from

Ross-shire, Aberdeenshire, Morayshire, Kincardine, Stuling,

Fiteshire, and Lancashire.

At Neweastleton, the low-lying grassy moors are credited with

the production of the black type of Grouse, while the other types

are found on the higher heather-ground.

‘The following specimen has been figured to illustrate the black

type of cock Grouse :—

Plate LXXIX. Under surface: black type in winter-

plumage.

Male Grouse, No. 723. Perthshire, 4.iv.07.

“ The third or white-spotted form has the feathers of the breast

and belly, and sometimes those of the head and upyerparts, tipped

with white. ‘The most typical examples of this variety are found,

as a rule, on the high grounds of the north of Scotland.”

This statement is again confirmed by the Grouse Inquiry’s

Collection, although an occasional white-spotted bird makes its

appearance farther to the south. Iti is, however, the exception in

the south, whether it be of the red or black or inter mediate type.

The most marked examples of this white-spotted form have

come from Caithness, Sutherland, and Inverness, while Dumfries,

Perthshire, and Yorkshire have each provided one or two very

fair examples.

At Dingwall birds are said to be most commonly dark red or

black with white beneath.

At Serafton, Middleham, the majority have white beneath, and

all ave dark red or bright red.

The predominance of white beneath is quite conspicuous in

a geographical arrangement of a large number of skins, as a

character of the north of Scotland, especially throughout the

Highlands.

‘The following specimen has been figured, illustrating the white-

spotted type of cock Grouse :-—

Plate LXXXI. Under surface: white-spotted form,

Male Grouse, No. 1377. Morayshire, 24.iv.08,

1910.] PLUMAGE OF THE RED GROUSE. 1025

Turning next to the female Red Grouse, no less than five

distinct types are described by Mr. Ogilvie-Grant :—

1. The red.

2. The black.

3. The white-spotted.

4, The buff-spotted.

5. The buft-barved.

The difficulty in sorting out hen Grouse into these classes is

that a single bird may fall under three headings at once. A hen

Grouse may be at once Buff-spotted, White-spotted, and Red or

Black, for the white spotting is an independent character and

may oecur on any type in the autumn-plumage of the breast and

abdomen, and this may also be definitely of the red or the black

type.

In the Grouse Inquiry’s Yollection, the first or red type is well

represented from all parts of the country, and follows very much

the same distribution as the red type of the cock Gronse.

Red examples were procured from the following areas :—

Sutherland (3), Argyll (9), Arran (1), Dumbarton (1), Cumber-

land (1), Westmorland (1), and Wales (3), all bright red birds ;

Ross-shire, all dark red; Inverness-shire, (3) very bright red and

(3) very dark red birds; Aberdeen, (3) very dark ved birds ;

Stirling, (4) red birds, with very fine black markings on the breast.

Perthshire, Moray, Kincardine, Dumfriesshire, Kirkcudbright,

Northumberland, Durham, and Yorkshire were all represented

by red hens, generally of the dark red type.

The following specimen has been figured, illustrating the red

type of hen Grouse :—

Plate LXXXVIT. Under surface: red type changing from

winter- to summer-plumage.

Female Grouse, No. 226. Roxburghshire, 22.v.06.

The second or black type of hen is certainly, as Mr. Ogilvie-

Grant says, extremely uncommon, and only one or perhaps two of

the Committee's birds should be included under this heading.

hyo others are, however, so dark as to come with difficulty under

the category of red birds.

Caithness produced a really black hen bird (No. 418), the sex

of which could not possibly have been determined from its plumage.

It appears to be an old hen, which has assumed male plumage.

Specimen No. 338 from Inverness is almost as dark a bird,

and No. 559 is a very dark reddish-black bird.

No. 414 from Dumbartonshire is similarly a case in which there

seems to be more black than dark red.

The following specimen has been figured to illustrate the black

type of female Grouse :—

Plate LXXXV. Under surface: black type in autumn-

plumage.

Female Grouse, No. 418. Caithness, 31.x11.06,

1026 DR. E. A. WILSON ON THE [June 14,

The third or white-spotted type is less rare, and occurs,

according to Mr, Ogilvie-Grant, much as it occurs in the male.

In the Committee’s Collection it is well represented by birds

from Sutherland (No. 1336); Ross-shire, a bird of the red type

(No. 176); and Inverness (No, 329).

Tt was less to be expected that examples both of the red type

and of a darker type, bordering on the black, should have been

met with in Yorkshire, Lancashire, and Westmorland. Single

examples were also procured in Dumfriesshire and Kincardine.

There is a fine Irish example from Co. Mayo in the British

Museum (Natural History), No. 99.12.1.1.

The fourth or buff-spotted type of hen Grouse, said to be

“much the commonest and most usually met with, has the

feathers of the upperparts spotted at the tip with whitish-buff.”

This type is generally distributed, and the Committee’s Collection

includes examples from Caithness, Sutherland, Ross, Perthshire,

Ayrshire, Kincardine, Dumfries, Northumberland, Yorkshire,

Westmorland, and Lanieashire,

The fifth or buff-barred form, according to Mr. Ogilvie-Grant,

“is met with in the south of Ireland, and resembles in winter

(autumn- -plumage) the ordinary female in breeding plumage,

having the upperparts coar sely barred with buff and black. Very

little is known of this last ‘variety, owing to the difficulty in

obtaining birds, except during the shooting season,”

Unfortunately, the repeated endeavours of the Grouse Inquiry

Committee to obtain specimens proved unsuccessful, and one hen

only was obtained from Donegal. This bird (No, 1217) was a

very typical example of the buff-barred type, and it certainly

differed from anything procured either in Scotland, England, or

Wales.

Plate XCVIII. Under surface: buff-barred form in autumn-

plumage.

Female Grouse, No. 1217. Donegal, Ireland, 17.x.07.

The nearest approach to it was to be found in four hens from

Selkirkshire and in hens from Inverness-shire, which were more

accurately described as buff-barred than as buff-spotted.

Single examples from Lanark, Midlothian, Roxburgh,

Haddington, and Northumberland might be referred to the buff-

barred type, and the females from Yorkshire were all rather of the

buff-barred type, but none of these birds had quite the same

markings as the Irish example. See Plate LXXXVJ., Female

Grouse, in full summer-plumage, Scottish buff-barred type.

Plates XOVIT., XCIX., and CI. represent abnormal varieties

of the Red Grouse and are drawn from specimens in the British

Museum (Natural History). They are described on p. 1033 in the

explanation of the Plates.

Two points in connection with the practical distinction of old

Grouse from young, and of cock Grouse from hens, are of

perennial interest both to the g gamekeeper and to the sportsman,

1910.] PLUMAGE OF THE RED GROUSE. 1027

No discussion is more apt to produce different opinions than that

which arises upon the age or the sex of certain stages of moulting

Grouse either at the luncheon hour upon the moor or in the

game-larder when the day’s bag has been overhauled, sorted out,

and hung upon the hooks. It must be at once admitted that

there are individual cases, by no means very rare, in which

it is almost impossible to tell the sex with certainty until the

bird has been cut open and the internal anatomy examined. In

these doubtful cases the only way to settle the point is to cut

the bird open down the middle of the abdomen, carefully turn

over the whole of the intestines from the right to the left—that

is, from the bird’s left side to the bird’s right side,—without

tearing the attachments, and then, having exposed to view the

flattened reddish portions of the kidney which lie closely packed

on each side into the inequalities of the backbone and pelvis, to

see whether the uppermost portion has overlying it an ovary or a

testis.

In the breeding-season, and in a breeding bird, there can be

no doubt whatever as to the sex, for the ovary is a conspicuous

bunch of more or less developed ova in the hen; and in the cock

the testis is a conspicuous round white object as large as the

kernel of a good-sized hazel-nut on each side of the backbone.

There is but one ovary, and it lies always on the left side of

the backbone of the bird. There are two testes, one lying on

each side of the backbone, the left one generally at a slightly

lower level than the right.

This development of the ovary only on one, the left, side is the

reason for advising the examination to be made as described

above, on the left side always. One testis or the ovary cannot

then be missed.

If the bird examined thus is not breeding, as may often be the

case with birds found dead of disease in April and in May, the

discovery of the ovary is still a matter of comparative ease, and

the discovery of the testes even easier. The testes are always

somewhat enlarged in the spring months, whether the bird be

diseased or not, and they may be the size of a pea or larger, and

will generally be white. The ovary may be small, but will always

be like a portion of hard cod’s roe, in which the ova, though no

bigger than a pin’s head, are distinct and numerous. The

undeveloped ovary of an adult female Grouse would about cover

a threepenny piece, but is always rather long and triangular in

shape instead of circular,

The oviduct in a breeding hen is a large and conspicuous part,

and may, of course, contain an egg with the shell in course of

formation, being pigmented in preparation for laying. The

oviduct in a barren bird or in a hen not in the breeding-season is

a very much less conspicuous object, and will be less easily found

than the small and undeveloped ovary.

If no ovary is seen on examination, and a very small blackish,

or whitish, or parti-coloured object is found in its place which is

1028 DR. E. A. WILSON ON THE | June 14,

suspected of being a testis, the intestines must then be gently

separated from their attachments about the middle line of the

back, and the other testis must be sought for in about the same

position on the opposite side.

Even in a young bird the ovary shows ova with sufficient

distinctness to make doubt as to its sex an impossibility ; but in

a very voung male bird the testes may be so small, and, being

very often quite black, may look so unlike what is expected that

both should be sought for : and found before arriving at a certain

conclusion as to sex.

It is easy, if the intestines are roughly handled and the

attachments torn carelessly away, to carry away the testes or the

ovary from their proper position and to remove them with the

intestinal attachments. The peritoneal folds are delicate and

require careful handling, and they overlie the generative organs

and the kidneys; but a very little practice will condble anyone to

do the necessary dissection with certainty and to arrive at an

irrefutable diagnosis as to sex.

It may be said that there is no other infallible means of

arriving at the sex of a Grouse at certain times of the year, for

it has so often happened that experienced and careful game-

keepers, who have handled Grouse for a lifetime, have sent in a

paper filled in for a certain specimen as a cock, when the specimen

has turned out to be a hen, and vice versd. The mistake is

unavoidable and excusable, for in certain individual Grouse in

the autumn-winter plumage there is no reliable characteristic

in the feathering or in the supraorbital comb (Pl. XCV. figs. 1

& 2), or in any external part of the bird, by which the sex can be

distinguished. In most Red Grouse, even in the vast majority,

the confusion of sex is not possible, for it is a matter of common

knowledge that for a great part of the year the cock and the

hen are so wholly unlike one another as to make it dffiieult for

anyone who did not know the birds to believe them to be of the

same species. Even in the summer months, when the cock puts

on a plumage closely simulating the breeding-plumage of the hen,

there is a difference in the general tone and colour, and confusion

is not likely. In the: autumn and winter it is comparatively easy

to mistake the sex of some individuals, when the nen has put on

her autumn-plumage for the winter and the cock has put on his

winter-plumage; certain individuals of opposite sex are then

indistinguishable, even to the practised eyes of the experienced

gamekeeper.

Generally speaking, the feathers of the head and neck give the

best indication as to sex in the autumn-winter plumage. In the

male the red colouring is, as a rule, far more unifor m than in

the female. In the male there is, as a rule, an absence of black

markings on these red feathers, except on the upper part of the

head, on the.crown, and nape of the neck. The cheeks are

generally a clear bronze- or chestnut-red colour; so are the

feathers of the chin, throat, foreneck, and upper breast, giving

1910.] PLUMAGE OF THE RED GROUSE. 1029

the bird a very rich uniform red colour all over the head and

neck. In the hen, as a rule, the whole of the feathers of these

parts are crossed by narrow black bars, which give her more of

the mottled and broken colouring, which the cock bird begins to

assume in the early summer when he puts on the first feathers of

his autumn-plumage.

The feathers of the chin are a very useful indication of sex

from August to November, practically throughout the shooting

season, for the chestnut-red feathers which can be found on the

chin of the cock Grouse in every month of the year will be

sought for in vain in the hen from August to November. Even

in December and January they are so imperfectly red as compared

with the same red feathers in the male that one may almost say

that red feathers are to be found on the chin of the hen only

from February to July, when they become conspicuous on

account of the contrast in colour with the increasing yellowness

of the breeding-plumage of the hen. These red feathers persisti

in the hen exactly as do the feathers of the lower breast and

abdomen, from her previous autumn-winter plumage.

This persistence of winter-plumage feathers on the chin, lower

breast, and abdomen is common to both cock and hen; but in the

cock they remain, as a rule, until replaced by the following winter-

plumage, persisting throughout the autumn-plumage change ;

whereas in the hen they are persistent only to June or July, and

are entirely replaced during the autumn-change.

Even when the autumn-plumage is put on, the yellow feathers

of the preceding breeding-plumage are almost always to some

extent persistent, and they are to be found in the chin of the hen

bird even though the throat and neck may be unusually red and

therefore unusually like those of a cock bird.

From January to May there is no possibility, as a rule, of con-

fusing the sexes. In June and July it is unlikely, but in August

and onwards to December it is sometimes quite easy to mistake

them, and the best guide is the persistence of feathers of the

preceding plumage such as occurs upon the chin in particular,

recollecting that the dominating plumage of the male is the

winter-plumage, while that of the female is the summer- or

breeding-plumage.

In the autumn, especially from September and October onwards,

there is the additional difficulty of distinguishing old birds and

young.

“Young birds in July resemble the adult female in breeding-

plumage in their general colour, but the flank-feathers of the

adult plumage begin to appear about this time. By the month

of November the young are generally not to be distinguished from

the adults.” This quotation is from Mr. Ogilvie-Grant’s account,

There is one sign of age in the majority of birds in the shooting

season, if it has not become obliterated—-namely, the mark of recent

shedding across the claws. Very often one may find the nails

or claws stilladhering to the toes, though ready to drop off, so that

Proc. Zoou, Soc.—1910, No. LX VII. 67

1030 " DR. BR, A. WILSON ON THE [June 14°

a gentle application of force removes them like small caps, ] eaving

the new shorter claws beneath, each marked by a groove where

the old claw was attached. This groove persists often for some

little time, and is as infallible a sign that the bird is at least

a year old, as the presence of the claw itself ready to drop off.

Young birds of the year do not shed their claws and therefore

“never have the groove.

There is another method of determining a bird’s age which is

‘often used as a rough indication upon the moor, namely, to pull

out the third primary-feather of the wing at its distal end. If

blood can be squeezed from the quill it is considered as a sign

that the bird is of the year. If no blood can be squeezed and the

more than a year old.

This reasoning is based upon the Selick ing facts: hte a

chicken produces its first set of primary wing- feathers they are all

very thin and weak, and soon become frayed out ; they have rather

‘narrow pointed ends, and are blackish-brown, ‘mottled with in-

complete buff bars. Nine of these little feathers can be counted in

series, and the weakest of all is the ninth counting from the distal

to the proximal end. The growing power, as judged by the size of

the root-sheaths of these feathers, increases from the ninth distally,

so that at one period of growth (F) the feather No. 4 is the longest,

then a little later (G) No. 3, then still later (H) No. 2. But No. 1

remains permanently shorter than No. 2.

By the time Nos. 1 and 2 bid approaching full prolate the

weak chicken-feathers Nos. 9, 8, 7, and so on have been shell: and

in their place have appeared strong feather-roots growing strong

round-ended uniform black primaries, instead of the weak, mottled,

more narrowly pointed chicken-primaries.

This shedding and replacement continues as far as No. 3 of ihe

chicken-primaries, but when No. 3 falls out Nos. 2 and 1 are

found to have grown into such long and strong feathers as to fall

into the series of new replacement feathers, and so they remain,

now fully grown, but can be recognised (KX) by their more pointed

ends having rather more of the buff markings on them than have

the remainder of the primaries just grown from No. 9 to No. 3’.

No. 3', moreover, being the latest of the whole series of primaries

to have emerged, is for a time the shortest one ; and, even when

as long as No. 2, will be still the last one grown and will therefore

upon withdrawal produce blood at the quill end when squeezed

between the thumb and the finger.

The following diagrams (text-fig. 150) show very roughly this

sequence of growth in the primaries of chick and pullet.

This sign of blood in the quill of the third primary is not an

infallible sign of youth, for it is evident that as soon as the feather

finishes its growth the quill becomes as hard and dry and bloodless

as all the others. ‘The only indication will then be a slight

difference in the shape and contour of the last two feathers as

indicated above, ,

1910.] | PLUMAGE OF THE RED GROUSE, 1031

Moreover, in September it is easy to find birds obviously adult

with claws attached, but on the point of being shed, and having

all the primaries moulted except the two most distal ones. The

Text-fig. 150.

4120 Daus old,

VAN

\ \\ ve {

\ e'

Diagrammatic representation of the growth of Primaries in the young Grouse.

third then will be found to be a short feather actively growmg

and if it is pulled out the growing root will be full of blood.

67*

1032 DR. E, A. WILSON ON THE [June 14,

Therefore not every bird that gives this sign is necessarily a

bird of the year.

The condition of the ends of the primaries will, however, be a

fair indication of age, and especially of the two most distal

feathers.

Another sign often used to test the age of a bird is the strength of

the lower jaw. The weight of the bird is allowed to hang with-

out support by holding the tip of the Jower bill only. The bone

of an old bird’s jaw easily stands this test, but the soft jaw of a

young bird of three or four months cannot carry its weight, and

the jaw either bends or breaks.

Yet another test is often tried by trying to crush in the skull

with the thumb and finger: in the young bird the soft skull

gives way readily, in the old bird it requires very considerable

force.

In judging at a moment’s notice whether a young Grouse chick

which rises to the dogs on the 12th is of a shootable age and

growth or not, the usual rule is probably as good a one as can be

found—namely, that there should be a very clear view of black

tail-feathers before the young bird is fit to kill.

It is only necessary, however, to look at a few young Grouse

chicks of various ages to be convinced that more than a little of

the black rectrices should appear before the bird is shot. Probably

most sportsmen will be guided better by the strength of the bird

upon the wing than by the black tail-feathers, however clearly

visible in a half-grown ‘chick.

In dissection, the age of an old bird is apparent, perhaps, as

plainly upon the table as elsewhere. The fibrous tissues all

toughen with age and use, and the bones become harder.

The grits of ‘the gizzard in an old bird seem to be larger and

more worn into rounded pebble shapes, and the reason for this

has been fully discussed elsewhere.

The question, therefore, as to whether a bird is less than a year

or more than a year old, may be answered, but it seems almost

impossible to judge more exactly the age of a bird by any sign

to be discovered either externally or internally.

EXPLANATION OF PLATES LXXVIII.—CI.

Pru. LXXVIII. Pair of Red Grouse in summer with young chicks.

, LUXXIX. Male grouse in full winter-plumage ; black type, underside. No, 723.

Perthshire, 4.iv.07.

» LXXX. Male grouse, in full winter-plumage; red type, underside. No. 407.

Glendoe, Inverness-shire, 7.xii.06.

,, LXXXI Male grouse, white-spotted bird of the red type; underside. No. 1877.

Morayshire, 24,iv.08. 173 oz.

"UXYXII. Male grouse, in full winter-plumage, red type carrying a few black-

centered feathers of the previous autummn-plumage ; upperside. No. 630.

Wales, 18.iii.07.

,, UXXXIIL. Male grouse, showing very markedly the beginning of the autumn-

plumage upon the head and neck; upperside. No. 1481. Inverness-

shire, 8.v.08. 20 ozs. ;

1910.] PLUMAGE OF THE RED GROUSE. 1033

Px. LXXXIV. Male grouse, changing from winter- to autumn-plumage ; red type,

»”

”

»”

”

underside. No. 915. Forfarshire, 4.vi.07.

LXXXV. Female grouse, in autumn-plumage, black type; underside. No. 418.

Caithness, 31.xii.06. 214 ozs.

LXXXVI. Female grouse, in full breeding-plumage, Scottish buff-barred type ;

upperside. No. 226. Roxburghshire, 22.v.06. 20} ozs.

LXXXVII. Female grouse, changing from winter- to summer-plumage, red

type; underside, No, 226. Roxburghshire, 22.v.06.

LXXXVIII. Female grouse in full summer-plumage, underside. No. 217.

Lanarkshire, 12.v.06.

LXXXIX. Female grouse showing the bare broody patch of skin on the abdomen,

and the beginning of a new growth of buff, broad-barred feathers in a

double line down the abdomen; underside. No. 996. Argyllshire,

14.vii.07. 16 ozs.

XC. Female grouse, red type, feathers from flanks. Natural size.

Figs. a & c (from No. 1864), 9 & x (from No. 226), & k (from No. 632), are

varieties of the spring’ flank-feathers.

Fig. 6 (from No. 575) is a flank-feather from a very black hen.

Fig. d (from No. 1864) is an example of what is termed fine-barred, dark-red

winter-plumage, with narrow black bars or lines on rather dark rufous

chestnut, the latter being slightly bleached towards the tip.

Figs. e & f (from No. 1864) and J & m (from No. 664) illustrate intermediate

stages of coloration, the feathers probably having broken through the

skin when winter-conditions prevailed, and having completed their

growth under summer:conditions.

Figs. i & n (trom No. 664) illustrate the reasoning: upon which is based

the view just mentioned ; of these two feathers there is no doubt that

nm was being grown much later than 7, and therefore more in summer-

conditions, producing summer breeding-plumage:

XCI. Female grouse, red type, upper tail-coverts showing effeet of wear; the

breakage has occurred in each case at the parts which are unpigmented or

show the smallest amount of dark pigment. About 14 times natural size,

XCII. Grouse, red type, feet showing winter-plumage.

Fig. 1. Right foot showing new winter-feathers and new nails (No. 1277).

» 2. Left foot showing full-feathered winter-plumage.

XCIII. Feet of grouse, showing replacement of nails.

Fig. 1. Right foot (No. 1148) with old nails ready to be shed:

pe ae ss im median vertical section.

» 3 Left ,, (No.1167). 34a, old nails; 34, new nails ;3c, shed nails.

» 4 Right ., (No. 1185) with new feathers and new nails.

XCIV. Heads of Grouse.

Fig. 1. Female. No. 61, 22.1.06.. Showing the black markings on the throat..

5, 2. Male. No. 545, 17.1.06. Showing the uniform chestnut throat.

XCV. Supraorbital comb in Grouse and Ptarmigan.

Fig. 1. Female Grouse (No. 1269; 8.11.08.).

» 2 Male » (No. 1268, 8.11.08.).

» 3 Male Ptarmigan.

XCVI. Supraorbital comb in Blaek-cock.

XCVII. Scottish male grouse, showing abnormal erythrism; underside. Loch

Sween, Argyllshire. No.19. 19.viii.92. British Museum (Nat. Hist.).

XCVIII. Female grouse of the Irish buff-barred type; underside. Donegal,

Ireland. No. 1217. 17.x.07. 22 ozs.

XCIX. Female grouse, abnormal yellow variety ; underside. Stornoway, Lewis,

Hebrides, 30.x.1900; British Museum (Nat. Hist.).

C, Female grouse, grey variety: underside. Perthshire, 6.x.89. British

Museum (Nat. Hist.).

CI. Female grouse, grey variety ; underside. Type of L. persicus G. R. Gray,.

but probably from Perthshire. British Museum (Nat. Hist.),

ibe i

mrad aan

i, 3< im

wa ¢ |

Feit tend

; bi

y ay, |

“4

Lo) %

Pid

‘

i at's

‘tap x

Acanthephyra, 646, 647.

debilis, 640, 641, 642,

643, 644, 645, 646,

650, 651.

pellucida, 640, 643.

Aeanthias, 621.

Acipenser, 595, 621.

sturio, 593, 626.

Acomys

cahirinus, 662, 663.

Addax, 548

naso ~ maeulatus,

910, 911,

Adelea

ovata, 679.

Adenota, 917, 984. .

908,

ob, 945, 930, 981, 934, |

976, 977.

/£goceros, 907.

ASluropus, 987.

/Epyceros, 845, 897, 907,

979, 977.

melampus, 891, 918,

919; 920,

Aglaophbenia

crispata, 799.

Alce. 967, %68, 971, 978,

985. 986.

alces, 958, 959,

machlis, 669.

Allodon, 764.

Amalopenzus, 640.

Amia, 596, 597.

cabva, 595, 626.

Ammodorecas, 846, 897

918, 984.

-clarkei, 917.

Ammodytes, 608.

tobianus, 607, 626.

Ammotragus, 866.

lervia, 862, 863.

Amieba

neleagridis, 703,

INDEX,

Ameceba

(Entameeba) dagopodis,

692, 702, 703, 708.

Anguilla, 602, 621, 625,

624.

vulgaris, 600, 626.

Antenella

secundaria, 800, 802,

822.

Antidoreas, 897, 984.

euchore, 898.

marsuptalis, 892, 893.

Autilocapra

americana, 935.

Antilope, 897, 908, 984.

bubalus, 9C0.

cervicapra, 849, 894,

895, 975, 976.

eleotragus, 913.

euchore, 898.

lalandia, 913.

leucoryx, 907.

oryx, 907.

perpusilla, 867.

pygarga, 893.

pygmed, 867.

scripta, 929.

sylvatica, 929.

Anurea

aculeata, 793, 794, 798.

— valga, 793, 799,

796, 797, 798.

.cochlearis, 793, 797,

798.

Argas

persicus, 699,

Avisteus

coruscans, 639.

Asplanchnt

brightwelli, 793 _

intermedia, 794, 198.

Atherina, 608, 622.

hayerti, GOT.

Axis, 939, 971, 985.

avis, 940, 948, 949,

950, 977, 981, 982.

porcinus, 950, 977, 981,

982.

(Hyelaphus) porcinus,

949, 950.

Bacterium

pullorum, 716.

septicemie gallinarum,

716.

Balena

biscayensts, 772, 773,

791. :

mysticetus, 771, 773,

731

| Balanophilus

unisetus, 786.

| Baleenoptera

borealis, 772, 773, 784,

786.

latirostris, 786.

musculus, 769, 772-

777, 781, 7838-787,

~789, 790, 792.

physalus, 785.

rorqual, 783.

rostrata, 783.

sibbaldii, 769, 772,773,

774, 775, 784, 786,

787.

Balistes

capriscus, 620, 627.

Belone, 591, 608, 621.

acus, 606. ;

Bibos, 932.

Blastocerus, 970, 971,

hezoarticus, 966.

campestris, 966.

dichoiomus, 966.

paludo.us, 966.

XAli INDEX.

Blennius, 618. Budoreas, 853, 857, 858, | Catodon

Bonneviella 985. colneti, 788.

taxicolor whitei, 856.

tibetanus, 852, 856.

grandis, 817.

Boocercus, 932.

euryceros, 931.

Bos, 928, 932, 977, 983,

Centriscus, 621.

scolopax, 605.

Cephalophus, 926, 982.

Callidina sp., 795. burchelli, 867,

986 Callionymus

bubalus, 934.

caffer, 984, 995.

— brachycerus, 993,

996, 997.

— cottoni, 996, 997.

-— nanus, 993, 995,

997.

— planiceros, 993, 995,

997.

— simpsoni, 993, 994,

995.

— thierryi, 992, 993.

— typicus, 997.

frontalis, 933.

gaurus, 933.

sondiacus, 933, 234.

taurus, 933.

Boselaphus, 845, 932,

934.

tragocamelus, 926, 927,

928.

Box, 618, 621.

salpa, 612, 615, 626.

Brachionus

angularis, 793, 798.

bakeri, 793, 797,

798.

hidentata, 797, 798.

caudatus, 793, 797,

798.

Jalcatus, 793, 797,

798.

Jorficula, 793, 797,

798.

militaris, 798, 796,

798.

pala, 793, 794, 796,

798.

— dorcas, 796, 798.

rubens, 793, 798.

tetracanthus, 708.

Bubalis

hoselaphus, 900.

caama, 900, 901, 902,

903, 905, 906.

cokei, 9U2.

jacksoni, 902.

lelwel, 9U2.

lichtensteini, 902, $06.

major, 902.

maurelanicus, SOO,

swayner, 902.

thierryi, 992.

tora, VO2,

lyra, 617, 627.

Calycella

nuttingi, 814.

oligista, 800, 802, 813,

825.

syringa, 813.

Cainelus, 848.

bactrianus, 9738.

dromedarius, 972,

973.

Cam panularia

attenuata, 831.

, eorrugata, 800, &01,

809, 816, 826, 830.

noliformis, 880.

raridentata, 799, 800,

801, 809, 810.

serrulata, 831.

Candona, 736.

Candonella, 741, 742.

Candonopsis, 741.

Cantharus, 618.

vulgaris, 612.

855, 863, 899, 982,

984.

egugrus, 865, 866

cylindricornis, 864,

865, 866.

Falconeri, 865, 866.

hircus, 865.

sthirica, 866.

958, 959, 960, 964,

967, 168, 971, 978.

capredlus, 997.

Capricornis, 853, 857,

085.

argurochetes,

856.

erispus. 854, 855.

sumatrensis, 856.

thar, 840, 858,

975.

a= Jamrachi, 855.

| Caprovis, 859.

Cariacus, 967. 968.

americanus, 962.

| Oathypna

leontina, 795, 798.

luna, 794, 795, 798.

ungulata, 799, 798.

Catodon

| australis, 78s.

855,

856,

Capreolus, 936, 939, 956,

dorsalis, 768, 871, 872,

873, 874, 875.

grimmi, 867.

maxwelli, 867, 872,

873, 874.

monticola, 867.

nigrifrons, 872, 873,

875.

perpusilla, 867.

ygmeus, 867.

iced. 867, 872,

873, 876.

Cephenomyia

ulrichii, 669.

Cepola

rubescens, 611.

Cercoleptes

caudivolvulus, 768.

Cervicapra, 917, 918.

arundinum, 918.

Julvorufula, 913.

redunea, 912, 913, 914,

916, 930, 984.

| Cervulus, 939, 947, 958,

Capra, 845, 852, 853, |

964, 967, 968, 971,

976, 982, 985.

bridgemani, 989.

crinifrons, 991.

fee, 991.

lachrymans, 989, 990,

OS].

muntjac, 950, 952, 953,

954, 956, 981.

reevesi, 954, 955, 956,

989, 990, 991.

sclateri, 989.

Cervus, 968, 971, 972,

978.

affinis, 942.

aristotelis, 840.

canadensis, 942.

— wardi, 987, 988,

989,

capreolus, 957.

cashmirianus, 942.

~- macneilli, 987.

duvauceli, 944, 946.

elaphus, 94), 942, 943,

944, 945, 946, 948,

957, 967, 982.

eldi, 945, 967, 969.

maral, 940, 942, 944,

945, 948, 950.

sika, 944, 945, 962.

unicolor dejeani, 987,

Cervus

wallichii, 942.

(Panolia) edi, 9H,

945, 948, 970.

(Pxeudaxis) sika, 942,

943, 948.

(Rucervus) duvauceli,

941, 943, 948, 950.

Chlamydoselachus, 624.

Chrysochloris, 763.

Cladocoryne

haddoni, 800, 891,

805.

Clarias sp., 598, 600.

Clupea

ptlchardus, 597, 626.

sprattus, 597.

Clytia

Johnstoni, 830.

noliformis, 826, 830,

831.

serrulata, 826, 831.

Cnidoglanis

megastomi, 599.

Coccidium, 708.

avium, 712.

cuniculi, 710.

lacazei, 679.

oviforme, 677, 679.

perforans, 677.

pfeifferi, 710.

proprium, 679.

tenellum, 674.

(Himeria) evium, 722.

(—) pfeiffert, 722.

Coendu

prehensilis, 980,

villosus, 980.

Ceenolestes, 766.

Conchoderma

auritum, 789.

virgatum, 789.

Conger, 621.

vulgaris, 602, 626.

Connochetes, 976.

gnu, 900, 904, 905,

906, 982.

Copeus

copeus, 793, 798.

Corydendrium

corrugatum, 803.

dendriformis, 803.

parasiticum, 803.

sessile, 800, 801, 802,

804, 806, 807, 825.

Crinia

levis, 627, 630, 631,

632, 634.

signifera, 627, 628, |

629, 6380, 631, 633,

654.

INDEX,

Crinia

tasmaniensis, 627.

Cristiceps, 618.

argentatus, 617, 627.

Cryptobia

helicis, 671.

Ctenacodon, 764, 765,

766.

Cuspidella

costata, 800, 802, £14, |

825.

Cypretta, 742.

| Cypria, 732, 738, 739.

Cypridinella

dlosvagi, 755.

Cypridopsella, 741, 742.

Cypridopsis

aculeata, 742.

bidentata, 746, 756,

759.

congenera, T47, 756,

costata, 748.

cunningtoni, 744, 748,

756.

gibha, TAT, 756, 759.

globulus, 742.

helvetica, 742.

monodonta, 745, 746,

756, 759.

ohesa, 742.

obliquala, 742, 756,

758.

picta, 742.

pusilla, 744, 755, 756,

759.

serrata,

S1a9:

sinuata, 748, 744, 745,

747, 748, 756, 759.

tumidula, 746, ‘747,

756, 759.

vidua, 741, 742.

villosa, 742, 744.

Cyprinotus, 751.

JSossulatus, 752.

Cypris

incongrueus, 751.

pyxidata, 751.

745, 756,

Dama, 9389, 943, 952,

9538, 956, 958, 971,

978, 980, 982.

dama, 950, 951, 954.

Damaliscus, 902, 907,

982. s

alhifrons, 900, 903,

998.

hunteri, 908.

jimela, 998.

Xxil

Damaliscus

korriguim,

993.

— jimela, 902, 903.

— tiang, 902.

lunatus, 900, 984.

phalius, 998, 999.

pygarga, 900, 903,

998.

902, 906,

Dasypus, 766.

Dendroceelum

lacteum, 670, 671.

Dicotyles, 979.

collaris, 980, 981.

Didelphys, 766.

Diphasia

digitalis, 800, 801, 802,

809, 821.

Dipneustes, 624.

Diurella

stylata,, 796, 798.

Dorcelaphus, 966, 971.

americanus, 939, 962,

964, 965.

— savannarum, 962.

Dorcotragus, 879, 881,

897, 982, 984.

megalotis, 878.

Doryceros, 964.

Dryotriorchis

spectabilis, '768.

Dynamena,

tubuliformis, 832.

| Kcheneis

naucrates, 616.

Echidna, 765, 766.

| Echiodon

dentatus, 604.

| Ectatops

rubriaceus, 839,

Eimeria

avium, 714,

716.

Jaleiformis, 677.

stiede, 677, 679, 710.

(Coecidium) avium,

672-692, 706, 710,

721, 729, 731.

(—) cuniculi, 710.

715,

(—) schubergi, 673,

677, 679, 686.

Elaphodus, ‘ 971, 980,

985, 991.

cephalophus, 955,

956.

michianus, 955, 9d6.

| Elaphurus, 946, 947,

967, 968, 970, 972.

davidianus, 945,

XXIV

Engrantis, 597,

encrasicholus, 596,

597.

Entamceba

coli, 703.

histolytica, 703.

Equus

chapmanni, 837.

Erinaceus, 765.

Eucervus, 966, 970,

71.

Eucehlanis

hyalina, 795, 798.

longicaudata, 793.

oropha, 79S, 748.

propaiula, 193,

798.

Budendrium

attenugtum, S00, 801,

804, 805, 815.

capillare, 805.

generalis, 800, 801,

R05.

galdivense, B04,

parvum, 816, 828.

Eulyes

amanda, 839.

795,

Faleo

nisus, 706.

Felis

bengalensis, 837.

caracal, 837.

concolor, 634.

yeofroyi, 769.

Fiber

sibethicus, 634.

Fierasfer, 619, 624.

acus, 604.

dentatus,

626.

604, 625,

Gadiculus, 609, 621.

argentatus, 608, 626.

Gadus, 621.

minutus, 608, 610,

Galaxias sp., GO4.

Galera

harbara, 768.

Galeus, 621.

Gasterosteus,

693,

aculeaius, 605.

Gazella, 897, 918, 984.

arahica, 887.

hennettii, Uf

R92,

colus, B87,

591,

889,

}

INDEX.

Gazella

cuvieri, 887, 889, 890,

892,

dama ruficollis, 89).

dorcas, 887, 888, 889,

890, 976.

qutturosa, 887, 893.

kevella, 887.

maried, B37.

mhor, 8&7.

muscalensis, B87, 839.

pelzelni. S87. :

picticaudata, 891, 898.

ruficollis, 887.

rufitrons, 887, 975.

semmeringsi, 887.

subgutturosa, 843, 887,

889, 890, 892,

893.

Gennadas, 640.

Georychus

zechi, 589,768.

Giraffa

caimelopardalis,

935, 956,

reticulata, 669.

Gobius

paganellus,

627.

Goniodes

telvaonis, 699,

Gorgon, 976.

albo-jubaius, 906.

taurinus, 9V0, 906.

Grus

japonensis, 837.

669,

615, 616,

Guevei

equatoridlis, 870, 875,

876.

maxwelli, 867, 868, |

869, 870, 871, 875,

. 876.

melanorheus, 868, 870,

871, 873, 876.

monticola, 870.

Guttera

pucherani, 706.

Gy paetus

barbatus, 769.

Hiemoproteus

mansoni, 692,

705, 707.

Halecium

beanir, R08.

parvulum, 808.

simplex, 800, 801, 807,

697,

tenellum, 800, S01,

ROR,

Halicornaria

hians lara, 827, 835,

836.

—, profunda, 825,

Halocordyle

coupert, 807.

Hebella :

calcaraia, 800, 801,

— contorta, 810,

crateroides, 800, 80),

R10,,

Hemicypris, 751.

Heuiitragus, &55, 863,

982, 983, 984.

hylocrius, 866.

jemlaicus, 865, 866.

Herpetocy pris, 750.

Heterocarpus

alphonsi, 639.

Heterocypris

dubia, TFA,

THO. 3

Heterodontus, 621.

phillippi, 62.

796,

Hippocamelus, 968, 969,

970, O71...

antisiensis, 966.

Aisuleus, 968.

Hippocampus

brevirostris, G06.

Hippotragus, 907, 908,

911, 982.

barhata, 909.

equinus, 909, 910.

veucophea, 909.

niger, IVY, 910, 976,

N77.

Hoplophorus

sp., 640.

grimaldii, 640,. 646.

Hydrelaphus, 956.

Hydropotes,

973, 985.

inermis, WO6.

Hyelaphus, 970, 985.

porcinus, 939,

Hyla

arhorea, 682, 633.

aurea, 027,

635, 634.

ewingti, 627, 628,

630, 622, 6388,

6p4.

peronti, 6:8,

Hylobates

lar, 768.

Hymenolep’s

microps, 730.

Hymenopus

hicornis, 839,

Hystrix, 764. ¢

952, 957,

628, 629, .

Tdia

pristis, 799, 800, 801,

802, 805, 806, 809.

810, 813, 815, 816,

817, 820, 822.

Tlyoeypris

alta, 753, 756, 760.

angulata, 753.

australis, 752. |

‘ gibba, 753.

propingua, 752, 756,

760.

Txodes

ricinus, 698, 705.

Kobus,

984.

defassa, 916.

ellipsiprymnus, 916,

kob, 916.

marie, 916. |

917, .918;,.,. 928;

Labrus |

festivus, 615, 626.

Latfvéa

contorta, 810.

dispolians, 816.

serrata, 8U0, 802, 814, |

815. |

venusta, 800, 801, 802, |

815, 816, 817, 825. |

Lagopus

scoticus, 692, 706, 707, |

731, 1000-10383.

Lama, 843.

vicugna, 980,

981.

Lepadogaster, 618.

Leptocephalus, 600.

brevirostris, 601, 626.

Leptonychotes

weddell?, 589.

Leucoeytozoon

caulleryi, 694, 695.

-lovati, 692, 693, 694,

696, 697, 705, 707,

729,

ziemanm, 695, 706,

Limnias sp., 795.

Limnicythere

obtusata, ‘754,

760.

Limnodynastes

dorsalis, 627,

O54.

' peronti, 628.

tasmaniensis, 627, 628,

629, 630, 633, 654. -

972,

756,

629,

| Lithocranius, 897, 918,

INDEX.

Limnotragus, 929, 945, |

gratus, 330, 931.

—- selousi, 931.

selousi, 837. It

Lithobius

Jorficata, 673.

984.

walleri, 895, 896.

Lobodon

carcinophagus, 589.

Lophius

piscatorius, 619, 627.

Lutra, 978.

Lycaon

pictus, 857.

Lycostomus

gestrot, 840.

Lytocarpus

pernarius, 799, 800,

802, 822.

pheniceus, 800, 801, |

802, 810, 817, 823. |

Lytoscy phus

marginatus, 811, 816.

Macropus, 7695.

Macroscelides, 765.

Madoqua, 881, 982,

984.

phillipsti, 874, 876, |

877, 879, 880.

saltiana, 876.

Mantis, 838.

Mastacembelus, 592, 619,

623, 624, 627.

Mazaina, 960, 966, 969,

971, 981, 985.

bricenti, 964, 965.

nemorivagus, 939, 962,

963, 964.

Meganyctiphanes

norvegica, 786.

Megaptera

longimana, 772.

Melophagus

ovinus, TO4, TOD.

Mephites, 594.

Microtus

inez, 636, 637. |

Johannes, 637, 688.

mandarinus, 637, 638. |

melanogaster, 637. |

(Eothenomys) — 22’, |

637.

Meeritherium, 764.

Molva, 609, 621.

vulgaris, 608, 626,

Monachus

alhiventer, 768,

XXV

Monocercomonas,

699,

700. 701, 708.

Mowttoeystis sp., 692, 703.

Monostyla

bulla, 798, '794, 797,

798.

lunaris, 79), 798.

Mormyrus

kannume, d96.

Mosehus, 844, 845, 893,

9X0. :

moschiferus, 937, 938.

Motella, 621.

Mugil, 608.

capito, 607.

Murena

helena, 603.

Mus

alevandrinus, 654, 656,

' 662, 664, 665.

decumanus, 652.

Jerdoni, 662, 665,

musculus, 665,

859, 889, 972.

— gentilis, 665.

— orientalis, 6609.

norvegicus, 638, 6952,

654, 656, 661, 662,

665.

rattus, 638, 651-657,

660, 661, 662, 664,

665.

— dalexandrinus, 652,

699.

— fectorum, 638, 632,

658, 659.

tectorum, 654, 655,

656, 660, 664, 568.

Musea

domestica, 718.

vomitoria, 718.

Myotis

blythi, 636.

myosotis, 636.

— ancilla, 636.

— omari, 636.

Myrus

tulgaris, 603, 626.

843,

Nawmorhedus, 857, 982,

985.

wgoral, 853, 854.

raddeanus, 854, 855,

Nemotragus, 856.

pygmaeus, Bb7.

Neotragus, 881.

Nephrops, 647.

Nerophis

equoreus, 606,

ophidion, 606,"

XXXVI

Nesotragus, 881.

livingstonianus, 831.

moschatus, 881.

Nirmus

cameratus, 698.

Noteus

quadricornis, 79), 797,

798.

— bhrevispinus, 79,

798.

stuhlmanni, 793, 708.

Notholea

labis, 796, 798.

Notops

lofuana, 794, 795, 798,

799.

macrourus, 793, 798.

Notopterus, 598.

horneensis, 597, 626.

Nototragus

melanotis, 882.

Numida

ptilorhyncha, 706.

Nyctipbanes

norvegica, 786.

Obelia

andersoni, 810.

(Ecistes

sp., 794.

mucicola, 795, 798.

Oecophylla

smaragdina, 669.

Oinmatophaga

rossi, 559.

Opercularella

lacerta, 800, £02,

812.

Ophidium

barbatum, 618, 627.

Oreamnos, 857. 8d.

montanus, 81, 852,

866.

Oreotragus, 845, 846, 972,

976, 984, 986.

oreotraqus, 885, 886.

Ornithomyia

legopodis, 662,

697, 704, 705.

Ornithorhynchus, 764,

765, 766.

Orthagoriscus, 624.

Oryx, 983.

algazel,

‘909.

heatrix. 908.

beisa, 907, 908, 909,

910, 982,

gqazella, 9O7.

Teucorye, 908, 909.

696,

837, 907,

INDEX,

Otis

ludwigi, 837.

vigorsi, 837.

Ourebia, 846, 874, 8&5,

982, 984.

montana, 882.

nigricaudata, 881, 882,

883, 884.

ourebi, 882.

scoporia, 882.

Ovibos, 846.

moschatus, 857, 858.

Ovis, 853, 899, 936, 978.

ammon, 859, 988.

— hodgsoni, 859.

ammonoides, 859.

aries, 859, 862.

canadensis, 859, 861.

lerva, 862.

MuUsiMonN,

862.

poli, 988.

tragelaphus, 86°.

vignei, 843, 859, 860,

861, 862, 889.

Ozanna, 907.

859, 861,

Panolia, 970, 971.

Pantholops, 840, 845,

846, 874, 593, 897,

899, 911, 977.

hodgsoni, 886, 898.

Paracypria, gen. nov.,

per

732.

claviformis, 739, 746,

758.

complanata, '734, 735,

736, 741, 756, 757.

conoidea, 734, 736, 756,

(Pte

curta, 741, 743,

760.

declivis, 738, 7385, 756,

TOs

deflexa, 736, 756, 757.

flexuosa, JTAQ, 756,

746,

758.

obtusa, 735, 736, 756,

757.

opaca, 739, 756, 758.

reniformis, 737, 738,

756, 758.

subangulata, 737, 756,

757.

tenuis, 733, 756.

Paracypridopsis, 742.

Paradoxurus

larvatus, 837,

Parypha

erocea, 829,

Pedalion

mirum. 793, 798.

Pedetes, 765.

Pediotragus

rufescens, 879.

tragulus, 879

Pelea, 917, 978, 82,

984.

capreolus, 911, 912,

914.

Pennaria

australis, 806, 829,

cavolinii, 806, 807,

cooper, 807.

disticha, 829.

— australis, 801, 806,

826, 830, 831.

Perigonimus

repens, 800, 801, 804.

Phacochcerus, 979, 980,

981.

Phaula

limbata, 840.

Philodina

emint, 798, 798.

Phyllium, 840.

Physalus

antiquorum, 7883.

Physeter

catodon, 788.

gibbosus, 788

macrocephalus, 769,

7172, 773, 781, 782;

788.

polyclystus, 788.

trumpo, 788.

Pionocypris. 742, 748.

Plagiaulax, 764, 765, 766,

767.

Pleuronectes, 615, 621,

624.

Plotosus, 599, 600, 621,

626.

Plumularia, 799.

sp., 802, 822, 827,

835.

alternata, 883.

buskii, 827, 882,

833.

cornucopieé, 833.

geminata, 834.

megalocephala, 827,

oO.

setacea, 800, 801, 802,

8)2, 822, 827, 831,

534.

strictocarpa, 822, 834.

Poephagus, 032.

Polyarthra

platyptera, 794, 798.

Polycelis

nigra, 671.

Polyeheles |

phosphorus, 639.

Polyedon, 595, 621,

624, |

spatula, 594, 626.

Potamocypris, 742.

¢ |

Presbytis

robinsoni, 635. |

Pronles |

daphnicola, 795. 798.

Procapra, 893, 897.

gutturosa, 892.

picticaudata, 892.

preewalskii, 892.

Procavia, 764.

Proceryvus

dimorphe, 943.

Pseudaxis, 943, 970, 971, |

975. |

sika, 943.

Pseudvis, 855, 868, 864. |

899, 978, 982, 984.

nayaur, 863, 866.

Pseudophryne

bibronii, 627, 630, 631, |

6384.

Psorospermium

avium, 674.

Pterodina

trilobata, 795, 798.

Pteropus, 767.

Ptilodus 763, 764, 765,

767, 768.

gracilis, 763.

Pudu, 965, 971, 978, 985. |

pudu, 967.

Querquedula

erecca, 706.

| Scatophaga

INDEX.

Rattulus

longiseta, 795, 798.

macerus, 795, 798.

stylatus, 794, Tb,

798.

Redunea, 977, 978, 982.

Rbachianectes

glaucus, 771, 773.

Rhizogeton

nudum, 826, 827, 828. -

Rotifer

vulgaris, 793, 798.

Rueervus, 970, 971, 975.

duvauceli, 943,

elephoides, 943.

Rupicapra, 845, 847, 848,

851, 853, 857, 893,

985.

rupicapra, 849.

— parva, 999.

— pyrenaica, 998, 999.

Rusa, 972, 985.

equina, 947.

unicolor, 946, 947.

| Saiga, 847, 893, 897, 899.

tartarica, 898.

Salpina

brevispina, 794, 798.

macracantha, 795, 798.

stercoraria, 683, 717,

718.

Schizocerea,

diversicornis, 793.

— homoceros, 793, 797,

798.

Scomber

scomber, 613, 626,

| Seorpena, 619.

scrofa, 616, 627.

Selaginopsis

ornata, 820.

Sergestes, 646, 647.

Rana

esculenta, 682.

temporaria, 033.

Rangifer, 939, 959, 962,

964, 967, 968, 971, |

981, 985.

tarandus, 960, 661.

— caribu, 960.

Raphicerus, 846, 874,

881, 885, 982, 984.

campestris, 879, 880,

881, 882, 885, 975.

Rattulus

bicristatus, 795, 798.

gracilis, 795, 798.

challengeri, 64AV, 641,

650, 651.

gloriosus, 640, 641.

Serinetha

ahbdominalis, 840.

Serranus, 618.

cabrilla, 611, 626.

Sertularella

bidens, 816.

cylindrica, 800, 817,

fell to

—- pusilla, 802, 818,

825.

integra, 818.

polyzonias, 800, 818, |

827, 851.

XXVIil

Sertularella

polyzonias cornuta, 802,

815.

— robusta, 818.

quadridens, 800, 801,

802, 809, 810, 816,

818, 819, 825.

Sertularia

luculosa, 821.

setacea, 834.

turbinata,

821, 822.

vege, 832.

versluysi, 822.

Siphonostomus

rondeletii, 605.

Smaris

alcedo, 612.

Solea

impar, 615.

lutea, 614, 615, 626.

monochir, 615.

Soleniopsis

dendriformis, 808.

Spirochwta

anodonte, 705,

anserina, 699.

duttont, 702, 706.

eherthi, 699, 706.

gallinarum, 699, 728,

lagopodis, 692, 697, 698,

699, 705, 728.

lovati, 692, 701, 702,

708,

recurrentis, 702, 706.

(Trypanosoma) balbi-

anit, TOD.

Stenocypris

chevreuri, 750.

malcolmsoni, TA0.

perarmata, TA,

796, 760.

Stenorhynchus

leptonyx, 589.

Stereognathus, 760.

Strepsiceros, 052.

Sylvicapra

abyssinica, 876.

coronata, 868, 873, 874,

875, 876.

— camphellie, 873.

grimmi, 872, 874, 876,

975.

mergens, 867, S76.

Syncheta

sp., 794.

oblonga, 795, 798.

Synguathus

acus, 605, 626,

Synodontis, 621.

schall, 600, 626.

800, 802,

755,

xxvili

Talpa, 765,

“Tapes, TOL.

aureus, 706.

Taurotragus,

930, 931,

983, 986.

oryx, 929, 932.

Tetraceros, 845, 876, 926,

928, 932, 973, 986.

quadricornis, 921, 922,

923, 924, 925, 975.

Tetramastix

opoliensis, 793, 798.

Thuiaria

robusta, 820.

inbuliformis, 827, 832.

vincta, 820.

Thylacinus

cynocephalus, 769.

Thylacoleo, 767.

Thyroseyphus

@qualis, 811, 812.

ramosus, 811.

regularis, 800, 801,

811, 812, 825,

simplex, 811.

torresii, 811.

tridentatus, 81).

vitiensis, 800, 801, 806,

808, 810, 811, 812,

813, 815, 821.

Trachinus, 613, 618.

899, 928,

934,

977,

‘

INDEX,

Tragelaphus, 921, 932, |

977

angasti, 930.

scriptus, 925, 929,

930.

— roualeyni, 930,

— seriptus, 922, 930,

931.

‘Tragulus

meminna, 979, 980.

pygmea, 867.

stanleyanus, 979, 980.

’ Triarthra

longiseta, 793, 798.

Tribonyx

mortiert, 769.

Trichomonas

batrachorum, 700.

eberthi, 692, 699, 700,

708.

intestinalis, 700.

Trichostrongylus

pergracilis, 717, 730.

Trigla, 618. °

lineata, 616, 627.

Triglyphus, 766.

Tritylodon, 766, 767.

longevus, 760,761, 762,

768.

Trypanoplasma, 670,

borreli, 671.

Trypanoplasma

(Cryptobia) dendroceli,

670.

Trypanosoma

eberthi, 699, 706.

Tubularia

, bethe, 829..

crocea, 826, 829, 830.

Turtur

humilis, 706.

Uranoscopus, 618.

scaber, 617.

Urenchelys, 603.

Viverra

civetta, 954.

_ Xenelaphus, 966.

Zeus, 621.

Jaber, 614, 626.

Zonocypris, 742.

costata, 748, 749, 756,

759.

elegans, 748.

levis, 749, 756, 760.

madagascariensis, 748,

+ — ++ ++ + + - Or'’_1*OMW} OO

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

No. 82.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

April 5th, 1910.

Prof. KE. A. Mincuiy, M.A., Vice-President, in the Chair.

The Minutes of the last Scientific Meeting were confirmed.

The Secretary exhibited a series of photographs taken in the

South Orkneys in 1908 by a correspondent of ‘The Times,’ and

kindly sent him by the Editor of ‘The Times.’ The photographs

included views of scenery and an interesting series of studies of

living Seals, such as the Sea-Leopard (Stenorhynchus leptonyx),

Weddell’s Seal (Leptonychotes weddelli), the White, or Crab-

eating Seal (Lobodon carcinophagus), and Ross’s Seal (Onumato-

phaga rossi).

Mr. R. I. Pococr, F.1.S., F.Z.8., exhibited a living specimen

of a Mole-Rat (Georychus) recently presented to the Society by

Capt. C. H. Armitage, who brought it from Ashanti. Mr. Pocock

remarked that the animal possibly belonged to the species from

Togoland described by Matschie as G. zechi, and certainly repre-

sented a form never previously exhibited in the Gardens,

Mr. R. H. Wurrrnouse, M.8c., introduced by the Secrerary,

gave an account of his Memoir on “The Caudal Fin of the

Teleostomi.” The paper dealt with the structure of the caudal fin

in about fifty different species of fishes, mostly Teleostei, and

representative of nearly all the sub-groups. After each sub-group

a few general remarks were added, and these were followed by a

short summary of results dealing with definitions and the taxo-

nomic value of the caudal fin.

* This Abstract is published by the Society at 3 Hanover Square, London,

W., onthe 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 Stxpence, or, if desired, sent post-free for the su: of Six

Shillings per annum, payable in advance.

— =

Mr. 'T. M. 8. Exeuisn read a paper, communicated by Mr. G.

A. Boutencer, F.R.S., V.P.Z.S8., entitled ‘Some Notes on Tas-

manian Frogs,” based on observations made during rather more

than two years’ residence in Tasmania.

The paper by Mr. F. E. Brepparp, M.A., F.R.S., “On the

Alimentary Tract of certain Birds, and on the Mesenteric

Relations of the Intestinal Loops,” announced for this evening,

was postponed owing to the indisposition of the Author.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, the 19th April, 1910, at half-past Hight

o'clock P.M., when the following communications will be

mace :

1. Srantey Kemp, B.A.

Notes on the Photophores of Decapod Crustacea.

2. J. Lewis Bonuore, M.A., F.LS., F.Z.S.

On the Varieties of Mus rattus in Egypt, with General Notes

on the Species having reference to Variation and Heredity.

3. G. E. BULLEN.

On an Example of Posterior Dichotomy in an Aylesbury

Duckling.

The following communications have been received :—

1. F. E. Bepparp, M.A., F.R.S.

On the Alimentary Tract of certain Birds, and on the

Mesenteric Relations of the Intestinal Loops.

2. Prof. G, O. Sars, C.M.Z.S.

Zoological Results of the Third Tanganyika Expedition,

conducted by Dr. W. A. Cunnington, F.Z.8., 1904-1905, —

Report on the Ostracoda,

3. Dr. R. Broom, D.Sc., C.M.Z.S.

On Tritylodon, and on the Relationships of the Multituber-

culata.

ee)

4, D. G. Livum, B.A.

Observations on the Anatomy and General Biology of some

Members of the larger Cetiacea.

H. B. Fantuam, D.S8c., F.Z.8.

(5) The Morphology and Life-History of Himeria (Coccidium)

avium: a Sporozodn causing a fatal disease among young

Grouse.

(6) Observations on the Parasitic Protozoa of the Red Grouse

(Lagopus scoticus).

(7) Experimental Studies on Avian Coccidiosis, especially in

relation to young Grouse, Fowls, and Pigeons.

(8) Observations on the Blood of Grouse.

Communications intended for the Scientific Meetings of the

ZOoLoGiIcAL Society or Lonpown should be addressed to

P. CHALMERS MITCHELL,

Secretary.

3 Hanover Square, Lonpon, W.

April 12th, 1910,

No. 838.

ABSTRACT OF THE PROCEEDINGS

oF THE

ZOOLOGICAL SOCIETY OF LONDON.”

April 19th, 1910.

Dr. 8. F. Harmer, M.A., F.R.S. Vice-President, in the Chair,

The Minutes of the last Scientific Meeting were confirmed.

The Sucrrvary read a Report on the additions that had been

made to the Society’s Menagerie during the month of March 1910.

On behalf of Mr. Ouprretp Tuomas, F.R.S., F.Z.S., two speci-

mens of the following new Monkey from the Malay Peninsula

were exhibited :—

PRESBYTIS ROBINSON], sp. 0.

Upper surface wholly white, scarcely relieved by a slight

admixture of darker hairs. Underside, hands, and feet darker,

greyish or greyish brown, varying in tone, but always darker

than the crown and back.

Hab. Trang, Malay Peninsula.

Type. Adult male in British Museum. Original number 3184,

Collected by H. C. Robinson, Esq., and presented by the Govern-

ment of the Federated Malay States.

On behalf of Mr. Oupriretp THomas, F.R.S., F.Z.S., a further

consignment of small mammals from China, collected by Mx. Mal-

colm Anderson for the Duke of Bedford’s exploration of Eastern

Asia, was exhibited, and attention drawn to their value for the

furtherance of zoological and geographical science.

The following forms were described as new :—

MYOTIS MYOSOTIS ANCILLA, subsp. n.

Smaller and with shorter ears than true myosotis. General

colour drab. Forearm of type 61 mm.

Hab. Shang-chow, 8.E. Shen-si. Z'yme. Male. Original

number 2082.

* This Abstract is published by the Society at 3 Hanover Square, London,

W., on the Tuesday following the date of Meeting to which it refers. It will

be issued, 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 Stapence, or, if desired, sent post-free for the sun of Six

Shillings per annum, payable in advance.

Microtrus NUX, sp. n.

Allied to MZ, inex, but darker and with a longer tail; 2nd and

3rd spaces of m* united.

Head and body 93 mm.; tail 59; hind foot 16°5.

Hab, Shang-chow, 8.E. Shen-si. 7'ype. Male. Original number

2089.

MICROTUS JOHANNES, Sp. 0.

Allied to I. mandarinus, but with smaller skull. Colour

wood-brown instead of drab.

Head and body 95 mm. ; tail 23; hind foot 17. Condylo-basal

length of skull 24°6.

Hab, N.W. of Ko-lan-chow, Shan-si. TZype. Male. B.M.

INO. Gol LL 78:

Mr. J. Lewis Bonuorr, M.A., F.L.S., F.Z.8., exhibited a yellow

variety of J/us rattws. This individual had been bred from two

wild-caught examples of J/us rattus tectorum from Egypt. The

long hairs on the upper parts, which are normally black, were

colourless and the shorter body-hairs had grey bases with yellow

tips. As far as he knew, this was the first recorded example of a

yellow rat, which was a colour quite unknown to fanciers and

was never met with by Crampe or Doncaster, who had conducted

colour-breeding experiments on Rats. Although it was not defi-

nitely stated, these experiments had probably been carried out with

Mus norvegicus. A careful search through the large series of

Mus rattus in the British Museum also failed to bring to light

any yellow forms.

Centenary of Philip Henry Gosse, PLS.

Born April 6, 1810; died Aug. 23, 1888.

The Secrerary exhibited the set of the works of Philip Henry

Gosse in the Society’s Library and gave a brief summary of his

contributions to science, upon which he placed a very high value.

The CHatrMAN and Prof. A. Sepawick, F.R.8., added their testi-

mony to his place in Zoological history ; Dr. Henry Woopwarp,

I. R.S., gave some personal reminiscences ; and Mr. EpMuND Gossk,

Librarian of the House of Lords, the son and biographer of Gosse,

thanked the Society for their notice of his father’s centenary.

Dr.W. T. Caiman, F.Z.S8., communicated a paper by Mr. Stanney

Kemp, B.A., entitled ‘“ Notes on the Photophores of Decapod

Crustacea,”

Mr. J. Lewis Bonnore, M.A., F.L.S., F.Z.S., read a paper on

the variations of J/us ratius, founded on an examination of the

forms of that species found in Egypt. The author poimted out

that on examination of the hind-foot measurements of a con-

siderable number he found that the curve showed three distinct

apices, and that two of these apices belonged respectively to the

two forms found in Egypt, these forms being also more easily

distinguished by their colour characteristics.

Mr. Bonhote in dealing with the Rats of this species from the

Oriental Region had some years ago subdivided them into three

subgroups, and it was now shown that the size of the feet typical

of the three Oriental subgroups corresponded with the three

apices in the curve of the Egyptian forms.

The author was inclined to think that these apices represented

centres of variation and were probably inherited as Mendelian

characters, for were this not the case the smallest apex would

have become swamped and a regular curve would result.

It was, however, evident that the small foot character was

present and ready to become the dominant form in a very short

time should conditions giving advantage to a small foot arise.

On comparing the curve of the hind feet of Mus norvegicus,

three apices were also observed, showing that in this species the

“hind foot character” was also present, but as there were no cor-

responding colow: differences it was impossible to tell to which

group any particular individual belonged,

The author drew the following conclusion, viz., that there was

considerable prima facie evidence that the size of the hind foot

and the colour of the hairs on the underparts were Mendelian

characters, and pointed out that the former character was also

found in another species, JZus norvegicus, and the latter in a third

species, Jus musculus,

On the presumption that these were true Mendelian character's

a clearer understanding was at once gained as to how species

might very quickly adapt themselves to altered conditions, and

how many very closely allied species, differing only in size or in

some definite but slight alteration of colour, could exist in the

same locality and yet maintain their characteristics intact and

show no merging of the one into the other.

Experiments in breeding the different forms were being under-

taken, but they were not as yet sufticiently advanced to permit of

any definite results being stated.

Mr. G. E. Bunten presented a paper, communicated by Mr.

JouN Hopkinson, F.Z.S., “Onan Example of Posterior Dichotomy

in an Aylesbury Duckling,” giving a detailed account of a dissec-

tion performed on a duckling having supernumerary legs. In

addition to a reduplicated pelvis and the usual condition of the

limbs presented in posterior dichotomy, it was found that the

specimen showed evidence of a further reduplication of the part

dichotomised,

The next Meeting of the Society for Scientific Business will

be held on Tuesday, the 3rd May, 1910, at half-past Hight

o'clock p.m., when the following communications will be

made :—

H. B. Fantuam, D.Sc., F.Z.8.

(1) The Morphology and Life-History of Limeria (Coccidium)

avium: a Sporozobn causing a fatal disease among young

Grouse.

(2) Observations on the Parasitic Protozoa of the Red Grouse

(Lagopus scoticus).

(3) Experimental Studies on Avian Coccidiosis, especially in

relation to young Grouse, Fowls, and Pigeons.

(4) Observations on the Blood of Grouse.

5. Prof. G. O. Sars, C.M.Z.S.

Zoological Results of the Third Tanganyika Expedition,

conducted by Dr. W. A. Cunnington, F.Z.S., 1904-1905,—

teport on the Ostracoda.

6. Dr. R. Broom, D.Sc., C.M.Z.S.

On Tritylodon, and on the Relationships of the Multituber-

culata.

The following communications have been received :—

1. F. E. Bepparp, M.A., F.R.S.

On the Alimentary Tract of certain Birds, and on the

Mesenteric Relations of the Intestinal Loops.

2. D. G. Litum, B.A.

“Observations on the Anatomy and General Biology of some

Members of the larger Cetacea.

3. C. F. Rousserer, F.R.M.S.

Zoological Results of the Third Tanganyika Expedition,

conducted by Dr. W. A. Cunnington, E. Z.S., 1904—1905.—

Report on the Rotifera.

Communications intended for the Scientific Meetings of the

ZooLOGICAL Sociery oF Lonpon should be addressed to

P. CHALMERS MITCHELL,

Secretary.

3 Hanover Square, Lonpon, W.

April 26th, 1910,

No. 84.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

May 3rd, 1910.

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

The Minutes of the last Scientific Meeting were confirmed.

The Secrerary exhibited a photograph of a Giraffe, kindly lent

him by Mr. Carl Hagenbeck. The animal was an adult female

3°30 metres in height, obtained in Gallaland, East Africa, and

was in the possession of Mr. Hagenbeck. It displayed a

reticulate coloration, in which the lighter areas were reduced to

narrow lines, clearly on the neck and body, but the hind-quarters

were blotched, whilst the spotting came lower down on the legs

than in the typical Giraffa reticulata. It possibly represented a

type intermediate between G. reticulata and GC. camelopardalis,

but it was to be remembered that the specimen was a female, and

that females did not exhibit the coloration so conspicuously as

old males.

Mr. E. E. Austen, F.Z.8., exhibited a specimen of a rare Fly

(Cephenomyia wrichit Brauer) from Kast Prussia, parasitic in the

larval stage in the throat of the Elk (Alces machlis Ogilby), and

remarked onits habits and those of allied species. This specimen

had been recently acquired by the Hon. Charles Rothschild and

presented by him to the British Museum.

Dr. R. A. O’Brisn, introduced by Prof. E. A. Mincury, M.A.,

F.Z.8., exbibited a series of lantern-slides illustrating the habits

of the North Queensland Green Tree-Ant (Oecophylla smarag-

dina %), and described the method of nest-building and bridge-

forming followed by these insects.

* This Abstract is published by the Society at 3 Hanover Square, London,

W.., on the Tuesday following the date of Meeting to which it refers. It will

be issued, along with the ‘ Proceedings,’ free of extra charge, to all Fellows who

subscribe to the Publications; butit may be obtained on the day of publication

at the price of Stwpence, or, if desired, sent post-free for the sum of Stix

Shillings per annum, payable in adyance.

Dr. H. B. Fanruam, F.Z.S., exhibited, on behalf of Miss A.

Porrer, B.Se., and himself, preparations and original drawings

of a new Trypanoplasma (Cryptobia) from the alimentary tract

of Dendrocelum lacteum. The parasites measure from 20 to

40 long approximately, They are biflagellate, the posterior

fiagellum forming the border of an undulating membrane. The

blepharoplast (kinetonucleus) is large and curved. It is proposed

to call the parasite 7’rypanoplasma (Cryptobia) dendroceli. There

is evidence of hereditary infection of the Planarian by the Zrypano-

plasma, for the parasites have been seen to enter the eggs of the

host. A Trypanosome-like flagellate was also seen on rare occa-

sions in the guts of Dendrocelum lacteum and Planaria nigra.

It is interesting to record, for the first time so far as is known,

the occurrence of a Trypanoplasma in a vegetable feeder like

Dendrocelum.

Dr. H. B. Fanrnam, F.Z.8., Protozoologist to the Grouse-

Disease Inquiry, read four papers relating to the Protozoa and

blood of Grouse, illustrated by lantern-slides made from the

original plates accompanying the memoirs.

The first paper, entitled “‘ The Morpholog ry and Life-History of

Eimeria (Coccidiwm) avium: a Sporozosn causing a fatal disease

among young Grouse,” gave a detailed account of the morphology

and life-history of the Coccidium which destroys the epithelial

lining of the duodenum and ceca of Grouse chicks, causing

enteritis accompanied by diarrhea.

The second paper, entitled ‘Observations on the Parasitic

Protozoa of the Red Grouse (Lagopus scoticus),” set forth the

author’s observations on some seven other Protozoa, parasitic

in the blood or in the digestive tract of Grouse. None of these

parasites, however, could ‘be said to be either numerous or very

harmful to the birds examined. The parasites were Leucocytozoin

lovati, in which schizogony in the spleen of the bird was recorded

for the first time, not only for Leucocytozodn lovati but for any

avian Leucocytozoon ; Hemoproteus mansoni and NSpirocheta

lagopodis in the blood ; Zrichomonas eberthi and Spirocheta lovati

in the ceca; and Amada lagopodis in the intestine and feces.

The Spirochetes and Ameeba of Grouse are new species. J/ono-

cystis spores are also found occasionally. All the parasitic

Protozoa mentioned were observed both alive and in stained

preparations.

The third paper dealt with ‘ Experimental Studies on Avian

Coceidiosis, especially in relation to young Grouse, Fowls, and

Pigeons.” The results of many and varied experiments were

recorded in this paper, relating to the time of ripening and

duration of infectivity of coccidian odeysts, their dispersal by

insect larve &e., and the effects of various reagents on the oueysts.

The distribution of the parasites within the host was given and

the results of the transmission of Grouse coccidiosis to young

fowls and pigeons were set forth.

“ Observations on the Blood of Grouse” were discussed by the

author in the fourth and last paper. The various blood-cells were

described, and the results of blood counts (both of red cells and

of leucocytes differentially) of healthy and diseased birds were

set forth. Both coccidiosis and strongylosis produce anemia, and

the presence of various parasites is also associated with numerical

differences in the leucocytic elements of the blood.

Dr. W. A. Cunninecron, F.Z.8., communicated a paper by

Prof, G. O. Sars, C.M.Z.8., containing the Report on the Ostra-

coda collected by the Third Tanganyika Expedition during

1904-1905.

Dr. A. Smita Woopwarp, F.R.S., communicated a paper by

Dr. R. Broom, D.Sc., C.M.Z.S., “On Zritylodon, and on the

Relationships of the Multituberculata.” The author had re-

examined the type and only known specimen of Zritylodon, and

in one or two points came to different conclusions from Owen

and Seeley. The large flat piece of bone which forms the upper

part of the snout, regarded by both Owen and Seeley as the

frontal, was believed to be the upper part of the nasal. The sup-

posed parietal was held to be the frontal. No distinct prefrontal

could be made out; but there was believed to be a large distinct

septomaxillary. The dental formula was believed to be 7@° m’,

instead of, as supposed by Owen, 2? m’°.

Gidley’s recent paper on Ptilodus was criticised at some length,

and an endeavour made to controvert his conclusion that Péilodus

is allied to the Diprotodont Marsupials.

It was held that while the Multituberculates are doubtless very

unlike the living degenerate Monotremes, they are more primitive

than the Marsupials and not at all closely allied to them, and

that till the evidence of their affinities is much greater than at

present they may well be left as an independent order.

The next Meeting of the Society for Scientific Business will

be held on Tuesday, the 24th May, 1910, at half-past Hight

o'clock p.M., when the following communications will be

made ;—

Ie 1D) (Gis Litera BN

Observations on the Anatomy and General Biology of some

Members of the larger Cetacea.

2. C. F. Rousserer, F.R.M.S.

Zoological Results of the Third Tanganyika Expedition,

conducted by Dr. W. A. Cunnington, F.Z.5., 1904-1905.—

Report on the Rotifera.

3. JAMEs Ritcuie, M.A., B.Sc.

The Marine Fauna of the Mergui Archipelago, Lower Burina,

colleeted by Jas. J. Simpson, M.A., B.Sc., and R. N. Rudmose-

Brown, D.Sc., University of Aberdeen, February to May, 1907.

—The Hydroids.

. James Rircute, M.A., B.Sc.

Hydroids from Christmas Island, Indian Ocean, collected by

C. W. Andrews, D.Sc., F.R.S., F.Z.S., m 1908.

—

The following communications have been received :—

1. F. E. Bepparp, M.A., F.R.S.

On the Alimentary Tract of certain Birds, and on the

Mesenteric Relations of the Intestinal Loops.

2. R.T. Pocock, F.L.S., F.Z.8.

On the Cutaneous Scent-glands of Ruminants.

3. R. LYDEKKER.

On a Pair of Wapiti Antlers and a new Muntjac.

Communications intended for the Scientific Meetings of the

ZooLoGIcAL Soctery or Lonpon should be addressed to

P. CHALMERS MITCHELL,

Secretary,

3 Hanover Square, Lonpon, W.

May 10th, 1910.

No. 85.

ABSTRACT OF THE PROCEEDINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.*

May 24th, 1910.

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

The Minutes of the last Scientific Meeting were confirmed.

The Secrerary read a Report on the additions that had been

made to the Society’s Menagerie during the month of April 1910.

Mr. Ernest Gipson, F.Z.S., exhibited a skin of Felis geoffroyt

and made some remarks on the distribution of the animal.

Mr. D. G. Linum, B.A., read a paper, communicated by Dr. H.

Gavow, F.R.S., F.Z.S., entitled ‘“ Observations on the Anatomy

and General Biology of some Members of the larger Cetacea.”

This paper was the outcome of seven weeks spent at the Irish

whaling station during the summer of 1909. The object of the

visit was to make a preliminary survey of the opportunities which

are now offered for a study of the larger Cetacea by the recent

establishment of whaling stations off the shores of the British

Isles. It was pointed out that hitherto our knowledge of whales

had been chiefly derived from the examination of stranded speci-

mens, which had often reached an advanced state of decay;

whereas it was now possible to obtain a plentiful supply of large

whales fairly near at hand which were sufficiently fresh for histo-

logical study. It was particularly desirable to increase our know-

ledge of the larger Cetacea before these much-hunted animals

became too scarce. A list was given of the species captured at

the Irish station during the two years of its existence, with notes

on the species Balenoptera musculus Linn., B. sibbaldiit Gray, and

Physeter macrocephalus Linn., which were seen by the author.

* This Abstract is published by the Society at 3 Hanover Square, London,

W.., on the Tuesday following the date of Meeting to which it refers. It will

be issued, along with the ‘ Proceedings,’ free of extra charge, to all Fellows who

subseribe to the Publications; but it may be obtained on the day of publication

at the price of Stapence, or, if desired, sent post-lree for the sum of Sia

Shillings per annum, payable in advance.

The paper also contained observations on the occurrence of

hairs in whales; the auditory organ of the Balenoptera; the

asymmetry of the odontocete skull; and a few remarks upon the

habits of whales. In conclusion, attention was called to the present

difficulty in obtaining information as to their method of copula-

tion, period of gestation, rate of breeding, &c., and a means of

overcoming this difficulty was suggested.

Mr. C. F. Rovsseter, F.R.M.S., read a paper, communicated

by Dr. W. A. Cunninerton, F.Z.S., on the collection of Rotifera

made by the Third Tanganyika Expedition, 1904-1905. Amongst

the specimens brought back by Dr. W. A. Cunnington were a

number of tubes containing fine surface plankton nettings from

Tanganyika and other lakes of that region. These the author

searched for Rotifera, and the result was contained in the present

paper. From Lake Tanganyika only 11 species were obtained,

all already known in other parts of the world; whilst the River

Lofu, which enters the lake at its south-western corner, yielded

23 species, one of which is a very remarkable new kind. In

Lake Nyassa only 6 species were found, and a single gathering of

Victoria Nyansa yielded 9 species. This collection is interesting

and important from the fact that no previous record of Rotifera

from Lake Tanganyika had been made, and very few species were

known from the Central African Region.

A complete list of species of Rotifera so far known from this

region was appended.

Mr. R. Kirxparrick, F.Z.8., communicated two papers by

James Rrrewig, M.A., B.Se., of the Royal Scottish Museum, on

the Hydroids of the Mergui Archipelago, collected by Mr. J. J.

Simpson and Dr. R. N. Rudmose Brown ; and on the Hydroids

of Christmas Island, collected by Dr. C. W. Andrews, F.R.S.

The chief interest of these two papers was faunistic. In the

former 30 species were recorded, in the latter 13. It was

apparent that the Hydroid fauna of the Eastern Indian Ocean,

of which hitherto little had been known, lacked distinctness, and

that its closest affinities were with the faunas of the neighbouring

Malay Archipelago and Australian areas.

The next Meeting of the Society for Scientific Business (closing

the Session 1909-1910) will be held on Tuesday, June 14th, 1910,

at half-past Hight o’clock p.m., when the following communications

will be made :—

1. R. I. Pococs, F.LS., F.Z.S.

On the Cutaneous Scent-glands of Ruminants.

2. R. LYDEKKER.

On a Pair of Wapiti Antlers and a new Muntijac.

3. R. LYDEKKER.

On Three African Buffaloes.

4. A. Caprera, C.M.Z.S.

On Two new Antelopes.

5. Dr. E. A. Wizson, F.Z.S.

The Plumage of the Grouse.

The following communication has been received :—

F. E. Bepparp, M.A., F.R.S.

On the Alimentary Tract of certain Birds, and on the

Mesenteric Relations of the Intestinal Loops.

Communications intended for the Scientific Meetings of the

ZooLocicaL Soctrry of Lonpon should be addressed to

P. CHALMERS MITCHELL,

Secretary.

3 Hanover Square, Lonpvoy, W.

May 31st, 1910.

THE ZOOLOGICAL SOCIETY OF LONDON.

Tars Society was founded in 1826 by Sir Sraurorp Rarrurs,

Mr. J. Sasryz, Mr. N. A. Viaors, and other eminent Naturalists,

for the advancement of Zoology and Animal Physiology, and for the

introduction of new and curious subjects of the Animal Kingdom,

and was incorporated by Royal Charter in 1829.

Patron.

HIS MAJESTY THE KING,

COUNCIL.

HIS GRACE THE DUKE OF BEDFORD, K.G., President.

Tue Eart or Atramont, F.8.A.

Grorce A. Bovurtencer, Ese,

F.RS., Vice-President.

Ricuarp H. Burne, Esa., M.A.

Lr.-Cotr. Srr R. Havetock

Cuaruzs, K.C.V.O., M.D.

AtrrepD H. Cocks, Hse., M.A.

F. D. Dawtrey Drewitt, Esa.,

MEA MD:

Cuartes Drummonp, Esa,

Treasurer.

Sir Epwarp Duranp, Br., C.B.

FREDERICK GiLuerr, Hse.

Sipney F, Harmer, Ese., M.A.,

Se.D., F.R.S., Vice-President. |

Sir Epuunp G. Loner, Br.

K. G. B. Meavze-Watpo, Esa.,

Vice-President.

Pror. Epwarp ALFRED MINCHIN,

M.A., Vice-President.

P. Caatmers Mircuett, Ese.,

MA. 6D Sc.» Hon LED

F.R.S., Secretary.

Apert Pam, Esa.

Aprian D. W. Pottocr, Esa.

OxprieLD THomas, Esa., F.R.S.

AUBYN Esa,

M.A.

A.Smira Woopwakp, Esea., LL.D.

F.R.S., Vice-President.

Henry Woopwarp, Esea., LL.D.,

F.R.S., Vice-President.

Trevor-Batrye,

The Society consists of Fellows, and Honorary, Foreign, and

‘orresponding 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, where all communications should be

sent, addressed to “*The Secretary,” is open from Ten till Five,

except on Saturdays, when it closes at Two p.m,

The Library, under the superintendence of Mr. F. H. Waterhouse,

is open daily at the above hours, except in September.

The Meetings of the Society for General Business are held on

the third Wednesday in every month of the year, except in

September and October, at half-past Four o’clock p.m.

The Meetings for Scientific Business are held 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 Gardens are open daily from Nine o’clock until Sunset.

Mr. R. I. Pocock, F.L.S., is the resident Superintendent and Curator

of Mammals and Reptiles. Mr. D. Seth-Smith is Curator of Birds

and Inspector of Works. The Prosectorium for Anatomical and

Pathological work is under the charge of Mr. Frank E. Beddard,

M.A., 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.

Frttows pay an Admission Fee of £5, and an Annual Contri-

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or a Composition of £45 in lieu thereof; the whole payment,

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No person can become a Fritow until the Admission Fee and

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Subscription for the year in which they are elected.

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Companions daily, upon signing their names in the book at the

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The Wire or Hussanp of a Frttow can exercise these privileges

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Every Fetxow is entitled to receive annually 60 undated Green

Cards, and, when no specific instructions are received, the supply

will be sent in this form. If preferred, however, 20 Green Cards

may be exchanged for a book containing 2 Orders for each

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may also be exchanged for 2 Buff Cards for the use of Children

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and Publications relating to Zoology in each year, for the sum of

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to withdraw his or her name upon giving notice in writing to the

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Ladies or Gentlemen wishing to become Fellows of the Society

are requested to communicate with the undersigned.

P. CHALMERS MITCHELL,

Secretary.

Regent’s Park, London, N.W.,

December, 1910,

MEETINGS

OF THE

ZOOLOGICAL SOCIETY OF LONDOG

" FOR

SCIENTIFIC BUSINESS.

1910.

Turspay, Fesruary 7 & 21 TuEspDAY, JUNE 13 & 27

ws Marcu 1 & 21 be OcroppR 24

3 APRIL 4 & 25 NovemMBER 7 & 21

‘9 May 9 & 23 |

The Chair will be taken at half-past Light o'clock in the Evening

precisely.

ZOOLOGICAL SOCIETY OF LONDON.

THE ZOOLOGICAL RECORD.

pes object of the ZoorogicaL Recorp is to give, by means of an

annual Volume, complete lists of the Works and Publications

relating to Zoology in all its branches that have appeared during

the year preceding the issue of the Volume, together with full

information as to the points they deal with, arranged in such a

manner as to serve as an Index to the literature of Zoology in all

parts of the globe, and thus to form a repertory that will retain its

value for the Student in future years.

The ‘ Zoological Record’ having been amalgamated with the

International Catalogue of Scientific Literature, Zoology, Volumes

from 43 onwards can now be obtained only from Messrs. Harrison

& Sons, except when purchasing complete sets from the Zoological

Society.

Under the scheme of amalgamation, Fellows of the Society, and

Institutions already on the subscription-lst, have the privilege of

subscribing at the old rate of 30s. per annum, which covers the

cost of carriage of the volume. The subscription becomes due on

July 1st. ineach year, and lapses if not paid by the Ist. of December

following.

The Society is able to supply comp/ete sets of the Record on the

following terms :—

Vols. 1 to 37, Price £14 10s. net.

Vols. 38, 39, and 40 at 10s. net.

Vol. 41 and onwards at 40s. each.

The prices for separate volumes are as follows :—

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Vols. 41 and 42 at 40s. each. The price of the ‘Zoological Record,’

Vol. 48 and subsequent volumes, published now by Messrs. Harrison

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Inpex Zootocicus. An alphabetical list of names of genera

and subgenera proposed for use in Zoology, as recorded in the

‘Zoological Record,’ 1880-1900; together with other names not

included in the ‘ Nomenclator Zoologicus’ of 8. H. Seudder. Com-

piled (for the Zoological Society of London) by CHartes Owrn

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public, 20s., or if sold with a set, 10s.

Divisions of the ‘ Zoological Record’ of Vols. 39 to 42 can be

supplied by the Society, but after Vol. 42 they can be had only of

Messrs. Harrison & Sons, 46 St. Martin’s Lane, W.C.

ey Ui ee

SEPARATE DIVISIONS OF THE ZOOLOGICAL RECORD,

Divisions of the ‘ Zoological Record,’ Vols. 39-42, containing

the literature of the years 1902-1905, may be obtained separately

as follows :—

30 0

List of abbreviations of journals, etc. 2 O net.

Special Records, viz. :—

I. General Subjects . DOr ae

II. Mammalia 2) 6

III. Aves ne) Peon ae 6 -On5:

IV. Reptilia and Batrachia.. 2) Gites

V. Pisces 2 (Gots

VI. Tunicata iy Orie

VII. Mollusca A: QU

VIII. Brachiopoda .. Le) Oinres

IX. Bryozoa L 0

X. Crustacea Dn Os

XI. Arachnida 2 OT

XII. Myriopoda 1 Gio

XIII. Insecta .. 12> Ovex

XIV. Echinoderma oy On ae

XV. Vermes.. a i'Or ae

XVI. Coelenterata .. il), snes

XVIL. Spongie rps

XVIII. Protozoa AI |

Index of new names of genera and subgenera. 2 0

Divisions from Vol. 43 onwards are now supplied by

Harrison & Sons, 46 St. Martin’s Lane, London, W.C.

P. CHALMERS MITCHELL,

Secretary.

Recent's Park, Lonpon, N.W.

December, 1910.

Messrs.

ZOOLOGICAL SOCIETY OF LONDON.

LIST OF PUBLICATIONS.

Tux scientific publications of the Zoological Society of London

are of two kinds—“ Proceedings,” published in an octavo

form, and “‘ Transactions,” in quarto.

According to the present arrangements, the ‘‘ Proceedings”

contain not only notices of all business transacted at the scien-

tific meetings, but also all the papers read at such meetings

and recommended to be published in the ‘‘ Proceedings” by

the Committee of Publication. A large number of coloured

plates and engravings are issued in the volumes of the

“ Proceedings,” to illustrate the new or otherwise remark-

able species of animals described therein. Amongst such

illustrations, figures of the new or rare species acquired in a

living state for the Society’s Gardens are often given.

The “ Proceedings” for each year are issued in four parts,

paged consecutively, on the first of the months of March,

June, September, and December. From January 1901 they

have been issued as two half-yearly volumes, indexed

separately.

An ‘“ Abstract of the Proceedings ” is published by the

Society on the Tuesday following the date of Meeting to

which it refers. It is 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 publi-

cation at the price of Sixpence, or, if desired, sent post free

for the sum of Six Shillings per annum, payable in advance.

The “ Transactions” contain such of the communications

made to the scientific meetings of the Society as, onaccount of

the nature of the plates required to illustrate them, are better

adapted for publication in the quarto form. They are issued

at irregular intervals.

Fellows and Corresponding Members, upon payment of

a Subscription of One Guinea before the day of the Anni-

versary Meeting in each year, are entitled to receive the

Society’s Publications for the year. They are likewise

entitled to purchase the Publications of the Society at 25 per

cent. less than the price charged for them to the Public. A

further reduction of 25 per cent. is made upon purchases of

Publications issued prior to 1881, if they exceed the value of

five pounds.

Fellows also have the privilege of subscribing to the

Annual Volume of the Zoological Record for a sum of 30s.

(which includes cost of delivery), payable on the Ist. of July

in each year; but this privilege is forfeited unless the

subscription be paid defore the 1st. of December following.

The following is a complete list of the publications of the

Society already issued.

TRANSACTIONS* OF THE ZOOLOGICAL SOCIETY OF LONDON.

4to. 19 vols. and Index. ie Preablion

Vol. I., containing 59 Plates.... (1833-35) .... £8138 6 .... £418 OF

eT ane (LBBB AL) oo cca 0 00, er

ee is 659s FT ve Chee 40) ak oe Oy ee AT oe

[vs 4 WE | ies es ead DROS): saan ae 8 2 6f

2 Bea as, G7. kt he: tf. CIEREEEO) ce aah ee 619 0

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SSR ee 78°, ».., (1869-72)'..:. 10 4 0...) Sate

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5) < es “ 00-278 pes. (USra-/7): cycle woe 1462 0

2 eerie e OB os cas, (187-79). 20. 10 00, 3...) oe

Index, Vols. 1-X. ...........00. (1888-79) .... 0.7 6 .... O70

Vol. XI., containing 97 Plates.. (1880-85) .... 912 0.... 1216 0

Me dul <\ T1G5. Vail, : (1888-00) tay ss heb t Bako 740

y eosin y+ 62) yt. (1801-05) wus. G& 8) Siu at One

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Habis Sigs p59 ef (SDR OO) aa ie ee 714 0

eas Rail i p ARS Mays even DOOL—1905)/ 22, cose ue , 20

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PROCEEDINGS OF THE COMMITTEE OF SCIENCE AND

CORRESPONDENCE OF THE ZOOLOGICAL SOCIETY OF

LONDON. 8vo. 2 vols. (Letterpress only). Frice te Pree oie

Parti. | esO=aL, alicvalsSvolik sects e ee wteore de. 6d: %, 0. Ose

Bi, Rl BeD: £ sirens tee ae ee 43, 6d: 2k) Om

PROCEEDINGS OF THE ZOOLOGICAL SOCIETY OF LONDON,

8vo. 15 vols. (Letterpress only) and Index. (Hirst Series.)

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Part I. 1833.1 vol. 8vo. 4s. 6d. .. 6s.¢ | Part IX. 1841.1 vol. 8vo. 4s. 6d. .. 6s.

BAT Meade, es Nea) aries. » 1842,

» III. 18385. i 4s, 6d. .. 6s. ay XI. 1843. + 4s. Gd... Gay

» LV. 1836. 5 4s, 6d. .. 6s. 3, SUL Sa4, . 4s. 6d. .. 6s.

V. 1837. * ABH OU aa. OB: » XIII. 1845. a 4s. 6d. .. 6s.

VI. 1838. * 4s. Gd. .. 6s. 5 ALV. 1846. 5 43. 6d, .. 63.F

» VII. 1839. 4s, 6d. .. 63. 5 ee Ve LET 43, Gd. .. 63.f

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Part XVI S468.40 vol) Byo;.48) 6d. ee.) (Oa. A wemieianrs £10 8 .. Slee

“ XVII. 1849. 3 ZenGaie ¥.0.72 OR, heieemieneas 1 0 8) ae

» &VIII. 1850. Ff 48: Gdw car YOSSI emer 1 8.56 L118 (OF

‘9 XIX. 1851. 39 ASSOC Oh caL Obs: Mobi artes 015 9 LES

be XX. 1852. . As BA ten tis Ate eens 015 9 1s 1 OF

+ XXI, 1853. ” BS Ody vai. SUBSE” watemitits\s 018 O lL: 4 OF

» XXIII. 1855. i As. iGd.0 ma PORE Sehees te LES) 4G Liss oF

“9 XXIV. 1856. 5 AS. ‘OY. he ROS Wie cnt LORS Li oy

a XXV. 1857. * Ag ADs: bis ial re einiete wits LOS LY oF

As XXVI. 1858. , ASAGQ: viitwa Oss etemianantarete ue 2 32 Ot

» &XVII. 1859. 9) Ae Da. ws OS a ah ae Patil 6) 2 2 30;

5, AXAVIIL. 1860. fs ASN6d. sie SOBA ate eiecests ah 2 2 0F

Index 1848-1860. - AgvGd, . jes. G8.

+ Out of print.

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PROCEEDINGS OF THE SCIENTIFIC MEETINGS OF THE

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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

of the Zoological Society of London. (Kighth Edition.) 8vo.

1883. Cloth, 4s. 6d.

List of the Vertebrated Animals now or lately Living in the Gardens

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1896. Cloth, 6s.

Catalogue of the Library of the Zoological Soeiety of London

(Fifth Edition.) 8yvo. 1902. Cloth, 6s.

THE OFFICIAL ILLUSTRATED GARDEN GUIDE—8th Edition

—with (1) a Railway and Street Map, showing a direct

route to the ‘‘ Zoo” from all parts of London and the Suburbs ;

(2) a Plan of the Grounds, showing at a glance the location of

the animals; (3) a short description of some of the principal

animals in the Colleetion (now containing over 3000 spe-

eimens), together with 48 Photographic Illustrations and

Index. Price 6d. in Stiff Paper Cover, postage 1jd., or in

Green Cloth Cover price 1s. 2d. post free.

P. CHALMERS MITCHELL,

Secretary.

Regent's Park, London, N.W..,

December, 1910.

These publications may be obtained at the Socrery’s Orrice,

at Messrs. Lonemans’ (Paternoster Low, £.C.), or through any

bookseller,

PratEe

LXXVIII.

LXXIX.

LXXX.

LXXXI.

LXXXII.

(XXXII.

XXXIV.

LXXXV.

XXXVI.

~XXXVITI.

LXXXVIII.

UXXXIX.

XC,

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XOII.

XCIII.

XCIV.

XCV.

XCVLI.

xcvVIl.

XCVITI.

XCIX.

Cl.

EIST’ OF PLATES.

1910, pp. 837-1033.

PaGn

Pair of Red Grouse in summer with young chicks .... \

Male Grouse, black type, in full winter-plumage

Male Grouse, red type, in full winter-plumage

Male Grouse, white-spotted bird of the red type

)

‘Male Grouse, red type, in full winter-~plumage with a |

few black-centerd feathers of the previous autumn- |

plumage

stake et alursal ales efeteial e's 8761650) %)8) OP SMT Cl OAei Gh Oe] BS 16/8) SiS

Male Grouse showing marked beginning of the autumn-

plumage on head RH DECIR Te ae Melee cin ew os

Male Grouse changing from winter- to autumn-pluma

Female ‘Grouse, black type, in autumn-plumage

Female Grouse in full breeding-plumage .. .

Female Grouse, red type, changing from w ieee es

summer-plumage |

eevee

aie: cle "ei6Ma a @ Vie 6) 44). o 6. « *) @\q of eye'S a) & (e000)

Female Grouse in full summer-plumage ....-+..----

Female Grouse showing bare patch of skin and ease

line of barred feathers on abdomen ......+-++.---

Female Grouse, red type, feathers from flanks ....-.

Female Grouse, red type, worn upper tail-coverts ....

Feet of Red Grouse: (1) New winter-feathers and

nails; (2) Full winter-plumage |

Feet of Grouse showing stages in moulting of nails ..

\ LOU

Heads of Grouse: (1) Female; (2) Male

Heads of: (1) Female Grouse; (2) Male Grouse ; |

and (3) Ptarmigan, showing supra-orbital combs. .

Head of Blackcock, showing supra-orbital comb

Male Grouse showing abnormal erythrism

Female Grouse, buff-barred type

Female Grouse, abnormal yellow variety ..-.-.-.---- |

Female Grouse, grey variety

Female Grouse, grey seeds

G. R. Gray) - ae

eeoceee tees

_ (ype « of L. s were us