COLLECTION
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WILLIAM SCHAUS
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TO: THE
NATIONAL MUSEUM

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PROCEEDINGS

OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY

OF LONDON.

1918, pp. 1-196,

witH 1 PuatTe and 32 TEx't-FIGURES.

PRINTED FOR THE SOCIETY,
SOLD AT ITS HOUSE IN REGENT’S PARK.
LONDON:

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

Ibe sss 0

OF THE

COUNLIL AND OFFICERS

OF THE

ZOOLOGICAL

SOCIETY OF LONDON,

1918.

Patron.
His Masgesty Tue Kine.

COUNCIL.
His Grace Tat Duke or Beprorp, K.G., F.R.S., President.

THe Hon. Cecrt Barina, M.A. |

Ricuarp H. Burne, Ksq.,
M.A.

Lt.-Cot. 8S. Monckton Cops-
MAN, M.D., F.R.S.

CHARLES DrumMonb, Ksq.,
Treasurer.

ALFRED Ezra, Esq., Vice-

President.
Carr. Haroup S. Ferauson.
Capt. Hucn 8S. GLADSTONE,
M.A.
Srpney F. Harmer. Ksq., M.A.,
Se.D., F.R.S., Vice-President.
Pror. James P. Hin, —D.Sc., |

F.R.S.

WiLuiAM HuntsMAN, Ksq.

Sir EpmMunp G. Lopsr, Br., |
Vice-President.

Pror. Ernest W. MacBripe,
D.Sc., LL.D., F.R.S., Viee-
President.

Con. Sir A. Henry McManon,
G.C.M.G., K.C.1.E.,G.C.V.O.,
CS:

P. CHatmers MircHeu, Esq.,
M.A., D.Se., LL.D, HORS,
O.B.E., Secretary.

ApRIAN D, W. Poxtiock, Esq.

THE LORD QUEENBOROUGH.

THE MaRrQugEss OF Siico, F.8.A.,
Vice-President.

AvuBYN Trevor-Barrye, Ksq.,
M.A.

AntHony H. WINGFIELD, Esq.

A. SmirH. Woopwarp, Ksq.,
LL.D.,. FUR.S.,. VicesBre-

sident.

PRINCIPAL OFFICERS.
P, Caoatmers Mircnett, M.A., D.Se., LL.D., F.R.S., O.B.E.,

Secretary.

R. I. Pococrn, F.RS., F.L.8., Curator of Mammals and
Resident Superintendent of the Gardens.

D. Seru-Suirn, Curator of birds and Inspector of Works.

Lieut. Epwarp G. BouLencer, Curator of Reptiles.

Prof. H. Maxwein Lerroy, Curator of Insects.

JOHN Barrow, Accountant.

W. H. Coun, Chief Clerk.

LIST’ OF CONTENTS.

1918, pp. 1-196.

EXHIBITIONS AND NOTICES.

The Secretary. Report on the Additions to the Society’s
Menagerie during the months of November and
WDeCemmlb ee Oli meckesuie diets tees coum tt ba vers okounenlne

Mr. D. Sers Sutra, F.Z.8., Curator of Birds. Exhibition
of, and remarks upon, a series of lantern-slides made
from photographs of Reptiles taken in the Gardens...

Dr. P. CHatmers MitcHeny, F.R.S., Secretary to the
Society. Communication of a letter from Mr. T. E.
Whitehead on the Wild Dingo ’-.........2...6..08e.eeekee

Dr. R. Broom, C.M.Z.8. Exhibition of, and remarks upon,
a New and a Rare Species of the Golden Mole.........

The Secretary. Report on the Additions to the Society’s
Menagerie during the month of January, 1918 ......

Dr. SmirH Woopwarp, F.R.S., V.P.Z.S. Exhibition of a
copy of an incised drawing of a hunted deer............

Professor Pouuron, F.R.S., F.Z.S. Communication of a
letter from Captain G. D. Hale Carpenter on an

African Civet attacking Human Beings ............ ee

Professor EK. W. MacBripr, M.A., D.Sc., F.R.S., F.Z.S.

An account, illustrated by lantern-slides, of his

recent investigations into the development of the
ee AW EME eh wea etch t ote) (104s eis logldsse atm sug renes

Mr. D. Sera Siru, F.Z.S., Curator of Birds. Exhibition
and description of skins of the Hoatzin, illustrated
Dyp laMbern-SliGed: gc, co arereac dtees itp se awemudenp sua cles

Page

187

188

189

190

LSM

Lou

192

PA
lv

Mr. C. Tare Reaan, M.A., F.RS., F.Z.S. Exhibition of
photographs of an Indo-Pacific Chetodont Fish.
(Text-figure 12). csccs Goth oe ee ee eee

The Secrerary. Report on the Additions to the Society’s
Menagerie during the months of February and
Marchi, -LO18) tn. cacp use om cetacean

Miss L. E. CoresmAn, Assistant Curator of Insects. Exli-
bition of specimens of an Kast- African homopterous
insect

ever e rere reese er ee ee ee eee eo eee seeeeeeesseroeseesereeeeeeseesere

Dr. A. S rea Woopwarp, F.R.S., V.P.Z.8. Exhibition of,
and remarks upon, the fossil rostral teeth of Hopr istis
DUCE TISUIS, 2 a. oes een ener een oe haulage eee eee

Mr. G. A. Boutencrr, F.R.S. Exhibition of the head of
a Congo Fish, Huan ocyon goliath *....:..:.: Pibsesigistaisicleci:?

Mr. D. Sera Smiru, F.Z.S., Curator of Birds. Exhibition

of, and remarks upon, a Zulu Head-dress..........2....

Dr. P. Cuatmers Mircuety, F.R.S., Secretary to the

Society. Remarks on an advertisement announcing

Fur Sales in the United States

Professor Woop-JoNrEs,*F.Z.8., Honorary Acting Prosector.
Exhibition of, and remarks upon, specimens from the

Prosectorium illustrating the effects of Rickets ‘

eee eee

PAPERS.

1. Skull of Rana tigrina Daud. By B. L. Buatia, M.Sc.,
and Baint PrasHuaD, M.Sc. (Assistant Professors of
Zoology, Government College, Lahore). (Text-

fiemres 1295) Skew se tedt. id AIO, ni ek eee sy ie

2. Description of a new Snake of the Genus Oigodon
from Upper Burma. By G. A. BouLEencsr, F.R.S.,
F.Z.8S. (Text-figure 1.)

eeoecevereescaeraere eer eseeeoe eee eeeeseoee

Page

pa2

193

194

194

195

195

195

196

3. Reptiles from the River Tajan Chasen: pie). By |

L, A. Lantz. (Plate I.)

Alphabetical List of Contributors

Vv

. On the External Characters of the Lemurs and
of Tarsius. By R. I. Pocock, F.R.S. (Text-
TOMATO MEENA ceo cavataia,s slsieaie sie.« «> deivisfenlee na Oy DRMt atte Welvelte'es
. A Classification of the Pyratip#®, subfamily Hypso-
TROPINA. By Sir Georce Hampson, Bart., F.Z.8. ...

. First Report on the Inheritance of Visible and
Invisible Characters in Silkworms. By its MAUDE
WS CLEGCHORNG: WeZicc5: Mla ig We WsOs, ses. caceseshccsatsare

. Notes on Cetacea stranded on the British Coasts during
1913-1917. By Srpney F. Harner, 8c.D., F.BS.,

F.Z.S., Keeper of Zoology in the British Museum
(Natur: al History)

. On the Variation of the Pit- Viper, Lachesis atrox. By
Miss Joan B. Procrer, F.Z.S. (Text-figures 1-5.)...

. Report on Deaths of Animals in the Gardens in 1917.
By J. A. Murray, M.D., B.Sc., F.Z.S., Director,

Imperial Cancer Research Fund, Pathologist to the
Society

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Page

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133

147

163

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Alm AC Bi ECA Le. tS 2
OF THE

CONTRIBUTORS,

_ With References to the several Articles contributed by each.

1918, pp. 1-196.
PP

Page
Buattia, B. L., M.Sc., and Barnt PrasHap, M.Sc.
Skull of Rana tigrina Daud. (Textefigures 1-9.) ...... ]
Boutencer, G. A., F.R.S., F.Z.8.
Description of a new Snake of the Genus Oligodon
GicomeUipper Burma, (lText-figure: 1) o5.cee..siek oscerwoet ees 9
Exhibition of the head of a Congo Fish, Hydrocyon
GI EOE em Neen reat eect cue Ole caacituad. by beak wv aewe 195
Broom, Dr. R., C.M.Z.S.
Exhibition of, and remarks upon, a New and a Rare
Bpectes oF the Golden NO. oir. cs cebcts sasscsdieeueectevsaas 189
CHEESMAN, Miss L. E., Assistant Curator of Insects.
Exhibition of specimens of an Kast-African homo-
[EET ROL) STO STECE a eine cre an ne 194
CLecHorn, Miss Mauve L., F.Z.S., F.L.S., F.E.S.
First Report on the Inheritance of Visible and
Invisible Characters in Silkworms ................ccc00ccene. 133

Vili

Page
Hampson, Sir Grorce, Bart., F.Z.S.
A Classification of the Pyratip%, subfamily Hypso-
PROPIN AD) |.) ctccs es seee ee ee ee Arne a eee DD
Harmer, Sipney F., Sc.D., F.RS., F.Z.S.
Notes on Cetacea stranded on the British Coasts
during 1913-1914). (35 20..3...$7. EF) DY. cai pecceereecen eee 147
era Ls A. + ag tie =
“Reptiles fron ihe River Dajan (Transcaspia ) “(Plate L) ll
MacBripg, Professor E. W., M. ae D. ay E.R.S., F.Z.S.
An account, illustrated by lantern-slides, of his recent
investigations into the development of the Sea-Urchin... 192
MitrcHE.LL, P. CHatmers, M.A., D.Sc., LL. D. a R, 5. » FZ. S;; ee
Secretary to the Society, |
Report on the Aadivion: se the Sonicty’s Monegene
during the months of November and December, 1917...) Be
Communieation of a letter from Mr. oO E. 3. Whitehead te
on the Wild Dingo ie uidinjeaicttaco® dol wesel ears Ny Bs a Rea ar sereeee 188
Report on the Additions to the Society’ S Menagerie if
during pa month of January, db alts Peicrie suonnadeeeee oede eo co. 190
Report on the Additions to the Society’s Menagerie
during the months of February and March, 1918 sa aaa 193
Remarks on an advertisement announcing Fur Sales
in the United States Raye Het ies eeeereee veered

Murray, J. A., M.D., BSc, BAS. Pathologist to the

Society.’

Report on’ Deaths of Animals in the cab .

Pocock, R. I., F.RB.S., F.LS., F.Z.8:, Curator of Mammals.:2"

On the External Characters of (aes ‘Lemurs’ and of.

Tarsiys.....(Text-figures 1-16.)......... Nd, FIO eas

“19

1X
Poutton, Professor, F.R.S., F.Z.S.

Communication of a letter from Captain G. D. Hale

Carpenter on an African Civet attacking Human Beings.
PrasHaD, Bartn1, M.Sc. See Buartia, B. L., M.Sc.

Procter, Miss Joan B., F.Z.S.

On the Variation of the Pit-Viper, Lachesis atrow.
(Mex ieures l—O.) iiss. sai op ssdimn cs eas ceneodaderestevessodechos

Regan, C. Tats, M.A., F.B.S., F.Z.S.

Exhibition of photographs of an Indo-Pacific Cheeto-
dont Fish. (Text-figure 1.) ..... ERC Pe eee Nore eS

Seru Smita, D., F.Z.S., Curator of Birds.

Exhibition of, and remarks upon, a series of lantern-
slides made from photographs of Reptiles taken in the
(ChATECISIINS) 0 Sa ser rae ete ee eee ge re ene

Exhibition of, and remarks upon, a Zulu Head-dress...

W oop-Jones, Professor, F.Z.S., Honorary Acting Prosector.

Exhibition of, and remarks upon, specimens from the

Prosectorium illustrating the effects of Rickets ............

Woopwarp, Dr. Sirs, F.R.S., V.P.Z.S.
Exhibition of a copy of an incised drawing of a hunted
OE Taree Perea Gee Vas csi sh cairn sama seueeeudnnetntstine nes aXe

Exhibition of, and remarks upon, the fossil rostral

teeth of Hopristis and Pristis .....0...00.c.c00ceccseeeeceeeee:

Proc. Zoo. Soc.—1918. b

Page

19]

163

196

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PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OV TITE

ZOOLOGICAL SOCIETY OF LONDON.

PAPERS.

1. Skull of Rana tigrina* Daud. By B. L. Buaria, M.Sc.,
and Batnr PrasHap, M.Se. (Assistant Professors of
Zoology, Government College, Lahore)f.

[Received August 8, 1917: Read February 5, 1918.]
(Text-figures 1-9.)

INDEX. Pages

SUMUCUUMEGMs fis fos acetate ates, 1-8

In a recent paper Nicholls (2) pointed out that Rana tigrina
~ Daud., which is generally used as a laboratory type throughout

India, differs markedly in several skeletal and other characters
from the common European forms (2. temporaria and R. escu-
lenta), of which a detailed description is generally given in the
ordinary English text-books. Nicholls had previously (1) pub-
lished a Note on the urostyle of &. tigrina and several other
anurous Amphibia, and in the first-mentioned paper he deals
with the vertebral column, the shoulder-girdle and sternum,
and the tenth spinal nerve; the skull of &. tigrina, which pre-
sents no less marked differences, however, has not received
consideration.

As far back as 1881, Parker (4), whilst working out the mor-
phology of the batrachian skull, published short descriptions of

* [Dr. G. A. Boulenger, F.R.S., informs me that the Lahore Frog has been
identified by him as belonging to the typical form.—Kp. P. Z. 8.
+ Communicated by Lieut.-Colonel J. STEPHENSON, D.Sc., I.M.S., F.Z.S.

Proc. Zoou. Soc.—1918, No. I. 1

2 PROFS. B. L. BHATIA AND BAINI PRASHAD ON THE

the skulls of several Indian species including 2. tigrina, but the
account of the latter is incomplete and suffers through having
been drawn up from the study of a single specimen. Both the
published description and the plate are inaccurate in several
-important respects, and therefore it was considered desirable to
work out in detail the anatomy of the skull of this common
Indian frog, and to correct the errors which have crept into the
otherwise excellent account given by him. For this purpose the
authors have prepared a large series of fresh skulls. and examined
them both in the wet and the dry condition, This has been
supplemented by an examination of the large number of skulls of
this type which are used for study by the students working in
the Government College laboratory.

The Cranium.

The skull in this frog is very much larger than in the two
common European species. An average-sized adult skull measured
40 mm. in the antero-posterior and 37 mm. in the transverse
direction, while one of the largest measured 45 mm. in the longi-
tudinal and 48 mm. in the transverse direction. The craniun,
which is wide behind, narrows somewhat anteriorly. The superior
surface is markedly arched, its most prominent point being a
little in front of the occipital region. All the bones associated
with the cranium, both investing and replacing, show a marked
development; thus the original cartilaginous structure has been
considerably reduced. As remarked by Parker (4), 2. tigrina
presents one of the most perfect examples of Batrachian cranial
architecture.

The Bones of the Cranium.

The Lxoccipital bones (text-figs. 1-4, eo.) bounding the foramen
magnum meet each other in the middle line ventrally, and leave
only a very smal] V-shaped area of unossified original cartilage
between their dorsal ends (all that is left uncovered of the
original tectum synoticum). ‘The tectum synoticwm never reaches
the superior border of the foramen magnum, and in the skull of
older specimens this little area also becomes ossified. The two
bones are seen meeting each other and the slight median pro-
jection on the posterior border of the fronto-parietals. In
conformity with the great strength and massive proportions of
this frog, the occipital condyles are large and are well seen in
both dorsal and ventral views of the skull (text-figs. 1-4).
Laterally, where the exoccipital meets the prootic it presents a
prominent bony ridge (processus mastoideus), there being a thin
strip of cartilage between the two bones in the young specimen
only.

The Prootic bones (text-figs. 1 & 7, po.) form a considerable
portion of the roof and anterior wall of the auditory capsule, and
extend forwards to form a portion of the inner wall of the orbit.
Dorsally each presents a quadrilateral area (vide text-fig. 1)

SKULL OF RANA TIGRINA. 5

Text-figure 1.

pm

Ge pees ad
Rana tigrina; dorsal view of skull.

-a.l,, alinasal process ; ¢.a., columella auris; e0., exoccipital; f., anterior fontanelle ;
f.o., fenestra ovalis ; fp , fronto-parietal; im., maxilla ; 2., nasal; p., palatine;
p.c., palatal cartilage ; pm., premaxilla; p.nl., prenasal process ; po., prootic ;
p.r., rostral process; p.rh., rhinal process; ps., parasphenoid; pt., pterygoid ;
q., quadrate cartilage ; q7., quadrato-jugal ; s., squamosal; se., sphenethmoid ;
sm., septo-maxillary ; s.2., septum nasi; sp., septum dividing the anterior part
of the sphenethmoid ; st., stapes; v., vomer. J, II, IIT, IV, V, VII, 1X, X,
refer to the foramina for the exit of the cranial nerves.

Text-figure 2.

ae

eo

y;
| LX, XA
R. tigrina ; ventral view of skuil.

For explanation of the letters see text-ng. i.

1*

4 PROFS. B. L. BHATIA AND BAINI PRASHAD ON THE

between the fronto-parietal and exoccipital on the inner side and
the inwardly directed horizontal flange of the squamosal, which
partly overlaps it, on the outer. Anteriorly the prootic bone
forms nearly the whole of the anterior wall of the auditory cap-
sule; a small area on the outer side of this anterior wall remains.
cartilaginous. On the inner side this anterior portion curves .
forwards to form the posterior part of the inner wall of the
orbit. In this angle is situated the foramen for the exit of
cranial nerves v. to vil. The foramen is completely surrounded

Text-figure 3.

H,IV

R. tigrina ; lateral view of skull.

Text-figure 4.

R. tigrina ; posterior aspect of skull.

For explanation of the letters see text-fig. 1.

by the prootic bone, and is not merely a notch completed below
by cartilage. The ventral and the posterior walls of the auditory
capsule are cartilaginous. Postero-laterally the prootic extends
to meet the exoccipitals; at. the junction of the two is a curved
irregular depression, the fossa tympanica, which lodges the
auditory ossicles and in which the foramen ovale is situated.
The suspensorium of the lower jaw is attached more externally
to the side of the auditory capsule. The styloid cartilage is
attached to the outer side of the cartilaginous portion of the
capsule.

SKULL OF RANA 'TIGRINA. 5

The Parasphenoid (text-figs. 2 & 6, ps.) is of the typical form,
but unusually strong. The transverse limb les under the
-occipito-auditory masses. Its posterior border is deeply concave
and the ends of the transverse limb are considerably wider than
the portion near the middle line. The posterior median process
of the longitudinal arm is short and often elegantly pointed.
‘The anterior longer longitudinal arm, besides forming the floor,
rises up on each side to form a portion of the lateral wall of the
‘cranium. About the middle it is broader than at the ends. The
outer edges articulate with the prootics, the cartilaginous portion
of the cranium, and the sphenethmoid. The cartilaginous por tion
of the side-wall of the cranium is velenoiahy much smaller than

the anterior sphenoidal portion.

Text-figure 5.

R. tigrina ; separated bones from the skull.

pm., premaxilla; m., maxilla; 7., nasal; fp., fronto-parietal ; s., squamosal.

The Fronto-parietals (text-figs. 1, 3, and 5, fp.) are adequately
described by Parker (4) in the following words :—

‘‘ Above, the fronto-parietals form a strong roof with a
notch in front, the remains of the frontal suture, but are
wholly coalesced beyond this; they end behind in two broad
wings which spread over the hinder region of the cranium
almost to the end. At first hollow in the middle, in the
postorbital region they develop a sagittal crest, which opens
out into two temporal wings. The temporal part dips into
the orbit and then rises over the ear-masses moulding on to
their sinuosities. The sides are notched, and the end has a
concave margin.”

6 PROFS. B. L. BHATIA AND BAINI PRASHAD ON THE

On removing the fronto-parietal it is seen that there is only
one anterior fontanelle (vide text-fig. 7, 7), bounded anteriorly
by a deep notch in the posterior margin of the sphenethmoid
and posteriovly by the cartilaginous cranium. The posterior:

Text-figure 6.

R. tigrina; separated bones from the skull.

v., vomer ; ps., parasphenoid; gj., quadrato-jugal; q., quadrate ; pt., pterygoid..

a

Text-figure 7.

prnl

\e oO.

R. tigrina; dorsal aspect of skull after removal of investing bones.

For explanation of the letters see text-fig. 1.

fontanelles usually found in other forms are absent. Parker, in
his description, stated ‘‘that the fontanelles are presumably like:
those of the lesser kinds,” but this is not the case.

SKULL OF RANA TIGRINA. 7

The Sphenethmoid (text-figs. 1, 2,3, 7, 8, and 9, se.) is more
strongly developed than in the European species, and extends
posteriorly almost to the region of the optic foramen. Anteriorly
it extends on either side so as to form a part of the anterior
boundary of the orbit (vide text-fig. 8); this feature is much
better marked in the adult specimens. In the average adult
specimen its front portion forms more than half the extent of the
nasal roof, floor, and the middle wall. In a dorsal view a lozenge
or diamond-shaped area of this bone (vide text-fig. 1) is left
between the nasals anteriorly and the anterior median notch of
the fronto-parietals behind.

Text-figure 8. Text-figure 9.

Se
R, tigrina.

Text-fig. 8.—Sphenethmoid, dorsal aspect after removal of investing bones.
se., sphenethmoid; p., palatine; p.c., palatal cartilage.

Text-fig. 9.—Sphenethmoid, anterior aspect.
I., aperture for olfactory nerve ; sp., septum.

The cartilaginous skeleton of the nose does not call for any
special remarks, except that in addition to the rhinal process
(text-fig. 7, p.rh.) there is a median prenasal rostrum (p.r.) in
continuation of the septum nasi (s.2.). A definite septo-maaillary
(sm.) 18 present on either side, extending horizontally from the
anterior wall into the nasal cavity, though Parker (4) regarded
them as mere ossifications in the nasal cartilage.

The Bones of the Face.

The Nasals (text-figs. 1, 3, & 5, 2.) are large broad-based
triangular bones, meeting each other in the middle line and
diverging posteriorly to enclose the diamond-shaped area of the
sphenethmoid referred to above, and to meet the anterior ends of
the fronto-parietals. Anteriorly also they extend as far forwards
as nearly to reach the nasal processes of the premaxille. The
apex or external angle of the triangular bone is drawn out and
extends outwards to meet the ascending process of the maxilla.
The Vomers (text-figs. 2 & 6, v.) meet each other in the middle

line posteriorly, but diverge anteviorly leaving a portion of the

8 ON THE SKULL OF RANA TIGRINA.

floor of the nasal capsule uncovered. The outer border presents
two notches, the posterior one bounding the posterior nares.
The vomerine teeth are situated in an oblique line along the
posterior border.

The Squamosals (text-figs. 1, 3, 4, & 5, s.) are strongly deve-
loped. From the posterior half of the cross-bar is given off a
horizontal shelf-like process covering the tegmen tympani and
extending inwards along its posterior border as far as the
junction of the prootic and the exoccipital.

The Pterygoids (pt.), Palatines (p.), Quadrato- ungelen (9.)-)s
Mazxille (m.), bones of the lower jaw and the hyoid apparatus
are of the usual type. An interesting feature, however, is the
presence of three well-marked depressions on the ventral side of
the upper jaw in its anterior portion (vide text-fig. 2). The
middle one of these depressions is between the two premaxille ;
it receives a corresponding median projection from the mento-
meckelians. The two lateral depressions are at the junction of
the premaxilla with the maxilla of each side; each accommodates
a strongly developed projection of the dentary of each side.

Literature cited.

1. NicHoLis, GEorcE E.—A Note on the Urostyle (0s coceygewm)
of the Anurous Amphibia. Proceedings of the Zoological
Society of London, 1915, p. 239.

2. NicHoLits, Georce E.—Some Notes upon the Anatomy of
Rana tigrina. Proceedings of the Zoological Society of
London, 1915, p. 603.

3. Parker, W. K.—On the Structure and Development of the
Skull of the Common Frog (Rana temporaria L.). Philo-
sophical Transactions of the Royal Society of London,
187 liwvel, VOI p: 37.

4, Parker, W. K.—On the Structure and Development of the
Skull in the Batrachia.—Part III. Philosophical Trans-
actions of the Royal Society of London, 1881, vol. 172,

Pele

ON A NEW SNAKE OF THE GENUS OLIGODON. 9

2. Description of a new Snake of the Genus Oligodon
from Upper Burma. By G. A. BouLEencER, F.R.S.,
E.2s.

(Published by permission of the Trustees of the British Museum.)
[Received January 1, 1918: Read February 5, 1918. ]
(Text-figure 1.)

OLIGODON HAMPTONI.

In1905 I described a new Oligodon * occupying an isolated
position in the genus, of which two specimens had been obtained

Text-figure 1.

Sar .
ty
or
age! |
fe
Sar
S

eine

on

Brats

ere
is Ns

Oligodon hamptoni, sp.n. X 1%.

at Mogok, Upper Burma, by the late Mr. Herbert Hampton.

_* Oligodon herberti Boulenger, Journ. Bombay N. H. Soe. xvi. 1995, p. 235,
pl. fig. 1.—The species has been rediscovered in Tonkin and notic2d as O. herberti,
var. eberhardti, by Pellegrin, Bull. Soc. Zool. France, xxxv. 1910, p. 30.

10 ON A NEW SNAKE OF THE GENUS OLIGODON.

Three years later the British Museum received another specimen
found at the same place by the same collector and which per-
tains to the same aberrant group, although unquestionably of a.
distinct species. I had put aside the specimen in the hope of
obtaining others; but as the hope must now be given up, I
propose to give a description of this handsome and very remark-
able Snake.

Nasal undivided; portion of rostral seen from above a little
longer than its distance from the frontal, penetrating rather far:
between the prefrontals; no internasals; frontal longer than its
distance from the end of the snout, shorter than the parietals ;
loreal small, longer than deep; one pre-and one postocular ;
temporals 1+1; five upper labials, second and third entering
the eye; three or four lower labials in contact with the anterior
chin-shields, which ave longer than the posterior. Scales in
15 rows. Ventvals 160; anal divided; subcaudals 32. A broad
yellow vertebral band, from the nape to the end of the tail,
between a pair of reddish brown, black-edged dorsal bands of
about the same width ; sides bluish grey, with two narrower dark
brown bands, the lower interrupted; head yellow with dark
brown markings: a spot capping the tip of the snout, a cres-
centic band from lip to lip through the eyes and across the
snout, an elongate spot on the frontal and on the suture between
the parietals, connected with a large occipital bifid spot which is.
continued as bands along the body, and an oblique band from the
parietal to the commissure of the mouth and below. Belly red,
with black bars occupying a whole shield or interrupted and
alternating ; lower surface of tail uniform red.

Total length 540, millim.; tail 70.

A single male specimen.

Although not longer than O. herberti, this is a much heavier
Snake, which must be regarded for the present as the largest and
handsomest of all the Oligodons.

Pi Z. S72 1918; LANTZ seals

Bale & Danielsson, Ltd.

LIZARDS FROM TRANSCASPIA.

ON REPTILES FROM THE RIVER TAJAN. “7 eh

8. Reptiles from the River Tajan (Transcaspia ).

By ce aaa Nize
[Received February 5, 1918: Read February 19, 1918. }

(Plate I.)

Tie small collection of reptiles which Lam about to describe:
was made from April to September 1914 by Mr. N. V.
Meriakri, who presented it to the Zoological Museum of the
Moscow University. Prof. G. A. Kojevnikof was kind enough
to entrust me with its study. ,

This material, although consisting of only 35 specimens, con-.
tains 16 species, one of which appears to le new. It thus affords.
interesting information on the herpetological fauna of the region
of the river Tajan, situated at the meeting of the Persian, Afghan,
and Transcaspian frontiers.

1, GYMNODACTYLUS MICROLEPIS, sp. n. (PI. I. fig. 1.)

3.6. Length of head and body (from snout to vent) 61, 60,
and 40 mm.

Head oviforimn, rather depressed. Snout much longer than the
diameter of the eye. Forehead slightly concave. Har-opening

small, elliptical. Proportion : song panos wed =()'27 to

0°30: average 0:28 length of head and body
5 © = 7 _ e
Body moderate, depressed. A well-marked lateral fold.
length of fore limb

= (42.

Limbs rather long. Proportion : lenerh See ada ae

to 0°43; average 0°43. Proportion : pesnyagelih oa Jara! Dea

=0°61 to 0:64; average 0°63. leneth of head and boy

Tail cyclotetragonal and slightly depressed at the base, becoming
cylindrical towards the end. Proportion : —— bone motaN
= 1-33 (1 spec.). length of head and body

Rostral broader than deep, with median clett above. Nostril
pierced between the rostral, the first supralabial, and 3 feebly
swollen nasals. 16 or 11 supralabials. Scales of snout, forehead,
and supraocular region polygonal, slightly convex, small in the
postnasal depression, enlarged in front of the eye. One enlarged
superciliary. Parietal and occipital regions covered with small,
roundish, feebly convex scales, intermixed with larger, more
convex, or slightly conical ones. On the temple a few large
moderately conical, but not keeled tubercles ; the other temporal
scales small, granular; in front of the ear-opening 2 or 3 small
tubercles.

Mental large, sub-triangular. 7 to 9 infralabials. 3 pairs
of chin-shields, the first »e¢ forming a suture behind the apex

* Communicated by G. A. BoutenaER, I.R.S., F.Z.S.

AD, MR. L. A. LANTZ ON

of the mental. Gular scales extremely small, roundish, scarcely
imbricate.

Neck with very small, juxtaposed, granular scales and longi-
tudinal rows of large, shghtly conical, but not keeled scales,
changing gradually along the back into 12 or 14 longitudinal
rows of moderate, elongated, trihedral tubercles; between the
median rows a row of small tubercles. The other scales of the
back larger than those of the neck, flat, feebly imbricate ; across
the middle of the body 62 to 65 dorsal scales (in a transverse line
passing between the tubercles).

Ventral scales small, cycloid, smooth, 35 to 38 in a transverse
row in the middle of the belly, 136 to 144 in the median line
from the mental to the vent.

Suprahumeral scales rather small, imbricate, more or less
rounded, smooth, or indistinctly keeled. Forearm covered with
scales like the dorsals, and with a few moderately keeled
tubercles. infrahumeral scales granular, juxtaposed. Infra-
radial scales like the ventrals.

Suprafemoral scales imbricated, pointed, smooth, or indistinctly
keeled on the inner side of the thigh; the other parts of the
thigh and leg covered with scales like the dorsals, intermixed
with large, moderately prominent trihedral tubercles. Infra-
femoral scales large, roundish, imbricate. Male with a continuous
series of 34 to 39 femoral and przanal pores. Infratibial scales
similar to the ventrals, but a little larger.

Tail covered above with transverse rows of very large, mode-
rately keeled spinose scales, decreasing in size towards the end of
the tail; between these rows of large scales about two rows of
small, imbricate, more or less distinctly keeled ones. On the
lower side of the tail, except at its base, a single row of enlarged
transverse plates.

Coloration grey above, with more or less distinct darker trans-
verse bands, which are disposed as follows :—one on the occipital
region, one on the neck, 4 to 7 on the back, about 12 on each
limb, 12 on the tai}. Lower parts white.

Comparative Notes.

The presence of a series of numerous femoral and preanal pores
show G'. microlepis to belong to the group of G. caspius Kichw.
It is especially closely allied to G. fedtschenkoi Str. and G’. longipes
Nik., agreeing with the former in proportions and with the
Jatter in most characters of scaling. J am greatly indebted to
Mr. W. A. Lindholm, who was so kind as to examine the types
and other material of G. longipes, which are preserved in the
Museum of the Petrograd Academy of Science *. Owing to his
notes, which complete the description given by Nikolskit, I
am able to state that these two species are quite distinct.

* Nos. 8809 (3 3), 8810 (1g, 2 2), 8811 (1 2), from Neh in Eastern Persia,
18. v. 1896, leg. N. A. Zarudny (types); Nos. 9191 (1@2), 91938 (1¢, 42), 9194
{1¢) from the country Zirkuh in Eastern Persia, 21.iv. to 6. v. 1898, leg. N. A.

Zarudny.
t+ Ann. Mus. Zrol. Acad. St. Petersb. 1897, p. 313.

REPLILES FROM THE RIVER 'TAJAN. 13.

Mr. Lindholm measured specimens of G. longipes and obtained
the following data :— .
I oh rete. 3 ¢
uength of fore limb ea a ee eee
Length of head and body 0°48 to 0°51; average 0°50 0°49 to 9°51; average 0°50
Length of hind limb
Length of head and body

= 0°69 to 0°73; average 0°71 0°65 to 0°70; average 0°68

The comparison with the corresponding data of G. microlepis-
shows the difference in the length of limbs*. Besides G. lon-
gipes has 12 to 15 supralabials, and the first pair of chin-shields
almost always forming a suture behind the mental; on one
specimen only out of 14 these plates are separated by two small
scales. As to the size of the tubercles of the head, neck, and
back, the scaling of the throat and belly, the number of femoral
and preeanal pores, the two species seem to agree.

There is a very interesting gradation in the characters of
scaling between the three species G, microlepis, G. fedtschenkot,
and G. caspius. G. microlepis has the smallest and the most
numerous scales, its tubercles are relatively feebly developed ;
G. fedtschenkoi forms the link between the foregoing and G. cas-
pius, which has the largest and the least numerous scales, and
very strongly developed tubercles. The following table con-
tains the most important distinctive characters of these three

species :—
G. microlepis. G. fedtschenkoi ¥. G. caspius t.
Tubercles of the temple .... moderately conical. conical, trihedral.
Before the ear-opening...... 2 or 3 small tubercles. 2 or 3 small tubercles. 1 large tubercle.
On the neck ................... slightly conical scales. moderately prominent very prominent large
trihedral tubercles. trihedral tubercles.

Tubercles of the back ...... moderate, prominent, large, prominent, not very large, very promi--

not spinose. spinose. nent, often spinose.
Gular scales .................. extremely small. very small, small.

Number ot scales in a line )
between the apex of the |
mental or the suture of > ~° 136 to 144. 128 to 131. 100 to 114,
the chin-shields and the |
analicletty Uerac.irt re. ss

Number of ventral scales
across the middle of the 35 to 38. 30 to 338. 24 to 29.
belly

Supratemorals on the inner 2small, smooth, or in- moderate, distinctly — large, strongly keeled.
side of the thigh ......... § distinctly keeled. keeled.

Number of femoral and 2 , oak
preanal pores. ....:....... ) SEL SUE E 26 §. 27 to 29 ||.

* Nikolski attributes as a distinctive character to G. ongipes the greater diameter
of the eye, which he supposes to be longer than the distance from eye to nostril ;
with this statement Mr. Lindholin does not agree.

+ Material: 2 ¢ from Samarkand.

t Material: 2 9 from Shemakha (Caucasus); 5 spec. (2 g, 19, 2 juv.) from
Sangatshaly near Baku (Caucasus): 3 spec. (1g, 22) from Askhabad (Transcaspia),
1 ¢ trom Anat (Transcaspia), and the @ described here trom the river Tajan.

§ 24 to 34 ventral scales and 30 to 37 pores, according to Nikolski, Fauna ot.
Russia. Reptiles I., Petrograd 1915, p. 78.

|| The maximum is 80, according to Nikolski, Joc. cit. p. 74.

14 MR. L. A. LANTZ ON

2. GYMNODACTYLUS casPIUs Hichw. (PI. I. fig. 3.)

1 ¢@. Length of head and body 65mm. 9/10 supralabials,
7/8 infralabials.. 14 longitudinal rows of dorsal tubercles. 65
“dorsal scales in a transverse row across the middle of the body.
101 scales in a line between the suture of the chin-shields and
the vent. 28 ventral scales across the middle of the belly.

3. AGAMA SANGUINOLENTA Pall.

4 specimens, agreeing in every respect with others from
Transcaspia (Askhabad, Anat, Bairam-Ah). Length of head and
body 83 mm. (<), 81 mm. (¢), 78 mm. (9), and 36 mm. (juv.).
14 to 17 supralabials, 15 to 17 infralabials. 43 to 47 gular scales
and 73 to 76 ventral scales in a line from mental to vent. 58 to
64 dorsal and ventral scales round the middle of the body. The

_young differs from the adults in having no spinose scales.

4, EREMIAS VELOX VELOX Pall.

2 specimens, entirely agreeing with other material from Trans-
-easpia (Askhabad, Bairam-Ali), Length of head and _ body
65 mm.(¢) and 34 mm. (juv.). 6 to 9 superciliaries. The large
-supraocular shields completely or almost completely separated by a
row of granules from the frontal and the postfrontal. Infranasal
not reaching the rostral. 6 anterior and 3 posterior supralabials,
6 to 8infralabials. 5 or 6 chin-shields in the young, the first 3 or 4
forming a suture. 22 or 23 gular scales in a line between the
suture of the chin-shields and the collar. 10 plates in the collar.
50 to 53 dorsal seales across the middle of the body. 30 trans-
verse rows of ventral plates, the longest of which consists of 13
to 15 plates. 20/21 femoral pores. Supracaudal scales strongly
keeled.

5. EREMIAS INTERMEDIA Str.

3 typical specimens. Length of head and body 55 mi. (@ ),
:37 and 38:5 mm. (juv.). In the 2 a granule between the
prefrontals. 6 to 8 superciliaries. The large supraocular shields
entirely separated by a row of granules from the frontal and the
postfrontal*, 6+1+2 to 4 supralabials. 7 or 8 infralabials.
26 or 27 gular scales in a line between the suture of chin-shields
and the collar. 11 or 12 plates in the collar. 47 to 50 dorsal
scales across the middle of the body. 29 or 30 transverse
rows of ventral plates, the longest of them consisting of 16 to
18 plates. 13 or 14 femoral pores on each side.

* I shall show in a more detailed publication that the subspecies transeaspica
Nik., which, according to the author, may be distinguished by this character, is
identical with the typical F. intermedia.

REPTILES FROM THE RIVER TAJAN. 15

6. Eremias (Mesauina) GurruLata Licht.

1 2. Length of head and body 49 mm. 5 superciliaries.
Row of superciliary granules beginning behind the 7th super-
ciliary only. Occipital as large as the interparietal. 4 anterior
and 3 posterior supralabials. 7 infralabials. 22 gular scales in a
line between the suture of the chin-shields and the collar. Collar
free, consisting of 9 plates. 40 dorsal scales across the middle
of the body. 28 transverse rows of ventral plates. 10 femoral
pores on each side.

i EUMECES SCHNEIDERI Daud.

1 spec. Length of head and body 109 mm. aS) and 63 min.
(hgr.). 6+1+3 supralabials in the ¢, 5+1-+42 only in the half-
grown specimen, the middle one being fused with the following
by forming a very large subocular. 8 infralabials. 4/5 or 5/8
nuchal plates. 25 or 26 dorsal and ventral scales round the
middle of the body. 68 scales in a line between the suture of
the chin-shields and the anal plates. .

8. Kumeces scuratus Theob.

3 spec. Length of head and body 122 mm. (¢), 124 and
116 mm. (both @). The head-shields offer many anomalies. In
one @ the right supranasal is divided; the other 2 has both
preefrontals divided into two unequal parts, and two loreals on
the left side. In the ¢ the parietals form a long suture behind
the interparietal. 4 to 6 superciliaries. 4 or 5+1+43 supra-
labials, the last being very small. 7 infralabials. 3/4 or 4/4
nuchal plates. 21 dorsal and ventral scales round the middle of
the body. 78 to 80 scales in a line between the suture of the
‘¢hin-shields and the anal plates.

2

9. MABUIA SEPTEMTANIATA Reuss.

2 spec. Length of head and body 82 mm. (¢) and 90 mm.
(2). Supranasals meeting in a point (d) or separated (2):
Prefrontals separated from each other, the internasal forming a
short suture with the frontal. 4 anterior and 2 posterior supra-
labials. 8 infralabials. 34 or 35 dorsal and ventral scales round
the middle of the body. 70 or 71 scales in a line between the
suture of the chin-shields and the vent.

10. VIPERA LEBETINA L.

1 spec.*. Length of head and body 520 mm. Tail incomplete.
10 supralabiils, 13/14 infralabials.. 25 longitudinal rows of
dcrsal scales. 121 ventral plates.

* It was impossible to ascertain the sex of the snakes, the viscera having been
removed.

16 MR. L. A. LANTZ ON

11. Boga rRIGgoNAtTUM Schneid.

2 specimens :—

Length of head and body............... 510 480 mm.

Length of tathy.cc0).).) icc). es ce a 108.4;

Number of supralabials ............... 8/9, the 3rd, 4th, and 5th entering-

Number of infralabials . ............... 11/12 [the orbit.
é 2+14+3

Memporalishield’s) case. hoctai. tA ee: 2+3 14343

Number of rows of dorsal scales...... 21

Number of ventral plates ............ 222 221

Number of pairs of infracaudals ... 86 84.

12. TAPHROMETOPON LINEOLATUM Brandt.

1 specimen. Length of head and body 790 mm. Length of tail
375 mm. 8 supralabials, the 4th and 5th entering the orbit.
11 infralabials. 2+1/2+3 temporal shields. 12 rows of dorsal
scales. 181 ventral plates. 121 pairs of infracaudals.

13. ZAMENIS RHODORHACHIS Jan. .

2 specimens :—

Length of head and body................ 715 630 mm.

Length of tat) ..21.3.: Se ees 6S 230 ,,

Number of supralabials ............... 9, the 5th and 6th entering the
Number of infralabials.................. 10 [ orbit...
Teamporal’shields cc sc0- of cape peace 2+5 to 8 irregular ones.
Number of rows of dorsal scales...... 19

Number of ventral plates................ 227 226

Number of pairs of infracaudals ... 121 17,

These specimens have no red stripe along the back.

14, ZAMENIS DIADEMA Schleg.

4 specimens :—

Length of head and body ............ 870 830 825 . 770mm.
Length oftailis.) Uaioe eee. He INeOdS 190 215 (140) ,,
Number of supralabials .:....)........ 11/12 ° 123% 1@Aa) 12s
Number of intralabials ... ............ 11/18 12 13/14 12
Number of rows of dorsal scales ... 27 29 27 27
Nuinber of ventral plates ............ 225 244 217 234
Number of pairs of infracaudals .... 76 83 83 | =

Behind the prefrontals a row of 3 accessory shields; in one
specimen the median one is fused with the right inter-

1
aii} frenals. 2 preoculars.

Supralabials more or less separated from the loreals and entirely
separated from the orbit by a row of small shields, the first of
which may reach the postnasal; there are 2 or 3 shields between
the supralabials and the loreals, 3 to 5 between the former and
the orbit, and, following them, 2 or 3. postoculars. Temple
covered with small irregular shields.

nasal. 3 (=) exceptionally 4 (

REPTILES FROM THE RIVER TAJAN. 17

15. NATRIX TESSELLATA Laur.

3 specimens :—

Length of head and body ................ 580 530 470 mm.
Dengthrotttatlact. ss cevustemaset. 165 - 166 195. .;
Si Bat Ge aaa
Namber of supralabials: x2... .....¢..... 8
Number of infralabials ..........:......: 10/11
Number of preoculars .................. 4, 2/3 2
siNumber of postoculars ........0.....-.: 4A 4 3
L——-~/- =)
Temporal shields . .. ....0:<... 002.00 «i s20%s: 14+2+2 or3
Number of rows of dorsal scales ...... 19
Number of ventral plates ............... 176 170 176
Number of pairs of infracaudals ...... 70 73 69

In the specimen having 2 preeoculars and 3 postoculars the 4th
and 5th supralabials enter the orbit.

16. Eryx minraris Pall.

1 specimen. Length of head and body 355 mm. Length of
~ tail 30mm. _ Internasals separated by the point of the rostral.
4 scales between the postnasals. Round the eye a circle of 13
small shields, the lowest of them much enlarged and reaching
the 6th supralabial, the two anterior ones a little enlarged (pre-
oculars), the others (supraoculars and postoculars) about equal
in size. Between the supraoculars 5 scales across the head.

ieee
Between the postnasals and the preeoculars 8 (H+t+4) small

loreal shields. 13 supralabials, the 3rd one being the highest.
20 infralabials, the first 3 or 4 enlarged, the others very small,
with larger shields below them.

EXPLANATION OF PLATE I.
Fig. 1. Gymnodactylus microlepis, sp. n., 6, River Tajan.

Fig. 2. G. fedtschenkoi Str., 6, Samarkand.
Fig. 3. G. caspius Kichw., 2, River Tajan.

Proc. Zoou. Soc.—1918, No. II. 2

ee, ae ae "1
ne Rta groit ata Bt tad
ee She nes

Pistia

eee te

THE EXTERNAL CHARACTERS OF THE LEMURS. 19

A, On the External Characters of the Lemurs and of
Tarsius. By R. 1. Pocock, F.R.S.

[Received March 5, 1918; Read March 5, 1918. ]
(Text-figures 1-16.)

TABLE OF CONTENTS.

Page
That mOdGtiOM. 2. tects eecgsee aren ee secon cnr ora ALO.
The Muzzle and the Rhinarium........................... 20
Wbtre wai erates hac: oa Novae ti cee ni thas seks iohataseenens Le
The Facial and Carpal Vibrissze........................ 24
The Glands of the Fore Limb..............0cc0000.002.0. Bd
(ines bands and! Weetsien.'cesavctttwie tek diction 27
Mes Sa DMN GU 4agee cosh as ate ev neared on aheceer asa. © 0
The Anus and its Glands...... 0.00.0 .....ccce ce cee eeeeeeeee 40
The External Genitalia of the Male .....0............... 42
The External Genitalia of the Female .................. 47
General Conclusions and Systematic .................. 06. 51

Introduction.

The materials upon which this paper is based are mainly the
lemuroid Primates which have died in the Zoological Gardens
during the past ten years or so. Representatives of practically
all the commonly imported menagerie species of the group have
passed through my hands in that time, namely, species belonging
to the genera Chiromys, Chirogaleus, Lemur, Galago, Perodicticus,
and Nyeticebus. For the loan of examples of Hemigalago and
Tarsius 1 am indebted to Prof. Wood-Jones, the Society’s
Prosector. I am also indebted to Mr. Oldfield Thomas and to
Prof. J. P. Hill for the chance to examine other specimens of
Tarsius. I have not, however, been able in all cases to see repre-
sentatives of both sexes of the species; and of many admitted
genera, notably Microcebus, Mixocebus, Lepilemur, and Loris, no
specimens have come to hand. This applies also to the three
genera of Indrisidee (Zndris, Propithecus, Lichanotus), which, like
Tarsius, seem to be intolerant of captivity even in their own
countries. The external characters of the Indriside, however,
have been tolerably fully described and illustrated in Milne-
Edwards and Grandidier’s great work on the Fauna of Mada-
gasear. From this I have freely borrowed. Unfortunately no
text accompanies the numerous plates on the various species of
Lemuride published in that work. Of other treatises dealing
with the Lemurs on a comprehensive scale the most important
is the paper by Mivart and Mure (Tr. Zool. Soc. vii. 1872), in
which some of the external characters of a few diverse types are
dealt with from the comparative point of view. * The rest of
the bibliography consulted consists mostly of special memoirs on
particular species, like Owen’s paper on Chiromys, Burmeister’s

on

20 MR, R,. I. POCOCK ON THE EXTERNAL

on Tarsius, Van der Hoeven’s on Perodicticus, Huxley’s on
Arctocebus, Beddard’s on Hapalenwur, and so forth.

Many of the facts dealt with im the following pages are
of course well known. Some characters, however, are here
described, I believe, for the first time; and I trust that the
collation of the facts and their comparative treatment may prove:
useful to future students of this group. .

In the matter of names [ have followed the conservative course
of using Lemur for the species to which it has been by common
consent assigned in all recent literature, although by the rules
of nomenclature, it has no right to a place in the Primates.
at all, but belongs by Storr’s very definite selection to the
Dermoptera, G'aleopithecus volans being its type-species. Even if
that selection be set aside, it appears to me that the “ indications”
of the 10th edition of the ‘Systema’ show that the species known
as Loris tardigradus is its type. This is clearly a case for the
‘* Fiat”? Committee on Mammalian generic names; and it is my
confidence that the Committee will see the wisdom of allowing
Lemur catta to stand as the type of Lemur, that induces me
to retain this generic name in its commonly accepted sense.
Similarly I have employed Chiromys for the Aye-Aye (mada-
gascariensis), although Daubentonia is the correct title for that
species. The Fiat Committee, I believe, has these names now
under consideration.

The Muzzle and the Rhinarium.

The bestial aspect of the face of the Lemuroid as compared
with the Pithecoid Primates is not due to the general elongation
of the jaws. In this character the Lemurs are surpassed by
Papio amongst the Pithecoids, It is due to two correlated
features, namely, the retention of the primitive moist glandular
rhinarium and the projection of the upper jaw supporting it.
beyond the level of the chin.

The rhinarium is naked to a varying extent on its dorsal side
and also beneath the nostrils laterally and in front. It is con-
tinued downwards in front as a strip of grooved naked skin
cleaving the upper lip to its inferior edge. At this point the lip
is adherent to the gum covering the premaxille, so that it is
incapable of protrusion.

Although the rbhinarium is tolerably similar throughout the
group, one or two variations may be pointed out. In the typical
Mascarene Lemurs, including Chirogaleus (text-fig. 1, A), the labial
extension of the rhinarium is comparatively short and the later-
ally extended infranarial portions shallow. In Perodicticus the
infranarial portion is deeper ; but in Vyeticebus it is not so. In
Galago crassicaudatus (text-fig. 1, B) the labial extension of the
rhinarium is « little longer and thinner than in Chirogaleus
and Lemur. In Hemigalago denidoffi it is remarkably long and
gradually widens above where it passes into the infranarial portion

’ CHARACTERS OF 'THE LEMURS AND TARSIUS.’ 24

of the rhinarium ; but in the length and shape of the rhinarium
G. senegalensis is intermediate between G. crassicaudatus and
H. demidoffi (text-figs. 1, C; 2, B).

In Chiromys the vhinarium is not so prominent; the nostrils
are longer, more slit-like, and separated in front by a narrower
septum. The infranarial portion is very deep towards the middle
line, reaching almost to the edge of the upper lip and making the
labial extension of the rhinarium appear very short. In this
genus also the frenum which ties the lip to the gum between the

Text-figure 1.

Biss
SA

ene

ss

.
y

YA,
Wy
Ay

A. Rhinarium of Chirogaleus major.

B. Rhinarium of Galago crassicaudatus.
C. Rhinarium of Hemigalago demidoffi.
D. Rhinarium of Chiromys.

E. Nose of Tarsius.

points of insertion of the incisor teeth is somewhat longer than in
typical Lemurs, so that the lip is capable of being protruded to a
slightly greater extent in the middle line. It is probable that
the variations of the rhinarium and upper lip, hke many of the
structural characters in Chiromys, are correlated with the rodent
dentition and peculiar method of feeding of this Lemur; but our
knowledge of the function of the rhinarium is too imperfect to
warrant more than a suggestion on this point (text-fig. 1, D).

22. MR. R. I. POCOCK ON TUE EXTERNAL

I have not been able to examine in a fresh state the rhinafium
of any species of Indriside.

The muzzle of Z'arstus is very different from that of all Lemurs..
The only trace of: the rhinarium, if such it can be called, that
remains is a narrow rim of naked skin surrounding the nostrils,
which are widely separated as in the Platyrrhine Pithecoid
Primates. The nose scarcely projects at all, and the muzzle is.
squarely truncated and deep, and the upper jaw hardly overhangs
the lower, so that in profile view the muzzle has a decidedly:
feline appearance, contrasting markedly with the generally
canine appearance of that of the true Lemurs. The long upper
lip is undivided and continuously hairy from side to side across-
the middle line, and its frenum is set higher above the incisor
teeth, suggesting that the lip is susceptible of partial protrusion
after the manner of the lip of the pithecoid Primates, but to a
lesser degree (text-figs. 1, E; 2, A).

The primitive muzzle of the lemuroid Primates, with its
rhinarium and adherent upper lip, is associated with the habit of
drinking by means of lapping. The specialised muzzle of the-
pithecoid Primates with aborted rhinarium and_ protrusible
upper lip is associated with the habit of drinking by means of
suction. But Varsius, although more resembling the pithecoids
in the structure of the muzzle, drinks, it is said, by the lapping

method*, This fact is full of interest in connection with the

view, supported by other considerations, that Zarsius links the-
Lemuroids and Pithecoids together.

The Karvy

In the species referred to Lemur the pinna of the ear is
tolerably uniform in shape and structure. It is small and con-
cealed to a greater or less extent by its own bairs and those of
the surrounding area of the head. Its superior posterior edge is.
not folded; but the anterior edge of the upper half forms a
strong ridge overlapping the anterior end of the simple shelf-like-:
supratragus (plica principalis) and descending below it and
vanishing in the capsule of the pinna above and within the
small lobate tragus. The antitragus is somewhat larger than

the tragus, sometimes much larger (Lemur catta); and the deep.

notch between them is approximately on a level with the external

auditory meatus. The flap of the ear behind the antitragus is.

marked with a depression, the anterior and posterior margins of
which are respectively the ridge running upwards from the anti-
tragus and the adjoining postero-inferior edge of the pinna.
The supratragus, forming the upper boundary of the capsule of

* H. Cuming, P. Z.S. 1838, p. 67. Mr. Cuming also makes the interesting
remark that when any object is put near a Tarsius, the animal “draws up the
muscles of the face similar to a Monkey and show its.... teeth.” I have never
seen Lemurs behave in this way.

+ Described in several types by Mivart and Murie, Tr. Zool. Soc. 1872

—_

CHARACTERS OF THE LEMURS AND TARSIUS. 23

the pinna, is set comparatively high up, approximately halfway
between the tragal notch and the upper edge of the pinna.

In Chirogaleus (text-fig. 2, C) the ear is similar to that of Lemur,
but the ear of Mfcrocebus is provided with a much larger pinna
which, judging from a living example, is ribbed above the capsule
and capable of being folded as in the Galagos. In the latter, as
is well known, the pinna is of great size. The portion of it just
above the capsule is ribbed and grooved and susceptible of
folding. The supratragus is more expanded and more flap-like

Text-figure 2.

Renita

Wy
YELLE,

C

A. Head of Tarsius.

B. Head of Hemigalayo demidoffi.
C. Head of Chirogaleus major.

1D. Head of Perodicticus.

Figures drawn from spirit specimens. All X $.

than in Lemur and the pouch, probably the homologue of the
bursa of the ear of the Carnivora, is set higher up than in the
Lemurs. Vycticebus and Perodicticus have the pinna no larger
relatively than in Lem, and it is simplified by the almost total
suppression of the tragal and antitragal thickenings; but, as in
Galago, the pouch is set high up and the supratragus is flap-like
and valvular (text-fig. 2, D). As Mivart and Murie pointed out,
there is a small fold of the integument above the supratragus in
Nycticebus, and a similar but better developed fold is developed

24 MR. R. I. POCOCK ON THE EXTERNAL

in Arctocebus. Huxley cites the presence of this fold as one of
the features distinguishing this genus from Perodicticus.

Although the ear of Chiromys is relatively as large as in the
Galagos, it is not ribbed and grooved above the capsule. The
supratragus is a thickened ridge as in Lemur; but the tragus is
not an angular projection as in that genus but a simple ridge,
and the notch between it and the well-developed antitragus is
comparatively deep and wide. Its lower rim, however, does not
extend downwards so low as the external auditory meatus, the
portion of the pinna just beyond this meatus being elevated as in
Carnivora, Ruminants, and many other Mammals.

The ear of Z'arsius is similar in all essential details to that of
the Galagos, except that the supratragus and the antitragus are
somewhat larger and more valvular (text-fig. 2, A).

The simplest type of ear in this group, and I suspeet the most
primitive, is that of Lemur and Chirogaleus, ears with a greatly
expanded and ribbed pinna and valvular .supratragus being
derivative and specialised structures. According to this view the
ear of Tarsius is the least primitive of all. It is gradationally
linked with the ear of Chirogaleus by the ears of Galago and
Microcebus.

7

In the development of the facial vibrissz * the most generalised
type I have examined is Chirogaleus major, where the mystacial,
superciliary, genal, and interramal tufts are all well developed
(text-fig. 2, C). There is a single genal tuft on each side set low
down behind the corner of the mouth. Most of the species
referred to Lemur resemble Chirogaleus except that the inter-
ramal tuft is absent; but in Z. variegatus it is usually, if not
always, retained, although of small size. The full complement of
tufts 1s present in Chiromys, but the vibrisse composing them
are generally shorter than in the typical Lemurs, and in two
cases the interramal tuft was reduced to a single vibrissa.

In the Galagos (Galago crassicaudatus, G'. senegalensis, and
Hemigalago demidoffi) the vibrisse are poorly developed as com-
pared with those of the typical Mascarene Lemurs, more par-
ticularly Chirogaleus major, with which the Galagos were at one
time associated. ‘The interramal tuft appears to be invariably
absent, and the genal tuft is set high up on the cheek a little
below and behind the posterior angle of the eye (text-fig. 2, B).
It resembles in position the upper genal tuft of the typical
Carnivora, whereas in Chirogaleus and Lemur the genal tuft
resembles in position the lower of the two tufts in that order.
In WNycticebus and Perodicticus the vibrisse are even less well
developed than in the Galagos, the genal tuft being suppressed
in the specimens examined (text-fig. 2, D). The genal and
interramal tufts are also absent apparently in Tarsius, although

The Facial and Carpal Vibrisse.

* P.Z.S. 1914, pp. 889-912.

CHARACTERS OF THE LEMURS AND TARSIUS. 25

Burmeister figures some vibrisse in front of the ear. These do
not, however, correspond in. position to the genal tuft of the
Galagos (text-fig. 2, A).

The prevalence in most orders of Mammalia of the complement
of tufts described above as occurring in Chirogaleus suggests that
the absence of one or more of the tufts is a derivative and not
a primitive feature. In this respect the Lorises, Pottos, and
Tarsius are more specialised than the Mascarene Lemurs.

The Carpal vibrisse in Lemurs have been studied by Beddard
and Bland-Sutton. Confirming and extending their observations,
I may add that I have found these tufts of tactile bristles in
Chirogaleus major, Hapalemur griseus, and in examples of the
following species of Lemur, namely, catta, variegatus, macaco,
mongoz, coronatus, rufiventer, albifrons, and many of the species,
subspecies or varieties grouped round the last. They are not
always easy to detect in the thick fur, and sometimes appear to
be wanting; but in such cases I suspect their absence is due to
moulting or to artificial removal by rubbing. I have not found
them in Chiromys, Nycticebus, Perodicticus, or Tarsius ; and, since
Bland-Sutton also noticed their absence in Perodicticus, it may
be inferred that their absence is characteristic of the Asiatic and
African lemuroids*.

The prevalence of these vibrissee in many orders of Mammals
suggests that they are a primitive Metatherian and EKutherian
character, a suggestion which involves the conclusion that thei:
absence in the above-mentioned Lemuroid genera is due to
‘suppression and is a derivative feature.

The Glands of the Fore Limb.

In Lemur catta, but in no other species referred to the genus
Lemus, there is a peculiar gland on the fore-leg, which was
figured and described by Biand-Sutton + and also figured by
Milne-Edwards and Grandidier. <A strip of black naked skin
extends from the palm of the hand over the wrist up the distal
third of the corresponding surface of the fore-leg. It ends
proximally in a smooth elliptical area, which is present even in
the newly-born young (text-fig. 3, A) In adult males the
elliptical area is raised into a swollen cushion-like pad composed
of white tissue, fatty in appearance and consistency and covered
with black skin. On the ulnar side of the pad a large, erect,
solid horny excrescenee is developed (text-fig. 3, B). This-varies
in size apparently with age; its apex is sometimes bifid, but
generally simple, and it is sometimes present on one limb and

* I have failed to detect the carpal vibrissse on dried skins of Indris and Propi-
thecus; and in both these genera the interramal tuft appears to be absent. Also
in the one skin of Indris available for examination the genal tufts are wanting,
whereas in a skin of Propithecus diadema these tufts are well developed and set
low down on the cheek as in Lemuride.

+ Proc. Zool. Soc. 1887, pp. 369-372. In 1863 Gray pointed out the presence of
this structure in L. catta and its absence in other species.

26 MR. R. I. POCOCK ON THE EXTERNAL

absent on the other. In females the excrescence is generally
absent or quite small. Only in rare cases of probably aged
individuals is it comparable in size to that of the males; and the
elliptical area is generally flat in the females.

Text-figure 3.

[eg
545 5
We OD 22a Ea
// \N S Wy) Wo SDA
Ws tp, oan — oe
i i ae 53 F kgs KEE Sa Ly Eh rb! Ne SAA
i Cs LZ Sa a OOS ~S EL ZS bn T=
ELLE ee . 4) Z a aes
Y ¢ Dig @ SSS SSE
= Sf ~ SOLS = =
MW see
AXA vv FN
NYA) V\MIRRSSeSSR
y 1} is NN
yi |

Ss Wi UM

Ky Re
am

SS : ae @ ;
WY seer?

iO O8 Or" ~_—

’ D

A. Fore-arm of Lemur catta, newly born, showing the extension of the naked
skin from the palm of the hand and the elliptical glandular area above the
wrist.

B. Section through the glandular area and the horny spur of the fore-arm of an.
adult male Lemur catta.

C. Superficial view of the shoulder-gland of an adult male Lemur catta.

D. Vertical longitudinal section through the same.

E. Vertical transverse section through the same.

F. Section through the shoulder-gland of adult female Lemur catta.

According to Bland-Sutton the ducts of numerous sweat-glands.
open upon the surface of the skin of the elliptical area. Secretion,
no doubt, exudes through the pores; but I have never succeeded
in squeezing any visible secretion to the surface by pressure.

The only use I have seen the Lemur make of this structure is.
to apply it to the tail by drawing simultaneously the applied
wrists along that organ from near the base to the point.
Possibly by this means the secretion is wiped on the long hairs.

CHARACTERS OF THE LEMURS AND TARSIUS.,. 27

to scent them. But so far as my observations go, this action is.
restricted to the male when stimulated to anger at the time
of rut.

Also in Lem catta, but in no other species referred to the
genus, there is a large gland in the male on the inner side of
the upper arm near the top of the. biceps muscle’ just below the
shoulder (text-fig. 3, C). It is a circular or elliptical mound-like:
elevation covered with short hair, except in the centre of the
summit where there is a small naked area carrying the orifice of
the gland which is usually shaped like a strongly curved crescent.
In section the gland is seen to be composed of a thick-walled
sac, the cavity of which is subdivided by ridges and outgrowths
of the walls, making, in a sense, a multilocular gland. It is the
partial blocking of the rounded orifice of the gland by one of
these outgrowths that gives the orifice its crescentic shape. The:
cavity of the gland is filled with strong-smelling brown sticky
secretion which under pressure can be “squeezed, hike a worm,
from the orifice (text-fig. 3, D, EK).

In the female this gland is not always developed, and when
present it consists of a small elevation covered with normally long
hair and having no cavity or trace of orifice (text-fig. 3, F).

The suggestion that these glands are modified mammary glands
is, I think, erroneous. At all events they coexist with the normal
pectoral mammary glands.

It is interesting to recall that Hapalemur also has a similar
gland on the shoulder and a somewhat similar gland above the
wrist. I have, however, seen these only on dried skins, and have:
nothing to add to the descriptions published by Beddard * and
Bland- Sutton +

But there are two points worth attention arising out of the
facts just mentioned. Despite the development of these glands
in Lemur catta and Hapalemur griseus, and in no other species,
these two Lemurs do not appear to be nearly related. Judging
from cranial and dental characters, the relationship of Hapalemur
griseus 18 with Prolemur simus, and of Lemur catta with the
other species usually assigned to the genus Len.

The second point is the coexistence in two otherwise dissimilar
genera of iwo sets of glands, one below the shoulder, the other
above the wrist. ‘This raises the very important question of
possible correlation in development between two or more struc-
tures, a question which opens a very wide field for research in
zoology.

The Hands and Feet.

In the species of Zemur the digits of the hand are longish and
slender, and free from webbing to approximately the same extent
asin Man. The pollex is the shortest of the series and is sepa-
rated by a wide space from the second, the base of which it

* Proc. Zool. Soc. 1884, p. 8393; and 1891, p. 450.
+ Proc. Zool. Soc. 1887, p. 369.

28 MR. R. I. POCOCK ON THE EXTERNAL

slightly overlaps when laid forwards. It can be extended at right
angles to the long axis of the palm, but is not truly opposable
since the large composite palmar pad—-the ‘‘ ball” of the thunb—
at its base is almost stationary. The succession of the digits in
length is 4, 3, 5, 2, but the fourth only exceeds the third slightly,
and the second and fifth are approximately equal. The palm is
longish, longer than wide, and passes proximally beyond the base
of the pollex. Of the four interdigital pads, the first or pollical
is confluent with the inner proximal (thenar), constituting the
“ball” of the thunb. The remaining three interdigitals form
a transverse trilobed cushion-like pad. Behind the fourth or
external interdigital, the external portion of the palm is occupied

Text-figure 4.

A. Foot and B. Hand of Lemur macaco; X 3.

1, 2, 3, 4, the intermediate pads; I, II, the proximal pads.

by the elongated external proximal (hypothenar) pad, which is
subdivided, the posterior expanded portion lying alongside the
posterior portion of the ‘ball’ of the pollex, from which it is
separated by a groove (text-fig. 4, B).

In its main features the hand only varies in minor particulars
in the different species examined, namely, Z. catta, macaco, albi-
frons, mongoz, variegatus, and coronatus, except that in L. varie-
gatus the palm is a little wider as compared with its length.

In the foot the hallux is of great length and thickness, 1s
capable of being extended at right angles to the sole, and is
opposable, the ‘ ball” of the hallux, consisting mainly of the
large, projecting first or hallucal interdigital pad, being movable

CHARACTERS OF THE LEMURS AND TARSIUS. 29:

and capable of being pressed against the second interdigital pad
and the elongated external proximal pad, closing up the depression
in the middle of the lower surface of the foot. For the rest, the.
digits and the pads of the foot resemble in a general way those
of the hand (text-fig. 4, A).

A point to notice is that in Lemur catta the naked sole is
extended proximally to the tip of the caleaneum or heel. In the
other species the heel is covered with hair, the hairy area being:
about one-third the length of the naked area, or a little more.

Text-figure 5.

A. Hand and B. Foot of Chirogaleus major ; nat. sized
C. Hand and D, Foot of Galago senegalensis; nat. size.

In Chirogaleus major the hands and feet are substantially
similar to those of Lemur, but the naked palmar and plantar.
surfaces are broader as compared with their length, and the
individual pads ave more sharply differentiated, and, judging by
their papillary ridges, endowed with greater tactile sensibility.
In the foot the hairy area of the heel is relatively longer, being

-30 MR. R. I. POCOCK ON THE EXTERNAL

-about two-thirds the length of the naked portion. In the speci-
men examined the third and fourth digits were approximately
equal both on the hands and feet (text-fig. 5, A, B).

The hands and feet of the Indriside, as figured and described

by Milne-Edwards and Grandidier, show some interesting pecu- —

harities suggesting more complete adaptation to arboreal life than
in the Lemuride. In Propithecus the digits of the hand are
hardly more webbed than in Lemur, but the palm is narrower,
especially posteriorly, where the pollex arises. The pads appear
to be very little differentiated, and the absence of the “ball” of
the thumb indicates a feeble grip for that digit.’ In Lichanotus
(Avahis) the ball of the thumb is better developed, but the palm
-of the hand is apparently longer than in Propithecus, the second

Text-figure 6.

A. Foot and B. Hand of Chiromys; X 3.

“digit, which is very short, being widely separated from the
pollex. The third, fourth, and fifth are long and united by
narrow webbing approximately to the ends of the first phalanges.
In Jndris also the hand is long and slender, with a wide space
between the long weak pollex and the second digit. The latter,
however, is not so short as in Lichanotus, and is united to the
third, as the third is to the fourth and the fourth to the fifth, by
integument permitting but slight divarication of these digits and
increasing in appearance the elongation of the palm.

In the feet there is less variation. The hallux is normally

-elongated, but is slender from base to apex without the muscular

-development seen in other Lemuroidea, and the digits are webbed

CHARACTERS OF THE LEMURS AND TARSIUS. 3

approximately to the distal ends of the first phalanges. In
Indris the interdigital webbing is somewhat deeper than in the
others and extends to an equal distance along all the digits, but
in Propithecus and Lichanotus it is deeper between the third
and fourth than between the second and third and the fourth
and fifth.

Some of the peculiarities of the hand of Chiromys, such as the
presence of claws and the modification of the third digit, are well
known (text-fig. 6, B). The palm is narrower than in the true
Lemurs. The “pollex is not truly opposable, but it is so long that
when turned forwards it overlaps the palm, asin Man. All the
other digits are long, even the second and fifth, which are sub-
equal, exceeding the length of the palm. The third is very long
and slender ; the metacarpal that supports it projects beyond the
palm, a unique modification which confers exceptional mobility
on the digit. Nevertheless this digit is shorter than the fourth,
which is nearly twice as long as the palm. Although a shallow
web joins these two digits at the base, the hand may be described
as unwebbed. In correlation with the grasping capacity of the
large claws, the digital pads are comparatively slightly expanded.

The feet are less modified than the hands and conform to those
of the Lemur-type, except that the digits are thinner, the hallux
is a little shorter and weaker, the second, third, fourth, and fifth
are a little longer and armed with claws correlated with narrower
pads. The heel is covered with hair as in all Lemurs except
L. catta (text-fig. 6, A).

In this genus the unique elongation of the metacarpal of the
third digit of the hand and the slenderness of the digit are
adaptations to feeding.

In Galago of the crassicaudatus and senegalensis types the hand
is wider than in Chirogaleus and its pads still better differentiated
and more prominent, especially the first, or pollical, intermediate.
All the normal six pads are distinguishable. . The internal
proximal is a small pad situated near the base of the outer border
of the thumb, and the external proximal is a rounded pad in
contact with the internal intermediate and not extending back-
wards to the wrist as in Chirogalews and the true Lemurs. A
wide space separates it from the internal proximal, which is also
set nearer the wrist. Similar differences obtain in the feet,
where all the six pads are clearly defined. In G. crassicaudatus,
monteirt and their allies the posterior part of the naked portion
of the foot, as Beddard pointed out, is covered with coarse close-
set papille, and the hairy area of the: heel is longer than in
Chirogaleus major, being about as long as the naked portion.
In G. senegalensis it is considerably longer (text-figs. 5, C, D;
htAc) By.

In Hemigalago the pads show some interesting differences
from those of Galago. In the hand the six pads are even more
sharply defined, and are arranged so as almost to encircle a large
naked submedian area of wrinkled skin. ‘The two proximal pads.

32 MR. R. I. POCOCK ON THE EXTERNAL

are almost in contact in the middle line, the external being elon-
gated and extending practically from the wrist up to the internal
intermediate. Behind the two proximal pads there is a short.

Text-figure 7.

A. Hand and B. Foot of Galago crassicaudatus.
1-4, the intermediate pads; I, II, the proximal pads.

area of naked wrinkled skin. The pollex is longer and more
prehensile than in Galago. The arrangement of pads on the
foot is similar, but the two proximal pads are relatively much
smaller than in the hand, and the internal is set farther from the

CHARACTERS OF THE LEMURS AND TARSIUS. 33

hallucal intermediate and is widely separated from the external.
In this genus there is no definite heel-like area behind the pads
as in Galago, the skin round the proximal pads being merely
soft and creased. ‘The hairy area of the foot back to the tip
of the caleaneum is nearly twice as long as the naked area (text-

ive. 9, A, B).
Text-figure 8.

ik

4

)
5
N

~~ «—)

A. Hand and B. Foot of Nycticebus; x }.
C. Hand and D. Foot of Perodicticus; x 3.

1-4, the intermediate pads; I, IT, the proximal pads.

The arrangement and distribution of the pads give a primitive
stamp to the hands and feet of Hemzgalago in the sense that they
recall very forcibly the pad-development seen in the extremities
of many Rodents, Insectivores, and Marsupials.

The hand of WVycticebus and Perodicticus differs in some

Proc. Zoot. Soc.—1918, No. III. 3

34 MR. R. I. POCOCK ON THE EXTERNAL

remarkable particulars from that of the other genera’ hitherto
noticed. It is capable of being turned at right angles to the axis
of the forearm. ‘The palm is short and broad, and the powerful
pollex can be set so far backwards that its long axis is practically
in the same line as that of the palm and of the fourth and
longest digit, thus giving the widest possible span to the extre-
mity, and ten the ‘pollex i is In this position the internal proximal
pad, constituting the proximal portion of the ‘“ ball” of the
thumb, lies nearly in a line behind the external proximal pad.
The pollex, moreover, is Incomparably more strongly opposable
than in any other genus of Lemuroid Primates, and surpasses
even the thumb of Man in that respect. When brought into
opposition, the composite ball of the thumb is pressed against the
second intermediate pad and the external proximal pad. Except
that the hallux is a little longer and stronger than the pollex,
the feet conform to the hands in type. In all other genera of
Primates the two extremities are dissimilar.

As is well known, the second digit of the hand in Vycticebus is
dwarfed and in Perodicticus is represented by an excrescence
upon the second intermediate pad. For the rest the hand of
Perodicticus is narrower with reference to its span, and the third
and fifth digits are tied basally by a shallow web to the fourth,
so that the three cannot be so w idely separated as in Nycticebus.
In the feet of both genera the heel is hairy and shorter than the
naked part of the sole, the second digit is dwarfed, afd the third
and fifth are basally webbed to the fourth in Perodicticus, free
in Vycticebus. The nails on both extremities are larger in the
former than in the latter genus (text-fig. 8, A, B, C, D).

The published descriptions of the extremities of Aretocebus
suggest that they differ from those of Perodicticus in having the
third, fourth, and fifth digits more completely webbed (see P. Z.S.
1864, pp. 316- 317, 319-: 320). Huxley, however, states that the
caleaneal tuberosity of the foot is naked and separated from the
padded portion by a narrow band of hair. But since the heel is
without exception covered with hair in all the remaining species
of arboreal * lemuroid Primates, I suspect that the nakedness
mentioned by Huxley was due to artificial rubbing in the specimen
he examined. It is not uncommon for the hair to be worn
off the heels in captive examples of common lemurs (L. albifrons,
etc.).

‘ the distinctuess and separation of the pads and their
encircling arrangement round a central palmar area, the hand of
Tarsius vecalls that of Hemigalago; but there ave certain differ-
ences. The hand and fingers of Varsius are relatively longer,
the pollex is smaller, closer to the second digit and not opposable,
the third digit is the longest of the series, slightly surpassing the
second and fourth, which are subequal. The first, or pollical,

* Lemur catta, which has the heels naked along the middle line, lives in
rocky hills.

CHARACTERS OF THE LEMURS AND TARSIUS. 35

intermediate pad is small and rounded, smaller than and in con-
tact with the internal proximal, which is elongated and surpasses
the external proximal in length. The second intermediate pad
is markedly larger than the others of that series. Finally, the
nails ave sharply pointed, compressed and convex from base to
tip, simulating claws, nearly the distal half being free from the
pad though normally resting upon it (text-fig. 9, C).

Text-figure 9,

A. Hand and B. Foot of Hemigalago demidoffi ; nat. size.
C. Hand and D. Foot of Tarsius; nat. size.

1-4, intermediate pads ; I-II, proximal pads.

In the feet the hairy area back to the heel is shorter than in
Hemigalago, and the naked padded area and digits longer. ‘The
plantar pads are peculiar. The hallucal, or first, intermediate is

large and prominent, and is fused to the internal proximal to
2%

36 MR. R. I. POCOCK ON THE EXTERNAL

form a single long pad which sometimes fuses proximally in the
middle line with the external proximal, which is itself united to
the external intermediate, forming a long pad extending along °
the outer edge of the sole of the foot. The second and third
intermediates are united to form a single large elongated pad,
broad distally, narrowed proximally, where it terminates in the
middle of the membranous area of the sole. Of digits two to five
the fourth is the longest as in Lemuroid Primates; but the third
has an erect claw like the second—a characteristic not found
in any Lemuroids, but probably primitive and forcibly recalling
the corresponding “ syndactyle” digits of many Marsupials (text-
fig..9, D).
The Sublingua.

There are one or two points to be recorded in connection with
the sublingua, an organ which is especially well developed in the
Lemuroid Primates. Typically it is a thin flat fibrous plate,
lyrate or leaf-like in form with free apex and free lateral margins.
It covers a considerable area of the lower surface of the tongue,
the frenum of which arises from an angular notch in the middle
of its posterior border. The pointed or truncated apex is ser-
rated or denticulated to a varying extent, and the underside is
strengthened by a fine median longitudinal ridge, with frequently
a smaller ridge close to it on each side, making three in all.

Beneath the sublingua the floor of the mouth is provided in
front with a pair of small soft flaps, the frenal lamelle, arising
at the bottom of the lingual frenum and continued backwards to
a varying extent as a free narrow edge towards the base of the
tongue. A similar and homologous structure is present in some
other Mammals—e. g., Canis, Pteropus, where, as in the Primates,
it overlies the orifices of the sublingual and submaxillary salivary
glands.

In the species usually referred to Lemur the sublingua ex-
hibits certain structual differences, which examination of a larger
number of individuals than I have seen may show to have
systematic value. For instance, in ZL. variegatus there are
three comparatively strong denticles at the tip and three corre-
spondingly strong keels below. In other species the tip is rather
serrulate and the keels weaker. On the other hand, I have noticed
differences in the shape of the sublingua within specific limits.
In a male of Z. coronatus, for example, it was lyrate with bulging
postero-lateral margins; in a female it was evenly attenuated,
the two margins gradually converging from near the base to the
tip. In the male again the frenal lamellae formed together a
broad semicircular flap; in the female they were narrower,
forming a nearly pavrallel-sided flap. In a male specimen of
L. albifrons albifrons and in a female of L. a. nigrifrons the
frenal lamellee were of the same shape as in the male J. coro-
»matus; but in examples of LZ. variegatus, macaco, catta, and mongoz
the lamellee were narrow and resembled more or less closely those

CHARACTERS OF THE LEMURS AND TARSIUS. oi

of the female Z. coronatus mentioned above (text-fig. 10, A-D, G).
In Chirogaleus major the narrowed sublingua is apically serrulate
and the frenal lamelle are narrow as in most Lemurs. The

sublingua of Hapalemur was briefly described by Beddard.

Text-figure 10.

. Lower side of tongue of Lemur coronatus.

. Sublingua of another specimen of the same species raised from the frenal
~ Jamellee.

. Lower side of distal end of tongue of Lemur variegatus with the tip of the

-sublingua showing beyond the frenal lamella.

. Lower side of tongue of the same without the frenal lamelle.

. Lower side of tongue of Galago crassicaudatus.

. The same of Nycticebus with supplementary frenal lamelle.

. Side view of tongue of Lemur coronatus.

. The same of Galago crassicaudatus.

e. b>

WMQdt

sl., sublingua; fJ., frenal lamellz.

38 MR. R. I. POCOCK ON THE EXTERNAL

The sublingua is also present, as Milne-Edwards and Grandidier
have shown, in the Indriside. It has a single median inferior
crest and three apical denticles. In Propithecus it hardly differs
in form from that of the Lemuride, being broad at the base
posteriorly, nearly parallel-sided, and attenuated at the apex.
In Jndris its lateral edges are rather strongly convex, the base
being narrowed like the apex. In Lichanotus it is very short,
its free lateral edge being apparently only about half as long
relatively as in Propithecus and Indris.

In Galago, Nycticebus, and Perodicticus the sublingua is broader
than in the Lemuride, especially at the apex, which is almost
truncated and armed with many denticles—i. e. 9 in a Galago
monterri—and its attachment to the tongue in the middle line is
set farther back so that a greater extent of its apex is free. I
also noticed 9 denticles at the tip in a specimen of Perodicticus
theanus. But in both Perodicticus and Galago: these denticles
form a continuous series, whereas in an example of Wycticebus,
with 7 denticles, the outer on each side was larger than the rest
and separated from them (text-fig. 10, E, F, H).

In Chiromys the sublingua is of a different type. It has a
free lateral margin, but towards its abruptly attenuated tip it is
closely adherent to the tongue. It is not denticulated, but ends
in front in a firm, deflected hook which is the anterior termi-
nation of a thick median keel or ridge, itself carrying two or
three teeth, and traversing the whole of the underside of the
sublingua back to the upper end of the lingual frenum. Just in
front of the point there is on each side of the sublingua a small,
probably glandular pouch, like a watch-pocket, with its orifice
looking forwards. The.frenal lamelle are present and narrow; but
I could not be sure of their exact form. In the example of this
genus I examined the sublingua is less cordate and the tongue
less rounded than depicted by Owen, who, moreover, did not
mention the small teeth on the carina or the pocket-like glands
near the base of the sublingua (text-fig. 11, A, B).

The sublingua of Z'arsius, described and figured by Burmeister,
is different from that of Lemurs. It is not so well developed, is
softer in consistency, and is defined from the tongue laterally by
a groove. In the middle it is provided with a rod-like thickening
which has a knob-like expansion at the apex. This rod, corre-
sponding to the median carina of the sublingua seen in Chiromys
and other Lemurs, was compared to the lytta of the tongue of
Canis by Burmeister. But the bifurcation of this rod and some -
other structures at the tip of the sublingua described by that
author I was unable to detect in the single example of the tongue
of Tarsius I examined. In this specimen the frenal lamelle
consist of a pair of longish slender processes, each tipped with a:
few small projections. At the proximal end of these processes
on the outer side the edge expands abruptly to form a flap with
a lobulate margin which extends far backwards along the sides
of the tongue. Burmeister described the frenal lamelle as the

CHARACTERS OF THE LEMURS AND TARSIUS. 39

“Unterzunge.” His figure represents this structure as an
undivided semielliptical lamina with evenly denticulated margin.
This does not agree with my observations (text-fig. 11, C—-E).

It has been suggested that the sublingua of the Lemurs acts as
a tooth-brush to clean the porrect, close-set, and comb-like series
of anterior mandibular teeth. Lemurs certainly use these teeth
to comb their own fur and that of their companions. I have
never seen them employ these teeth for any other purpose, and

Text-figure 11.

. Side view of the tongue of Chiromys.

Lower view of the same.

Lower view of the tongue of Tarsius raised from the floor of the mouth.

The same extracted with the frenal lamelle and the fringe attached.

. Side view of the tongue of Tarsius, showing the comparatively slght
differentiation of the sublingua and the well-developed frenal lamella with
backwardly extending lobulated fringe.

A
B.
C.
D,
E

since there is frequently a rapid movement of the tongue after
the combing action, I do not doubt that the suggestion as to
the function of the sublingua is correct; and the suggestion is
strengthened by the structure of the sublingua in Chiromys,
where the median keel ends in a firm hook well adapted to all
appearance for removing particles lodged between the two rodent
incisor teeth of the lower jaw.

40 MR. R. I. POCOCK ON THE EXTERNAL

In connection with the fur-combing use of the pectiniform
mandibular teeth in the true Lemurs, it must be remembered
that the spatulate finger-tips and short nails of these animals
deprive the digits of the scratching power they possess in ordinary
mammals with narrow finger- ns and comparatively long claws
or nails. Only one digit in the Lemurs is functional as a
scratcher, namely, the second of the hind-foot, which is short
with small terminal pad and long semi-erect claw. Thus is
established a most interesting correlation of characters in the
Lemurs :—namely, the uselessness of the fingers for scratching
the fur, the modification of the anterior mandibular teeth to sub-
serve that end, and the differentiation of the sublingua to keep
these teeth clean from scurf and hair. Now in Varsius the lower
mandibular teeth are not modified to form a comb. Possibly
they are employed for cleaning the coat, but their structure
makes them less lable to be clogged, and at the same time less
efficient instruments for the purpose than the corresponding teeth
of the typical Lemurs. Possibly, perhaps probably, for these
reasons, the sublingua is less differentiated and two of the digits
of the hind-foot, the second and third, are set apart as scratchers
and are capable of acting in unison, almost like the corresponding
united digits of the syndactylous Marsupials.

The sublingua of Chiromys probably serves mainly the purpose
of keeping the gnawing-teeth free from woody fibre; and, for
anything I know to the contrary, it may also cleanse them of
hair and scurf. But I am not aware whether these teeth are
used as a comb or not. At all events, in the case of this genus it
is interesting to note that the absence of the typical lemurine
dental comb is accompanied by well-developed claws and by fur
of a very different texture from that of ordinary Lemurs, since
it consists of a hght underwool covered by long coarse hairs.
Possibly the usefulness of the claws in combing this fur has been
one of the principal guiding factors in their evolution from nails.

The Anus and its Glands.

In all the Lemuroids examined, with one exception, the anus
is situated in the normal position below the joint of the tail so
that the base of that organ, when depressed, closes over it. But
in Chirogaleus major it is placed below, or on the distal side of
the joint of the tail, so that when that organ is raised the anus
is carried up on its base (text-fig. 16, A).

Normally in this group, that is to say in Vycticebus, Perodic-
ticus, Galago, Chiromys, the Indriside apparently, and in Lemur
catta, variegatus, and coronatus, the perineal and circumanal areas
are covered with hair; but in L. macaco, L. albifrons, and the
various forms such as nigrifrons, rufus, fulvus, cinereiceps, asso-
ciated with albifrons, these regions are covered in both sexes
with nearly naked, folded, wrinkled, and glandular skin, and
similar naked skin extends for an inch or so along the root of

s

CHARACTERS OF THE LEMURS AND TARSIUS. 4]

the tail (text-figs. 12, A; 15, E). In other species referred to
this genus, i.e. Z. mongoz and L. rubriventer, the glandular area
is much less differentiated, the skin being more closely covered
with fine, short hair and less distinctly folded and wrinkled.

Text-figure 12.

eS
BA Uy \
e YY M!
D “Wigs
Zn ip,
j f Vy
Hpsif;' 4 Wy
LaR i, Y Hf
vy fa’ y iy
Wy es Vibe
Ey Fy, \ “7 Lf
HH it ne af!
aH on A
Mh i)

I! We f ip
AS
Ns

\
i)

SS = RE Yy

ye

BS ‘
wv

NY YY re " : \\
\\ YY, i iN \
AY NY i fA IN

LAS
- ay iH Cc

A. Anal and genital area of male Lemur albifrons, showing the nearly naked
glandular skin stretching from the root of the tail to the scrotum.

B. The same of Lemur mongoz, showing the hairy but partially wrinkled skin
round the anus.

C. The same of female Lemur mongoz, showing the absence of wrinkled skin
round the anus, the long grooved clitoris (c/.), and the orifice of the urethra
(u.) below that of the vagina (v.).

D. The same of male Lemur catta, showing the hairy unwrinkled skin round the
anus and the naked scrotum.

42, MR. R. I. POCOCK ON THE EXTERNAL

The only example of Z. rubriventer examined, namely a female,
had the skin of the circumanal area distinctly but not strongly
wrinkled. On the other hand, some females of Z. mongoz show
no sign of wrinkling, others show traces of it, and in some
males the wrinkling is rather strongly pronounced. <A complete
gradation therefore in the development of the glandular area can
be traeed from ZL. coronatus through LZ. mongoz to L. macaco and
L. albifrons (text-fig. 12, A—C).

The paired anal glands, so well known in some orders of
Mammals, e. y. the Redentia and Carnivora, appear to be absent
in almost all Lemurs. The only genus in which I have found a
trace of them is Chiromys, where they are represented by a pair
of small, shallow invaginations, one on _ each side of the anal
orifice.

The Haternal Genitalia of the Male.

According to Milne-Edwards and Grandidier each of the three
genera of Indriside may be characterised by the structure of the .
penis and baculum. The penis appears to be short, subcylin-
drical, and apically truncated, and the baculum is distally
biramous in all cases.

In Lichanotus the epithelium is striated, and there are on each
side of the organ near the middle of its length two strong
recurved spines set one above the other. The baculum is greatly
expanded at its proximal end and gradually narrows from the
thickening to the middle of its length. From that point the two
sides diverge gradually to the apices of the two branches, which
themselves diverge evenly at about an angle of sixty degrees,
each branch being rather less than one-third of the length of the
whole bone.

In Jndris the epithelium of the penis is irr egularly reticulated,
and on each side of the organ there is a patch of rather small
spines set in three irregularly vertical rows which extend also to
the underside. The baculum differs from that of Lichanotus in
being thicker in the middle of its length, without any proximal
(or basal) expansion, and in having the two branches much less
divergent and a little longer, each slightly exceeding one- third
the length of the entire bone (text-fig. 14, C).

In Propithecus the epithelium of the penis is grooved and beset
with minute spicules. The baculum differs from that of Jndris
in having the branches very long, each being more than half the
length of the entire bone, longer, that is to say, than the stalk
instead of shorter as in /ndris and Lichanotus.

The penis of Chirogaleus major (text-fig. 13, I, K, L) is rather
short and broad, nearly parallel-sided, with an ovate extremity
giving it a somewhat linguiform appearance from the lower or
upper view. It is longitudinally grooved and closely punctured,
the punctures possibly marking the position of minute spicules

CHARACTERS OF THE LEMURS AND TARSIUS. 43

rubbed off. The orifice of the urethra is situated above the pad
covering the tip of the baculum, a character in which the penis

Axe mQe so

wOoWAReH

Text-figure 13.

. Penis of Galago crassicaudatus trom below with the frill folded over the tip

of the glans.

. Tip of the glans of the same with the frill spread.
. End of the penis of the same from the side with the tip protruding from the

frill.

. Penis of Galago senegalensis from below.

. Tip of the glans of the same showing the absence of the frill.
. Penis of Nycticebus from the side.

. Tip of the glans of the same to compare with B and E.

. The samme of Perodicticus.

Penis of Chirogaleus major from below.
Tip of penis of the same showing the orifice of the urethra above the tip of the
baculum.

. Penis of the same from the side with a probe passed down the urethra.

End of penis of Lemur macaco from below.

. The same from above.
. Penis of Lemur albifrons from the side.
. Tip of penis of the same,

44 MR. R. I, POCOCK ON THE EXTERNAL

differs from that of all the Lemuroid Primates examined by me.
Above the orifice there is a transverse glandular depression
overlapped by a thick flap of epithelium. A somewhat similar
crescentic flap half encircles the bacular pad laterally and below.
The baculum (text-fig. 14, A, B) itself is a little longer than that
of an adult example of Lemur albifrons more than twice the size
of Chirogaleus major. Seen from the side the baculum shows a
slight sinuous curvature, the distal half being depressed. Seen
from above or below, the main part of the shaft is straight and
subcylindrical, but its apical fourth is divided into two curved
branches, a right and left, which diverge at first, then converge
so as almost to meet apically in the middle line, circumscribing
an ovate space. The urethra, which runs along the underside of
the main portion of the baculum, passes through this space at
the point of bifurcation of the baculum. Hence it comes about
that the orifice of the urethra lies above the pad which covers the
juxtaposed tips of the two branches.

In being distally biramous the baculum of Chirogaleus resembles
that of the Indrisine Lemurs, a point of great interest. It is
most like that of Lichanotus, but has the base less expanded and
the branches curved and convergent apically instead of widely
divaricated. According to Beddard Hapalemur also has a
bifid baculum. I do not know the course of the urethra with
regard to the baculum in the Indriside, nor is Beddard quite
clear on this point in his description of the penis of Hapalemur,
but since he states that the urethra opens at the posterior end
of a groove marking on the glans penis the forking of the
baculum, it may be inferred that the orifice of the urethra is
beneath the tip of the baculum as in typical Lemurs and not
above it as in Chirogaleus.

On one of the plates of his unfinished work Milne-Edwards
illustrates the male genitalia of a Lemur which, although un-
named in the legend, is clearly shown to be L. catta by the naked
scrotum. Four bacula are figured on the same plate. They
may have been taken from specimens representing four distinct
species or from four examples of Z. catta. If the latter, the
figures attest a certain amount of individual variation in this
bone, particularly in width at the base and expansion and
curvature at the tip. There the matter must rest.

In the species I have examined, namely L. catia, macaco,
albifrons, and coronata, there is very little variation in the penis
and baculum. The penis (text-fig. 13, M-P) is subcylindrical
and armed in the middle of its length with many reversed spines
which are mostly of smal] size; two or more pairs, however, on
each side are much larger than the others, but these large spikes
are not always either symmetrically placed or numerically iden-
tical on the two sides. The orifice of the urethra is terminal
and opens just beneath the tip of the baculum. The baculum is
a comparatively short and slender rod with a larger proximal and

CHARACTERS OF THE LEMURS AND TARSIUS. 45

a smaller, sometimes incipiently bilobed distal expansion. As
compared with the size of the animals, the baculum in the
species referred to Lemur is smaller than in any other genus
(text-fig. 14, H).

In Galago senegalensis the penis is long (text-fig. 13, D, E).
Its narrow and cylindrical, proximal portion gradually expands
distally to a considerable extent, and its terminal portion is
attenuated. The orifice of the urethra, opening beneath and
behind the hardened rounded pad covering the tip of the baculum,
is provided with a small inferior and two small lateral labia.
Except at the distal and proximal ends the epithelium of the
penis is thickly beset with comparatively coarse reversed spines

Text-figure 14.

oe
Bias eS SSS

i a a
Cie
: ame
J H

A. Baculum of Chirogaleus major from the side, the line indicating the course of
the urethra.

B. The same from above showing the apical bifurcation.

C. Baculum of Indvis trom above (copied from Milne-Hdwards).
D. The same of Galago senegalensis from the side.

{. The same of Galago crassicaudatus.

F. The same of Nycticebus.

G. The same of Perodicticus.

H. The same of Lemur albifrons.

I. The same of Chiromys.

(All the figures twice nat. size.)

approximately equal in size. The baculum (text-fig. 14, D) is
long and slender, being actually nearly twice as long as that of
an adult male Lemur altifrons. It is practically straight and
gradually attenuated from its broad base, but there is a slight
sinuosity in its distal third and a slight ad gradual thickening
at the apex which is blunt.

46 MR. R, I. POCOCK ON THE EXTERNAL

In Galago crassicaudatus and monteiri (text-fig. 13, A-C) the
penis differs considerably from that of G. senegalensis. It is
clavate in form, being gradually incrassate from the base to the:
blunted tip. The orifice is just below the apex of the baculum,
and the two are encircled by a frill of grooved, wrinkled epithe-
lium, forming a sort of secondary prepuce, which is attached by a
frenuin to the lower lip of the orifice and encloses a glandular
space. ‘The spines covering the penis are bidentate or tridentate,
and much smaller than in G. senegalensis. The baculum (text-
fig. 14, KE), although actually longer than in G. senegalensis, is
approximately the same shape and relatively about the same size,

In Vycticebus the penis (text-fig. 13, F, G) is much shorter than
in Galago and smooth and of tolerably even thickness through-
out. The callous pad at the tip of the baculum is large, rounded
and prominent, and overhangs the orifice of the urethra, which
lies in a glandular space bounded laterally and below by a pair
of labia practically as in G. senegalensis. The baculum (text-
fig. 14, F) is a nearly straight rod, only about two-thirds the
length of that of G. senegalensis, although the animal itself. is
much bigger.

In Perodicticus the penis (text-fig. 13, H) is short and smooth
and like that of NVycticebus except that the tip of the baculum
and the orifice of the urethra are surrounded by a complete hood
or frill of wrinkled epithelium enclosing a circular glandular
space. The baculum (text-fig. 14, G) in the specimen examined
is shorter than that of the example of Vycticebus, but decidedly
thicker at the base inferiorly and with its upper edge a little
more sinuous.

According to Huxley's description (P. Z.S. 1864, p. 334) the
extremity of the glans penis in Arctocebus closely resembles that
of Perodicticus except that the encircling hood is bifid in the
middle line below. He states, however, that the baculum is °75
of an inch in length, about twice as long, that is to say, as in my
example of Perodicticus; although, judging from the dimensions
of the limbs, his animal was considerably the smaller of the two.

In Chiromys the penis is rugulose, wrinkled, and very slightly
narrowed distally for three-fourths of its length, then somewhat
abruptly attenuated to the apex, where the upturned tip of the
baculum terminates; but just below this point there is a little
soft, curved, subcylindrical process upon which it seems probable
the orifice of the urethra opens. When the prepuce is reflected
to its fullest extent, the basal fourth of the exposed portion of the
penis is seen to be provided with five probably glandular, longi-
tudinal grooves, one in the middle line above and two on each
side, one above the other, the ventral middle line being occupied
by the frenum. The baculum (text-fig. 14, 1) is tolerably stout and
longer than in any genus of Lemuroid known to me. Allowing for
its curvature, it is as long relatively as in G'alago crassicaudatus.
When seen from the side it is markedly sinuous, its upper edge

CHARACTERS OF THE LEMURS AND TARSIUS. ‘A7

being concave close to the base, then convex, then strongly con-
cave owing to the upcurling of the distal fourth of its length.
The ventral side is curved in correspondence. No other
Lemuroid known to me has a strongly upcurled tip to this bone.

In connection with the scrotum there is one fact to be re-
corded. In Lemur catta this sac is always naked (text-fig. 12, D).
In all other species of Lemuroid Primates it is clothed with hair
normally (text-figs. 12, A, B; 16, A). Occasionally the postero-
inferior portion is naked, as I have noticed in one or two specimens
of Galago; but I have no doubt that in these cases the absence
of the hair was due to rubbing.

The External Genitalia of the Female.

In the standard text-books of Mammals published even as
recently as Max Webev’s in 1904, it is stated that the urethra
traverses the clitoris in the Lemuroidea, This is not true of any
Madagascar Lemur [ have examined, and applies only to the
Asiatic and African forms. In the Madagascar species the
urinary orifice opens at a varying distance between the vaginal
aperture and the apex of the clitoris, generally much closer to
that orifice and only in one case, Lemur catta, a little nearer the
tip of the clitoris.

Milne-Edwards and Grandidier have shown that the clitoris
varies considerably in form in the three genera of Indriside.
In Lichanotus (Avahis) it is long, pendulous, and narrow, and
parallel-sided in its distal two-thirds, but expanding proximally
towards the orifice of the vulva. From the orifice of the urethra,
situated a little below the vulva, a groove extends towards the
tip of the clitoris, and at the extreme tip of the latter is placed
the aperture of a glandular depression.

In Propithecus the clitoris is much shorter and thicker, with
a broadly rounded distal end carrying a glandular orifice. The
aperture of the urethra lies approximately midway between the
orifice of this gland below and of the vagina above. In /ndyis
the distal end “of the clitoris is gradually and widely expanded
laterally and extended considerably beyond the orifice of the
gland, but the orifice of the urethra is situated even nearer to
the vulva than in Lichanotus. '

In the species usually referred to the genus Lemur there is
considerable variation in the structure of the genital area of the
female.

In Lemur varius the clitoris is a short, thick, fleshy excrescence
with a blunt apex and somewhat cordate in shape, rising from
the centre of a tolerably large area of naked skin. Its free
posterior surface is marked by a median groove or rima defined
laterally by a pair of thick labia. When the latter are separated
the urinary channel, a gutter with thin elevated margins, is
displayed. In a mated female this gutter is seen to lead from

48

A

b.

C.

MR. R. I. POCOCK ON THE EXTERNAL

Text-figure 15.

=
‘

‘
y i
é
j

= VE
S Ce

“SN

. Anal and genital area of female Lemuw catta, showing the thick clitoris
protruding between the labia of the vuiva.

The same of female Lemur variegatus, showing the short cordate vulva arising
from a naked area of skin.

Vulva of the same enlarged, showing the lateral grooves and the orifices of the
urethra and vagina covered by membrane (hymen) in the virgin female.

ID. Side view of the same vulva.
E. Anal and genital area of female Lema albifrons, showing the circumanal

oa
ed ad

glands and the long attenuated clitoris.
The svme of Lemur coronatus, showing the long clitoris and the absence of
circumanal glands.

x. Vulva of the same on larger scale, showing the supplementary labia beneath

the orifice of the vagina.

cl., chtoris ; w., orifice of urethra; v., orifice of vagina.

CHARACTERS OF THE LEMURS AND TARSIUS. 49

the orifice of the urethra, which is just below that of the vulva ;
but in a virgin female the two orifices are eoncealed by a flap of
membrane (hymen), below the edge of which the open channel
extends, narrowing in its course to the apex of the clitoris. The
sides of the clitoris are marked by a couple of deep, probably
glandular, grooves (text-fig. 15, B, C, D).

In Lemur catta the clitoris is a thick elongated excrescence

Text-figure 16.

¥,

2S a

A. Anal and genital area of male Chirogaleus major, showing the position of the
anus on the root of the tail.

B. The same of female Hemigalago demidoffi, showing the long pendulous clitoris
below the vulva.

C. The same of female Galago crassicaudatus with the labia of the vulva spread
open.

D. Vulva of the same with the labia closed.
E. Vulva of the same from the side.
F, Anal and genital area of female Tarsius, showing the labia of the vulva
partially spread open.
G. The same from the side with the labia closed.
el., clitoris; U., labium of vulva; w., orifice of urethra; v., orifice of vagina.

Proc. Zoot. Soc.—1918, No. IV. 4

50 MR. R. I. POCOCK ON THE EXTERNAL

with the orifice of the vulva at its base and that of the urethra
about one-third of the distance from the tip—much lower, that
is to say, than in Z. variws. The urinary channel runs from the
urethral orifice to the apex of the clitoris. The orifice of the
vulva is bounded on each side by a pair of thick labia, which,
when separated, diverge from above that orifice and inferiorly
disappear in the area of naked integument from which the
clitoris rises (text-fig. 15, A):

In L. macaco, albifrons, fulvus, mongoz, rufiventer, and coro-
natus, the clitoris is narrow, elongated, attenuated towards the
apex, and arises from an area of naked or nearly naked skin. It
is channelled to the apex from the orifice of the urethra, which
opens a short distance below that of the vagina. It varies toa
certain extent in length within specific limits, e. g. L. albifrons,
and is rather exceptionally long, less attenuated and narrower
at the base in Z. coronatus than in the others. Moreover, in the
single example of this species examined, two pairs of elongated
lamine, constituting labia, diverged obliquely just below the
orifice of the vagina (text-figs. 12, C; 15, E, F, G).

In the Asiatic Lemuroids the clitoris differs from that of the
Madagascar forms in:being traversed by the urethra which opens
at its tip.

In Galago crassicaudatus the clitoris is elongated, thick from
before backwards in profile view, and parallel-sided and com-
pressed from the posterior aspect : at its tip there is a depression,
probably glandular, almost encircling the urethral orifice, and
comparable to the similarly situated gland in the Indriside.
The vulva consists of two turgid labia, of which the opposed
surfaces are provided with several small fine lamine, running
from the elges of the labia inwards towards the orifice of the
vagina (text-fig. 16, C, D, E).

In Hemigalago the clitoris is relatively much longer and
thinner than in G. crassicaudata. The distal portion appears to
be strengthened by a baculum and the tip is provided with a
glandular depression encircling the end of the bone. The two
labia at the sides of the orifice of the vagina are relatively
smaller than in G. crassicaudatus (text-fig. 16, B).

In NVyeticebus the clitoris is short and thick, resembling the
prepuce of a penis. There is a glandular depression at the tip.
The orifice of the vagina is a wide transverse rima just at the
base of the clitoris. The general appearance of the external
genitalia is very different from that of Galago and Henugalago.
Possibly the specimen examined was a virgin female.

The female external genitalia of Zarsius are unlike those of
the Asiatic and Mascarene Lemurs, but in the single example
examined recall rather those of the Old World pithecoid Primates.
‘he vulva is a laterally compressed, elongated excrescence, with-
out pendulous apex. The rima is short and bounded by a pair
of labia concealing the small clitoris and the orifices of the

CHARACTERS OF THE LEMURS AND TARSIUS, 5]

arethra and vagina, which are comparatively close together. It
is significant that the vulva of this genus as a whole is less like
the vulva of the Galagos than that of the true Lemurs. It
approaches most nearly. the vulva of Z. varius (text-fig. 16, F, G).

General Conclusions.

The conclusions of general interest which have suggested them-
selves in the course of the investigations above detailed are
Subjoined in the order of their importance :—

1. In the case of Tarsius the structure of the upper lip and of
the nose severs the genus completely from the Lemurs and brings
it into line with the Pithecoid Primates. Taking this character
in conjunction with the nature of the placenta, the presence of
the postorbital partition, and other well-known features, 1t seems
that Hubrecht was quite right in removing Yursius from the
Lemurs and placing it in the higher grade of Primates. I pro-
pose to give practical expression to this view by dividing the
Primates into two great series. For the first, comprising the
Lemurs, the old name STREPSIRHINI is available. For the second,
comprising Zarsius and the Pithecoidea, I suggest the title
Harnornini. The Haplorhini will contain two divisions, the
‘Tarsioidea and the Pithecoidea *. |

2. With Zarsius eliminated from the Lemurs, the Strepsirhini

may be divided into the Chiromyoidea and the Lemuroidea. It
has been the fashion of late years to depreciate the characters of
Chiromys. Dr. Standing indeed gave the genus merely subfamily
rank under the Indriside, the latter being equivalent to the
Lemuride. But that classification sacrificed the characters of
Chiromys to the hypothesis that the genus is a specialised off-
shoot of the Indrisoid stock. That may be true. Nevertheless,
the specialisation has proceeded so far, and in so many directions,
that it appears to me impossible to dispute the claim that the
Lemuride and the Indriside are much more nearly akin to each
other than either is to the Chiromyide. In the description of
Chiromys the peculiarities of the teeth generally distract attention
from the curious cranial modifications correlated with the rodent
dentition, such as the immense size of the premaxille, which
reach the lachrymals and exclude the reduced maxilla from con-
tact with the nasals; also the absence of the bony ridge closing
the glenoid behind and the longitudinal extension of the man-
dibular condyle, two correlated characters subservient to the
back and forth movement of the mandible well known in the

* I prefer this name to Anthropoidea because in ordinary terminology the title
“anthropoid,” reasonably according to its meaning, has hecome restricted to the
man-like Apes. A marmozet can hardly be called “ anthropoid,” with any appr oach
to the real meaning of the word. But amarmozet and a man are alike “ pithecoid.”’

4*

5? MR. R. I. POCOCK ON THE EXTERNAL

Rodentia. These cranial features distinguish Chiromys from all
Lemurs ; and when taken in conjunction with the teeth, with the
peculiarities of the hands and feet, and of the sublingua, they out-
weigh, in my opinion, the known differences between the true
Mascarene Lemurs (Lemuride and Indriside) and the Asiatic
and African Galagos, Pottos, and Lorises.

3. With regard to the Galagos, Pottos, and Lorises, I am only
acquainted with one invariable cranial character distinguishing
them from the Lemuride and Indriside. This was pointed out
by Forsyth Major and has been briefly expressed by Mr. Gregory *
as follows :—In the Asiatic forms the ectotympanic is enlarged
and external to the bulla of which it forms the outer wall. In
the Mascarene forms the ectotympanic is inclosed within the
bulla, forming a ring or horseshoe. To this difference may be
added the one pointed out above in connection with the clitoris,
which in the Asiatic genera is traversed by the urethra, whereas
in the Mascarene forms the urethra opens above the tip of the:
clitoris.

In view of these facts, I should divide the Lemuroidea into
two series, for which Mr. Gregory’s names Lemuriformes for the
Lemuride and Indviside, and Lorisiformes for the Loriside (or
Nycticebide) and the Galagide may be adopted. Similarly
for the subdivision of the Lemuride I follow Mr. Gregory in
relegating the genera to two subfamilies, the Lemurine and
Chirogaleine ; but I cannot agree with him that LHapalemur
belongs to the Chirogaleine. That genus appears to me to be
essentially a Lemurine, its inclusion in the Chirogaleinz spoiling
the definition of the subfamily.

4. As a matter of minor interest it is quite clear that the
genus Lemur as generally admitted and as recognised in this
paper is susceptible of division into two or three genera. L. catta,
for instance, differs from the other species in having the glands
on the fore-limb and the naked heel and scrotum, and also
in the structure of the vulva. JZ. variegatus 1s also peculiar in
the structure of the vulva. Furthermore, the Galagos of the
G. senegalensis-type may be distinguished by the structure of the
penis from that of the crassicaudatus-type. Generic names
appear to be available for these subdivisions of Lemur and
Galago; but I do not propose to enter into that question now.

My views above set forth differ in so many particulars from
those of Mr. Gregory that it may be interesting to tabulate our
classifications side by side, omitting those he adopts based upon
extinct genera, which, so ‘lems as I am aware, do not materially
affect the arrangement of recent forms.

* Bull. Geol. Soc. Amer. 26, pp. 432-436, 1915. See also Bull. Amer. Mus. Nat.
Hist. 35, pp. 266-267, 1916.

CHARACTERS OF THE LEMURS AND TARSIUS. D3

Gregory, 1915-1916. | Pocock, 1918.

‘Order PRIMATES. | Order PRIMATES.

Subord. LEMUROIDEA. Grade STREPSIRHINI.

Series LEMURIFORMES. Subord. LEMUROIDEA.

Fam. LEMURID. Series LEMURIFORMES.

Subfam. Lemurine. | Fam. LEMURID#.
” Chirogaleine. Subfam. Lemurine.
Fam. INDRISID2. | 4 Chirogaleine.
», CHIROMYIDS. | Fam. InDRISID=.
Series LoRIsIFORMES. Series LoRISIFORMEs.

Fam. Loristp%. Fam. Lorisipm.

Subfam. Lorisine. ‘
» GALAGID2.

Galax ine.
as J Subord. CHTROMYOIDEA.

= ae 4 | es
Series TARSIIFORMES. | Fam. CHIROMYID®.

Fam. TARSIIDA. |
Grade HAPLORHINI.

‘Subord. ANTHROPOIDEA. Subord. TARSIOIDEFA.
|

» PITHECOIDEA.

Apart from the removal of Zarsius from the Lemuroidea and
the union of the Loriside and Galagide in a group equivalent to
the Lemuride and Indriside combined, my classification agrees
in the main with the classification published in Flower and
Lydekker’s ‘Mammals,’ and in other works of about that period.
‘Of more recent classifications it appears to me that Gadow’s
(Class. of the Vertebrata, 1898, pp. 52-53), although brief, ex-
presses the facts with the nearest approach to the truth, especially
in the primary division of the Primates into the three suborders
Lemures, T'arsii, and Simie.

S}

-

\

wr
oy |

ON THE PYRALIDH, SUBFAMILY HYPSOTROPINA.

5). A Classification of the Pyrauip®, subfamily Hypso-
TROPINA. By Sir Grorce Hampson, Bart., F.Z.S.

[Received February 19, 1918: Read March 5, 1918. ]

Proboscis aborted or absent ; palpi upturned, oblique or down-
curved, the males of the species with the maxillary palpi brush-
like and contained in a hollow of the labial palpi usually having
the palpi oblique in the male and downcurved in the female;
maxillary palpi small and filiform, well developed and more or
less dilated with scales at extremity, or brush-like and contained
in a hollow of the labial palpi; frons smooth, with tuft of hair,
or prominences of various forms; antenne of male ciliated,
laminate, serrate or pectinate with uniseriate branches, the
basal joint often dilated and the shaft often downcurved at base
with a ridge of scales in its sinus; the build slender; tibize with
all the spurs present; abdomen smoothly scaled. Fore wing
narrow; vein lq@ separate from 16; le absent; veins 2, 3
rarely stalked; vein 4 often absent or stalked with 5, 3 and 5
stalked or from the cell; 6 from below upper angle; 7 absent ;
8,9 and often 10 stalked or 9 absent, rarely 10 also absent,
8 given off before or after 10; 11 from cell. Hind wing with
the median nervure pectinated on upper side; veins la, b,c
present; 2 from near or well before angle of cell; veins 3 and 4
sometimes gent or 3 and 5 stalked or from the cell, 4 often
absent, or 3, 4, 5 all present, 4 stalked with 5 or from ceil oper
stalked or from upper angle of cell; 8 anastomosing with 7 or
approximated to it but free.

The larvee of very few species are known; these live on
Graminacee and pupate in the ground in a chamber formed
by agglutinated particles of the. soil; Hmmalocera depressella,
however, is injurious to sugar-cane, the larva feeding in its.
roots.

The subfamily is a development from the Anerastiane—Ane-
rastia Hiibn. Verz. p. 367 (1827), type dignella Hubn.—from
which it differs only in the proboscis being absent or aborted
and non-functional.

The types of the new species are in the British Museum, the
species of which the types are in the collection are marked with
a ¥, and those not in the collection with a *.

No quotations from German authors published since Aug. Ist,
1914 are inserted. ‘ Hostes humani generis.”

SIR GEORGE HAMPSON ON THE

“p.lao0] DULuT

“‘pysohjog ‘wyouobyy “piwvs—Q “vuLogvg “DMOZ
gama
“prqUodfoosuy “DUD T aa — ‘prspgydorg
“smd oeesyc ema
‘pdoujzojdvg ~pdougOvy ya ‘DAforg “sapnayzougy ambpidy
‘phoquy “india
“DIQWOLfLSSOT aang ‘pdouqouvpny — “DLpWaLnDT
‘paaspung = “wshayoopoyyy aan
sdngenw, ania onde “PIDU0zLOyD | “pdo.wgosdh Fy
“ajjanaouamg
ee
wndlopoaayTnr
“PJOSDADNT — “pau, "p0qa) ca, Lposa yr
oma ‘pdo.uqoa'T ‘DUSDABULT, , “DULDIS
‘paquag” sapieaa “DADaD mais “DL2)0) aaiinitng

‘WNICGOULOSAA HHL FO ANYDVOTAHG

PYRALIDA, SUBFAMILY HYPSOTROPIN®E.

Key to the Genera.

A. Hind wing with veins 3 and 4 absent.
a. Fore wing with vein 8 from 9 before 10, 3 and 5 stalked ......
b. hae wing with vein 8 from 9 after 10.

Fore wing with veins 3 and 6 stalked.

b},

a Palpi uptur ned

b2, Palpi oblique, straight |

c?, Palpi downcurved

Fore wing with veins 3 and 5 ‘fro om 1 the Celle.

a’, Palpi upturned
b?. Palpi porrect.

a’, Fore wing with vein 9 absent, 11 stalked with 8

and 10

63, Fore wing with vein 9 pr -esent, ‘11 from cell .

c. Fore wing with vein 10 from the cell .

B. Hind wing with vein 3 present, 4 ern
a. Fore wing with vein 4 absent.

a, Fore wing with veins 9 and 10 absent, 8 from cell, 3 and

b1. Fore wing with vein 9 absent, ‘10 from ‘cell, 3 and 5

a

5 from cell

fom cell e600 s0c0s0s

Fore wing with vein 8 from 9 before 10, 3 and 5 stalked...
Fore wing with vein 8 from 9 after 10.

a*, Fore wing with veins 3 and 5 stalked.

a®, Palpi upturned.

a‘, Frons with rounded prominence ......

b+. Frons without prominence.
a, Palpi reaching to above vertex of head, the 2nd
joint slenderly scaled .

6°, Palpi not reaching to above vertex of. head, ‘the

2nd joint broadly scaled ..

53, Palpi oblique or downcurved.
a*, Frons with corneous ome proses to two slight
points in front .

b4. Frons without corneous plate

b?. Fore wing with veins 3 and 5 from cell.

a’, Hind wing with veins 3
63, Hind wing with veins 3 and 5 from the cell.
a‘, Palpi upturned

and 5 stalked

b4. Palpi downcurved
ey ore wing with vein 10 from the cell, 8 and 9 stalked.
Hind wing with veins 3 and 6 stalked.
Pos Frons with flattened corneous plate, rather trifid in

front and produced to short lateral points, a cor-
...... Raphimetopus,

neous plate below the frons .........

63. Frons with pointed conical prominence ae

57

Ceenochroa,
[p. 58.

... Menuthia, p. 60.
.... Calera, p. 59.
. Statina, p. 59.

. Tinerastia, p. 61.

Coenotropa, p.89.

| Mesodiphlebia,

paGl.

. Calamotropa,

[p. 91.

. Tampa, p. 62.

. Bandera, p. 89.

. Hosidia, p. 63.

. Dembea, p. 64.

Ceara, p. 63.

. Leotropa, p. 64.

. Alamosa, p. 65.

Navasota, p. 68.

. Hypsotropa,

ped.

. Sudania, p. 88.

. Rhodochrysa,

c3, Frons with long truncate corneous es with raised
rim at extremity ..
d3, Fyons with rounded | prominence with raised rim at
EXE MOND Varin ates See ott See ce gate a cadet Cagis

e°. Frons without prominence

b*. Hind wing with veins 3 and 5 from the cell.
a3. Hind wing with vein 8 anastomosing with 7
63. Hind wing with vein 8 not anastomosing with 7

b. ee wing with vein 4 present.

a ore Wing with vein 10 stalked with 8.
Fore wing with veins 4, 6 stalked or from a point.

Sas Hind wing with vein 2 from or from close to angle

of cell.

a‘. Frons with conical prominence
b+. Frons without prominence.
a’. Frons with long pointed tuft of hair ...............
b>, Frons without tuft of hair

[p. 88.

r

(pets.

Chortoneca,
[p. 80.
Metacrateria,
[p. 79.
Prinanerastia,
[p. 80.

. Rhinaphe, p. 82.

Laurentia, p. 90.
Epidauria,p. 91.

. Fossifrontia,

Lp. 107.
Aurora, p. 106.

. Commotria,

[p. 107.

58 SIR GEORGE HAMPSON ON THE

63, Hind wing with vein 2 from well before angle of cell.
a+, Frons with rounded PYOMINENEE 20. .......00-0.00.-.-.. Stboga, pe 2,
64. Frons without prominence.
a. Fore wing with veins 4, 5, hind wing with veins

3 and 5 stalked ee ...... Ematheudes,
. Fore wing with veins 4, 5, hind 1 wing ‘with veins [p. 114.
3 and 5 from angle Shed. eee Baptotropa,

62. Fore wing with veins 4, separate, [p. 116
a’, Hind wing with vein 2 from near angle of cell......... Biafra, p. 114.
63, Hind wing with vein 2 from well before angle of cell. [p. 116..

a+. Hind wing with vein 8 anastomosing with 7 ...... Ethiotropa,

b+, Hind wing with vein 8 not anastomosing with 7 ... Patna, p.117.
b!, Fore wing with vein 10 from the cell.
a*. Fore wing with veins 4, 5 stalked.

a’, Hind wing with veins 3 and 5 stalked .................. Saluria, p. 93.
6%, Hind wing with veins 3 and 5 from the cell ............ Prophtasia,
62. Fore wing with veins 4, 5 from the cell. [p. 104..
a’, Hind wing with veins 3 and 5 stalked; frons with
truncate conical prominence sete ceccee ss Megalophota,
63, Hind wing with veins 3 and 5 from the cell. pte z,
at, Frons with prominence hollowed out in front ...... Martia, p. 118.
64, Frons with disk of scales. .......c0-...0. seehe-ec2e-00 ten COUSCOPPONDECS
[p. 118.
c*. Frons without disk of scales or prominence .......... Critonia, p. 119.
C. Hind wing with vein 4 present. :

a. Fore wing with vein 10 stalked with 8. [p. 121.
a, Hind wing with vein 8 anastomosing with 7) ............... Monoctenocera,
b!, Hind wing with vein 8 not anastomosing with 7............ Saborma, p. 122.

6. Fore wing with vein 10 from the cell.

a‘, Fore wing with veins 2, 3 stalked «...............-..c:-0c20505. OS@eia, py 123.

61, Fore wing with veins 2, 3 from the cell.
a*, Frons with small rounded prominence with a corneous

plate-helow it: 22.2001... Si ah ee A, os eon oyaae
63, ot without prominence. : fp, 123.
Fore wing with veins 4, 5 stalked ............ . Polyocha, p. 124.
a Fore wing with veins 4, © fromthe eelli.. tc... e Emmalocera,
p. 126.
”
Genus CaiNOCcHROA.
Type.
Cenochroa Rag. N. Am. Phye. p. 20 (1887) ...... californiella.
Petaluma Hulst, Trans. Am. Ent. Soc. xvu.
1, 215 (B80) ashen aan eee ilibella.

Proboscis absent; palpi downcurved, about three times length
of head and thickly scaled; maxillary palpi minute and filiform ;
frons with conical prominence, thickly scaled above; antenne
of male Jaminate and almost sunpls, Fore wing narrow and
elongate, the apex rounded; vein 2 from close to ‘angle of cell ;
3and 5 strongly stalked, 4 absent; 6 from well below upper
angle; 8, 9, 10 stalked, 8 from before 10; 11 from cell. Hind
wing with vein 2 from close to angle of cell; 3 and 4 absent,
5 sometimes not reaching the termen; 6, 7 from upper angle ; ,
8 stron ely anastomosing with 7.

(1) *CCENOCHROA ILLIBELLA.

Anerastia illibella Hulst, Ent. Am, il. p. 138 (Oct. 1887);
Rag. Rom. Mém. viii. p. 419, pl. 45. f. 9; Dyar, Cat. Lep. N..
‘Am. p. 440.

Cenochroa puricostella Rag. N. Am. Phyc. p. 20 (Dec. 1887).

U.S.A., Texas, Arizona.

PYRALIDH, SUBFAMILY HYPSOTROPIN &. 59:

(2) Ca:NoCHROA INSPERGELLA,

Cenochroa inspergella Rag. N. Am. Phye. p. 20 (1887); id.
Rom. Mém. viii. p. 419, pl. 45. f. 10; Dyar, Cat. Lep. N. Am.
p. 440.

U.S.A., Texas, Colorado, Arizona.

(3) *Ca:NOCHROA CALIFORNIELLA.

Cenochroa californiella Rag. N. Am. Phye. p. 20 (1887) ; id..
Rom. Mém. viii. p. 420, pl. 45. £.8; Dyar, Cat. Lep. N. Am.
p. 441.

U.S.A., Colorado, California, Arizona ; Mexico, Presidio.

Auctorum.
Cenochroa monomacula Dyar, Pr. U.S. Nat. Mus. xlvil. p. 348:
MEY oteaarerr srs pteraae onan seve ie 2s.) Seaemakiee er ee PANAMA.
Genus CALERA.
Type.
Calera Rag. Nouv. Gen. p. 50 (1888) ............ punctilimbella.

Proboscis aborted and minute; palpi downcurved, extending
about twice the length of head and smoothly scaled ; maxillary
palpi minute and filiform; frons smooth; antennze of male
somewhat laminate and ciliated, the basal joint somewhat en-
larged and flattened on outer side. Fore wing long and narrow,
the apex rounded; vein 2 from near angle of cell; 3 and 5
strongly stalked, 4 absent; 6 from below upper angle; 8, 9, 10
stalked, 8 from beyond 10; 11 from cell. Hind wing with
vein 2 from close to angle of cell; 3 and 4 absent; 6, 7 stalked ;
8 anastomosing strongly with 7.

CALERA PUNCTILIMBELLA.

Calera punctilimbella Rag. Nouv. Gen. p. 50 (1888); id. Rom.
Mém. viii. p. 417, pl. 40. f. 20; Dyar, Cat. Lep. N. Am. p. 441.

U.S.A., N. Carolina, Iowa, Texas.

Genus STaTINA.
Type.
praune Bar. N; Ami. Phye. p. 19 (188i)... 2... roseotinctella.

Proboscis aborted and minute; palpi downcurved, extending
about two and a half times length of head and rather roughly
sealed; maxillary palpi minute, filiform; frons with tuft of
scales ; antenne of male Jaminate and ciliated, the shaft thickened
with scales above at base, the basal joint long, dilated and
flattened. Fore wing long and narrow, the apex rounded;
vein 2 from near angle of cell; 3 and 5 strongly stalked,
4 absent; 6 from below upper angle; 8, 9, 10 stalked, 8 from
beyond 10, 11 from cell. Hind wing with vein 2 from close to

60 SIR GEORGE HAMPSON ON THE

angle of cell; 3 and 4 absent; 6, 7 from upper angle; 8 anasto-
mosing strongly with 7.

(1) *SraTina GAUDIELLA.

Statina gaudiella Hulst, Trans. Am. Ent, Soc. xvii. p. 216
(1890); Rag. Rom. Mém. viii. p. 415, pl. 51. f.5; Dyar, Cat.
Lep. N. Am. p. 440.

U.S.A., Texas.

(2) STATINA ROSEOTINCTELLA.

Statina roseotinctelia Rag. N. Am. Phye. p. 19 (1887); id.
Rom. Mém. viii. p. 416, pl. 47. f. 7; Dyar, Cat. Lep. N. Am.
p. 440.

U.S.A., Florida, Texas.

(3) TSTATINA PUNCTILINEELLA.

Statina punctilineella Hmpsn. Rom. Mém. viii. p. 416, pl. 52.
tls 901).

S. Brazit.

(4) TSTATINA BIFASCIELLA,

Statina bifasciella Hmpsn. Rom. Mém. viii. p. 416, pl. 52.
f. 8 (1901).

U.S.A., Texas.

(5) STATINA RHODOBAPHELLA.

Statina rhodobaphella Rag. Nouv. Gen. p. 50 (1888); id. Rom.
Mém. viii. p. 417, pl. 45. f. 6.

CELEBES, Sangir I.; N. GuINEA ; QUEENSLAND.

(6) TSTATINA ROSINELLA, sp. Nn.

Q. Head and thorax purplish pink; abdomen whitish suf-
fused with red-brown, dorsally fulvous towards base; antenne,
pectus, and legs whitish tinged with red-brown. Fore wing
purplish pink, the veins streaked with white; a brown shade
below basal half of cell. Hing wing ochreous white.

N. Nigeria, Minna (Macfie), 1 2 type. Hap. 16 mm.

(7) TSTATINA CASHMIRALIS.

Statina cashmiralis Hmpsn. J. Bomb. Nat. Hist. Soc. xv.
p. 20 (1901).

KASHMIR.

Genus MENUTHIA.
Type.

Menuthia Rag. Nouv. Gen. p. 50 (1888) ...... nee: nanella.

Proboscis absent ; palpi upturned to far above vertex of head,
slender and smoothly scaled; maxillary palpi filiform; frons

PYRALIDAH, SUBFAMILY HYPSOTROPINAE. 61

smooth ; antenne of female almost simple. Fore wing long and
narrow, the apex rounded; vein 2 from angle of cell; 3 and 5
stalked, 4 absent; 6 from below upper angle; 8, 9, 10 stalked,
8 from beyond 10; 11 from cell. Hind wing with the cell short ;
veins 2 and 5 stalked, 3 and 4 absent; 6, 7 from upper angle ;
8 strongly anastomosing with 7.

* MENUTHIA NANELLA.

Menuthia nanella Rag. Nouv. Gen. p. 50 (1888); id. Rom.
Mém. viii. p. 418, pl. 40. f. 22.
ZANZIBAR.
Genus TINERASTIA.

a0 Type.
Tinerastia Wimpsn. Rom. Mém. vii. p. 414 (1901)... fissirella.

Proboscis absent ; palpi upturned to rather above vertex of
head and nearly smoothly scaled; maxillary palpi minute and
filiform; frons with rounded prominence; antenne of male
laminate and ciliated, the basal joint rather long. Fore wing
rather long and narrow, the apex rounded ; vein 2 from towards
angle of cell; 3 and 5 from angle, 4 absent ; 6 from below upper
angle; 8, 9, 10 stalked, 8 from beyond 10; 11 from cell. Hind
wing with veins 2 and 5 from angle of cell, 3 and 4 absent; 6, 7
stalked; 8 anastomosing strongly with 7.

(1) +TINERASTIA DISCIPUNCTELLA.

Menuthia discipunctella Hmpsn. Moths Ind. iv. p. 52 (1896) ;
id. Rom. Mém. vii. p. 414, pl. 52. f. 10.

CEYLON.

(2) PTINERASTIA FISSIRELLA.

Menuthia fissirella Himpsn. Moths Ind. iv. p. 52 (1896)¢. 1d:
Rom. Mém. viii. p. 414, pl. 52. f. 9.

CEYLON.
Genus MESODIPHLEBIA.
: ce : ae Type.
Mesodiphlebia Zell. Hor. Soc. Ent. Ross. xvi.
Dito Miia bene het seniea aren ine n tio’ wedeh sue cob bes crassivenia.

Proboscis absent; palpi porrect, about twice the length of
head, almost straight and thickly scaled; maxillary palpi well
developed and slightly dilated with scales; frons with conical
tuft of hair; antenne of male laminate and ciliated, the shaft
somewhat curved at base and thickened with scales in the sinus,
the basal joint rather elongate and flattened. Fore wing long
and narrow, the apex rounded; vein 2 from near angle of cell;
3 and 5 separate, 4 absent; 8, 9, 10 stalked, 8 from beyond 10;
11 from cell. Hind wing with veins 2 and 5 from angle of cell;
3 ea absent ; 6, 7 from upper angle; 8 anastomosing strongly
with 7.

“62 SIR GEORGE HAMPSON ON THE

(1) *MEsoDIPHLEBIA DELIQUELLA.
Anerastia deliquella Gell. Isis, 1848, p. 861; Rag. Rom. Mém.

vill. p. 413, pl. 45. f. 5.
Anerastia trinotella Berg, An. Soc. Arg. 1885, p. 275.

ARGENTINA.

(2) fMESODIPHLEBIA CRASSIVENTA.

Anerastia crassivenia Zell. Hor. Soc. Ent. Ross. xvi. p. 251,
pl. xii. f. 52 (1881); Rag. Rom. Mem. viii. p. 415, pl. 40. f. 16.

COLOMBIA.

(3) MESODIPHLEBIA STRICTICOSTELLA,

Mesodiphlebia stricticostella Rag. Ann. Soc. Ent. Fr. 1887,
‘p. 260; id. Rom. Mém. viii. p. 413, pl. 40. f. 15.

N. NIGERIA; SUDAN.

(4) TMESODIPHLEBIA ROSELLA.

Calera rosella Hmpsn. Moths Ind. iv. p. 53 (1896); id. Rom.
Mém. viii. p. 412, pl. 52. f. 7.

Mapras, Nilgiris.

(5) TMESODIPHLEBIA OCHRACEELLA, Sp. Nn.

2. Head and thorax brownish ochreous; pectus, legs, and
‘abdomen white suffused with brownish ochreous. Fore wing
uniform brownish ochreous. Hind wing white with a faint
ochreous tinge.

ARGENTINA, Corrientes, Goya (Perrens), 1 2 type. Hap.
ZOcm mM.

Auctorum.
Calera albicostella Grossbeck, Bull. Am. Mus. Nat. Hist. xxxvii.
pal San Lica yspec i peomer nace ay oes U.S.A., Florida.
Genus TAMPA.
, Type.
Tampa Rag, NevAm, Phyempoloi 830) ee... dimediatella.

Proboscis aborted and small; palpi obliquely upturned to well
above vertex of head and thickly scaled; maxillary palpi well
developed and dilated with scales; frons smooth; antenne of
male ciliated. Fore wing long and narrow; the costa highly
arched, the apex rounded, the termen very oblique; vein 2 from
close to angle of cell; 3 and 5 separate, 4 absent; 6 from below
upper angle; 9 and 10 absent; 11 from close to 8. Hind wing
with vein 2 from near angle of cell; 3 and 5 strongly stalked,
4 absent; 6, 7 from upper angle; 8 anastomosing with 7 to
near apex.

PYRALIDH, SUBFAMILY HYPSOTROPIN &. 63

TAMPA DIMEDIATELLA.

Tampa dimediatella Rag. N. Am. Phye. p. 20 (1887); id. Rom.
Meém.-viil. p. 411, pl. 45. f.4; Dyar, Cat. Lep. N. Am. p. 440.

_U.S.A., Florida, Texas.

Genus CEARA.
Type.
Ceara Rag. Nouv. Gen. p. 45 (1888) ............... discinotella.
Proboscia absent; palpi upturned to rather above vertex of
head and moderately scaled ; maxillary palpi small and filiform ;
frons smooth, with large conical tuft of scales ; antenne of male
laminate and ciliated, the shaft with small tuft of scales above
at base. Fore wing narrow, the apex rounded: vein 2 from
angle of cell; 3 and 5 stalked, 4 absent; 6 from below upper
angle; 8, 9, 10 stalked, 8 from beyond 10; 11 from cell. Hind
wing with vein 2 from near angle of cell; 3 and 5 strongly
stalked, 4 absent; 6, 7 stalked; 8 strongly anastomosing with 7.

(1) *GEARA DISCINOTELLA.

Veara discinotella Rag. Nouv. Gen. p. 45 (1888); id. Rom.
Mém. viii. p. 368, pl. 39. f. 3.
S. BRraAziu.

(2) TCEARA ALBIFASCIATA, sp. n.

3. Head white; thorax white tinged with reddish brown ;
abdomen white tinged with brown and dorsally with fulvous
yellow towards base; pectus and legs white tinged with brown.
Foré wing with white costal fascia narrowing to a point before
apex and with vein 12 defined above and below by grey-brown,
the area below it to just below the cell and vein 2 white suffused
with grey-brown, leaving the veins streaked with white; the
inner area pale pink with a white streak on vein 1. Hind
wing white, tinged with brown except on inner area,

Br. KE. Arrica, Eb Urru (Betton), 1 dg type. Hxp. 26 mm.

Genus Hosipta.
Type.

Hosidia Hmpsn. Rom. Mém. viii. p. 408 (1901). ochrinewrella.

Proboscis absent ; palpi downcurved, extending about one and
a half times length of head and roughly scaled ; maxillary palpi
minute and filiform; frons smooth, with conical tuft of hair;
antenne of male ciliated, the shaft with sinus and tuft of scales
at base. .Fore wing narrow, the apex rounded; vein 2 from
close to angle of cell; 3 and 4 stalked, 5 absent ; 6 from below
upper angle; 8, 9, 10 stalked, 8 from before 10; 11 from cell.
Hind wing with vein 2 from close to angle of cell; 3 and 5
strongly stalked, 4 absent; 6, 7 stalked: 8 anastomosing strongly
with 7.

64 SIR GEORGE HAMPSON ON THE

* HosIDIA OCHRINEURELLA.

Hosidia ochrineurella Hmpsn. Rom. Mém. viii. p. 409, pl. 45.
flee Oss):

NaAtaAt.

Genus DEMBEA.
Type.

Dembea Rag. Nouv. Gen. p. 45 (1888) ............ venulosella.

Proboscis aborted and minute; palpi upturned to about vertex
of head and thickly scaled; maxillary palpi minute and slightly
dilated with scales; frons with large rounded prominence thickly
clothed with scales ; antenne of male laminate and ciliated, the
shaft with ridge of scales at base. Fore wing narrow, the apex
rounded; vein 2 from towards angle of cell; 3 and 5 shortly
stalked, 4 absent; 6 from below upper angle; 8, 9, 10 stalked,
8 from beyond 10; 11 from cell. Hind wing with vein 2 from
angle of cell; 3 and 5 stalked, 4 absent; 6, 7 stalked; 8 anas-
tomosing strongly with 7.

+D&EMBEA VENULOSELLA.

Dembea venulosella Rag. Nouv. Gen. p. 45 (1888); id. Rom.
Mém. viii. p. 368, pl. 44. f. 19.

N. Niceria; AByssintA; Br. C. AFrica.

Genus Lrorropa, nov.

Type, L. phenicias.

Proboseis absent ; palpi obliquely upturned to about vertex of
head and moderately scaled ; maxillary palpi minute and filiform ;
frons smooth, rounded; antenne of male laminate and ciliated,
with a sinus at base containing a double ridge of scales. Fore
wing narrow, the apex rounded; vein 2 from towards angle of
cell; 3and 5 stalked, 4 absent; 6 from below upper angle; 8, 9, 10
stalked, 8 from beyond 10; 11 from cell. Hind wing with vein 2
from just before angle of cell; 3 and 5 very strongly stalked,
4 absent; 6, 7 stalked; 8 strongly anastomosing with 7.

(1) tLeorroPa PHENICIAS, sp. n.

3. Head and thorax purplish pink; abdomen pale brownish
grey dorsally tinged with fulvous; antenne brownish ; pectus,
legs, and ventral surface of abdomen, except towards extremity,
brownish tinged with purplish pink. Fore wing purplish pink,
the veins streaked with white; a dark brown fascia through
upper part of cell to apex, the costal area towards apex white.
Hind wing brownish white ; a slight brown terminal line.

Sierra Leone (Clements), 1 3d type. Hap, 22 mm.

(2) tLEOTROPA SARCINA, Sp. 0.
¢. Head and thorax white mixed with ochreous brown;
abdomen white, dorsally faintly tinged with fulvous towards

PYRALIDH, SUBFAMILY HYPSOTROPINAE, 65

base ; pectus and legs white suffused with brown. Fore wing
white tinged with purplish pink, the veins streaked with white ;
the costal area white narrowing to a point at apex and with vein 12
defined on each side by brown, a brown shade below the costal
area. Hind wing white with a faint ochreous brown tinge on
costal area and in submedian interspace.

Br. E. Arrica, Teita (Jackson), 1 3 type. Hap. 16mm.

(3) fLEOTROPA PAPUANENSIS, Sp. n.

3S. Head, thorax, and abdomen whitish suffused with reddish
brown, the last dorsally tinged with fulvous towards base. Fore
wing whitish irrorated with reddish brown, the costal area rather
whiter, narrowing to a point at apex and with slight brown
streak below it. Hind wing white slightly tinged with brown,
the cilia white.

Durcu N. Guinea, Ron [. (Doherty), 1 3 type. Hap. 14 mm.

Genus ALAMOSA.
Typ

Alamosa Hmpsn. Rom. Mém, vill. p. 369 (1901). piper ae

Proboscis aborted and minute; palpi downcurved, extending
about twice the length of head and thickly scaled; maxillary
palpi minute and filiform; frons with flattened corneous plate
produced to two slight points in front and thickly scaled above :
antenne of male laminate. Fore wing narrow, the apex rounded;
vein 2 from towards angle of cell; 3 and 5 stalked, 4 absent; 6
from below upper angle; 8, 9, 10 stalked, 8 from beyond 10;
11 from cell. Hind wing with vein 2 from just before angle of
cell; 3 and 5 strongly stalked, 4 absent ; 6, 7 from upper angle ;
8 str ongly anastomosing with 7.

(1) ALAMOSA PIPERATELLA.

Alamosa piperatella Hmpsn. Rom. Mém., viii. p. 369, oa D1.
f, 25 (1901). ;

U.S.A., Colorado, New Mexico.

(2) *ALAMOSA BIPUNCTELLA.

Alamosa bipunctella Barnes & McD. Contr. Nat. Hist. Lep.
Ne Am, 11. 4, p-184, pl. 1. f. 7 (1913). ;

U.S.A., Florida.

Genus NAVASOTA.

Type.
Navasota Rag. N. Am. Phye. p. 18 (1887) ............ hebetella.

Proboscis aborted and minute; palpi extending about the
length of head and moderately scaled, the 2nd joint oblique, the
3rd porrect ; maxillary palpi minute and filiform; frons smooth
with conical tuft of hair; antenne of male laminate and ciliated,

Proc. Zoou. Soc.—1918, No. V. 5

66 SIR GEORGE HAMPSON ON THE

the shaft with sinus at base containing a ridge of scales. Fore
wing narrow, the apex rounded; vein 2 from towards angle of
cell; 3 and 5 stalked, 4 absent ; 5 from below upper angle ; 8, 9,
10 stalked, 8 from beyond 10; 11 from cell. Hind wing with
vein 2 from close to angle of cell; 3 and 5 strongly stalked,
4 absent ; 6, 7 stalked; 8 anastomosing with 7.

(1) *NAVASOTA HEBETELLA.

Navasota hebetella Rag. N. Am. Phye. p. 18 (1887); id. Rom.
Meém. vill. p. 369, pl. 39. f.5; Dyar, Cat. Lep. N. Am. p. 439.
US.A., Texas.

(2) rNAVASOTA PERSECTELLA, sp. 0.

Q. Head and thorax purplish red; abdomen white, dorsally
suffused with fulvous yellow towards base; pectus and legs
white tinged with rufous. Fore wing purplish pink; the costal
area creamy white, narrowing to a point at apex, vein 12 defined
by the costa above it being purplish red and by a fine streak
below, the costal area defined by a red-brown fascia below ;
veins 4, 3, and 1 slightly streaked with white. Hind wing white
slightly tinged with ochreous brown.

Timor, Dili (Doherty), 1 2 type. Hap. 16 mm.

(3) +NavasorA HEMAPHAELLA, Sp. 0.

@. Head and thorax dusky crimson ; abdomen Fureoues the
base of dorsal area fulvous. Fore wing dusky crimson with a
slightly defined rather paler costal fascia attenuated to a point
before apex; a very slight irroration of black scales and traces of
a terminal series of points. Hind wing ochreous tinged with
pale fuscous. ;

LovuisIaDEs, St. Aignan I. (J/eek), 1 9 type. Hap. 24 mm.

(4) fNAVASOTA LEUCONEURELLA, Sp. Nn.

¢. Head and thorax dusky crimson ; abdomen ochreous
white. Fore wing crimson; the veins streaked with white, the
subcostal and median nervures strongly so and the former
defined on lower side by brown suffusion from base to apex.
Hind wing yellowish white, the costal area tinged with fuscous.

Uaanpa, Kampala (Ansorge) ; TransvaaL, White R. (Cooke),
1 $6; Narat, Estcourt (Hutchinson), 1 g type; Carr Cotony,
Transkei (Miss F. Barrett), 1b. Hap. 24 mm.

(5) tNAVASOTA CHIONOPHLEBIA, Sp. n.

Head and thorax purplish red mixed with some ochreous
white; abdomen rufous, dorsally fulvous yellow on basal half ;
pectus and legs whitish suffused with rufous. Fore wing purplish
pink, the veins strongly streaked with white. Hind wing white
tinged with rufous.

~

PYRALID-E, SUBFAMILY HYPSOTROPIN. 67

S. Niceria, Yorubaland, Ogbomoso (Sir G. Carter), 1 9 ;
N. Nicerta, Zungeru (Simpson, Macfie), 2 2, Minna (Macfie),
13,1 9 type. Hap. 16-22 mm.

(6) {NAVASOTA SYRIGGIA, Sp. Nn.

@. Head and thorax rufous tinged with purplish pink, the
metathorax ochreous white; abdomen fulvous yellow; pectus
‘and legs white, the latter tinged with purplish rufous. Fore
wing pale purplish pink, the veins slightly streaked with white.
Hind wing white with a faint ochreous tinge.

ARGENTINA, Tucuman, Los Vasquez (Dinelly), 1 Q type. Hap.
‘24 mm.

(7) NAVASOTA DISCIPUNCTELLA, Sp. N.

2. Head, thorax, and abdomen white mixed with brown, the
last faintly tinged with fulvous yellow towards base of dorsum.
Fore wing white trrorated with fascous brown and some pale
purplish-pink scales ; a black discoidal point; a slight blackish
jpostmedial shade from vein 2 to inner margin.

N. Niceria, Minna (Macfie),1 2 type. Hap. 20 mm.

Auctorwim.
Navasota myriolecta Dyar, Pr. U.S. Nat. Mus. xlvi. p. 347

OM ite on cate, ta toe teanasdan ge amen erveh ti ee ase TE ANAMA,
Genus Hypsorropa.
. Type.

Hypsotropa Zell. Isis, 1848, p. 591 2.0.00... ee
Heosphora Meyr. P. Linn. Soc. N.S.W. vii.

asd en (leks) iene Se weno eee evn ae Sears psamathella.
Seleucia Rag. Ann. Soc. Ent. Fr. 1887, p. 259. | semiresella.
Peoria Rag, IN. Am. Phyec: p. 19) (1887) .......:: approximella,
Ambala Rag. Nouv. Gen. p. 45.(1888) ......... Suscostrigella.
Lymira Rag. Nouv. Gen. p. 46 (1888) ......... semirosella.
Tiarra Rag. Nouv. Gen. p. 46 (1888) ............ pusillella.
Socora Rag. Nouv. Gen. p. 46 (1888) ............ tenurcostella.
Talamba Rag. Nouv. Gen. p. 47 (1888) ......... tenuinervella.
Wekiva Hulst, Trans. Am. Ent. Soc. xvu.

[oe lO (UO OU) eeeceiiy or egkea venshos a gees wah sees lutercostella,
Tinitinoa Dyar, Pr. U.S. Nat. Mus. xlvii.

OS ee eo Mp ania, Stay cans gee oeasncVowss conc phyrdes.

Proboscis aborted and minute; palpi typically obliquely up-
‘turned to far above vertex of head, the 2nd joint fringed with
scales in front; maxillary palpi shghtly dilated with scales ;
frons smooth, with tuft of scales; antenne of male typically
somewhat laminate and ciliated, the shaft slightly curved at
base. Fore wing narrow, the apex rounded ; vein 2 from towards
‘angle of cell; 3.and 5 separate, 4 absent; 6 from below upper
angle; 8, 9, 10 stalked, 8 from beyond 10; 11 from cell. Hind

5

68 SIR GEORGE HAMPSON ON THE

wing with vein 2 from just before angle of cell; 3 and 5 strongly
stalked, 4 absent; 6, 7 shortly stalked ; 8 strongly anastomosing
with 7.

Sect. J. Palpi of male upturned.

A. Palpi of male hollowed out to contain the brush-like maxillary palpi; the-
antennee with sinus and ridge of scales at base of shaft.

a. (Ambala), Antenne of male with rather long uniseriate branches, the apex
ciliated.
(1) *Hypsorropa CONTRASTELLA.

Ambala contrastella Rag. Nouv. Gen. p. 45 (1888); id. Rom..
Mem*-viii. p, 370, pl, 38. 1. 11.

NATAL.

(2) Hypsorropa FUSCOSTRIGELLA.

Ambala fuscostrigella Rag. Nouv. Gen. p. 45 (1888); id. Rom.
Mem. viit.p. 371, pl. 38. £10:

PUNJAB.

(3) HypsorroPA SUBCOSTELLA, sp. Nn.

Head and thorax white, the tegule and patagia suffused with
red-brown ; antenne red-brown; palpi irrorated with red-brown ;:
abdomen white, usually irrorated with brown and dorsally suf-
fused with fulvous yellow towards base; legs irrorated with
brown. Fore wing white with a faint pink tinge, and strongly
irrorated with brown; an ochreous-brown fascia through the.
cell from base to apex, extending to just below the cell. Hind
wing white with a taint ochreous tinge.

N. Niceria, Zungeru (Mache, Simpson), 1 5, 2 2 type;
Br. EK. Arrica, N. Kavirondo, Maramas Disty., Ilala (WVeave),.
26. Hap. 20-28 mm.

(4) Hypsorropa PERVITTELLA, Sp. nN.

Head and thorax white mixed with red-brown; abdomen
white tinged with red-brown and dorsally suffused with fulvous
yellow towards base. Fore wing white thickly irrorated with
red-brown ; a red-brown fascia through the cell from base to.
apex, the veins below it streaked with white. Hind wing white.
with the costa tinged with brown towards apex.

PungaB, Ajmere, 1 ¢ ; Bompay, Deesa (Vurse), 1 ¢, 1 2
type. Hap. 20 mm.

6. Antenne of male with uniseriate branches below the sinus only, then»
serrate.
(5) HypsorropA HETEROCERELLA.

Hypsotropa heterocerella Hinpsn. Moths Ind. iv. p. 54 (1896) ;-
id. Rom. Mem, vii. p.. 371, pl..92.f. 1.

KASHMIR ; PUNJAB.

PYRALIDEH, SUBFAMILY HYPSOTROPIN®. 69

ce. Antenne of male somewhat laminate and ciliated to base.

(6) HypsorroPa OCRACEELLA, sp. Nn.

3. Head and thorax brownish ochreous mixed with some
‘white; abdomen ochreous white, dorsally tinged with fulvous
yellow towards base. Fore wing brownish ochreous; a dark
brown fascia along median nervure ; a diffused antemelial line
from cell to inner margin ; a diffused dark postmedial line tend-
ing to form streaks in the interspaces, oblique to vein 6, then
inwardly oblique; a slight dark terminal line. Hind wing
white tinged with ochreous brown and with a dark terminal ine
to submedian fold.

Timor, Oinainisa (Doherty), 1 g type. Hap. 14 min.

-B. Maxillary palpi of male filiform.
a. Antenne of male with sinus at base of shaft containing a ridge of scales.
a’, (Socora). Palpi of male with the 2nd joint obliquely upturned and thickly

scaled, the third joint porrect.
(7) *HypsoTROPA TENUICOSTELLA.
Socora tenuicostella Rag. Nouv. Gen. p. 46 (1888); id. Rom.
-Mém. viii. p. 374, pl. 39. f. 6
GAMBIA.

b’. (Lymira). Palpi of male slender, the 3rd joint upturned.

(8) *HypsorTROPA SEMIROSELLA,

Seleucia semirosella Rag. Ann. Soc. Ent. Fr. 1887, p. 259; id.
Rom. Mém. viii. p. 373) pl. 38. £.9; Staud. Cat. Lep. pal. 1
‘p. 83.

SYRIA.

(9) Hypsorropa ILLECTALIS.

Addyme illectalis Wik. xxx. 959 (1864); Rag. Rom. Mém. vil.
ip. 373, pl. 38, f. 8.
Seleucia costatella Rag. Nae Gen. p. 46 (1888).

BoRNEO; CELEBES.

(10) TH yYPsoTROPA BISCRENSIS, sp. 0.

Head and thorax white tinged with red-brown; abdomen
creamy white,-dorsally suffused with ochreous brown towards
base. Fore wing white tinged with ochreous except a white
costal fascia narrowing to a point at apex, defined below by a
red-brown fascia; the median nervure and veins 5, 3 slightly
‘streaked with white. Hind wing creamy white.

AuerriA, Biskra (Walsingham), 1 3, 2 2 type, Hammam-es-
‘Salahin (Walsingham), 13. Hap. 22-28 mm. —

(11) rHypsorRoPA RHODOCHROELLA, sp. n.
Head and thorax deep rose-red; abdomen ochreous yellow,

‘70 SIR GEORGE HAMPSON ON THE

dorsally fulvous yellow towards base; antenne fulvous yellow ;:
pectus, legs, and ventral surface of abdomen ochreous yellow
suffused with brown. Fore wing deep rose-red, the costa and
veins strongly’ streaked with white, the interspaces of costal
area tinged with brown. Hind wing ochreous white slightly
tinged with brown. :

Hab. Ucanna, Mulema (Doggett), 1 3,-Ketoma (Doggett),
1 $; Br. C.Arrica, Mt. Mlanje (Weave), 7 ¢, 3 2 type; TRraAns-.
vAAL, White R. (Cooke), 1 6; Nava, Estcourt (Mutchinson),
26. Hep. 20-28 mm.

(12) }Hypsorropa ALBIVENALIS.

Ambala albivenalis Hmpsn. J. Bomb. Nat. Hist. Sod. xxi.
p- 1250 (1912).
CEYLON.

6. Antenne: of male without sinus at base of shaft containing a ridge of scales.
al, (Hypsotropa). Palpi long; antennee of male with the shaft thickened
at base.

(13) *HyPsorropa LUTEICOSTELLA.

Hypsotropa lutercostella Rag. N. Am. Phyc. p. 19 (1887) ; id.
Rom. Meém. vii. p. 376, pl. 39. f. 9; Dyar, Cat. eps iN,
p. 439.

U.S.A., Florida.

(14) TH yPsorrRoPA OCHRICOSTELLA, sp. n.

Q. Head and tegule dark red-brown; thorax purplish red ;,
abdomen fulvous yellow; pectus, legs, and ventral surface of
abdomen red-brown. Fore wing deep purple-red ; a pale ochreous
yellow costal fascia narrowing to a point at apex, the costa
tinged with purple-red and irrorated with a few dark scales to.
beyond middle, the costal fascia defined by diffused dark brown
below. Hind wing reddish brown.

MASHONALAND (Dobbie), 1 2 type. Hap. 20 mm.

(15) tHyPsoTROPA CHIONORHABDA, sp. n.

Head and thorax black-brown mixed with some purple-grey,.
especially on dorsum of thorax ; antenne whitish tinged with red-
brown; abdomen pale reddish brown with darker segmental
lines ; pectus, legs, and ventral surface of abdomen grey suffused
with blackish, the tibie and tarsi in front white. Fore wing
purplish grey irrorated with black-brown; a silvery white costal:
fascla narrowing to a point at apex and leaving the costal edge
black towards base, defined below by a broad black-brown fascia
with diffused lower edge. Hind wing grey suffused with reddish.
brown and with darker terminal line.

©. Nigeria, Yorubaland (Sin G. Carter) 11d, lO ae
Nicerta, Zungeru (Macfie), 1 ¢; Ueanpa, Gondokoro (Reymes-
Cole), 1 3d, Ketoma (Doggett),2 5 ; Br. C. Arrica, Mt. Mlanje-

PYRALIDH, SUBFAMILY HYPSOTROPIN &. 71

(Neave), 5 3, 3 2, Luchenya R. (Veave), 6 5, 2 2 type, Ruo
Valley (Weave), 1 5. Hap. 16-24 mm.

(16) Hypsorropa INFUMATELLA.

Hypsotropa infumatella Hmpsn. Rom. Mém. viii. p. 377, pl. 39.
f3( 1901).

TRANSVAAL; NATAL.

(17) *Hypsorropa UNIPUNCTELLA.

Hypsotropa unipunctella Rag. Nouv. Gen. p. 47 (1888); id.
Rom. Mém. vii. p. 377, pl. 38. f.12; Staud. Cat. Lep. pal. ii. p. 12.
EK. Siperia, Amurland.

(18) +H YPsSoTROPA FUSIFASCIATA, sp. n.

©. Head and thorax whitish suffused with red-brown ; abdo-
men fulvous yellow, the ventral surface white tinged with red-
brown. Fore wing whitish tinged with red-brown and irrorated
with blackish; a white costal fascia irrorated with blackish,
rather diffused below, the costal edge blackish towards base ;
a black discoidal point; the terminal area suffused with blackish
except at costa; cilia with a blackish line near base. Hind
wing whitish strongly suffused with brown ; a fine dark terminal
line and line near base of cilia.

Cryton, Kandy (Green), 1 9 type. Hap, 18 mm.

(19) *Hypsorropa ZOPHOPLEURA.

Hypsotropha zophopleura Turner, Pr. R. Soc. Queensl. xviii.
ped? (1903).
(QUEENSLAND.

(20) fH ypsorropa PURPURELLA, sp. n.

Head and thorax purple-pink; antenne fulvous yellow; palpi
tinged with fuscous ; abdomen fulvous yellow; pectus, legs, and
ventral surface of abdomen greyish suffused with fuscous, the
claspers of male white. Fore wing purple-pink irrorated with
black, the costa brownish ; a black spot in the cell near base;
antemedial spots in the cell and on vein 13; obliquely placed
spots at upper and lower angles of cell; a subterminal series of
four spots, the spot at discal fold further from termen ; a terminal
series of black points. Hind wing ochreous white; a terminal
black line; cilia blackish at tips.

Br. C. Arrica, Mt. Mlanje (Veave), 9 ¢, 3 2 type. Hap.,
6 20, 2-22 mn;

(21) Hypsorropa LIMBELLA.

Hypsotropa limbella Zell. Isis, 1848, p. 591; Herr.-Schaff.
Eur. Schmett. iv.\p. 110, Tin. -f,. 38; Rag. Rom. Mém. viii.
p. 3/6; Staud. Cat. Lep. pal. ii. p. 12.

S. France; [rary ; Datmatia; Asta Minor.

72 SIR GEORGE HAMPSON ON THE

(22) Hypsorropa SYRIACELLA.

ITypsotropa syriacella Rag. Nouv. Gen. p. 46 (1888); id. Rom.
Mém. vii. p. 377, pl..39. f. 7; Staud--Cat. Lep. pal. 11. joi:

SYRIA.

(23) *Hypsorropa SOLIPUNCTELLA
Sines solipunctella Rag. Boe Mém. viii. p. 377, il 43.
oe:

(24) *Hypsorropa PAUCIPUNCTELLA.

Hypsotropa paucipunctella Rag. Bull. Soc. Ent. Fr. 1895,
p. cli; id. Rom. Mém. viii. p. 378, pl. 51. f. 10; Staud. Cat. Lep.
Palais eo.

Asta Minor, Taurus.

(25) *HypsorrRopa ICHORELLA.

Anerastia ichorella Led. Verh. zool.-bot. Ges. Wien, 1855,
p. 221, pl. 3. £.8; Rag. Rom. Mem. vili.-p. 3/8, pl’corr es
‘Staud, Cat. Lep. pal. p. 13.

SYRIA.

(26) HypsorropA VULNERATELLA.

Epischnia vulneratella Zell. Isis, 1847, p. 769: Rag. Rom.
Mém. vii. p. 378, pl. 39. f, 11; Staud. Cat. Lep. pal. 13.9, ha:

Anerastia ostrinella Lah. Bull. Soc. Vaud. vi. Contr. Faun. Sic.
p. 396 (1861).

Hypsotropa roseostrigella Rag. Rom. Mém. viii. p. 379, pl. 39.
£: L2 (1901): |

SARDINIA; Sicity; Daumatrya; SYRIA.

(27) fHyYPsoTROPA SCELETELLA.

Anerastia sceletella Zell. Stett. Ent. Zeit. 1867, p. 404; Hmpsn.
Moths Ind. iv. p. 54; “Rag. Rom. Mém. viii. p. 379, pl. 44. f. 22.

BENGAL, Calcutta.

(28) Hypsorropa VERTHEIMSTEINI.

Hypsotropa wertheimsteini Rebel, Rovart. Lep. xx. p. 171
(1913).

HUNGARY.

(29) fHyPsorRoPA PUNCFINERVELLA, sp. Nn.

Head and thorax pale ved mixed with whitish; abdomen
creamy white, dorsally fulvous yellow towards base ; palpi rufous,
white below towards base : pectus and legs creamy white, the latter
irrorated with some red-brown. Fore wing pale red slightly

PYRALIDA, SUBFAMILY HYPSOTROPINE. 13

irrorated with dark brown, the veins and costal edge white; a
minute antemedial black spot on vein 1 and postmedial spots on
veins 3, 2, 1; a terminal series of slight blackish points. Hind

wing ochreous white. : |
CryLon, Haputale (Alston), 1 ¢ type, Kegalle (Alston), 1 9,
Hambantota (Pole),1 9. Hap. 14-16 mm.

(30) HypsorroPA ADUMBRATELLA.

Hypsotropa adumbratella Rag. Nouv. Gen. p. 47 (1888); id.
Rom. Mém. viii. p. 380, pl. 38. f. 13.

GamBiA; Natrat; Cape CoLony.

b'. (Tiarra). Palpishort; antennee of male with the shaft curved at base.

(31) *HypsorRoPa PUSILLELLA. :

Tiarra pusillella Rag. Nouv. Gen. p. 46 (1888) ; id. Rom. Mém.
vill. p. 372, pl. 39. f. 4.
ZANZIBAR,

‘Sect. IJ. Palpi of male porrect,

A. (Talamba). Palpi of male hollowed out to receive the brush-like maxillary
palpi; antenne with sinus at base of shaft containing a ridge of scales.

(32) Hypsorropa TENUINERVELLA.

Talamba tenuinervella Rag. Nouv. Gen, p. 47 (1888); id. Rom.
Mem. viii. p. 387, pl. 40. f. 25; Hmpsn. Moths Ind. iv. p. 55.

PungaB, Kangra; Mapras, Nilgiris.

B. Maxillary palpi filiform.
a. Anteanz of male with sinus at base of shaft containing a ridge of scales.

al, (Tinitinoa). Antenne of male pectinate with long uniseriate branches
to middle.

(33) *HypsorroPpA PHYRDES.

Tinitinoa phyrdes Dyar, Pr. U.S. Nat. Mus. xlvu. p. 348
(1913).

PANAMA.

b’. (Heosphora). Antenne of male laminate.

(34) *HypsorROPA PAPUASELLA.

Heosphora papuasella Rag. Nouv. Gen. p. 47 (1888); id. Rom.
Mem; exxill.. py 382, pl, 39: f. 14.
N. Guinea.

(35) *HypsorRoPA SABULETELLA.

Anerastia sabuletella Zell. Lep. Micr. Caffr. p. 73 (1852); Rag.
Rom. Mém. viii. p. 383, pl. 39. f. 18.
Cape CoLony.

74 SIR GEORGE HAMPSON ON THE

(36) tHyPsorTROPA CREMORICOSTA, sp. n.

3. Head and thorax ochreous, the head and tegule suffused
with purplish red ; antenne ochreous yellow ; abdomen ochreous,.
dorsally fulvous yellow towards base; pectus, legs, and ventral
surface of abdomen purplish red. Fore wing pale flesh-red,
deepening in colour towards the costal fascia which is ochreous
white, narrowing to a point at apex, slightly defined by brown
below and leaving the costal edge brown towards base. Hind
wing ochreous white, the costal area and termen except towards.
tornus deeper ochreous.

U.S.A., Colorado, Colorado Springs (Cockerell), 1 3g type.
Hap. 24 mm.

(37) tHypsorRoPA NIVEICOSTA, Sp. n.

@. Head and thorax white suffused with rufous except on
dorsum of thorax; abdomen ochreous white, dorsally fulvous
yellow towards base ; pectus, legs, and ventral surface of abdomen
white suffused with rufous. Fore wing pale flesh-red irrorated
with a few red-brown scales and deepening to rufous towards
the costal fascia, which is snow-white narrowing to a point at
apex and leaving the costal edge rufous towards base. Hund
wing white with a faint ochreous tinge.

ARGENTINA, Gran Chaco, Florenzia (Wagner), 1 2 type. Hap.
20 mm.

(38) FHypsorroPa PERIPH#A, sp. 0.

Q. Head and tegtile white tinged with red-brown; thorax
pale purplish pink; abdomen white dorsally tinged with fulvous
yellow towards base ; pectus, legs, and ventral surface of abdomen
white tinged with red-brown. - Fore wing pale purplish pink; a
pure white costal fascia 1rrorated with afew brown scales towards.
costa, narrowing to a point at apex and defined below by dark
red-brown, diffused below. Hind wing white with a faint
ochreous tinge. |

N. Nigeria, Minna (Macfie), 2 9 type, Borgu, Yelwa Lake
(Migeod),1 2. Hap. 16 mm.

(39) THYPSOTROPA TRIPARTALIS, sp. n.

@. Head white suffused with rufous; thorax pale purplish
pink; abdomen ochreous yellow; pectus aud legs ochreous.
yellow, the fore legs tinged with brown. Fore wing with narrow
pure white costal fascia leaving the costal edge brown towards
base, the area to discal fold red-brown and the area below it
bright purplish pink. Hind wing white with a faint ochreous
tinge.

Formosa, Takow (Wileman), 1 2 type. Hap. 18 mm.

(40) fHypsorropa LATERCULELLA.
Anerastia laterculella Zell. Stett. Ent. Zeit. 1867, p. 4033.

PYRALIDH, SUBFAMILY HYPSOTROPINE. 75-

Hmpsn. Moths Ind. iv. p. 55; Rag. Rom. Mém. viii. p. 383,.
pl. 44. f. 21.

BENGAL.

(41) THyPsorROPA ROSESCENS, Sp. n.

3. Head and thorax purplish red-brown, the latter with the-
dorsum grey-brown ; abdomen fulvous yellow; pectus, legs, and
ventral surface of abdomen whitish tinged with brown. Fore.
wing pale grey-brown suffused with crimson-red, rather browner
towards the costal fascia, which is white slightly irrorated with
pale red and narrowing to apex. Hind wing white tinged with,
ochreous brown.

Cryton, Nawalapitiya (Green), 1 ¢ type. Hxp. 20 mm.

(42) *Hypsorropa STEREOSTICHA.

Hypsotropha stereosticha Turner, Pr. R. Soc. Queens]. xix.
p. 41 (1905).

QUEENSLAND, Thursday I.

(43) Hypsorropa ICASMOPIS.
_ LHypsotropha icasmopis Turner, Pr. R. Soc. Queensl. xvii. p. 116:
(1903).

(JUEENSLAND.

(44) Hypsorropa EURYZONA.

Hypsotropa euryzona Meyr. Ent. Mo. Mag. xix. p. 256 (1882) ;
Rag. Kom. Mem. viii. p. 382, pl. 39. f. 13.

S. AUSTRALIA.

(45) HypsorropA NIPHOPLEURA.

Hypsotropha niphopleuwra Turner, Pr. R. Soc. Queensl. xxiv.
pe ti 1913).

N. AUSTRALIA.

(46) fH yPpsorroPA DIAPH.ZA, sp. n.

9. Head and thorax black-brown, the frons and metathorax
rufous; antenne rufous; palpi black with a greyish gloss;
abdomen grey suffused with brown; pectus, legs, and ventral
surface of abdomen grey suffused with blackish. Fore wing
whitish ; the costal edge black with a flesh-red fascia below it to
beyond middle, the area below the cell black with a whitish
streak along vein 1. Hind wing whitish strongly suffused with.
fuscous brown, the cilia whiter with a dark line near base.

Br. C. Arrica, Mt. Mlanje (Weave), 1 9 type. Hap. 24 min.

(47) HypsorroPpaA DYSEIMATA.

Hypsotropha dyseimata Turner, Pr. R. Soc. Queensl. . xxiv.
pelle (1913):

Timor Laur; N. AUSTRALIA.

76 SIR GEORGE HAMPSON ON THE

(48) Hypsorropa QUADRIPUNCTELLA,

Hypsotropa quadripunctella Hmpsn. J. Bomb. Nat. Hist. Soc.
xu. p. 307 (1897); id. Rom. Mém. viii. p. 384, pl. 39. f. 16.
Pungas, Oude; Borneo; Puno Laur.

(49) HypsorropaA PSAMATHELLA.

Anerastia psamathella Meyr. Proc. Linn. Soc. N.S.W. iv.

p. 234 (1879); Rag. Rom. Mém. viii. p. 384, pl. 39. f. 15.
TAnerastia nitens Butl. Trans. Ent. Soc. 1886, p. 440.
(JUEENSLAND; N.S. WALES.

(50) THYPSOTROPA MONOSTIDZA, Sp. .

2. Head and thorax ochreous with a faint purplish pink
tinge; abdomen ochreous with a fulvous yellow tinge; pectus,
legs, and ventral surface of abdomen ochreous with a brownish
tinge. Fore wing ochreous faintly tinged with purplish pink
and irrorated with dark brown ; a minute blackish diseoidal spot.
Hind wing white tinged with ochreous especially towards termen.

SIERRA LEONE (Clements), 1 9 type. Aap. 18 mm.

(51) rHypsoTRoPpA GRAPTOPHLEBIA, Sp. n.

3. Head and thorax white mixed with purplish red-brown ;
abdomen white slightly tinged with brown; pectus, legs, and
ventral surface of abdomen white suffused with red-brown.
Fore wing white suffused with purplish red-brown and slightly
irrorated with dark brown, the veins white defined on each side
by fine dark brown streaks. Hind wing white tinged with
‘ochreous brown.

MASHONALAND, Salisbury (Aarshall), 1 d type. Hap. 20 mm.

(52) TH YPSOTROPA POLYACTINIA.

Heosphora polyactinie Hmpsn. Rom. Mém. vii. p. 384, pl. 52.
fa 2 (1901):

UGANDA; MASHONALAND; TRANSVAAL; NATAL.

(53) TH ypsorropA ENDORHODA, sp. Nn.

Q. Head and thorax ochreous suffused with purplish pink ;
abdomen ochreous white, dorsally fulvous yellow towards base ;
pectus, legs, and ventral surface of abdomen ochreous white
suffused with red-brown. Fore wing with the costal half to
median nervure and vein 5 ochreous, tinged with purplish pink
towards costa and with slight pale streaks on the veins; the
inner half purplish pink with slight white streaks on the veins.
Hind wing white tinged with ochreous.

N. Ruopesia (Coryndon), 1 9 type. Hap. 24 mm.

(54) HypsorropA RAMULOSELLA.
Heosphora ramulosella Rag. Bull. Soc. Ent. Fr. 1895, p. cii; id.

PYRALIDA, SUBFAMILY HYPSOTROPIN. (i:

Rom. Mém. viii. p. 385, pl. 52. f. 3; Staud. Cat. Lep. pal. ii.
p. 12.
SYRIA.

(55) Hypsorropa LEUCOPHLEBIELLA.

Heosphora leucophlebiella Rag. Nouv. Gen. p. 47 (1888); id..
Rom, Mém. viii. p. 385, pl. 39. f. 17.

Br. C. ArricA; MASHONALAND; TRANSVAAL; Natal; CAPE
CoLony.

(56) HypsorropA RHODOSTICHA.

Hypsotropha rhodosticha Turner, Pr. R. Soc. Queensl. xviii.
pr ule: (1903).
QUEENSLAND.

b. (Peoria). Antenne of male without sinus and ridge of scales at base of shaft
(57) THypsorRoPA POLYSTICTELLA, sp. 0.

¢. Head, thorax, and abdomen whitish suffused with rufous.
Fore wing ochreous white suffused with rufous in the inter-
spaces leaving the veins pale; the costal edge black towards
base; a slight shade formed by black irroration below the costa
to apex ; a small antemedial black spot on vein 1, a small spot
on lower discocellulars and an oblique postmedial series of four
small spots on veins 5, 3, 2, 1; a terminal series of small black
spots. Hind wing white tinged with ochreous; a fine fuscous
terminal line and slight brown line near base of cilia.

Br. C. Arrica, Mt. Mlanje (Weave), 2 35 type. Hap. 20 mm.

(58) * Hypsorropa ACNIDIAS.

Hypsotropha acnidias Turner, Pr. R. Soc. Queensl. xviii. p. 117
(1903)
. QUEENSLAND.

(59) rHypsorropA APPROXIMELLA.

EHurhodope approwimella Wik. xxxv. 1722 (1866); Rag. Rom.
Mém. viii. p. 386, pl. 40. f. 24; Dyar, Cat. Lep. N. Am. p. 439.

Anerastia hwmatica Zell, Verh. zool.-bot. Ges. Wien, 1872,
p: 555.

Anerastia roseatella Pack. Ann. N. Y. lye. Nat. Hist. x.
p20 (1870).

U.S.A., Massachusetts, N. York, Pennsylvania, Ohio, Illinois,
N. Carolina, Texas, Colorado.

(60) rHypsorroPA LEUCOCRASPIS, sp. n.

6. Head, thorax, and abdomen white suffused with ochreous .
brown. Fore wing pale ochreous brown, the costal edge white.
Hind wing white tinged with ochreous brown especially on apical
area,

ARGENTINA, Corrientes, Goya (Perrens), 1 3 type. Lxp.20 mm.

8

78 SIR GEORGE HAMPSON ON THE

(61) *HypsorROPA BIPARTELLA.

Peoria bipartella Rag. N. Am. Phye. p. 19 (1887); id. Rom.
Mem. viii. p. 386, pl. 40. f. 33 ; ieee Cat. Lep. N. Am. p. 439.

U.S.A., N. Carolina.

Auctorum.

Peoria albidella Hulst, Can. Ent. xxxiii. p. 178 (1900); Dyar,
Cat. ep. No Am 2p) 439 2), ee U.S.A., California.
Hypsotropha laropis Turner, Pr. R. Soc. Queensl. xxiv. p. 173
(ROPES) iiahde hc coteteeeoneeee Eeeeee (Queensland.
i neurica Turner, Pr. R. Soc. Queensl. xxiv. p. 113
CEOS Votan, oe chanel cts ere N. Australia.

my castella Pag. Zoologica, xxix. p. 163 (1900).

Bismarek Arch.

xenus RAPHIMETOPUS, nov.

Type, FR. spinifrontella.

Proboseis aborted and minute; palpi downcurved, extending
-about three times length of head and thickly scaled; maxillary
palpi minute and filiform; frons with long flattened corneous
plate, somewhat trifid in front and produced to short lateral
processes, a corneous plate below the frons; antennze of male
laminate. Fore wing narrow, the apex rounded, the termen
obliquely curved; vein 2 from close to angle of cell; 4 absent ;
5 from above angle; 6 from below upper angle; 8, 9 stalked;
10, 11 from cell. Hind wing with vein 2 from just before angle
of cell; 3 and 5 stalked, 4 absent; 6, 7 shortly stalked; 8 closely
approximated to or anastomosing with 7,

(1) RAPHIMETOPUS SPINIFRONTELLA.

Anerastia spinifrontella Rag. Nouv. Gen. p. 48 (1888); id
Rom. Mém., viii. p. 399, pl. 40. f. 2; Hmpsn. Moths Ind. i
p. 96.

Bompay, Sind.

(2) RAPHIMETOPUS ABLUTELLA.

Anerastia ablutella Zell. Isis, 1839, p. 17; Herr.-Schaff. Hur.
Schmett. iv. p. 109, Tin. f. 89; Rag. Rom. Mém. vii. p. 403;
Hmpsn. Moths Ind. iv. p. 56; Staud. Cat. Lep. pal. 11. p. 12.

+ Anerastia stigmatella Rag. Nouv. Gen. p. 49 (1888); id. Rom.
Mém. viii. p. 403, pl. 40. f. 4.

Anerastia bimaculata Rag. Nouv. Gen. p. 49 (1888); id. Rom.
Mém. viii. p. 404, pl. 40. f.5; Hmpsn. Moths Ind. iv. p. 56.

Anerastia majorella Roths. Nov. Zool. xx. p. 138 (19138).

Spain; Siciry; AugertaA; N. Nigeria; Conco, Eeypr; Da-
MARALAND; TRANSVAAL; ORANGE R. Cotony ; ADEN; Asia M1nor;*
- Persia; W. TURKESTAN; JAPAN; KaAsumirn; PunjAB; MADRAS.

' PYRALIDH, SUBFAMILY HYPSOTROPINE. ahd

Auctorum.

Anerastia korbi Cwwadja, Ivis, xxiv. p. 117 (1910).
CL UHLCULGS Sere PE A CASPIAN SBA.

Genus METACRATERIA, nov.

Type, JI. pulverulella.

Proboscis absent; palpi downcurved, extending about twice
the length of head and thickly scaled; maxillary palpi minute
and filiform; frons with long truncate corneous prominence with
raised rim at extremity, a flattened plate below the frons ;
antenne of male laminate and ciliated. Fore wing rather
narrow, the apex rounded, the termen evenly curved ; vein 2 from
near angle of cell; 4 absent; 5 from above angle; 6 from below
upper angle; 8,9 stalked; 10, 11 from cell. Hind wing with
vein 2 from just before angle of cell; 3 and 5 stalked, 4 absent ;
6, 7 shortly stalked; 8 approximated to but not anastomosing
with 7.

(1) METrACRATERIA METALLACTIS.

Anerastia metallactis Meyr. Trans. Ent. Soc. 1887, p. 622;
Rag. Rom. Mém. viii. p. 399, pl. 40. f. 11.
N.S. WALEs.

(2) METACRATERIA PULVERULELLA.

Anerastia pulverulella Hmpsn. Moths Ind. iv. p. 56 (1896) ;
id. Rom. Mém. viii. p. 400, pi. 52, f. 4.

CEYLON.

(3) TMETACRATERIA PERIRRORELLA, Sp. n.

Q. Head and thorax red-brown tinged with grey and slightly
irrorated with fuscous ; abdomen grey suffused with red-brown ;
palpi below, pectus, legs, and ventral surface of abdomen whitish
tinged with red-brown. Fore wing whitish suffused with red-
brown and thickly irrorated with black; small spots formed by
black irroration at upper and lower angle of cell; cilia with a
whitish line at base. Hind wing whitish strongly suffused with
reddish brown, the cilia whiter.

MASHONALAND (Dobbie), 2 2 type. Hap. 28 mm.

(4) METACRATERIA MIASTICTA, Sp. n.

9. Head, thorax, and abdomen whitish suffused with red-
brown. Fore wing whitish suffused with red-brown and slightly
irrorated with black, especially on the veins; a small black spot
at lower angle of cell. Hind wing white tinged with red-
brown.

Mexico, Presidio (Forrer), 1 2 type, Godman-Salvin Coll.
Hep. 24 mum.

80 SIR GEORGE HAMPSON ON THE

Genus PRINANERASTIA *,

Type, P. lotella.

Proboscis absent ; palpi downcurved, extending about twice
the length of head and thickly scaled; maxillary palpi minute
and filiform ; frons with rounded prominence with slight raised
rim at extremity, a corneous plate below the frons; antennee of
male laminate and minutely ciliated. Fore wing narrow, the apex
rounded, the termen evenly curved; vein 2 from towards angle of
cell; 4 absent; 5 from above angle; 6 from below upper angle ;
8, 9 stalked; 10,11 from cell. Hind wing with vein 2 from
just before angle of cell; 3 and 5 stalked, 4 absent; 6, 7 shortly
stalked ; 8 closely approximated to but not anastomosing with 7.

(1) PRINANERASTIA LACTEALIS.

Lymire lactealis Roths. Nov. Zool. xx. p. 138 (19138).
Enosima albicostalis Roths. Nov. Zool. xx. p. 1388 (1913).
Lymire strictipennis Roths. Nov. Zool. xxii. p. 237 (1915).
ALGERIA ; TUNIS.

(2) PRINANERASTIA NITIDICOSTELLA.

Anerastia nitidicostella Rag. Ann. Soc. Ent. Fr. 1887, p. 259
id. Rom. Mém. viii. p. 399, pl. 40. f. 12; Staud. Cat. Lep. pal. ii.
p12,

S. Russia.

(3) PRINANERASTIA LOTELLA.

Tinea lotella Hiibn. Kur. Schmett., Tin. f. 344 (1817); Dup.
Lép. Fr. x. pl. 283. £. 6; Herr.-Schaff. Hur. Schmett. iv. p. 100,
Tin, ff. 90-92; Leech, Pyr. pl. 10. f.3; Rag. Rom. Mém., viii.
p. 397; Staud. Cat. Lep. pal. 1. p. 12.

Tinea miniosella Zinck. Germ. Mag. iii. p. 126 (1818).

Tinea pulverella Hubn. Kur. Schmett., Tin. f. 454 (1823),

Britain ; FRANCE: GERMANY; Austria; Huncary ; Swirzer-
LAND; W. Russta; AstA Minor; Persia; W. TURKESTAN.

(4) *PRINANERASTIA INCARNATELLA,

Anerastia incarnatella Rag. Ann. Soc. Ent. Fr. 1887, p. 259;
id. Rom. Mém. viii. p. 398, pl. 38. f. 15; Staud. Cat. Lep. pal. 11.
palz:

S. Russa.

Genus CHORTONGCA, nov.

Type, C. leucocraspia.

Proboscis aborted and minute ; paipi of male typically obliquely
upturned to above vertex of head and thickly scaled, hollowed
out to receive the brush-like maxillary palpi, of female down=

* The type of Anerastia Htibn. is dignella Hiibn.

PYRALIDA, SUBFAMILY HYPSOTROPIN&. 81

curved and about twice the length of head; frons with pointed
conical prominence; antenne of male typically with rather long
uniseriate branches, the shaft with sinus at base containing a
ridge of scales. Fore wing narrow, the apex rounded, the termen
evenly curved; vein 2 from towards angle of cell; 3 and 5 from
angle, 4 absent ; 6 from below upper angle; 8,9 stalked; 10, 11
from cell. Hind wing with vein 2 from just beforeangle of cell ;
3 and 5 stalked, 4 absent; 6, 7 shortly stalked; 8 closely
approximated to 7 but not anastomosing with it.

Sect. I. Palpi of male obliquely upturned and hollowed out to receive the brush-
like maxillary palpi; antenne pectinate with uniseriate branches, the shaft
with sinus and ridge of scales at base.

(1) TCHORTONGCA LEUCOCRASPIA, sp. n.

Head and thorax whitish suffused with rufous ; abdomen white
tinged with rufous and dorsally suffused with fulvous yellow
towards base; palpi white in front; pectus, legs, and ventral
surface of abdomen white tinged with rufous. Fore wing
creamy white slightly tinged with rufous; a white costal fascia,
narrowing to a point at apex, slightly irrorated with rufous and
defined below by a rather diffused rufous streak; the veins
white defined on each side by fine rufous streaks ; a fine rufous
streak in submedian fold. Hind wing semihyaline white.

AucEeRIA, Hammam-es-Salahin (Walsingham). 2 ¢,3 Q type.
Exp. 30-34 mm.

Larva. White with numerous red-brown lines interrupted at
the incisures, the head and Ist thoracic somite wholly red-
brown. Feeds in the stems of a reed-like grass.

Szot. II. Palpi of male obliquely upturned and not hollowed out to receive the
maxillary palpi, which are filiform; antenne of male laminate and ciliated
with slight sinus at base of shaft, but no ridge of scales.

(2) }CHORTONGCA MINIMELLA.
Meliarpha minimella Hmpsn. Rom. Mém. viii. p. 392, pl. 52.

f. 11 (1901).

Borneo, Pulo Laut; CreLeses, Talaut I.; N. Avsrraia.

Sect. III. Palpi of male downcurved and extending about three times length of
head, not hollowed out to receive the maxillary palpi which are filiform ;
antenne of male laminate and ciliated, with a sinus at base of shaft con-
taining a ridge of scales.

(3) *CHORTONGCA MINORALIS.
Arerastia minoralis Lower, Tr. R. Soc. 8. Austr. 1903, p. 52.
QUEENSLAND.

(4) CHorToNgCA EURYSTICHA.

Anerastia eurysticha Turner, Pr. R. Soc. Queensl. xviii. p. 119
(1903).

Proc. Zoou, Soc.—1918, No. VI. 6

82 SIR GEORGE HAMPSON ON THE

Hypsotropha niphosema Turner, Pr. R. Soc. Queensl. xxiv.
Poll2 Clos):

QUEENSLAND; N. AUSTRALIA.

Genus RHINAPHE.

Type.

Lhinaphe Berg, Bull. Soc. Imp. Nat. Mose. xlix. i

2y, Ds Zo k QLSL4s) -c.-so esas ae ua eee eens signaticollis.
Ampycophora Meyr. Pr. Linn. Soc. N.S.W. vii.

py lOS CUS82) es: caeaecaene creck nee on eR EeEe apotomella.
Comorta Rag. Nouv. Gen. p. 48 (1888) ......... nigricostalis.
Maliarpha Rag. Nouv. Gen. p. 48 (1888) ...... separatella. .
Homosassa Hulst, Trans. Am. Ent. Soc. xvii.

De ZT (ISO rh so pce ea nau eee ella.

Enosima Rag. Rom. Mém. viii. p. 389 (1901)... neesimella.
Ampycodes Hmpsn. Rom. Mém. vii. p. 393

(LOO Tie fvtecrecate tees Cae R ET ie oan tatters ae Te pallrdicosta.
Hrythphlebia Hmpsn. Rom. Mém. viii. p. 393 ,
CO Oda) aes heeiad 22.2) 20 Re See eee enervella.

Proboscis absent ; palpi typically downcurved, extending about
three times length of head and thickly scaled; maxillary palpi
filiform ; frons smooth and rounded; antenne of male typically
laminate and ciliated. Fore wing narrow, the apex rounded, the
termen obliquely curved; vein 2 from near angle of cell; 4
absent; 5 from above angle; 6 from below upper angle; 8, 9
stalked; 10, 11 from cell. Hind wing with vein 2 from just
before angle of cell; 3 and 5 stalked, 4 absent; 6, 7 shortly
stalked ; 8 closely approximated to or anastomosing with 7.

Srcor. I. Palpi of male hollowed out to receive the brush-like maxillary palpi.
A. Palpi of male upturned.
a. (Ampycophora). Antenne of male uniserrate.

(1) + RHINAPHE APPROXIMELLA, Sp. 0.

®. Head and thorax purplish red, the vertex of head and
dorsum of thorax whitish tinged with red; abdomen white,
dorsally tinged with fulvous yellow towards base ; pectus white ;
legs red-brown. Fore wing whitish suffused with purplish pink,
leaving the veins whiter and defined on each side by fine
purplish-red streaks; a white costal fascia tinged with purplish
pink at costa, harrowing to a point at apex, defined below by
a narrow dark brown streak giving off a fine streak above vein 6.
Hind wing white, the costal area and termen tinged with brown.

(JUEENSLAND, Peak Downs, 1 2 type. Hap. 24 mm,

(2) RHINAPHE APOTOMELLA.

Pempelia apotomella Meyr. Pr. Linn. Soc. N.S.W. iv. p. 224
(1879); Rag. Rom. Mém. viii. p. 388, pl. 40. f. 17.

PUNJAB; CEYLON; SELANGOR; PHILIPPINES; CELEBES, Sangir I.
Timor, Oinainisa, Dili ; QUEENSLAND.

PYRALIDH, SUBFAMILY HYPSOTROPINE. 83

b. (Comorta). Antenne of male laminate.
(3) *RHINAPHE PLINTHINA.

Anerastia plinthina Turner, Pr. R. Soc. Queensl. xix. p. 43
(1905).

QUEENSLAND; N. AUSTRALIA.

(4) TRHINAPHE CASTANEALIS.

Anerastia castanealis Hmpsn. J. Bomb. Nat. Hist. Soc. xxi,
p. 1251 (1912).

Br. C. Africa; TRANSVAAL; CEYLON; BALI.

(5) TRHINAPHE CELSELLA.

Araxes celsella Wik. xxvii. 193 (1863); Hmpsn. Moths Ind.
iv. p. 55; Rag. Rom. Mém. viii. p. 405, pl. 40. f. 7.

CEYLON ; PHILIPPINES.

(6) TRHINAPHE NIGRICOSTALIS.

Trachonitis nigricostalis Wik. xxvii. 40 (1863); Hmpsn.
Moths Ind. iv. p. 57; Rag. Rom. Mém. viii. p. 389, pl. 39. f. 22.
Comorta atricostella Rag. Nouv. Gen. p. 48 (1888).

Gambia; 8. Nigeria; TRANSVAAL; CEYLON, BurMA; ANDA-
MANS; Borneo; Fst.
(7) RuINAPHE HOLOPH@A.

Ampycophora holophea Turner, Pr. R. Soc. Queensl. xix.
p. 42 (1905).

QUEENSLAND; N. AUSTRALIA.

B. Palpi obliquely upturned. ‘ q
a. Antenne of male with short uniseriate branches, the apical part ciliated.

(8) RHINAPHE VECTIFERELLA.

Enosima vectiferella Rag. Rom. Mém. vui. p. 391,.pl. 42. £. 24
(1901).

Br. E. Arrica; Ucanpa; Br. C. Arrica; TRANSVAAL 3
Mapacascar; Comoro Is.

‘b. (Hnosima). Antennee of male laminate.

(9) *RHINAPHE VENELLA.

Enosima venella Hmpsn. Rom. Mem. viii. p. 391. pl. 40. £. 19
(1901),

CocHIN CHINA.

(10) RHINAPHE NEESIMELLA.

Enosima neesimella Rag. Rom, Mém. viii. p. 390, pl. 43. f. 22
(1901).
JAPAN; COREA.
6*

84 SIR GEORGE HAMPSON ON THE

(11) RaInaPHEe FLAVESCENTELLA.

Enosima flavescentella Hmpsn. Rom. Mém. viii. p. 390, pl. 40.
f. 18°(1901). :

Formosa ; C. CHINA.

c. (Ampycodes). Palpi downcurved in both sexes; antenne uniserrate and
ciliated, with a large sinus at base of shaft containing a ridge of scales.

(12) }RHINAPHE STICTELLA.

Anerastia stictella Hmpsn. J. Bomb. Nat. Hist. Soc. xvii.
p. 259 (1908).

PUNJAB.

(13) fRHINAPHE PALLIDICOSTA.

Anerastia pallididicosta Hmpsn. Moths Ind. iv. p. 57 (1896) ;
id. Rom. Mém. viii. p. 393, pl. 39. f. 23.

AssaAM; BurMaA.

(14) *RHINAPHE HAPLOSCHEMA.

Ampycophora haploschema Turner, Pr. R. Soc. Queensl. xviii.
p. 117 (1908).

QUEENSLAND.

Secr. II. Palpi of male not hollowed out to receive the maxillary palpi, which are
filiform.

A. (Maliarpha). Palpi of male obliquely upturned; the antenne laminate and
ciliated, with a large sinus at base of shaft containing a ridge of scales.

(15) *RHINAPHE SEPARATELLA.

Maliarpha separatella Rag. Nouv. Gen, p. 48 (1888); id. Rom.
Mém. viii. p. 392, pl. 39. f. 21.

CAMEROONS.

B. Palpi downcurved in both sexes.

a. (Erythphlebia). Antennz of male laminate and ciliated, with a large sinus at
base of shaft containing a ridge of scales.

(16) RHINAPHE ENERVELLA.

Erythphlebia enervella Hrapsn. Rom. Mém. viii. p. 394, pl. 39.
f. 24 (1901).

N. Guinea; Louisiane Is. ; QuEENSLAND; W. AUSTRALIA.

(17) RuINAPHE VIRGINELLA.

Anerastia virginella Meyr. Proc. Linn. Soc. N.S.W. iv. p. 233
(1879); Rag. Rom. Mém. viii. p. 394, pl. 40. f. 8.

QUEENSLAND.

PYRALID®, SUBFAMILY HYPSOTROPINA. 85

6. Antenne of male bipectinate, without sinus and ridge of scales at base of
shaft.
(18) RHINAPHE BISERIELLA.
Anerastia biseriella Hmpsn. Rom. Mém, viii. p. 397, pl. 52.
f. 18 (1901).
QUEENSLAND.

c. (Rhinapha). Antenne of male laminate and ciliated, without sinus and
ridge of scales at base of shaft.

(19) }RHINAPHE HEMIRHODELLA.

Anerastia henirhodella Hmpsn. Rom. Mém. viii. p. 402, pl. 52.
f. 12 (1901).

S. BRaziu.

(20) RHINAPHE LOTRICELLA.

Anerastia lotricella Zell. Isis, 1848, p. 861; Rag. Rom. Mém.
viii. p. 401, pl. 8. f. 21.
S. BRAZIL.

(21) * RHINAPHE LEUCOTAENIELLA.

Anerastia leucoteniella Rag. Nouv. Gen. p. 48 (1888) ; id. Rom.
Mém. viii. p. 401, pl. 40. f. 3.

JAPAN,

(22) TRHINAPHE SANGIRENSIS, Sp. n.

@. Head and thorax dark red-brown ; abdomen grey tinged
with brown and dorsally fulvous yellow towards base; pectus
and legs grey suffused with brown. Fore wing deep purple-
pink; a white costal fascia narrowing to apex, defined below
by diffused dark red-brown and with the costal edge and vein
12 purple-pink. Hind wing whitish tinged with red-brown
especially on costal area; a fine brown terminal line except
towards tornus.

CeesBes, Sangir I. (Doherty), 1 2 type. Hap. 18 mm.

(23) RHINAPHE BRUNNEOVITTELLA.

Anerastia brunneovittella Rag. Nouv. Gen. p. 49 (1888); id:
Rom. Mém. viii. p. 401, pl. 40. f. 6; Hmpsn. Moths Ind. iv.
p. 56.

C. Cu1na; Pungasp; BomBay; CEYLON.

(24) }RHINAPHE PHHOSTROTELLA, Sp. n.

@. Head and thorax creamy white mixed with red-brown ;
abdomen creamy white, dorsally fulvous yellow towards base ;
* pectus and legs creamy white. Fore wing creamy white irrorated
with pinkish brown; a creamy white costal fascia with hardly

86 SIR GEORGE HAMPSON ON THE

any brown irroration on it, narrowing to apex and defined below
by diffused brown. Hind wing white.
Cryton, Puttalam (Pole), 1 9 type. Hap. 22 mm.

(25) RHINAPHE SYSSEMA.

Anerastia syssema Turner, Pr. R. Soc. Queensl. xxiv. p. 114°
(1913). :

QUEENSLAND; N. AustTrAuiIA; W. AUSTRALIA.

(26) RHINAPHE LATERITIELLA, Sp. n.

Q. Head and thorax grey-brown mixed with some white ;
abdomen whitish suffused with red-brown, the basal segment
white. Fore wing reddish brown, darker towards the pure
white costal fascia narrowing to apex. Hind wing white slightly
tinged with brown.

B. Niger1a, Yorubaland, Ogbomoso (Sir G'. Carter), 1 Q type.
Hap. 20 mm. 3

(27) *RHINAPHE ELLA.
* Ephestia ella Hulst, Ent. Am. ui. p. 138 (1887); Rag. Rom.
Mém. viii. p. 400, pl. 40. f. 1; Dyar, Cat. Lep. N. Am. p. 440.
U.S.A., Florida. °

(28) RHINAPHE TALIELLA,

Anerastia taliella Hmpsn. Rom. Mém. vui. p. 402, pl. 53. £. 17
(1901). |

(JUEENSLAND.

(29) + RHINAPHE VENILINEELLA, Sp. n.

Q. Head and thorax white irrorated with reddish brown;
abdomen white tinged with red-brown, dorsally fulvous yellow
towards base. Fore wing white thickly irrorated with reddish
brown, the veins white defined on each side by fine brown
streaks. Hind wing white.

SupDAN, Port Sudan (Waterfield), 1 2 type. Hap. 24 mm.

(30) *RHINAPHE INFUMELLA.

Anerastia infumella Rag. Ann. Soc. Ent. Fr. 1887, p. 260; id.
Rom. Mém. vii. p. 402, pl. 38. f. 16; Staud. Cat. Lep. pal. 1.
Oe Lee.

PERSIA,

(31) *RHINAPHE CONSPERSELLA.

Anerastia conspersella Rag. Rom. Mém. viii. p. 404, pl. 40
f. 13 (1901).

U.S.A., Colorado,

PYRALIDH, SUBFAMILY HYPSOTROPINA. 87

(32) *RHINAPHE SEEBOLDI.

Anerastia seeboldi Rag. Ann. Soc. Ent. Fr. 1894, p. 177; id.
ire, xi. pl dot. 7 5 Staud. Cat. Lep. pal. p. 12.
SPAIN.

(33) TRHINAPHE ENDONEPHELE, Sp. 0.

6. Head, thorax, and abdomen white slightly tinged with
red-brown and irrorated with dark scales, the last dorsally
fulvous yellow towards base; antenne brownish. Fore wing
white irrorated with blackish and faintly tinged with red-brown ;
a rather diffused rounded blackish antemedial spot on vein 1; a
terminal series of blackish points. Hind wing white faintly
tinged with ochreous brown ; a slight brown terminal line.

Braziu, Rio Janeiro, 2 ¢ type. Hap. 20 mm.

(34) PRHINAPHE IGNETINCTA, Sp. 0.

2. Head and thorax ochreous white suffused with fiery red ;
abdomen ochreous. Fore wing ochreous suffused with fiery red
and on costal half tinged with purplish pink; oblique diffused
antemedial and medial blackish bars on inner area ; two slight
dark diseoidal points; an indistinct oblique brown postmedial
line; a terminal series of slight dark points. Hind wing semi-
hyaline white tinged with ochreous especially towards termen.

ARGENTINA, Sta. Fé, Ocampo (Wagner), 2 2 type, Buenos
Ayres (Wilkinson), 1 9. Hap, 22 mm.

(35) *RHINAPHE MICTOCHROELLA.

Anerastia mictochroella Rag. Nouv. Gen. p. 49 (1888) ; id. Rom.
Mem. viii. p. 404, pl. 40. f. 10.

ARGENTINA, Goya.

(36) TRHINAPHE FURVIMACULA, Sp. n.

®. Head and thorax ochreous faintly tinged with rufous ;
abdomen ochreous, dorsally fulvous yellow towards base. Fore
wing ochreous faintly tinged with rafous and irrorated with
dark brown scales; an oblique slightly sinuous red-brown almost
medial line with a conical fulvous-red spot on its outer side
in submedian interspace ; a small dark brown spot at lower angle
of cell; an indistinct brown postmedial line, oblique below
vein 5. Hind wing ochreous white suffused with purple-brown
especially on terminal half except towards tornus ; cilia whiter.

ARGENTINA, Tucuman, Los Vasquez (Dinelly), | 2 type. Hap.
28 mm.

(37) RHINAPHE SIGNICOLLIS.

Rhinaphe signicollis Berg, Bull. Soc. Imp. Nat. Mose. xlix, 2,
p. 233 (1874); Rag. Rom. Mém. viii. p. 405, pl. 40. f. 9.

ARGENTINA, Gran Chaco, Florenzia, Buenos Ayres,

88 SIR GEORGE HAMPSON ON THE

Auctorum.
Anerastia viphimela Lower, Tr. R. Soc. S. Austr. 1903, p. 52,
ni, Ay CUslaneals: Jae eee (JUEENSLAND.
Fe ablepta Turner, Pr. R. Soc. Queensl. xxiv. p. 114
(1913). ?% Metacrateria. N. QUEENSLAND ;

N. AUSTRALIA.
argosticha Turner, Pr. R. Soc. Queensl. xxiv. p. 115

(1913), nr. f. enervella 2... 0.02. N. AUSTRALIA.
* anemopsis Turner, Pr. R. Soc. Queens). xxiv. p. 116
(T9103); 2 mr. Miccelsella cecastaeee N, AUSTRALIA.
ss baliora Turner, Pr. R. Soc. Queens]. xxiv. p. 116
(L913), (nr. di. plinthid, Vee ee (QUEENSLAND.
= acrophea Turner, Pr. R. Soc. Queensl. xxiv. p. 117
(1913). @Sect. Hrythphlebia ....... N. AUSTRALIA.
“ pleurochorda, Turner, Pr. R. Soc. Queensl. xxiv.
p- 117 (1913). @Sect. Hnosima ... QUEENSLAND.
. erasmia Turner, Pr. R. Soc. Queensl. xxiv. p. 117
(1913). ?Sect. Hrythphlebia......... (QUEENSLAND.

‘* ephestiella Viard, Bull. Soc. Ent. Fr. 1913, p. 82.
Basses ALPES.

Genus SUDANIA.

Type.
Sudania Hmpsn. Rom. Mém. viii. p. 380 (1901)... — sebeostella.

Proboscis aborted and minute; palpi upturned to just above
vertex of head, the 2nd joint moderately scaled, the 3rd moderate ;
maxillary palpi filiform ; frons smooth, with conical tuft of scales ;
antennee of male with short uniseriate branches. Fore wing
rather narrow, the apex rounded; vein 2 from near angle of cell ;
3 and 5 from angle, 4 absent; 6 from below upper angle; 8, 9,
10 stalked, 8 from beyond 10; 11 from cell. Hind wing with
vein 2 from before angle of cell; 3 and 5 from angle, 4 absent ;
6, 7 stalked ; 8 not anastomosing with 7.

*SUDANIA SUBCOSTELLA.

Sudania subcostella Hmpsn. Rom. Mém. viii. p. 381, pl. 51.
f; IE Gl90L).
GABOON.
Genus RHopocHRysa.

Type.
Rhodochrysa Hmpsn. Rom. Mém. viii. p. 387 (1901). seperbella.

Proboscis aborted and minute; palpi downcurved, extending
about twice the length of head and thickly scaled; maxillary
palpi filiform ; frons smooth, with conical tuft of scales; antennz
of male laminate and minutely ciliated, the shaft slightly curved
at base but without ridge of scales. Fore wing narrow, the apex
rounded, the termen obliquely curved ; vein 2 from towards angle
of cell; 3.and 5 separate, 4 absent; 6 from below upper angle ;
8,9, 10 stalked, 8 from beyond 10; 11 from cell. Hind wing

PYRALIDH, SUBFAMILY HYPSOTROPINA. 89

with vein 2 from just before angle of cell; 3 and 5 from angle,
closely approximated towards origin, 4 absent; 6, 7 shortly

stalked; 8 closely approximated to 7 but not anastomosing
with it.

RHODOCHRYSA SUPERBELLA.

Rhodochrysa superbella Hmpsn. Rom. Mém. viii. p. 387, pl. 39.
Fo bas(l901).

TRANSVAAL; NATAL.

Genus Ca@NoTROPa, nov.
Type, C. limitella.

Proboscis absent; palpi downcurved, extending about three
times length of head and thickly scaled ; maxillary palpi minute
and filiform ; frons with rounded prominence; antenne of female
minutely ciliated. Fore wing long and narrow, the apex rounded ;
vein 2 from towards angle of cell; 4 absent; 5 from above angle ;
6 from below upper angle; 8, 10, 11 stalked, 9 absent. Hind
wing with vein 2 from towards angle of cell; 3 and 4 absent ;
6, 7 from upper angle; 8 anastomosing strongly with 7.

{C@NOTROPA LIMITELLA, Sp. Nn.

2. Head and thorax red-brown mixed with some flesh-white ;
abdomen red-brown, dorsally fulvous towards base. Fore wing
pale purplish pink; a white costal fascia slightly irrorated with
purplish red, narrowing to apex and defined below by a rather
diffused dark brown streak. Hind wing semihyaline white tinged
with flesh-colour, the costal area suffused with brown.

PARAGUAY, Sapucay (foster), 1 2 type. Hap. 22 mm.

Genus BANDERA.
Type.

Bandera Rag. N. Am. Phye. p. 19 (1887)......... binotella.
Osceola Hulst, Smith, List N. Am. Lep. p. 85

(1891), non descr. nec Baird, Rept. 1853 ...... perlepidella.
Chipeta Hulst, Can. Ent. xxiv. p. 62 (1892)...... perlemidella.

Proboscis aborted and minute; palpi downcurved, extending
about twice the length of head and roughly scaled; maxillary
palpi well developed, filiform ; frons smooth and rounded ; an-
tenn of male minutely ciliated, the shaft shghtly curved at base.
Fore wing narrow, the apex routers vein 2 from towards angle
of cell; 3 and 5 om a point or separate ; 6 from below upper
angle; 9 absent; 10,11 from cell. Hind wing with vein 2 from
just before angle of cell; 3, 5 strongly stalked, 4 absent ; 6,7
from upper angle; 8 anastomosing strongly w ich ie

(1) BANDERA BINOTELLA.
Anerastia binotella Zell. Verh. zool.-bot. Ges. Wien, 1872, p. 108 :

90 SIR GEORGE HAMPSON ON THE

Rag. Rom. Mém. viii. p. 409, pl. 44. f. 23; Dyar, Cat. Lep.
N. Am. p. 440.
U.S.A., Texas, New Mexico.

(2) *BANDERA PERLEPIDELLA.

Chipeta perlepidella Hulst, Can. Ent. xxiv. p. 62 (1892); Dyar,
Cat. Lep. N. Am, p. 441.

U.S.A., Florida.

(3) BANDERA GUPIDINELLA.

Bandera cupidinella Hulst, Ent. Am. iv. p. 119 fae ; Rag.
Rom. Mém. viii. p. 410, pl. 45. f.3; Dyar, Cat. Lep. N. Am.
p. 440.

U.S.A., Colorado, New Mexico.

(4) * BANDERA SUBLUTEELLA.

Bandera subluteella Rag. N. Am, Phye. p. 19 (1887); id. Rom.
Mém. viii. p. 410, pl. 40. fe 21; Dyar, Cat. Lep. N. Am. p. 440.

U.S.A., Colorado, Giiearena. New Mexico.

Auctorum.

Bandera virginella Dyar, Proc. Ent. Soc. Wash. x. p. 116
(UOOS) fikcatceanc: sone teres U.S.A., Washington.
carneella Barnes & McD. Contr. Nat. Hist. “Lep.

N. Am. ii. 4. p. 184, pl. i. f. 5 (1913).
U.S.A., Florida.

tke)

Genus LAURENTIA.
Type.
Laurentia Rag. Nouv. Gen. p. 49 (1888) ............ inclarella.

Proboscis absent; palpi of male upturned to vertex of head,
thickly sealed and hollowed out to receive the brush-like maxillary
palpi, of female long and downcurved; frons smooth, with tuft
of scales; antenne of male ciliated, the shaft with sinus at base
of shaft containing a ridge of scales. Fore wing narrow, the
apex rounded, the termen obliquely curved ; vein 2 from towards
angle of cell; 3 and 5 from angle, 4 absent ; 6 from below upper
angle; 8,9 stalked; 10,11 from cell. Hind wing with vein 2
from before angle of cell : ; 3and 5 from angle, closely approximated
towards base, 4 absent; 6, 7 shortly stalked; 8 anastomosing
with 7.

(1) fLAURENTIA ALBIVENELLA, sp. 0.

' $. Head and thorax creamy white mixed with some red-
brown; abdomen white, dorsally tinged with fulvous yellow
towards base; legs tinged with brown. Fore wing creamy
white irrorated with some dark brown, the veins white defined

PYRALIDH, SUBFAMILY HYPSOTROPIN&. ‘91

on each side by fine dark brown streaks, the median nervure
more strongly streaked with white; a dark brown shade through
upper part of cell and thenee to termen below apex; the inter-
spaces between veins 4 and 2 shaded with brown; cilia chequered
with brown except at base. Hind wing white, the costal area
and termen tinged with brown. .

Formosa, Takow (Wileman), 1 3 type. Hap. 22 mm.

(2) *LAURENTIA INCLARELLA.

Laurentia inclarella Rag. Nouv. Gen. p. 49 (1888); id. Rom.
Mém. viii. p. 408, pl. 38. f. 14.

JAVA.
Genus CALAMOTROPA, nov.

Type, C. pulverivena.

Proboscis absent; palpi downcurved, about two and a half
times length of head and thickly sealed; maxillary palpi well
developed and slightly dilated with scales; frons with large
pointed conical prominence clothed with rough scales ; antenne
of female minutely ciliated. Fore wing long and narrow, the
apex rounded; vein 2 from towards ‘angle. of cell; 3 and 5
separate, 4 absent ; 6 from below upper angle; ; 8,9 stalked ; 10;
1] from cell. Hind wing with vein 2 from just below angle of
cell; 3 and 4 absent; 6, 7 shortly stalked ; 8 not anastomosing
with 7.

TCALAMOTROPA PULVERIVENA, Sp. 0.

@. Head and thorax ochreous slightly tinged ih brown ;
pectus, legs, and abdomen ochreous white. Fore wing ochreous
with diffused fuscous irroration along the veins except on costal
area; a slight black point at ion angle of cell. Hind wing

ochr ure eine: the costal area and termen tinged with brown.

W. AUSTRALIA, ss R. (Clements), 3 9 type. Hap.
24 mm.

Genus EpPrpAurRtia.
Type.
Kpidauria Rag. Rom. Mém. vii. p. 405
(LS STONS a ere ges oe eee ee eee an ee .. transversariella.

Proboscis absent ; palpi downcurved, extending about three
times length of head and thickly scaled ; maxillary palpi filiform ;
frons smooth, with tuft of scales; antenne of male laminate.
Fore wing narrow, the apex, rounded, the termen evenly curved:
vein 2 from towards angle of cell; 4 absent; 5 from above angle; 6
from below upper angle; 8,9 stalked; 10,11 from cell. Hind
wing with vein 2 from before angle of cell; 3 and 5 from angle ;
4 absent; 6, 7 from upper angle; 8 approximated to but not
anastomosing with 7,

92 SIR GEORGE HAMPSON ON THE

(1) TEprpAURIA PERFASCIELLA, sp. n.

2. Head and thorax whitish suffused with rufous ; abdomen
whitish suffused with rufous and with obscure dark dorsal seg-
mental bands. Fore wing whitish suffused with pale purplish
red and slightly irrorated with blackish, the costal area rather
whiter; a rather diffused black fascia from base through upper
part of cell to termen below apex, forking in end of cell on vein 5
to well beyond the cell; vein 1 white at middle with rather
diffused short black streaks above and below it; the postmedial
line represented by a slight black mark above vein 6, a very
oblique line between veins 5 and 2 and a short streak above
vein 1; a terminal series of minute rather V-shaped biack marks.
Hind wing whitish suffused with reddish brown; cilia ochreous
white with a slight brown line near base.

STERRA LEONE (Clements), 1 9 type. Hap. 28 mm.

(2) *EPIDAURIA TRANSVERSARIELLA.

Anerastia transversariella Zell. Isis, 1848, p. 588; Herr.-Schiff.
Kur. Schmett. iv. p. 109. Tin. f. 33; Rag. Rom. Mém. viii. p. 406,
pl. 38. f. 1/7 ; Staud, Cat. Lep. pal. 1i. p. 12.

Daumatia ; Corru; Asta MInor.

(3) TEPIDAURIA CHIONOCRASPIS, Sp. 0.

2. Head white tinged with purplish red; thorax pale purplish
red; pectus, legs, and abdomen white tinged with brown. Fore
wing with narrow silvery white costal fascia, the area below it
dark chocolate-brown to median nervure and on terminal area
diffused on its lower side; the rest of wing pale purplish red.
Hind wing white with a slight ochreous tinge, the costa tinged
with brown.

Br. C. Arrica, Mt. Mlanje (eave), 1 2 type. Hap. 24 mm.

(4) EPIDAURIA PHCENICEELLA.

Epidauria pheniceella Rag. Bull. Soc. Ent. Fr. 1895, p. ciii;
id. Rom. Mém. viii. p. 407, pl. 52. f.5; Staud. Cat. Lep. pal. ii.
peolZ:

AstA Minor; Syria.

(5) *EPIDAURIA DISCELLA.

Epidauria discella Rag. Rom. Mém. ii. p. 407, pl. 40. f. 14
(1901); Stand, .Cat. Lep: pal. a. .p. 12:

MESOPOTAMIA.

(6) *EPIDAURIA STRIGOSA.

Anerastia strigosa Staud. Hor. Soc. Ent. Ross. xv. p. 225
(1879); Rag. Rom. Mém. viii. p. 407, pl. 45. f. 2; Staud. Cat.
Lep. pal. il. p. 12.

DatmatiA; AstA Minor; Syria; EH. Siperta.

PYRALIDA, SUBFAMILY HYPSOTROPIN#. 93

(7) TEPIDAURIA SUBCOSTELLA, Sp. Nn.

®. Head, thorax, and abdomen whitish mixed with brown, —
the last dorsally fulvous yellow towards base; pectus, legs, and
ventral surface of abdomen whitish suffused with dark brown.
Fore wing whitish suffused with ochreous brown, the costa with
a white streak below it to beyond middle defined below by a dark
brown streak extending to apex. Hind wing whitish suffused
with brown, the inner area whiter.

Yunnan, Teng Yeuk (//obson), 1 9 type. Hap. 26 mm.

Genus SALURIA.

Type.
Saluria Rag. Ann. Soc. Ent. Fr. 1887, p. 258. maculivittella.

Powjadia Rag. Nouv. Gen. p. 42 (1888) ...... sepicostella.
Baroda Rag. Nouv. Gen. p. 42 (1888) ......... paucigraphella.
Goya Rag. Nouv. Gen. p. 13 (1888) .. albivenella.
Pe Be den id ae Nouv. Gen. p. 43 (1888,

Ps) macrella.
Cayuga Hulst, “Ent. Am. iv. '?. “116 (1888,

Sept.) .. Setter eons agemmatella,
Atascosa Hulst, “Trans. Am. Ent. Soc. xvii.

p. 210 (1890) . - glareosella.
Parramatta Eopen. Rom. Mem. viii. ae 366

GEOON) 23... ensiferella.
Ollia Dyar, J. N.Y. Ent. Soc. xii. P. “107

et OL eae santaritella.
Humoorea mele Insec. Inscit. Meustr. v. 'p. 91

(Ee ae anchridis.

Proboscis aborted and minute; palpi of male typically down-
curved, extending about three times length of head and thickly
scaled, the 3rd joint moderate; maxillary palpi filiform; frons
smooth with slight tuft of hair; antenne of male typically with
long uniseriate branches and without sinus and ridge of scales at
base of shaft. Fore wing long and narrow, the apex rounded,
the termen evenly curved; vein 3 from close to angle of cell;
4, 5 strongly stalked; 6 from below upper angle; 8, 9 stalked ;
10, 11 from cell. Hind wing with vein 2 from close to angle of
cell; 3 and 5 strongly stalked, 4 absent; 6, 7 shortly stalked ;
8 typically not anastomosing with 7.

Sort. I. Palpi of male hollowed out to receive the brush-like maxillary palpi.
A. Antenne of male with a sinus at base of shaft containing a ridge of scales.
a. Antenne of male pectinate with long uniseriate branches.
a’, Palpi of male oblique.

(1) SaLURIA ERODELLA.

Powjadia erodella Rag. Nouv. Gen. p. 48 (1888); id. Rom.
Mém. viii. p. 347, pl. 37. f. 5; Hmpsn. Moths Ind. iv. p. 59.

Poujadia parviplumella Hmpsn. Moths Ind. iv. p. 59 (1896).

PunsgaB; BEna@AL; Mapras; CEYLON.

94 ' SIR GEORGE HAMPSON ON THE

b’. (Baroda). Palpi of male downcurved,
(2) SALURIA CTENUCHA.
Poujadia ctenucha Turner, Pr. R. Soc. Queensl. xxiv. p. 118
(1913).
N. AustTRALtiA.

(3) *SALURIA FLAMMELLA.

Baroda flammella Hmpsn. Rom. Mém. viii. p, 348, pl. 51. f. 12
(1901).

GAMBIA.

(4) TSALURIA CARNESCENS, sp. 0.

Head and thorax. pale rufous; abdomen white tinged with
rufous, dorsally fulvous. yellow towards base; pectus and legs
white suffused with rufous. Fore wing with the inner half pale
purplish pink shading to rufous towards the white costal fascia,
narrowing to apex ; the costal edge pale purplish red oh basal
half. Hind wing white with an. satoone, tinge.

ARGENTINA, Sta. Fé, Ocampo (Wagner), 1 3 type, Cratt Chaco,
Florenzia (Wagner), 1.2... Hap. 28-30 mm.

(5) fSALURIA -PSAMMATELLA, Spo Ty

3. Head, thorax, and abdomen white tinged with ochreous

brown; antenne red-brown, white towards base. Fore wing

white tinged with egatcous a own, the oy area aga ee

fine aaa fer mene ey
Q. Fore wing more strongly suffused with ochreous brown
leaving a distinct white costal fascia narrowing to apex.
ALGERIA, Hammam-es-Salahin (Walsingham), 1 3, 1 @ type.
Hap. 26-32 mm.

(6) ¢SALURIA PAUCIGRAPHELLA.

Baroda paucigraphella Rag. Nouv. Gen. p. 48 (1888); id. Rom.
Mém. viii. p. 348, pl. 38. f. 5: Hmpsn. Moths Ind. iv. p. 60.

BENGAL, Calcutta ; Bompay, Kutch; Mapras.

b. Antenne of male pectinate with short uniseriate branches; palpi oblique.

(7) TSALURIA HEMIPHALIS.

Saluria hemiphealis Hmpsn. J. Bomb. Nat. Hist. Soc. xxi.
pe lZol (oi).
CEYLON.

(8) TSALURIA SEMIROSELLA, Sp.n. — .
Head whitish suffused with rufous; thorax pale purplish pink

PYRALID#, SUBFAMILY HYPSOTROPINA. 95.

with some rufous on shoulders; abdomen white tinged with red-
brown, dorsally fulvous yellow towards base; pectus and legs
white tinged with red-brown. Fore wing with the inner half
pale purplish pink, the veins finely streaked with white; the
cell and area beyond it red-brown below the white costal fascia
narrowing to apex; the costal edge red-brown towards base, then
pale pink to beyond middle. Hind wing white tinged with red-
brown, the cilia whiter with a red-brown line near base.

Sierra Leone (Clements), 1 do, 2 2; Uaanpa, Nakwai Hills
(Lowe), 1 3, Gondokoro (Heymes-Cole), 1 g type. Hap. 16-

20 mm.

c. Antenne of male strongly serrate ; palpi downcurved.

(9) TSALURIA MINUTELLA.

Saluria minutella Hmpsn. J. Bomb. Nat. Hist. Soc. xv.,p, 20
(1903).

BomBAY ; CEYLON, sp. n..

(10) TSALURIA NILGIRIENSIS, sp. n.

3. Head and thorax pale rufous ; abdomen white tinged with
rufous; antenne with some black in the ridge of scales ; pectus,
legs, and ventral surface of abdomen brown. Fore wing with the
inner half creamy white suffused with rufous varying io purplish
pink; the upper part of cell and area beyond it brown below the
white costal fascia narrowing to apex; the costal edge brown
towards base, then irrorated ache rufous; a rather oblique post-
medial series of short dark streaks on veins 6 to 1 and a terminal
series of sight dark points. Hind wing white tinged with brown
especially on costal area, the cilia whiter with a slight brown line
near base.

Mapras, Nilgiris, Pykara (Andrewes), 4 ¢ type. Hup. 26-
28 mm.

d. (Pouwjadia). Antenne of male laminate ; palpi oblique.

(11) SALURIA SEPICOSTELLA.

Poujadia sepicostella Rag. Nouv. Gen. p. 42 (1888); id. Rom.
Mém. vii. p. 345, pl. 36. f. 2; Hmpsn. Moths Ind. iv. p. 58.
Formosa ; PunsaB: BorneéEo, Pulo Laut.

(12) SALURIA CLARICOSTELLA.

Pouwjadia claricostella Rag. Nouv. Gen. p. 42 (1888); id. Rom.
Mém. viii. p. 345, pl. 36. f. 26.
GoLp Coasr; Br. C. AFRICA,

(13) TSALURIA GLAREOSELLA.
Anerastia glareosella Zell. Verh. zool.-bot. Ges. Wien, 1872,

96 SIR GEORGE HAMPSON ON THE

p- 593; Rag. Rom. Mém. viii. p. 345, pl. 37. f£. 19; Dyar, Cat.
Lep. N. Am. p. 438.

U.S.A., Texas.

(14) TSALURIA STICTOPHORA, Sp. n.

Head and thorax reddish brown; abdomen whitish tinged
with brown, dorsally fulvous yellow towards base; antenne of
. male with the ridge of scales black; pectus, legs, and ventral
surface of abdomen white mixed with brown. Fore wing whitish
suffused with brown and thickly irrorated with dark brown; a
white costal fascia slightly irrorated with dark brown and rather
diffused below; a small round black discoidal spot; an indistinct
rather diffused blackish postmedial line not quite reaching the
costa ; cilia with a fine white line at base. Hind wing whitish
suffused with brown. ;

MASHONALAND, Salisbury (Varshall), 1 $, 19 type. Hap.
20 mm.

(15) SALURIA SPURCELLA.

Powjadia spurcella Rag. Nouv. Gen. p. 42 (1888); id. Rom.
Mém. vii p. 346, pl. 37. f.4; Hmpsn. Moths Ind. iv. p. 59.

PungaB; BomBaAy; MApras; Burma; LABUAN.

(16) TSALURIA INFICITA.

Acrobasis inficita Wik. xxvii. 30 (1863); Hmpsn. Moths Ind.
iv. p. 58; Rag. Rom. Mém. viii. p. 346, pl. 44. f.18. —

TRANSVAAL; Mapras ; CEYLON.

(17) tSALURIA FLAVICOSTA, sp. n.

Head rufous; thorax grey-brown, the tegule dorsally rufous ;
abdomen grey-brown, the 2nd and 3rd segments dorsally fulvous
yellow, the anal segment pale yellow; antenne brown; palpi
yellow with black patch on 2nd segment above ; pectus, legs, and
ventral surface of abdomen grey-brown and whitish. Fore wing
dark brown glossed with grey; a narrow pale yellow costal fascia,
the costal edge dark towards base. Hind wing pale grey-brown,
the cilia whitish with a brown line near base.

SrerrA LEONE (Clements), 135; N. Nigeria, Zungeru (Macfie),
1 ¢ type. Hap. 16 mm.

(18) *SaALURIA FLOSCELLA.

Atacoea floscella Hulst, Trans. Am. Ent. Soc. xvii. p. 210
(1890); Rag. Rom. Mém. viii. p. 346, pl. 51. f. 4; Dyar, Cat.
Lep. N. Am. p. 438.

U.S.A., Texas.

(19) TSALURIA STICTELLA, sp. n.
do. Head and thorax brownish ochreous; abdomen ochreous

PYRALIDE, SUBFAMILY HYPSOTROPIN &. 97

white, dorsally fulvous yellow towards base; palpi white below
towards base; pectus, legs, and ventral surface of abdomen
white, the legs tinged with brown. Fore wing white tinged with
ochreous brown except the costal area to discal fold, irrorated
with black scales, the costal and inner areas less irrorated; small
obliquely placed black antemedial spots in and below the cell and
a cdiscoidal spot; a postmedial series of black points on veins 6
to 1 and a terminal series of slight black points. Hind wing
white tinged with ochreous brown.
Bananas, Andros (Lonhote), 1 ¢ type. Hep. 16 mm.

(20) SALURIA OCHRIDORSELLA.

Powadia ochridorsella Rag. Nouv. Gen. p. 42 (1888); id. Rom.
Mém. viii. p. 347, p. 37. f. 1; Hmpsn. Moths Ind. iv. p. 58.

PungaB; BeneaL; CEYLON.

(21) TSALURIA ROSELLA.

Poujudia rosella Hmpsn. Moths Ind. iv. p. 59 (1896); id. Rom.
Mém. viii. p. 347, pl. 51. f. 20.

Mapbras, Nilgiris.

(22) PSALURIA VERECUNDELLA.

Poujadia verecundella Rag. Rom. Mém. viii. p. 347, pl. 37. f. 2
(1901).

CotomBiIA ; 8. BrRAziL; ARGENTINA.

(23) TSALURIA LENTISTRIGELLA, Sp. Nn.

Head and thorax white, the teguiz except dorsally and. the
patagia slightly tinged with meroue: abdomen white slightly
tinged with rufous, dorsally fulvous yellow towards base. Fore
wing chalky white, the veins defined by slight streaks of red-
brown scales and the inner area slightly irrorated with red-
brown; a faint rufous shade below median nervure; a small
rather diffused round blackish spot on vein 1 and slight obliquely
placed postmedial spots on veins 2 and 1; a fine punctiform
blackish terminal line; cilia chequered with blackish near base
except towards tornus. Hind wing white with fine brown ter-
minal line except towards tornus.

SreRRA LEONE (Clements), 1 6; Goup Coasr, Bibianaha
(Spurrell), 23,192 type. Hap, 22- 24 mm.

(24) TSALURIA DESERTELLA, sp. n.

@. Head and thorax white faintly tinged with rufous ;
abdomen white, dorsally fulvous yellow towards base. Fore
wing white tinged with ochreous, the cilia whiter. Hind wing
white.

N, Austrauia, Alexandria (Stalker), 3 29; W. AusTRALta,
Sherleck R. (Clements), 19 type. Hap, 22-32 mm.

Proc, Zoou, Soc,—1918, No. VII. 7

\

98 SIR GEORGE HAMPSON ON THE

B. (Humoorea). Antenne of male laminate, without sinus and ridge of scales
at base of shaft.

(25) *SALURIA ANCHRIDIS.

Eumoorea anchridis Dyar, Insec. Inscit. Mentr. v. p: aL (1917).
Br. GUIANA.

(26) ¢SALURIA FLAVIPURPURELLA, Sp. Nn.

¢. Head and thorax rufous mixed with ochreous; abdomen
ochreous rufous, dorsally fulvous yellow towards hase; pectus
and legs white suffused with brown; ventral surface of abdomen
brown, whitish at base. Fore wing with creamy white costal
fascia narrowing to apex, the costal edge blackish at base, defined
below by dark brown followed by deep purple-pink to just below
the cell and vein 4, the inner half of wing yellow. Hind wing
ochreous white, the apical area tinged with brown.

Paraguay, Sapucay (Foster), 1g type. Hap. 16 mm.

Srcr. II. Maxillary palpi of male filiform.
A. Antennse of male with a sinus at base of shaft containing a ridge of scales.

a. Antenne of male pectinate with rather long uniseriate branches to about
half length ; palpi downcurved.

(27) TSALURIA PARANENSIS.
Pectinigeria paranensis Hmpsn. Rom. Mém. vin. p. 355, pl. 5d.

f. 6 (1901).
S. BRAZIL.

(28) *SALURIA MUSHELLA.
Pectinigeria muscella Schaus, A. M.N. H. (8) xi. p. 239 (1913).
Cosra Rica.

(29) TSALURIA TENUICOBTA, Sp. Nn.

Head and thorax rufous tinged with pale pink; abdomen
ochreous, dorsally fulvous yellow towards base ; pectus, legs, and
ventral surface of abdomen whitish suffused with red-brown.
Fore wing pale purplish red sparsely irrorated with black; a
narrow white costal fascia detined below by blackish to beyond
middle; a terminal series of slight dark points. Hind wing
white, tinged with ochreous brown on costal area and termen.

Paracuay, Sapucay (fester), 13,19 type. Hap. 18 mm.

(30) SALURIA PLEUROSTICHA.

Hypsotropha pleurosticha Turner, Pr. R. Soc. Queensl. xviii.
p- 115 (1903).

QUEENSLAND.

(31) fSALURIA NEURICELLA, sp. n.

Head and thorax rufous mixed with some whitish ; abdomen

PYRALIDH, SUBFAMILY HYPSOTROPIN A, 99

ochreous white, dorsally fulvous yellow towards base; pectus and
legs white suffused with brown. Fore wing rufous tinged with
purplish pink, the veins streaked with white. Hind wing whitish
suffused with ochreous brown, the cilia whiter.

QUEENSLAND, Peak Downs, 1¢,19 type. Hap. 22 mm.

b. Antennse of male laminate and ciliated.
a'. (Goya). Palpi oblique, the 3rd joint porrect.

(32) SaLURIA ALBIVENELLA.

Goya albivenella Rag. Nouv. Gen. p. 43 (1888); id. Rom. Mém.
vin. p, 349, pl. 37..f. 6:

S. Braziu; ARGENTINA.

(33) *SALURIA VARICOSELLA.

Goya varicosella Rag. Nouv. Gen. p. 43 (1888); id. Rom. Mém.
Taliep. o00, pl, of. t. 7.
GAMBIA.

(34) }SALURIA PROLEUCELLA, sp. n.

3. Head and thorax white suffused with red-brown ; abdomen
white tinged with red-brown, dorsally fulvous yellow towards
base. Fore, wing white tinged and irrorated with red-brown, the
veins finely streaked with white; a white costal fascia slightly
irrorated with red-brown, narrowing to apex. Hind wing creamy
white, the costal area faintly tinged with brown.

Goup Coast, Accra (Sir G'. Carter), 13 type. Hap. 22 mm,

(35) TSALURIA GRAMMIVENA, Sp. n.

Head, thorax, and abdomen white suffused with red-brown ;
pectus, legs, and ventral surface of abdomen white irrorated with
red-brown. Fore wing white thickly ivrorated with red-brown,
the veins strongly streaked with white and defined on each side
by fine red-brown streaks; cilia whitish. Hind wing white
tinged with red-brown.

N. Ausrrauia, Alexandria (Stalker), 1 § ; W. AvusrrRaLtia,
Sherlock R. (Clements), 1 9 type. Hap. 26 mm.

b’. (Pectinigeria). Palpi downcurved.

(36) SALURIA NIGRITELLA.

Pectinigeria nigritella Rag. Rom. Mém. viii. p. 353, pl. 37. f. 13
(1901).

Gapoon ; N. RHODESIA.

(37) *SALURIA ARDIFERELLA.

Altoona ardiferella Hulst, Ent. Am. iv. pl. 118 (1888); Rag.
Rom. Mém. vin. p. 506, p. 24, f. 14: Dyar, Cat. Lep, N. Am.
p. 439, |

7*

100 SIR GEORGE HAMPSON ON THE

Aurora nigromaculella Hulst, J. N. Y. Ent. Soc. viii. p. 224
(1900); Dyar, Cat. Lep. N. Am. p. 438.
U,S.A., Texas, Colorado, California, N. Mexico, Arizona.

(38) SALURIA DEVYLDERI.

Pectinigeria devylderi Rag. Nouv. Gen. p. 43 (1888); id. Rom.
Mem. vii. p. 354, pl. 37. f. 10.

N. Reopesta; DAMARALAND; NATAL.

(39) SALURIA ARCTICOSTELLA.

Pectinigeria arcticostella Rag. Nouv. Gen. p. 438 (1888) ;
Rom. Mém. viii. p. 354, pl. 37. f. 9.

N. NicGErIA ; ZANZIBAR.

(40) *SaLURIA FURVELLA.

Pectinigeria furvella Rag. Nouv. Gen. p. 43 (1888); id. Rom.
Mém. viii. p. 355, pl. 37. f. 11.

MADAGASCAR.

(41) SALURIA MACRELLA.

Pectinigeria macrella Rag. Nouv. Gen. p. 44 £1888); id. Rom.
Nem: Vill. p. 390, pl oi.t slo. |
Br. C. Africa; NArAat.

(42) SALURIA SUBCARNELLA.

Pectinigeria subcarnella Rag. Nouv. Gen. p. 44 (1888); id.
Rom. Mém. vii. p. 356, pl. 37. f. 15.
N. Nigeria; TrRAnsvAAL; Natau; Cape CoLony.

(43) SALURIA GEMMATELLA.

Spermatophora gemmatella Hulst, Ent. Ain. ii. p. 134 (1887) ;
Rag. Rom. Mém. viii. p. 356, pl. 37f. 12; Dyar,Cat. Lep:
N. Am. p. 438.

U.S.A., Illinois, Colorado, California.

(44) TSALURIA RHODOPHMA, Sp. 0.

2. Head and thorax vinous purple ; abdomen fulvous tinged
with brown except dorsally towards base ; palpi, pectus, legs, and
ventral surface of abdomen red-brown. Fore wing vinous purple
with a slight whitish streak on subcostal nervure tending to
fork at extremity and defined below by a dark fascia extending
through the cell to apex. Hind wing ochreous white tinged with
brown especially towards costa.

TRANSVAAL (Pead), 19 type. Hap. 24 mm.

(45) TSALURIA SUBCOSTELLA, Sp. nh.

Q. Head and thorax deep rufous mixed with some whitish
especially on metathorax ; abdomen fulvous with slight whitish

PYRALIDA, SUBFAMILY HYPSOTROPINA. 101

segmental lines and the extremity whitish; pectus, legs, and
ventral surface of abdomen whitish suffused with red-brown.
Fore wing whitish tinged with red-brown; the costal edge dark
brown to well beyond middle ; a white costal fascia narrowing to
apex, defined below by a dark brown fascia. Hind wing ochreous
white tinged with brown especially towards costa.

Br. C. Arrica, Mt. Mlanje (Weave), 19 type. Hxp. 30 mm.

(46) TSALURIA DISTICTELLA, sp. n.

3. Head and thorax whitish, the vertex of head, tegule and
patagia suffused with red-brown; antenne tinged with dark
brown ; palpi with some blackish scales at side of 2nd joint ;
abdomen white, dorsally fulvous yellow towards base; legs tinged
with red-brown. Fore wing white tinged with flesh-pink; a
pure white costal fascia narrowing to apex and defined below by
rufous which is rather diffused below; an antemedial red-brown
point on vein | and medial point in submedian fold. Hind wing
white.

QUEENSLAND, Brisbane, 1 ¢ type Hap. 20 mm.

(47) TSALURIA RUFELLA, Sp. n.

®. Head and thorax rufous; abdomen whitish suffused with
brown, dorsally fulvous yellow towards base; pectus and legs
whitish suffused with rufous, Fore wing uniform rufous. Hind
wing ochreous white.

BenGaL, Oudh (Pilcher), 12 type. Hap. 26 mm.

(48) TSALURIA OPIFICELLA.

Anerastia oprficella Zell. Stett. Ent. Zeit. 1867, p.406; Himpsn
Moths Ind. iv. p..60; Rag. Rom. Mém. viil.’p. 357, pl. 44. f. 16.

KASHMIR ; BENGAL; CEYLON; BuRMA.

(49) *SALURIA BREVICULELLA.

Saluria breviculella Hmpsn. J. Bomb. Nat. Hist. Soe. xii. p. 808
1898); id. Rom. Mém. viii. p. 358, pl. 37. f. 14.

] : |

BomBay.

B. Antenne of male without sinus and ridge of scales at base of shaft.
a. (Parramatta). Hind wing with vein 8 anastomosing with 7; antenne of
male laminate and ciliated.
(50) SALURIA DICHROELLA.
Saluria dichroella Rag. Ent. Am. v. p. 118 (1889); id. Rom.
Mém. viii. p. 363, pl. 39. f. 1; Dyar, Cat. Lep. N. Am. p. 439.
U.S.A., Texas.

(51) *SALURIA NEOTOMELLA.

Hucarphia neotomella Meyr. Proc, Linn. Soc. N.S. W. iv. p. 226
(1879); Rag. Rom. Mém. viii. p. 363, pl. 37. £ 25.

N.S. W ALES.

1OZ SIR GEORGE HAMPSON ON THE

(52) *SALURIA OSTREELLA.

Saluria ostreella Rag. N. Am. Phyc. p. 18 (1887); id. Rom.
Mém. viii. p. 362, pl. 38. f.6; Dyar, Cat. Lep. N. Am. p. 438.

U.S.A., Arizona.

(53) TSALURIA TETRADELLA.

Anerastia tetradella Zell. Verh. zool.-bot. Ges. Wien, 1872,
p- 552; Rag. Rom. Mem. vii. p. 362, pl. 42. f. 23; Dyar, Cat.
Lep. N. Am. p. 439.

U.S.A., Texas, Colorado.

(54) SALURIA HOLOCHROA.

Powadia holochroa Turner, Pr. R. Soc. Queens]. xviii. p. 121
(1903).

W. AUSTRALIA; VICTORIA.

(55) *SaALURIA CANCELLIELLA.

Saluria cancelliella Rag. Nouv. Gen. p. 44 (1888); id. Rom.
Meém: vill: p, 363, pl. ova, 2a.

BRAZIL.

(56) *SALURIA ENSIFERELLA. _

Eucarphia énsipherella Meyr. Proc. Linn. Soc. N.S. W. i.
p. 209 (1878); Rag. Rom. Mém. viii. p. 366, pl. 39. f. 2.

N.S. Wags.

6. Hind wing with vem 8 not anastomosing with 7.
a’. (Saluria), Antenne of male pectinate with long uniseriate branches.

(57) *SALURIA PECTIGERELLA.
Saluria pectigerella Rag. Ann. Soc. Ent. Fr. 1887, p. 259; id.

Rom. Mém. viii. p. 360, pl. 37. f. 24; Staud. Cat. Lep. Pal. i.
p. La:

W. TURKESTAN.

(58) SALURIA MACULIVITTELLA.

Saluria maculivittella Rag. Ann. Soc. Ent. Fr. 1887, p. 258;
id. Rom. Mém. viii. p. 361, pl. 38. f. 7; Staud. Cat. ep. pal.
i. plo.

Saluria armeniella Rag. Nouv. Gen. p. 44 (1888).

ALGERIA; Tunis; ARMENIA; Cyprus; W.'TURKESTAN ; CEYLON.

(59) TSALURIA PULVEROSA.

Powadia pulverosa Hmpsn. Moths Ind. iv. p. 60 (1896); id.
Rom. Mém. viii. p. 361, pl. 37. f. 3.

SIND,

PYRALIDH#, SUBFAMILY HYPSOTROPIN&. 108

(60) *SALURIA MAGNESIELLA.

Saluria magnesiella Rag. Nouv. Gen. p. 44 (1888); id. Rom.
Mem. vii. p.360, pl. 37. f. 18; Staud. Cat. Lep, pal. 11 p. 13

W. TURKESTAN.

b’. Antenne of male serrate and ciliated.

(61) TSALURIA MESOMELANELLA, sp. n.

@. Head purple-red and black-brown ; thorax purple-red and
whitish ; abdomen whitish suffused with brown, dorsally rufous
towards base; pectus, legs, and ventral surface of abdomen
whitish suffused with red-brown and dark brown. Fore wing
whitish tinged with purplish pink, the costa and termen deep
.purple-crimson, the medial area suffused with black except at
costa, and the costa defined below by black; an antemedial black
point on vein 1 with a purple-pink shade below it on inner
margin; the inner edge of the medial black area obliquely curved
and its outer edge bent inwards below submedian fold; cilia
pale fuscous. Hind wing white tinged with brown especially ab
termen, the cilia with a fine brown line near base.

TRANSVAAL, White R. (Cooke), 12 type. Hap. 28 mm.

(62) SALURIA TRIPARTELLA.

Saluria tripartella Rag. Rom. Mém. vii. p. 359, pl. 39. f. 19
(1901).
TRANSVAAL; Nata; BAsutroLAND.

(63) SALURIA CALLIRHODA,

Powadia callirhoda Turner, Pr. R. Soc. Queensl. xvill. p. 120
(1903).
(QUEENSLAND.

(64) TSALURIA INTERPUNCTELLA, Sp. n.

Head, thorax, and abdomen ochreous suffused with rufous ;
pectus, legs, and ventral surface of abdomen reddish ochreous
mixed with brown, Fore wing ochreous suffused with rufous ;
the veins white defined on each side by fine black-brown streaks,
the submedian fold and vein | defined by fine rufous streaks ;
the costal edge black towards base; an antemedial blaek point on
vein |, a point in lower angle of cell, an oblique postmedial series
in the interspaces, Incurved beiow submedian fold and a terminal
series. Hind wing ochreous with a fine brownish terminal line
except towards tornus.

MASHONALAND (Dobbie), 2 5,2 2 type; TRANSVAAL, Kranspruit
(Janse), 12, Pretoria (Distant), 19. Hap. 20-24 mm.

(65) TSALURIA INSIGNIFICELLA, sp. N.

@. Head and thorax white mixed with red-brown ; abdomen
white tinged with brown, dorsally fulvous yellow towards base.

104 SIR GEORGE HAMPSON ON 'THE

Fore wing whitish tinged with red-brown, especially towards
costa; the costa narrowly white. Hind wing white tinged with
red-brown.

PARAGUAY, Sapucay (foster), 1G type Hap. 18 mm.

Auctorum.
Atascosa yuudricolorella Dyar, Proc. Ent. Soc. Wash. vi. p. 114
(1904)0. Sect? howadia’ (71s Orewa U.S.A., N. Mexico.

Powadia pimella Dyar, J. N. Y. Ent. Soc. xiv. p. 31 (1906).
| U.S.A., Arizona.
Pectinigeria pamponerella Dyar, Proc. Ent. Soc. Wash. x. p. 10%

(OOS) eaten te teks Sheet ae Gee ree cee eee U.S.A., Colorado.
Ollia par vella Dyar, J. N. Y. Ent. Soc. xiv. p. 31 (1906), Seen.
DSO EURUD, a Ne Sess sieehcokt cad Soe ae SNe EE U.S.A., Texas.
Ollia honoponerella Dyar, Proc. Ent. Soc: Wash, x. p: 107
(E908). SSCChs SOlUIO ae. a0 eee U.S.A., Arizona.
Ollia santaritella Dyar, J. N. Y. Ent. Soc. xii. pp. 107-8 (1904).
PSCC SS COL UTIL sie oie Reruietc rae 4 ore ape nectar ee are eee U.S.A., Arizona.
Poujadia leuconeura Turner, Pr. R. Soc. Queensl. xxiv. p. 118
(19U3). Gseetaramalia o. 1c eee N. Australia.
Pectiniyeria violodis Dyar, Pr. U.S. Nat. Mus. xlvii. p. 347
(UG1'S) dnc bigPencesanele secs ene nenan a ren hee ee PANAMA.
Powjadia cyttarella Dyav, Pr. U.S. Nat. Mus. xlvii. p. 347
(LD V8) a sak vtse cseceveuis bleny Canker suchen a ieetr et CL Tt eee PANAMA.

Genus PROPHTASIA.
Type.
Prophtasia Rug. Ann. Soc. Ent. Fr. 1887, p. 259... platycerella.

Proboscis aborted and minute; paipi typically obliquely up-
turned, the 3rd joint porrect; maxillary palpi filiform; frons
smooth, with shght tuft of scales; antenne of male typically
laminate, the shaft slightly curved at base and without distinct
ridge of scales. Fore wing long and narrow, the apex rounded,
the termen evenly curved ; veins 2 and 3 typically from angle of
cell; 4, 5 stalked; 6 from below upper angle; 8, 9 stalked ; 10,
ll from cell. Hind wing with vein 2 from well before angle
of cell; 3 and 5 from angle, typically approximated for some
distance, 4 absent ; 6, 7 shortly stalked; 8 typically not anasto-
mosing with 7.

Secr. I. Fore wing with veins 2, 3 from a point at angle of cell; hind wing with
vein 8 not anastomosing with 7.

(1) PRopHTASIA PLATYCERELLA.

Prophtasia platycerella Rag. Ann. Soc. Ent. Fr. 1887, p. 259 ;
id. Rom. Mém. viii. p. 252, pl. 37. f. 8; Staud. Cat. Lep. pal.
Isao snlids

ARMENIA.

PYRALID, SUBFAMILY HYPSOTROPIN#. 105

Sect. II. Fore wing with vein 2 from well before angle of cell, 3 from before angle ;
hind wing with veins 3 and 5 not approximated towards base.

A. Hind wing with vein 8 anastomosing with 7; palpi obliquely upturned and
hhardly reaching to vertex of head, the maxillary palpi filiform ; antennee of
male with the shaft slightly curved at base and without distinct ridge of
scales,

(2) {PROPHTASIA SPHALMATELLA, Sp. .

3. Heal and thorax black-brown tinged with grey ; abdomen
brownish ochreous, dorsally fulvous 9 yellow towards base, ventrally
grey-brown. ove wing black- brown mixed with some grey,
especially on inner area; a pure white fascia on costal area,
leaving the costa black-brown and not reaching the apex; an
indistinct diffused dark antemedial spot on vein 1 and oblique
postmedial line. Hind wing whitish suffused with brown; a
darker terminal line and white line at base of cilia.

MASHONALAND, Salisbury (Marshall), 1d type. Hxp. 28 mm.

(3) TPROPATASIA EPITEUXIS, sp. n.

3. Head and thorax glossy black-brown ; abdomen grey suf-
fused with brown; palpi white below towards base ; pectius, legs,
and ventral surface of abdomen black-brown, the femora andl
tibie streaked with white. Fore wing glossy black-brown, the
inner area tinged with grey; a pure white costal fascia, narrow-
ing to apex; an indistinct diffused dark antemedial line from cell
to inner margin and oblique postmedial line. Hind wing grey
suffused with fuscous; a slight punctiform dark terminal line and
white line at base of cila.

Burma, Hsipaw (de Nicéville), 25 type. Hap. 24-28 mm.

(4) TPROPHTASIA GLAUCOPHA, Sp. 0.

Head and thorax grey-brown tinged with white; abdomen
white suffused with grey-brown ; palpi white in front except
towards tips ; peetus, legs, and ventral surface of abdomen white
mixed with some brown. Fore wing pale grey-brown, black-
brown towards the rather broad white costal fascia which does
not narrow to apex. Hind wing creamy white slightly tinged
with brown.

Kasumir (Pilcher), 1 6 3; Punsas, Kangra (Dudgeon), 1 3,
Moghal Sarai (Betton), 1 3; Brnean, Oudh (Pilcher), 1 ¢ ; Sinp,
Karachi reo): 19; Manpras, Belganm (Watson), lg type.
Hap. 18-22 mm.

(5) fPROPHTASIA AMPHICHEA, Sp. N.

Head and thorax grey-brown nuxed with white; abdomen
white tinged with brown; pectus and legs white mixed with
brown. Fore wing pale purplish brown mixed with white, the
costal area broadly white with slight brown irroration ; indistinct
diffused dark curved ante- and postmedial lines except towards

106 SIR GEORGE HAMPSON ON THE

”

costa ; two slight blackish discoidal spots. Hind wing white
tinged with brown ; a slight dark spot at upper angle of cell and
fine terminal line; cilia whiter.

SreRRA Leone (Clements), 1 ¢,49; N. Nigeria, Minna (Macfie),
1d type, Zungeru (Simpson), 12. Hap. 16-20 mm.

B. Hind wing with vein 8 not anastomosing with 7.
a. Palpi of male obliquely upturned to above vertex of head and hollowed out
to receive the brush-like maxillary palpi, the 3rd joint porrect ; antennze
serrate with sinus at base of shaft containing a large ridge of scales.

(6) FPROPHTASIA PYROSTROTA, Sp. n.

3. Head and thorax creamy white suffused with rufous ;
abdomen, pectus, and legs creamy white. Fore wing ochreous
white irrorated with fiery red, the costal fascia only defined by a
red-brown fascia below it from base to apex ; minute antemedial
black stveaks on median nervure and vein 1 and an oblique post-
medial series; two slight black discoidal spots ; a terminal series
of black points. Hind wing creamy white.

: PungaB, Kangra Valley, 4500' (Dudgeon), 1 ¢ type. Hap.
6 mm.

5. Palpi of female downcurved and extending about three times length of head.

(7) PROPHTASIA BISTRIATELLA.

Cayuga bistriatella Hulst, Trans. Am. Ent. Soc. xvii. p. 209
(1890); Dyar, Cat. Lep. N. Am. p. 438.

Peoria discostrigella Dyar, Proc. Ent. Soc. Wash. vi. p. 115
(1904).

U.S,A., N. Mexico, Arizona.

Genus AURORA.

Type.
Aurora Rag. N. Am. Phyenp, 18 (183i) en .. longipalpalla.

Proboscis aborted and minute; palpi porrect, extending about
four times length of head, thickly fringed with hair, the 2nd
joint slightly downcurved ; maxillary palpi slight and filiform ;
frons sinooth, with long pointed tuft of hair; autenne of female
minutely ciliated. Fore wing long and narrow, the apex rounded,
the termen obliquely curved ; vein 3 from elose to angle of cell ;
4,5 strongly stalked; 6 from below upper angle ; 8,9, 10 stalked ;
11 from cell. Hind wing with vein 2 from close to angle of cell ;
3 and 5 stalked, 4 absent; 6, 7 shortly stalked ; 8 anastomosing
with 7.

+AURORA LONGIPALPELLA.

Aurora longipalpella Rag. N. Am. Phye. p. 18 (1887); id. Rom.
Mém. viii. p. 337, pl. 44. f. 2; Dyar, Cat. Lep. N. Am. p. 437.

U.S.A.

PYRALIDH, SUBFAMILY HYPSOTROPIN. 107

Genus FosstFRONTIA.
7 . . Y .4 eee Cc = Type.
Fossifrontia Hmpsn. Rom. Mem. viii. p. 388
GOO De is ec nctaet ns. make Sines cebsatease a Ne othe leuconeurella.

Proboscis absent; palpi upturned to about middle of frons,
thickly scaled and hollowed out to receive the brush-like maxillary
palpi; frons with truncate conical prominence hollowed out in
front ; antenne of male uniserrate and ciliated, the shaft with
large sinus and ridge of scales at base; fore femora with tuft of
hair above, the mid tibie fringed with long hair on outer side.
Fore wing narrow, the apex rounded, the termen evenly curved ;
vein 3 from before angle of cell; 4.5 stalked; 6 from below
upper angle; 8, 9, 10 stalked; 11 from cell; the male with small
fold on underside from medial part of costa fringed with large
seales and with some androconia below it above, in and below
the cell. Hind wing with vein 2 from towards angle of cell ;
3 and 5 from angle and closely approximated for some distance,
4 absent; 6, 7 shortly stalked ; 8 closely approximated to 7 but
not anastomosing with it.

FOssIFRONTIA LEUCONEURELLA.

Fossifrontia leuconeurella Hmpsn. Rom. Mem. vii. p. 339,
pl. 52. f. 19 (1901).

QUEENSLAND, Cooktown, Cedar Bay.

Genus COMMOTRIA.

Type.
Commotria Berg, An. Soc. Arg. xix. p. 278 is
(USS ON Gre tekk Beth ad tS aottapustinn ans ded invenustella.
Mangala Rag. Nouv. Gen. p. 41 (1888)......... crassiscapella.
Tolima Rag. Nouv. Gen. p. 41 (1888) ......... oberthuri.
Altoona Hulst, Ent. Am. iv. p. 116 (1888) ... opacella.

Volusia Hulst, Trans. Am. Ent. Soe. xvii.

p. 206 (1890), nee Rob. Desv. Dipt. 1830... roseopennella.”
Trivolusia Dyar, Cat. Lep. N. Am. p. 438

(GIES) i acces aaah ween crete yn ara we crna'n Sinaia nem ae bees roseopennella.

Proboscis absent; palpi downcurved, extending about three
times length of head and thickly scaled ; maxillary palpi shght
and filiform; frons rounded and with short tuft of hair; antenne
of male typically unipectinate, the apical part ciliated, the shaft
with ridge of scales above at base. Fore wing long and narrow,
the apex rounded, the termen obliquely curved; vein 3 from
angle of cell; 4, 5 stalked; 6 from below upper angle; 8, 9, 10
stalked; 11 from cell. Hind wing with vein 2 from angle of
cell; 3 and 5 strongly stalked, 4 absent; 6, 7 shortly stalked ;
8 anastomosing with 7.

108 SIR GEORGE HAMPSON ON THE

Srct. I. (Commotria.) Antenne of male unipectinate, the apical part ciliated.
(1) CoMMoTRIA MESIELLA, sp. D.

Head and thorax purple-pink mixed with some whitish ; abdo-
men pale fulvous yellow, ochreous white at base and extremity ;
pectus, legs, and ventral surface of abdomen whitish tinged with
purple. Fore wing purple-pink, the veins white defined on each
side by fine brown streaks, the white on median nervure stronger
and more strongly defined by black-brown below; a minute black-
brown spot at upper angle of cell and point at lower angle. Hind
wing whitish tinged with ochreous brown.

Br. C. Arrica, Mt. Mlanje (Weave), 93, 3 2 type. Hap.

20-22 mm.

(2) CoMMOTRIA LATICOSTELLA.

Commotria laticostella Hmpsn. Rom. Mem. vii. p. 3438, pl. 52.
f..14.(1901).

Brazit, Amazons.

(3) COMMOTRIA INVENUSTELLA.

Commotria invenustella Berg, An. Soc. Arg. xix. p. 278
(1885); Rag. Rom. Mém. vii. p. 343, pl. 36. £. 24.

S. Brazin; ARGENTINA.

(4) ComMMorRIA ARRHABDELLA, sp. 0.

®. Head and thorax pale red-brown ; abdomen whitish tinged
with red-brown, dorsally fulvous yellow towards base; pectus.
and legs whitish tinged with red-brown. Fore wing pale flesh-
red slightly irrorated with dark scales; a terminal series of slight
dark points.

Perv, R. Pachaya, 192 type. Map. 24 mm.

Sect. I]. (Tolima.) Antenne of male serrate.
(5) *COMMOTRIA ROSEOPENNELLA.

Volusia roseopennella Hulst, Trans. Am. Ent. Soc. xvii. p. 206

(1890); Rag. Rom. Mém. viii. p. 340, pl. 51. f. 3; Dyar, Cat.
Lep. N. Am. p. 438.

U.S-A.; Florida.

(6) *CoMMOYRIA OBERTHURI.

Tolima oberthuiit Rag. Nouv. Gen. p. 41 (1888); id. Rom.
Mem. vin. p. 341, pl. 38. f 4.

COLOMBIA.

(7) *COMMOTRIA OPACELLA.

Anerastia opacella Huist, Ent. Am. ui. p. 138 (1887); Rag.

Rom. Mém. vii. p. 341, pl. 36. f. 23; Dyar, Cat. Lep. N. Am.
p. 438.

U.S.A., Texas.

PYRALIDA, SUBFAMILY HYPSOTROPIN &. 109

(8) TCOMMOTRIA PHYCITELLA.

Tolima phycitella Rag. Nouv. Gen. p. +1 (1888); id. Rom. Mém.
vill. p. 341, pl. 44. f. 17.

GOLD Coe

Sect. III. (Mangala.) Antenne of male laminate and ciliated.

(9) FCOMMOTRIA TRIPARTELLA, Sp. n.

2. Head and thorax purple-pink ; abdomen white aeons
fulvous yellow towards base ; pectus, legs, and ventral surface of
abdomen white tinged with purphsh pink; anal tuft yellow
below. Fore wing with white costal fascia narrowing to apex
leaving the costal edge dark towards base, then pinkish, defined
below by a dark reddish-brown fascia, the inner half. of wing
rose-pink, whiter towards inner margin. Hind wing white.

N. Cura, Pekin (/ughes), 1 9 type. Hap. 20 mm.

(10) }CoMMOrRIA NEURIAS, Sp. n.

Head and thorax white suffused with red-brown ; abdomen
white, dorsally tinged with fulvous yellow towards base ; pectus
and legs white tinged with brown. Fore wing white tinged with
reddish brown, the veins white defined on each side by slight
dark brown streaks; the costal area defined below by a brown
shade; obliquely placed almost medial dark brown points in
submedian fold and on vein 1; a dark point just above lower
angle of cell; obliquely placed postmedial dark poimts on veins 6
to 2 and in submedian fold and-+a point nearer the termen
on vein 1; a terminal series of black points. Hind wing white
slightly tinged with ochreous brown.

N. NiceriA, Zungeru (Macfie), 1g; Br. C. Arrica, Mt. Mlanje
(Neave), 13,19 type. Hap. 16 mm.

(11) TCOMMOTRIA ERYTHROGRAPTA, Sp. n.

+. Head and thorax white tinged with rufous; abdomen
white tinged with ochreous yellow ; pectus and legs white mixed
with red-brown. Fore wing white tinged with rufous, the veins
and submedian fold white Heaned on aa side by slight rufous
streaks; the costal area defined below by a dark brow streak
and a dark streak below basal half of median nervure; obliquely
placed almost medial dark points helow the cell and on vein 1; a
dark point just above lower angle of cell and obliquely placed
postmedial dark points on veins 6 to 2, Hind wing white with
a slight ochrecus tinge.

Br. C. Africa, Katungas (de Jersey), 1 3 type. Hap. 20 mm,

(12) fCoMMOTRIA RUFIMEDIA, Sp. nh. ,

So. Head and thorax white tinged with red-brown ; abdomen
creamy white. Fore wing white faintly tinged with ee and
irrorated with fuseous; a rufous shade below median nervure ;

110 SIR GEORGE HAMPSON ON THE

a minute antemedial black spot on vein 1 and minute discoidal
spot; an oblique postmedial series of black points on veins 4 to 2
and a point on vein 1; a terminal series of black points. Hind
wing white tinged with ochreous.

Br. C, Arrica, Mt. Mlanje (eave), 1d type. Hap. 16 mm.

(13) TCoMMOTRIA RHODONEURA, Sp. n.

Q. Head and thorax rufous; abdomen whitish tinged with
rufous; palpi whitish below ; fore legs red-brown ; mid and hind
legs whitish tinged with rufous. Fore wing yellowish white, the
veins streaked with purplish pink and the costal area suffused
with purplish pink; a black anteinedial point on vein 1, a
discoidal point and postmedial points on veins 4, 2,1; a terminal
series of black points. Hind wing white tinged with ochreous
rufous.

TRANSVAAL, White R. (Cooke), 19 type. Hap. 20 mm.

(14) fCoMMOTRIA RUFIDELINEATA, Sp. Nn.

Head and thorax whitish suffused with rufous; abdomen
creamy white, dorsally fulvous yellow towards base; pectus and
legs white tinged with red-brown. Fore wing ochreous white,
the veins defined on each side by fine rufous streaks, the costal
area defined below by a slight rufous shade; two antemedial
blackish points on vein | and obliquely placed postmedial points
on veins 4 to 1; a terminal series of black points. Hind wing
ochreous white.

Br. E. Arrica, Nairobi (Anderson), 1 2 type; Br. C. AFrRica,
Mt. Mlanje (Weave), 1d. Hap. 24 mm.

(15) fComMorRIA MIOSTICTA, sp. n.

3. Head and thorax white tinged with rufous; abdomen
creamy white, dorsally fulvous yellow towards base ; pectus, legs,
and ventral surface of abdomen white suffused with brown. Fore
wing white tinged with rufous, the veins white defined on each
didi by fine r ufous streaks; the costal edge blackish towards base ;
the costal area defined below by a slight red-brown shade; a
black point just above lower angle of cell. Hind wing white
slightly tinged with ochreous.

SrERRA LEONE (Clements), 1¢ type. Hap. 22 mm.

(16) *ComMorRIA CRASSISCAPELLA.

Mangala crassiscapella Rag. Nouv. Gen. p. 41 (1888) ; id. Rom.
Mém. viii. p. 342, pl. 37. f. 17.
SUDAN.

(17) }CoMMOTRIA PHENICTIAS, sp. n.

¢. Head and thorax whitish suffused with rufous; abdomen
yellowish white; pectus and legs whitish suffused man brown.
Fore wing white very thickly irrorated with deep purple-pink,

PYRALIDH, SUBFAMILY HYPSOTROPINA. 111

the lower part of cell and the area just beyond it much whiter,
the costa tinged with brown; a small black-brown discoidal spot ;
the veins towards termen with slight brown streaks. Hind wing
yellowish white, the costa tinged with brown towards apex.

N. Nigeria, Zungeru (Macfie), 1 o ; UGanna, Katesa (Betton),
1g type. Hap. 20-22 mm.

(18) TCoMMOTRIA ROSELLA, Sp. Nn.

Head and thorax bright rose-pink ; abdomen ochreous; pectus
whitish; legs and Shaemen pink. Fore wing bright rose-pink
mixed atti some whitish except on costal area, the median
nervure and veins beyond the cell with fine deep pink streaks; a
small deep pink spot on upper discocellular ; cilia fuscous at apex.
Hind wing pale ochreous, the costa and cilia at apex tinged with
pink,

Ab. 1. Fore wing with a dark reddish-brown shade along sub-
costal nervure and thence to apex, the spot on upper discocellular
dark brown.

Br. C. Arrica, Mt. Mlanje (Weave), 4 3, 52 type. Hap.
22-26 mm.

a

(19) fTCoMMOTRIA ALBINERVELLA, sp. ni.

2. Head and thorax pale purplish pink; abdomen ochreous
white, dorsally fulvous yellow towards base; pectus, legs, and
ventral surface of abdomen whitish tinged with pink. Fore wing
pale purple-pink, the veins streaked with white ; the costal area
white tinged with pink, narrowing to apex and defined below by
a slight brown shade; a slight dark point at upper angle of cell.
Hind wing white with a slight ochreous tinge.

Ruopesta, Bulawayo (Hyles), 1 9 type. Exp. 20 mm.

(20) TCOMMOTRIA VENOSELLA, Sp. 1.

¢. Head, thorax, and abdomen whitish suffused with rufous ;
fore femora and tibize black-brown in front. Fore wing whitish
suffused with purplish rufous and slightly irrorated with blackish
in the interspaces ; the veins prominently streaked with white.
Hind wing ochreous whitish, with a slight reddish-brown terminal
line except towards tornus.

Br. C. Arrica, Mt. Mlanje (Neave), 1d type. Hap. 30 mm.

(21) fCoMMOTRIA RHODOCHROA, Sp. n.

3. Head and thorax pale rose-pink ; abdomen ochreous white,
dorsally fulvous yellow towards base; pectus, legs, and ventral
surface of abdomen whitish suffused with red-brown. Fore wing
pale rose-pink, the costal area with a slight red-brown tinge; the
veins streaked witlf white and defined on each side by fine deeper
pink streaks. Hind wing ochreous white.

Naat, Tugela R., Bonds’ Drift (Reynolds), 1 3 type. Hap.
26 mm, |

ay SIR GEORGE HAMPSON ON THE

(22) rCOMMOTRIA CASTANEIPARS, Sp. Nh.

?. Head, thorax, and abdomen deep red-brown ; hind tibiee
white towards base. Fore wing with narrow white costal fascia
narrowing toa point before apex, the area below it deep chestnut-
brown to median nervure and vein 2, the inner area purplish
pink. Hind wing red-brown.

Br. C. Arrica, Mt. Mlanje (Weave), 12 type. Hap. 22 mm.

(23) TCOMMOTRIA PROPH ELLA, Sp. Nn.

2. Head and thorax pale flesh-red ; abdomen ochreous ; palpi
tinged with brown; pectus and legs whitish suffused with brown.
Fore wing pale flesh-red, the costal area broadly suffused with
brown, extending on basal half to median nervure; the veins
beyond the cell finely streaked with brown, Hind wing ochreous
white.

Br. C. Arrica, Mt. Mlanje (Weave), 19 type. Hap. 22 mm.

(24) TCOMMOTRIA PHLEBICELLA, Sp. n.

2. Head and thorax pale flesh-pink mixed with some whitish ;
abdomen whitish, dorsally tinged with fulvous-yellow towards
base ; pectus, legs, and ventral surface of abdomen white tinged
with brown. Fore w ing pale flesh-pink, the costal area broadly
suffused with brown, extending to the median nervure towards
base ; the costal edge and veins finely streaked with white. Hind
wing ochreous white.

MASHONALAND, Salisbury (J/arshall), 292 type. Hap. 26 mm.

(25) tCoMMOTRIA ENERVELLA, Sp. Nn.

3. Head and thorax whitish suffused with pale flesh-red ;
ubdomen ochreous white. Fore wing whitish suffused with pale
purplish pink and the costal half tinged with brown ; the veins,
except on inner area, streaked with white and defined on each
side by fine brown streaks. Hind wing white tinged with brown
except towards base and inner margin.

Formosa, Takow (Wileman), 1 ¢ type. Hap. 18 mm.

Genus SIBOGA.
Type.
Siboga Hmpsn. Rom. Mém. viti. p. 338 (1901)......... Salsella.

Proboscis aborted, and minute; palpi upturned, the 2nd joint
reaching to vertex of head and hollowed out to receive the brush-
like maxillary palpi, the 3rd joint moderate and porrect ; frons
with conical prominence; antenne of male typically serrate and
ciliated, the basal joint elongate, the shaft with double ridge of
scales at base above enclosing wu hollow. Fore wing long and
narrow, the apex rounded, the termen obliquely curved ; vein 3
from close to angle of cell; 4, 5 strongly stalked; 6 from ‘helow
upper angle ; 8, 9 9,10 stalked; 11 from cell, Hind wing with

PYRALID®, SUBFAMILY HYPSOTROPINAE. 1B:

vein 2 from well before angle of cell; 3 and 5 stalked, 4 absent ;
6, 7 shortly stalked; 8 not anastomosing with 7.

Sect. I. Antenne of male unipectinate with moderate branches, the apical part
ciliated.

(1) SrpoGA ALBIMEDIELLA, sp. n.

Head and thorax white tinged with pink and brown; abdomen
ochreous, the first three segments ochreous on dorsum. Fore
wing pale pink, the veins streaked with white; the costal area
tinged with brown; a white fascia from base to termen above
median nervure and vein 5. Hind wing white slightly tinged
with ochreous especially towards termen.

Pungas, Simla, 1 ¢ type. Hap. 26 mm.

Sect. IT. (Siboga.) Antenne of male serrate and ciliated.

(2) *SrpoGA FALSELLA.

Hypsotropha falsella Snell. Midden-Sumatra Lep. p. 82 (1880) ;
Rag. Rom. Mém. viii. p. 338, pl. 38. f. 3.

SUMATRA.

Sect. IIJ. Antenne of male laminate and almost simple.

(3) TSIBOGA DIALEUCELLA, Sp. n.

6. Head and thorax white, suffused with rufous except on
vertex of head and dorsum of thorax; abdomen creamy white,
dorsally fulvous yellow. towards base ; palpi white in front; pectus
and legs white slightly tinged with red-brown. Fore wing white
suffused with rufous; the veins white defined on each side by
fine rufous streaks, the median nervure defined below by a rufous
fascia ; a white fascia through the cell, then narrower along
discal fold to termen. Hind wing ochreous white.

Kasumir, Goorais Valley (Leech), 1d type. Hap. 24 mm.

(4) TSIBOGA ZEAVORA, Sp. N.

@. Head, thorax, and abdomen whitish suffused with rufous,
the last dorsally fulvous yellow towards base. Fore wing whitish
tinged with rufous, the veins defined on each side by streaks
formed of red-brown scales, the cell with two streaks in it; the
inner margin irrorated with red-brown; a terminal series of
prominent black points. Hind wing ochreous white with a
terminal series of brown striz except towards tornus.

Mauay States, 1 9 type. Aap. 30 mm.

Larva feeds on maize.

Proc. Zoou, Soc.—1918, No. VIII. 8

114 SIR GEORGE HAMPSON ON THE

Genus EMATHEUDES.
Type.

EHmatheudes Zell. Stett. Ent. Zeit. 1867, p. 385 ... punctella.

Proboscis aborted and minute; palpi downcurved, extending
about three times length of head and thickly scaled; maxillary
palpi minute and filiform; frons with large tuft of scales;
antenne of male minutely serrate and ciliated. Fore wing
narrow, the apex rounded, the termen evenly curved; vein 3
from close to angle of cell; 4,5 stalked; 6 from below upper
angle; 8, 9, 10 stalked; 11 from cell. Hind wing with vein 2
from well before angle of cell; 3 and 5 stalked, 4 absent; 6, 7
shortly stalked ; 8 not anastomosing with 7.

(1) EmaTHEUDES PUNCTELLA.

Chilo punctella Treit. Schmett. Kur. ix. 2, p. 268 (1833); Dup.
Lép. Fr. pl. 273. f. 4; Herr.-Schiff. Hur. Schmett. iv. p. 108,
Tin. f. 85; Rag. Rom. Mém. vii. p. 333; Staud. Cat. Lep. pal.
li. p. 13, .

S. France; Spain; Corsica; Itaty; Sicity; Morocvwo ;
GREECE; TURKEY ; Cyprus; AstA MINoR; SyRta.

(2) *EMATHEUDES PSEUDOPUNCTELLA.

Kmatheudes pseudopunctella ate Nouv. Gen. p. 40 (1888) ; ale!
Rom. Mém. viii. p. 334, pl. 36. f. 22; Staud. Cat. ion pal. 1: pels,
_ Sykra, Probably an aberration of 4’, punctella.

(3) EMATHEUDES STRAMINELLA.
EKmatheudes straminella Snell, Tijd. v. Ent. 1872, p. 107, pl. 8.
es ae

Gampia; SrerrA Leones; Ancora; Br. C. Arrica; Porru-
GUESE KE. AFRICA.

(4) PEMATHEUDES LENTISTRIGALIS.

Eimmatheudes lentistr igalis Hmpsn. Tr. Zool. Soc. xix. p. 134,
pl. iv. f. 65 (1909).

Goutp Coast; N. Nigeria; Br. EK. ArricaA; Ucanpa; Br. C.
AFRICA.
(5) EMATHEUDES PALEATELLA.

Ematheudes puleatella kag. Nouv. Gen. p. 40 (1888); id. Rom.
Mém. viii. p. 334, pl. 36. f. 20.

Br. E. Arrica; Ucanpa; Br. C. Arrica; MAsHONALAND;
TRANSVAAL; NaraL; BASUTOLAND; CaprE CoLony.
(6) KMATHEUDES TUNESIELLA.

Ematheudes tunesiella Rag. Iris, v. p. 298*(1892); id. Rom.
Mém. vii. p. 335, pl. 42. f. 22 ; Staud. Cat. Lep. pal. ii. p. 13,

S, Ivany ; TUNIS; SYRIA; W. TURKESTAN,

PYRALIDA, SUBFAMILY HYPSOTROPIN®. 115

(7) EMATHEUDES CRASSINOTELLA.

Ematheudes crassinotella Rag. Nouv. Gen. p. 41 (1888); id.
Rom. Mém. viii. p. 335, pl. 35. f£. 26.

ZANZIBAR; Br. EK. ArricA; MASHONALAND; NATAL.

(8) *KMATHEUDES VARICELLA.

Ematheudes varicella Rag. Ann. Soc, Ent. Fr. 1887, p. 258 ;
id. Rom. Mém. viii. p. 336, pl. 35. f. 27; Staud. Cat. Lep. pal.
Me ps le.

ARMENIA; W. TURKESTAN.

(9) * RMATHEUDES VITELLINELLA.

Ematheudes vitellinella Rag. Ann. Soc. Ent. Fr. 1887, p. 258;
id. Rom. Mém. viii. p. 336, pl. 49. f. 22; Staud. Cat. Lep. pal.
top. 13.

Asta Minor, Georgia.

(10) *EMATHEUDES EUCHLYTELLA.
Ematheudes euchlytella Rag. Nouv. Gen. p. 41 (1888); id.
Rom. Mém. viii. p. 387, pl. 36. f. 21.

ARGENTINA.

Genus BraFra,
: Type.
Biafra Rag. Nouv. Gen. p. 40 (1888)............... concinnella.

Proboscis aborted and minute; palpi downcurved, extending
about three times length of head and moderately scaled; maxil-
lary palpi slightly dilated with scales ; frons smooth, with large
pointed tuft of hair; antenne of male minutely serrate and
ciliated, the basal joint rather long, the shaft with double ridge
of scales at base enclosing a hollow, Fore wing very narrow,
the apex rounded, the termen obliquely curved ; vein 3 from near
angle of cell; 4, 5 separate ; 6 from below upper angle; 8, 9, 10
stalked ; 11 from cell. Hind wing with vein 2 from near angle
of cell; 3 and 5 stalked, 4 absent; 6, 7 shortly stalked; 8 anas-
tomosing with 7.

(1) BrarRA CONCINNELLA,

Biafra concinnella Rag. Nouv. Gen. p. 40 (1888); id. Rom.
Mém. viii. p. 330, pl. 38. f. 2.

N. Nicgerta; Br. C. Arricad; MASHONALAND; TRANSVAAL;
NATAL.

(2) TBIAFRA RHODINELLA. '
Biafra rhodinella Rag. Nouv. Gen. p. 40 (1888); id. Rom.
Mém. viii. p. 381, pl. 44. f. 20.

Goutp Coast; N. Nigeria; MAsHoNALAND; TRANSVAAL.
&*

116 SIR GEORGE HAMPSON ON THE

Genus ErHIorRopA, nov.

Type, #. pyromerella.

Proboscis aborted and minute; palpi porrect and _ slightly
downcurved, extending about twice the length of head, the 2nd
joint fringed with rough scales below, the 3rd moderate ; maxil-
lary palpi strongly dilated with scales; frons smooth, with
pointed tuft of hair above; antennz of male somewhat laminate
and minutely ciliated, the basal joint long, the shaft with ridge
of scales above at base. Fore wing long and very narrow, the
apex rounded, the termen obliquely curved; veins 3, 4 stalked
on one side of the specimen from a point on the other; 5 sepa-
rate; 6 from below upper angle; 8, 10, 11 stalked, 9 absent.
Hind wing with vein 2 from well before angle of cell; 3 and 5.
from angle and approximated for some distance, 4 absent; 6, 7
shortly stalked ; 8 anastomosing with 7.

KTHIOTROPA PYROMERELLA, Sp. N.

3. Head, tegule, and base of patagia fiery red, the rest
of thorax brownish ochreous; abdomen ochreous white, dorsally
fulvous yellow towards base; pectus, legs, and ventral surface of
abdomen whitish suffused with brown. Fore wing with whitish
costal fascia narrowing to apex, the costal edge fiery red to
beyond middle, defined below by a black-brown fascia; the rest
of wing fiery red. Hind wing white tinged with ochreous
brown.

N. Niceria, Akassa (Lugard), 1 3 type. Hap. 22 mm.

Genus BAPTOTROPA, nov.

Type, &. tricolorella.

Proboscis aborted and minute; palpi porrect and_ slightly
downcurved, extending about three times length of head, the
2nd joint fringed with rough scales above, the 3rd moderate ;
maxillary palpi minute and filiform; frons smooth and rounded,
without tuft of hair; antenne of male strongly uniserrate, the
basal joint long, the shaft with double ridge of scales at base
enclosing a hollow. Fore wing narrow, the apex rounded, the
termen evenly curved; vein 3 from before angle of cell; 4, 5
from angle; 6 from below upper angle; 8, 9, 10 stalked; 11
from cell. Hind wing with vein 2 from well before angle of
cell; 3 and 5 from angle, 4 absent; 6, 7 shortly stalked; 8
anastomosing with 7.

+BAPTOTROPA TRICOLORELLA.

Patna tricolorella Hmpsn. J. Bomb. N. H. Soc. xii, p. 308
(1899); Rag. Rom. Mém. vii. p. 340, pl. 52. f. 15.

Assam, Khasis.

PYRALIDH, SUBFAMILY HYPSOTROPIN&. lays

Genus PATNA.
Type.

laing Rag, Nouv: Gen. p. 39 (1888).....:......40 eboricostella.

Proboscis absent; palpi porrect and almost straight, extending
about three times length of head, the 2nd joint slightly fringed
with hair above towards extremity, the 3rd moderate ; maxillary
palpi small and filiform; frons smooth and rounded, without
tuft of hair; antenne of female almost simple. Fore wing
rather narrow, the costa typically almost straight, the apex
rounded, the termen evenly curved; vein 3 from near angle of
cell; 4,5 from angle.and approximated for a short distance; 6
from below upper angle; 8, 9, 10 stalked; 11 from cell. Hind
wing with vein 2 from well before angle of cell; 3 and 5 from
angle, 4 absent; 6, 7 from upper angle; 8 not anastomosing
with 7.

(1) PaTNA EBORICOSTELLA.

Patna eboricostella Rag. Nouv. Gen. p. 39 (1888); Hmpsn.
Moths Ind. iv. p. 57; Rag. Rom. es Villueps sa0, ple od: te 1.

SIKHIM; BHUTAN.

(2) fPATNA VENATELLA, sp. n,

Q. Head and thorax pale rufous ; palpi whitish except above ;
pectus, legs, and abdomen whitish tinged with red-brown. Fore
wing white, the veins, discal fold in the cell and the submedian
fold defined by fine purplish-pink streaks; a minute dark brown
spot just above lower angle of cell; a terminal series of black
pomts. Hind wing white slightly tinged with ochreous.

Br. E. Arrica, Tagas (Letion), 1 9, Aios (Betton), 1 2 type.
Hep. 32 mm.

(3) TPATNA BRUNNEICOSTELLA, Sp. n.

@. Head and thorax pale flesh-pink; abdomen ochreous
white; palpi, pectus, legs, and ventral surface of abdomen
whitish suffused with +ed-brown. Fore wing white, the costal
area tinged with brown, the area below the cell and vein 2
suffused with pale pink; the veins white, those beyond the cell
defined on each side by fine pink streaks, the median nervure and
vein 2 defined below by stronger streaks; a dark point in lower
angle of cell. Hind wing ochveous white.

TRANSVAAL, White R. (Cooke), 1 2 type. Hap. 32 mm.

Genus MEGALOPHOTA, nov.

Type, J. leonella. |

Proboscis absent; palpi obliquely upturned, the 2nd joint
reaching to above vertex of head, dilated and hollowed out to
receive the brush-like maxillary palpi, the 3rd minute ; frons

118 SIR GEORGE HAMPSON ON THE

with long truncate conical prominence; antenne of male pecti-
nate with rather long uniseriate branches to near apex, the shaft
with large sinus at base containing a double ridge of scales
enclosing a hollow. Fore wing rather long and narrow, the apex
rounded, the termen evenly curved; vein 2 from well before
angle of cell; 3 from before angle; 4, 5 from angle; 6 from
below upper angle; 8, 9 stalked; 10, 11 from cell. Hind wing
with vein 2 from well before angle of cell; 3 and 5 stalked,
4 absent; 6, 7 shortly stalked ; 8 not anastomosing with 7.

MEGALOPHOTA LEONELLA, Sp. N.

3. Head and thorax ochreous white with a slight red-brown
tinge on shoulders, the antennal tufts black on inner side;
abdomen ochreous ; pectus and legs ochreous white tinged with
red-brown. Fore wing ochreous white irrorated with brown,
the costal area slightly irrorated to near apex, Hind wing
ochreous white.

SrERRA LEONE (Dudgeon), 1 3 type. Hap. 20 mm.

Genus MARTIA.

Type.

Martia Rag, N. Am. Phyc. p. 18 (1887) 2... 2.42.4 arizonella.
Urula Hulst, Can. Ent, xxxu. p. 175 (1900) ...... arizonella.

Proboscis absent ; palpi with the 2nd joint porrect, extending
about twice the length of head, the 3rd rather oblique, long,
slender, and somewhat acute at extremity; maxillary palpi
slight and filiform; frons with large truncate conical prominence;
antenne of male minutely serrate and with fascicles of long cilia.
Fore wing rather narrow, the apex rounded, the termen evenly
curved; vein 2 from long before angle of cell; 3 from before
angle; 5 from above angle; 6 from below upper angle; 8, 9
_ stalked; 10, 11 from cell. Hind wing with. vein 2 from long
before angle of cell; 3 and 5 from angle, approximated for a
short distance, 4 absent; 6, 7 from upper angle; 8 not anasto-
mosing with 7.

MARIA ARIZONELLA.

Martia arizonella Rag. N. Am. Phye. p. 18 (1887); id. Rom.
Mém. viii. p. 367, pl. 38. f. 20; Dyar, Cat. Lep. N. Am. p. 439, .
Orula incongruela Hulst, Can. Ent. xxxil. p. 175 (1900);
Dyar, Cat. Lep. N. Am. p. 437.

U.S.A., Colorado, Arizona.
Genus D1scoFRONTIA.

Type.
Discofrontia Hmpsn. Rom. Mém. viii. p. 350 (1901). normella.

Proboscis aborted and minute; palpi upturned, the 2nd joint
reaching to vertex of head and moderately scaled and flattened,

PYRALIDA, SUBFAMILY HYPSOTROPIN#. aS,

the 3rd moderate; maxillary palpi slight and filiform; frons
broad, with a disk of concentric white scales converging to middle;
antenne of male strongly serrate, with a large sinus and ridge of
scales at base of shaft. Fore wing rather narrow, the apex
rounded, the termen evenly curved; vein 2 from well before
angle of cell; 3 from before angle; 4,5 from angle: 6 from
below pee angle; 8, 9 stalked; 10, 11 from cell. Hind wing
with vein eon betare angle of cal: 3 and 5 from angle,
4 absent ; 6, 7 from upper angle ; ; 8 not anastomosing with 7.

* DISCOFRONTIA NORMELLA.

Iiscofrontia normella Hmpsn. Rom. Mém, viii. p. 350, pl. 52.
f. 20 (1901).
NAtAu.

Genus CRITONIA,

Type.
Critonia Rag. Bull. Soc. Ent. Fr. 1890, p. cexiv. subconcinnella.
Singhalia Himpsn. J. Bomb. Nat. Hist. Soc. xii.

p. 809 (1898); id. Rom. Mém. vu. p. 351
(LGA) Ee erences a neers ree ener or ee eae mr reme aereT: sarcoglauca.

Proboscis aborted and minute ; palpi typically downcurved,
extending about three times length of head and moderately
scaled; maxillary palpi small and filiform; frons smooth and
with slight tuft of hair; antenne of male typically serrate and
fasciculate, the shaft with sinus at base containing a large ridge
of scales. Fore wing rather long and narrow, fer apex rounded,
the termen evenly curved; vein 3 from before angle of cell;
4, 5 from just above angle; 6 from below upper angle - 8, 9
stalked ; 10, 11 from cell, 10 approximated to 8, 9. Head: wing
with veins 3 and 5 from angle of cell and approximated for a
short distance, 4 absent; 6, 7 shortly stalked ; 8 not anastomos-
ing with 7,

Y)

Sect. I. Palpi of male obliquely upturned, the 2nd joint hollowed out to receive
the brush-like maxillary palpi, the 3rd short; antenna laminate, with large
sinus and ridge of scales at base of shaft.

(1) fCRITONIA PHAONEURA, Sp. n.

3. Head and thorax whitish suffused with red-brown ; abdo-
men ochreous white, dorsally fulvous yellow towards base ;
pectus and legs whitish tinged with brown. lore wing whitish
tinged with red- brown, the veins except on inner area ehleck ct
Hedned on each side by fine white streaks ; the costal area white,
narrowing to apex and defined below by a red-brown shade, the
costal edge brown to beyond middle and the veins on costal area
finely streaked with black; a minute antemedial black spot on
vein ],a point at middle of submedian fold and minute post-
medial streaks above vein 2 and on vein 1 ; cilia flesh-white with

120 SIR GEORGE HAMPSON ON THE

series of minute black streaks near base except towards tornus.
Hind wing whitish tinged with brown especially on costal area
and at termen.

Formosa, Banshorio (Wileman), 1 d type. Hap. 18 mm.

(2) TCRITONIA PROMELENA.

Critonia promelena Hmpsn. J. Bomb. Nat. Hist. Soc. xii.
p- 809 (1898); id. Rom. Mém. viii. p. 366, pl. 51. f. 23.

SIKHIM.

(3) TCRITONIA PURPUREOTINCTA.

Critonia purpureotincta Hmpsn. Moths Ind. iv. p. 61 (1896) ;
id. Rom. Mém. viii. p. 365, pl. 51. f. 22.

SIKHIM; Buurtdn.

(4) TCRITONIA HOLORHODA.

Critonia holorhoda Hmpsn. J. Bomb. Nat. Hist. Soc. XVill.
p. 259 (1908).
CEYLON.

Sect. II. Maxillary palpi of male filiform.

A. (Critonia). Antenne of male serrate, with larger sinus and ridge of scales
at base of shaft; palpi downcurved and about three times length of head.

(5) *CRITONIA SUBCONCINNELLA.

Critonia subconcinnella Rag. Bull. Soc. Ent. Fr. 1890, p. cexiv ;
id. Rom. Mém. vii. p. 365, pl. 6. f. 20.

BuRMA.

(6) TCRITONIA LEUCOPLEURA, Sp. n.

3. Head and thorax whitish suffused with purplish pink;
abdomen whitish suffused with ochreous brown ; antenne with
the tuft blackish; palpi, pectus, and legs whitish .tinged with
brown. Fore wing whitish suffused with purplish pink, the
veins streaked with blackish, vein 1 only towards termen; the
costal area pure white, narrowing to apex and defined below by
a blackish shade; the costal edge dark to beyond middle and the
interspaces of terminal area with slight dark streaks except
towards tornus. Hind wing whitish, the costal area broadly
suffused with brown.

BasutToLanD, Maseru (Crawshay), 1 3d type. Hap. 26 mm.

(7) TCRITONIA ROSEISTRIGELLA.

Critonia roseistrigella Hmpsn. Moths Ind. iv. p. 61 (1896); id.
Rom. Mém. viii. p. 365, pl. 51. f. 21.

Mapras, Nilgiris ; Puinipprnes, Luzon.

PYRALIDH, SUBFAMILY HYPSOTROPINA. Al

‘ (8) CRITONIA OCHRACEALIS.

Critonia ochracealis Hmpsn. J. Bomb. Nat. Hist. Soc. xxi.
p- 1251 (1912).

PungaB, Kangra; MaAnpras, Nilgiris.

(9) *CRITONIA HILGERTI.
Pectinigeria hilgerta Roths. Nov. Zool. xxii. p. 236.

ALGERIA.

B. Antenna of male laminate and without sinus and ridge of scales at base of

shaft.
a. (Singhalia.) Palpi of male with the 2nd joint obliquely upturned to vertex
of head and thickly scaled, the 8rd porrect, long and blunt.

(10) TCRIvONTA SARCOGLAUCA.

Critonia sarcoglauca Hmpsn. Moths Ind. iv. p. 60 (1896); id.
Rom. Mém. viii. p. 351, pl. 51. f. 24.

CEYLON.

b. Palpi of male downcurved and extending about three times length of head.
(11) CrrroniA RHODESSA.
Saluria rhodesso Turner, Pr. R. Soc. Queensl. xviii. p. 120
(1903).
QUEENSLAND.

(12) TCRITONIA SARCOIDA, sp. N.

Head and thorax purplish pink, the head and shoulders
tinged with brown; abdomen ochreous white, dorsally fulvous
yellow towards base; pectus, legs, and ventral surface of abdo-
men ochreous tinged with brown. Fore wing purplish pink, the
costal area tinged with brown; the veins, discal fold in the cell,
and submedian fold white defined on each side by fine purplish-
pink streaks, the median nervure rather more strongly streaked
with white and with a brown streak below it. Hind wing
ochreous white, the costal area tinged with brown.

Br. HE. Arrica, Njora (Cholmley), 1 ¢; Br. C. Arrica,
Mt. Mlanje (Veave), 3 9 ; Porrucurse E. Arrica, Kola Valley
(Veave), 3 9 ; MASHONALAND, Salisbury (J/arshall), 1 2 ; TRANs-
VAAL (Pead), 1 3 type. Hap. 28-34 mim.

Genus MoNnocTENOCERA.
Type.
Monoctenocera Hmpsn. J. Bomb. Nat. Hist. Soc. xii.
p- 310 (1898) ; id. Rom. Mém. viii. p. 311 (1901). brachiella.

Proboscis aborted and minute ; palpi obliquely upturned, the
2nd joint reaching to just above vertex of head and hollowed out
to receive the brush-hke maxillary palpi, the 3rd short and

122 SIR GEORGE HAMPSON ON THE

thickly scaled ; frons smooth, with large tuft-of hair; antenne’
of male unipectinate, typically with very short branches, the
apical part serrate, the shaft with large sinus and ridge of scales
at base; mid and hind tibie typically fringed with long hair.
Fore wing narrow, the apex rounded, the termen evenly curved ;
vein 3 from close to angle of cell; 4, 5 strongly stalked; 6
from below upper angle; 8, 9, 10 stalked; 11 from cell. Hind
wing with vein 3 closely approximated to 4, 5 for some distance ;
4,5 strongly stalked; 6, 7 shortly stalked; 8 anastomosing
with 7.

Seor. I. Antenne of male with the branches long; mid tibize fringed with hair at
base only, the hind tibie at extremity only.

(1) MonocrENOcCERA LEUCANIA.

Catagela leucania Feld. Reis. Nov. pl. 137. f. 13 (1874);
Himpsn. Moths Ind. iv. p. 63; id. Rom. Mem. viii. p. 312, pl. 51.
fee hie

Hab. Manvras, Nilgiris; TRAVANCORE ; CEYLON.

Sect. II. Antenne of male with short branches towards base, then serrate: mid and
hind tibie fringed with long hair throughout.

(2) MonocrenocEra BRACHIELLA.

Polyocha brachiela Hmpsn. J. Bomb. Nat. Hist. Soc. xii.
p- 310 (1898) ; id. Rom. Mém. vil. p. 312, pl. 36. f. 6.
Tab. StKKIM; Bencat, Calcutta; Borneo.

Genus SABORMA.

Type.
Saborma Rag. Nouv. Gen. p. 37 (1887) ............ Jorcipella.

Proboscis aborted, minute; palpi of male upturned to about
vertex of head, slender, typically hollowed out to contain the
brush-like maxillary palpi; frons smooth; antenne of male
typically strongly serrate, with sinus and large double ridge of
scales at base. Fore wing narrow, the apex rounded, the termen
evenly curved ; vein 3 from close to angle of cell; 4, 5 strongly
stalked; 6 from below upper angle; 8, 9, 10 stalked; 11 from
cell. Hind wing with vein 3 approximated for some distance to
4, 5 which ave strongly stalked; 6, 7 shortly stalked; 8 not
anastomosing with 7.

Sect. I. Palpi of male with the 2nd joint hollowed out to receive the brush-lke
maxillary palpi; antenne serrate.
(1) *SABORMA FORCIPELLA.

Saborma forcipella Rag. Nouv. Gen. p. 87 (1888): id. Rom.
Mem. viii. p. 310, pl. 35. f. 22,
SUMATRA.

PYRALIDA, SUBFAMILY HYPSOTROPINA. “123

(2) *SABORMA VICINA.

Anerastia vicina Saalm. Ber. Senck. Ges. 1879, p. 307; id. Lep.
Madag. p. 511; Rag. Rom. Mém. viii. p. 309, pl. 42. f. 21.

MADAGASCAR.

Secor. If. Maxillary palpi of male filiform; antenne laminate and ciliated.

(3) TSABORMA PAPUACOLA, Sp. h.

6. Head and thorax whitish tinged with red-brown ; abdomen
whitish tinged with brown, dorsally fulvous yellow towards base ;
antenne with the tuft blackish on inner side; pectus and legs
whitish tinged with brown. Fore wing white tinged with
ochreous brown, the veins, discal fold in the cell, and the sub-
median fold white defined on each side by streaks formed of
blackish scales; a terminal series of black points. Hind wing
ochreous white with terminal series of dark points and strie.

Durcn N. Guinea, Mimika R. (Wollaston), 3 3 type. Hap.
26-30 mm.

Genus OSACIA.
Type.
Osakia Rag. Rom. Mém. viii. p. 318 (1901) ......... lineolella.

Proboscis aborted and minute; palpi obliquely upturned to
about vertex of head, moderately scaled; maxillary palpi each a
minute brush of scales; frons smooth and rounded ; antenne of
mnale ciliated, the shaft with sinus and double ridge of scales at
base; tibie fringed with hair. Fore wing rather narrow, the
apex rounded, the termen evenly curved ; veins 2 and 3 stalked
from before angle of cell; 4, 5 separate ; 6 from below upper
angle; 8, 9 staiked; 10,11 from cell. Hind wing with vein 3
from angle of cell; 4 and 5 strongly stalked ; 6, 7 strongly stalked ;
8 not anastomosing with 7.

*OSACIA LINEOLELLA.

Osakia lineolella Rag. Rom.’ Mem. viil. p. 319, pl. 43: f. 21
(1901). :

JAPAN.
Genus RAGoNOoTIA.
Type.
Ciris Rag. N. Am. Phyc. p. 17 (1887), nec Grote,
Wiep SO neo erace seme a eee cranuah canted tee hayes dotalis.
Ragonotia Grote, Can. Ent. xx. p. 75 (1888)............ dotalis.

Proboscis aborted and minute; palpi downturned, about three
times length of head and rather broadly fringed with scales
below; maxillary palpi filiform; frons with small rounded pro-
minence with corneous plate below it ; antenne of male ciliated,
the basal joint large. Fore wing long and narrow, the apex
rounded, the termen obliquely curved; vein 2 from towards
angle of cell; 3 and 5 from close to angle; 6 from below upper

124 SIR GEORGE HAMPSON:ON THE

angle; 8, 9 stalked; 10,11 from cell. Hind wing with vein 2
from close to angle of cell; 3 approximated for some distance to
4, 5 which are strongly stalked, or 4 rarely absent; 6, 7 stalked;
8 not anastomosing with 7.

RAGONOTIA DOTALIS.

Anerastia dotalis Hulst, Trans. Am. Ent. Soc. xii. p. 164 (1886) ;
Rag. Rom. Mém. viii. p. 329, pl. 38. f. 19; Dyar, Cat. Lep. N.
Am. p. 437.

Ciris discigerella Rag. N. Am. Phye. p. 17 (1887).

U\S.A., Colorado, Arizona.

Genus PoLyocHa.

Type.
Polyocha Geller, Isis, 1848, p. 876............... sanguinariella.
Polyochedes Chretien, Bull. Soc. Ent. Fr. 1911,
Pabst cette Alen ancien Mee ees eteeee ee stipella.

Proboscis aborted and minute; palpi typically downcurved,
extending about three times length of head, the 2nd joint
moderately scaled, the 3rd rather long and naked; maxillary
palpi dilated with scales; frons smooth, with tuft of scales ;
antenne of male typically laminate and without sinus and ridge
of scales at base. Fore wing long and narrow, the apex rounded,
the termen obliquely curved ; vein 3 from close to angle of cell;
4, 5 strongly stalked; 6 from below upper angle; 8, 9 stalked ;
10, 11 from cell. Hind wing with vein 2 from close to angle of
cell ; 3 approximated for some distance to 4,5 which are strongly
stalked ; 6, 7 shortly stalked; 8 not anastomosing with 7.

Secor. I. Palpi of male obliquely upturned ; antennee serrate, with sinus and ridge of
scales at base of shaft.

(1) TPOLYOCHA PLINTHOCHROA, sp. n.

Head and thorax bright rufous mixed with some ochreous ;
abdomen ochreous, dorsally fulvous yellow towards base; antenn
ochreous. Fore wing bright rufous with a faint purplish gloss,
some ochreous in lower part of cell and below and just beyond
the cell, the veins remaining rufous; a narrow yellowish-white
costal fascia, tapering to a point just before apex. Hind wing
ochreous white, the costal area and termen tinged with red-
brown.

TRANSVAAL, Karina (Cooke), 1 ¢ type, White R. (Cooke), 1 9.
Hap. 26 mm.

(2) fPoLYocHA LEUCOPLEURELLA.

Emmatocera leucopleurella Rag. Nouv. Gen. p. 38 (1888) ; id.
Rom. Mém. viii. p. 317, pl. 44. f. 15.
GouLp Coast; 8. & N. Nigeria; Kasumir; MApRas.

PYRALIDH, SUBFAMILY HYPSOTROPIN®. 125

(3) TPOLYOCHA GENSANALIS.

Emmatocera gensanalis South, Trans. Ent. Soc. 1901, p. 405,
pl. xiv. f. 30.

CoREA.

Sect. IT. Palpi of male downcurved ; antenne without sinus and ridge of scales at
base of shaft.

A. (Polyochodes). Antenne of male pectinate with uniseriate branches.

(4) *PoLyocHA STIPELLA,.
Polyochodes stipella Chretien, Bull. Soc. Ent. Fr. 1911, p. 13.
ALGERIA.

B. (Polyocha). Antenne of male laminate.

(5) TPoLYocHA CINERELLA.

Polyocha cinerella Hmpsn. Moths Ind. iv. p. 62 (1896); id.
Rom. Mém. viii. p. 328, pl. 55. f. 4.

PunsaB; BENGAL.

(6) PoLyocua VENOSA.

Kpischnia venosa Zell. Isis, 1847, p. 31; Herr.-Schiff. Schmett.
Kur. iv. p. 109; Rag. Rom. Mém. viii. p. 327; Staud. Cat. Lep.
pal... p13.

CYPRUS; SYRIA.

(7) PoLYOCHA SANGUINARIELLA.

—Polyocha sanguinariella Zell. Isis, 1848, p. 876; Rag. Rom.
Mem. vili. p, 327, pl. 8; f. 20.
‘Br. C. AFricA ; MASHONALAND; T’RANSVAAL ; MADAGASCAR.

(8) PoLYocHA VESCULELLA.

Polyocha vesculella Rag. Nouv. Gen. p. 39 (1888); id. Rom.
Mem. viil. p. 323, pl. 36. f. 11; Hmpsn. Moths Ind. iv. p. 63.

Manpras, Palni Hills; Travancore.

(9) *PoLYOCHA FLAGRANTELLA.

Polyocha flagrantella Rag. Rom. Mem. viii, p. 323, pl. 44. f. 24
(1901).

MADAGASCAR.

(10) TPoLYOCHA STRIGIVENELLA.

Polyocha strigivenella Hmpsn. J. Bomb. Nat. Hist. Soc. xil.
p. 310 (1898); id. Rom. Mém, viii. p. 322, pl. 51. f. 19.
BuRMA,

126 SIR GEORGE HAMPSON ON THE

(11) *PoLyocHA NEUROPTERELLA.

Polyocha neuropterella Rag. Ann. Soc. Ent. Fr. 1887, p. 258 ;
id. Rom. Mém. viii. p. 322, pl. 35. f. 23; Staud. Cat. Lep. pal. ii.
pe 13.

W. TuRKESTAN.

A P )LYOCHA FOUCARTI.

Polyocha foucarti Rag. Ann. Soc. Ent. Fr. 1887, p. 258; id. Rom.
Mém. viii. p. 322, pl. 35. f. 24; Staud. Cat. Lep. pal. ii. p. 13.

ALGERIA.

(13) PPoLYocHA ACHROMATELLA, Sp. nN.

2. Head and thorax white tinged with ochreous ; abdomen
white, dorsally fulvous yellow towards base; pectus and legs
ochreous white. Fore wing pale ochreous, the veins white, less
distinctly so on costal area. _ Hind wing white.

N.S. Wates, Broken Hill (Lower), 3 9 type. Hxp. 28 mm.

(14) *PoLyocHA DETRITELLA.

Polyocha detritella Rag. Nouv. Gen. p. 39 (1888); id. Rom.
Mém. vill. p. 326, pl. 36. f. 14; Hmpsn. Moths Ind. iv. p. 63.

FU NIAB TA:

(15) PTPoLYOCHA FUSCICOSTELLA, Sp. Nn.

@. Head and thorax glossy fuscous brown ; abdomen fuscous
brown, pale red-brown at sides and extremity ; pectus and legs
pale red-brown, the tarsi fuscous brown. Fore wing pale
red-brown, the costal area broadly glossy fuscous brown and
the inner basal area tinged with fuscous brown. Hind wing
ochreous white, the costal area tinged with brown.

N. Nicrria, Zungeru (Macfie), 2 9 type. Hap. 24 mm.

Genus EMMALOCERA.

Type.
Emmalocera Rag. Nouv. Gen. p. 38 (1888)...... lewcocincta.
Lodiana Rag. Nouv. Gen. p. 38 (1888) ......... umbrivittella.

Papua Rag. Bull. Soc. Ent. Fr. 1889, p. eexx... datilimbella.

Proboscis aborted and minute; palpi of male typically obliquely
upturned to above vertex of head, the 2nd joint hollowed out to
receive the brush-like maxillary palpi, the 3rd short and porrect;
frons smooth, obliquely flattened; antenne of male typically
with short uniseriate branches, the basal joint large, the shaft
with large sinus and ridge of scales at base. Fore wing rather
long and narrow, the apex rounded, the termen evenly curved ;
vein 3 from near angle of cell, 5 from just above angle; 6 from
below upper angle; 8, 9 stalked; 10, 11 from cell, Hind wing

PYRALIDH, SUBFAMILY HYPSOTROPIN &. 127

with vein 2 from well before angle of cell; 3 approximated for
some distance to 4,5 which are strongly stalked; 6, 7 shortly
stalked ; 8 not anastomosing with 7.

Szot. I. Palpi of male upturned, the 2nd joint hollowed out to receive the brush-
like maxillary palpi; antennze with sinus and ridge of scales at base of shaft.

A. Antenne of male with rather long uniseriate branches.

(1) TEMMALOCERA ORNATELLA.

Polyocha ornatella Hmpsn. J. Bomb. Nat. Hist. Moe. xv. p. 21
(1903). .

PUNJAB..

(2) f}EMMALOCERA PULVEREALIS.

Polyocha pulverealis Hmpsn. J. Bomb. Nat. Hist. Soc. xv.
p. 20 (1903).
ASSAM.

(3) TEMMALOCERA ENDOPYRELLA, Sp. n.

3. Head ochreous tinged with purplish pink, the antenne
ochreous; thorax bright purplish pink ; abdomen fulvous yellow ;
pectus and legs ochreous tinged with red-brown. Fore wing
with narrow pure white costal fascia, the rest of wing bright
purplish pink, suffused with red-brown to median neivure and
veins 4. Hind wing ochreous white, the costa shghtly tinged
with brown.

Assam, Khasis, we, type. Hap. 26 mm.

(4) EMMALOCERA LONGIRAMELLA.

Emmalocera longiramella Hmpsn. Rom. Mém. vii. p. 315,
pl. 52. f. 16 (1901).

(QNUEENSLAND.

(5) *EMMALOCERA RADIATELLA.

Hinmatlocera radiatella Umpsn. Rom. Mém. vil. p. 315, pl. 52.
f. 21 (1901).
()UEENSLAND,

(6) TEMMALOCERA ACTINOLEUCA, Sp. Nn.

o. Head and thorax pale purplish pink mixed with white ;
abdomen ensamy white, dorsally fulvous yellow towards base ;
antenne white; frons, palpi, pectus, and legs white suffused with
rufous. Fore wing pale purplish pink, the veins, two streaks in
the cell and one in submedian fold white. Hang wing white
faintly tinged with ochreous.

SIERRA Leone E (Clements), 1 ¢ type. Hap. 24 mm,

128 SIR GEORGE HAMPSON ON THE

(7) TEMMALOCERA DEPRESSELLA.

Melissoblaptes depressella Swinh. P. Z. 8. 1885, p. 876, pl. 57.
f.5; Hmpsn. Moths Ind. iv. p. 63; id. Rom. Mém. viii. p. 324,

[Se eLiyuiee
Polyocha saccharella Dudgeon, J. Bomb. Nat. Hist. Soe. xvi.

p. 405 (1905).
ADEN ; PunsJAB; BENGAL; BomBay. The larva feed on the
roots of sugar-cane.

(8) TEMMALOCERA STRIGICOSTELLA.

Polyocha strigicostella Hmpsn. P. Z. S. 1896, p. 270; id. Rom.
Mem, vill..p.-315, pl. 31. £. 18.

ADEN.

B. (Emmalocera). Antenne of male with short uniseriate branches. ‘

(9) TEMMALOCERA LEUCOCINCTA.

Crambus leucocinctus Wik. xxvii. 169 (1863); Hmpsn. Rom.
Mém. viii. p. 316, pl. 36. f. 9.
Hmmalocera crenatella Rag. Nouv. Gen, p. 38 (1888).

SINGAPORE; BORNEO; PHILIPPINES.

C. (Papua). Antennz of male laminate, serrate towards base.

(10) FEMMALOCERA SANGUIFUSALIS,
Polyocha sangwfusaiis Hmpsn. P. Z. 8. 1910, p. 493, pl. xl.
eae |
N. Ruopesia.

(11) PEMMALOCERA AURIFUSELLA.

Crambus aurifusellus Wik. xxxv. 1756 (1866); Hmpsn.
Moths Ind. iv. p. 62; id. Rom. Mem. viii. p. 317, pl. 36. f. 18.

Kasumir; Pungap; Bompay; MaAnpRAS.

(12) EMMALOCERA BIFIDELLA.
Polyocha bifidella Wileman, ‘Trans. Ent. Soc. 1911, p. 357,
pl oli 22:

JAPAN.

(13) TEMMALOCERA POLYCHROELLA, Sp. 0.

Head and thorax ochreous mixed with fiery red; abdomen
othreous ; antenne of male with the tuft blackish on inner side ;
palpi, pectus, legs, and ventral surface of abdomen ochreous
tinged with red-brown. Fore wing with narrow creamy white
costal fascia leaving the costal edge red-brown towards base,
defined below by a fiery rufous streak to beyond middle, the area
below it red-brown to median nervure and vein 4; the inner

PYRALIDA, SUBFAMILY HYFSOTROPIN®. 129

half of wing yellow thickly irrorated with fiery red, the terminal
half of inner margin tinged with brown.

W. Arnica (Dudgeon), 1 3,1 9; 8. Nigeria, Mama (Dudgeon),
1 gd type. Hap., g 22, 9 28 mm.

(14) EMMALOCERA UMBRICOSTELLA.

Emmalocera wmbricostella Rag. Nouv. Gen. p. 38 (1888) ; id.
Rom. Mém. viii. p. 316, pl. 36. f. 10; Hmpsn. Moths Ind. iv.
p: 62;

Correa; C. Cuina; Srkuim ; Borneo, Pulo Laut; PHILIPPINES ;
JAVA; FLORES; BALI.

(15) EMMALOCERA LUCIDICOSTELLA.

Emmalocera lucidicostella Rag. Nouv. Gen. p. 38 (1888); id.
Rom. Mém. viii. p. 316, pl. 35. f.20; Hmpsn. Moths Ind. iv. p. 62.

PunsaB; BENGAL; CEYLON; SUMATRA.

(16) EMMALOCERA ANERASTICA,
Nephopteryx anerastica Snell. Veth’s Midden-Sumatra, Lep.
p. 81 (1880); Rag. Rom. Mém. viii. p. 317, pl. 36. f. 8.

SIERRA LEONE; Formosa; PungaB; NIcoBARS; JOHORE;
SELANGOR ; SINGAPORE; SuMATRA; BorNEo, Pulo Laut; PHI.ip-
PINES ; Louis1ApDE Is., St. Aignan I.

(17) EMMALOCERA LATILIMBELLA. ;

Papua latilimbella Rag. Bull. Soc. Ent. Fr. 1889, p. eexx; id.
Rom. Mém. viii. p. 313, pl. 36. f. 7.

Polyocha rhabdota Turner, Pr. R. Soc. Queens}. xviii. p. 122
(1903).

Polyocha achrosta ‘Turner, Pr. R. Soc. Queensl. xviii. p. 122
(1903).

N. GUINEA; QUEENSLAND.

D. Antenne of male laminate, not serrate towards base.

(18) EMMALOCERA LAMINELLA.

Emmalocera laminella Hmpsn. Rom. Mém. viii. p. 318, pl. 51.
£2. (1901).

SreRRA LEONE; Br. E. Arrica; Br. C. AFRICA.

Sect. II. Palpi of male downcurved ; maxillary palpi filiform.
A. Antenne of male with sinus and ridge of scales at base of shaft.
a. Antenne of male with long uniseriate branches.

(19) EmMMALOCERA VARIEGATELLA.
Polyocha variegatella Rag. Nouv. Gen. p. 39 (1888); id. Rom.
Mém. viii. p. 326, pl. 36. f. 16; Hmpsn. Moths Ind. iv. p. 63.
PUNJAB.
Proc. Zoot, Soc.—1918, No. LX. s)

130 SIR GEORGE HAMPSON ON THE

(20) fEMMALOCERA TRICOLORALIS.

Polyocha variegatella Hmpsn. J. Bomb. Nat. Soe. Hist. xii
p. 320 (nec Rag.).

Polyocha tricoloralis Hmpsn. J. Bomb. Nat. Hist. Soc. xv.
p. 20 (1903).

S1kkiM; PHILIPPINES.

(21) EMMALOCERA DIVERSELLA.

Polyocha diversella Hmpsn. J. Bomb. Nat. Hist. Soc. xii.
p. 310 (1898); Rag. Rom. Méin. viii. p. 324, pl.’vi. f. 21.

Manpras, Nilgiris.

(22) *EMMALOCERA COSTELLA.

Polyocha costella Rag. Nov. Gen. p. 39 (1888); id. Rom. Mém.
vill. p. 326, pl. 36. f. 15.

GAMBIA.

(23) EMMALOCERA CREMORICOSTA.

Polyocha cremoricosta Rag. Bull. Soc. Ent. Fr. 1895, p. cii; id.
Rom. Mém. viii. p. 325, pl. 51. f. 9; Staud. Cat. Lep. pal. ii.
pinks:

AstA Minor; Syria.

(24) +EMMALOCERA ERYTHRINELLA,

Polyocha erythrinella Rag. Nouv. Gen. p. 38 (1888); id. Rom.
Mém. viii. p. 323, pl. 44. f. 14.

N. Niageria; ABysstntaA; Br. E. Arrica; Br. C. AFRICA.

(25) *EMMALOCERA CARNATELLA.

Polyocha carnatella Rag. Nouv. Gen. p. 39 (1888); id. Rom.
Mém. viii. p. 325, pl. 35. f. 25; Hmpsn. Moths Ind. iv. p. 63.
PUNJAB.

(26) * HMMALOCERA MONOCHROMELLA.

Polyocha monochromella Rag. Nouv. Gen. p. 39 (1888); id.
Rom. Mém. vii. p. 325, pl. 36. f.13; Staud. Cat. Lep. pal. ii.
D.ld.

(27) fEMMALOCERA EREMOCHROA, sp. n.

@. Head and thorax whitish suffused with ochreous brown ;
abdomen creamy white, dorsally fulvous yellow towards base ;
pectus, legs, and ventral surface of abdomen white tinged with
brown. Fore wing pale ochreous, the costal half and terminal
area suffused with brown, the cell and veins beyond it with
some whitish irroration, the interspaces beyond the cell with

PYRALID#, SUBFAMILY HYPSOTROPINA. 131

slight ochreous streaks, the area below the cell with slight red-
brown irroration. Hind wing creamy white.

W. AusrrattiA, Sherlock R. (Clements), 19 type. Hap.44 mm.

b. (Lodiana). Antenne of male serrate.

(28) EMMALOCERA UMBRIVITTELLA.

Lodiana umbrivittella Rag. Nouv. Gen. p. 38 (1888); id. Rom.
Mém. viii. p. 319, pl. 35. f. 19; Hmpsn. Moths Ind. iv. p. 62.
Polyocha venosella Wileman, Trans. Ent. Soc. 1911, p. 357.

JAPAN, Yezo; PuNJAB; SIKKIM; ASSAM.

(29) EMMALOCERA ALBICOSTALIS.

Lodiana albicostalis Hmpsn. Trans. Ent. Soc. 1900, p. 375;
Staud. Cat. Lep. pal. 11. p. 13.

PALESTINE; PUNJAB.

B. Antenne of male without sinus and ridge of scales.

(30) EMMALOCERA SUBFASCIATELLA.
Polyocha subfasciatella Rag. Ann. Soc. Ent. Fr. 1887, p. 258 ;
id. Rom. Mém. viii. p. 328, pl. 36. f. 17; Staud. Cat. Lep. pal. 1.
p. 13:
ARMENIA; PERSIA.
Auctorum.

Polyocha rhodesie Strand, Archiv. f. Naturg. 75. 1, 3, p. 384
(CUS IONS) iialeates a hed eet a nner Ren Pe ee ere N, RuHopeEsia.

GENERA AUCTORUM.
Barberia affinitella Dyar, Proc. Ent. Soc. Wash. vii. p. 39

GUSWD) emt na phe”, cosscc so ausbers se eeurees US.A., Texas.
Cabnia myronella Dyar, J. N. Y. Ent. Soc. xii. p. 108 (1904),
MAS LUT Laren oe na acuca vadinaeseeeaNe U.S.A., Washington.
Fondoukia translucidella Chretien, Bull. Soc. Ent. Fr. 1911,
Oe ied INE LUA ane een renee eo renee See eee ALGERIA.
Schenectadia merilesella Dyar, Pr. U.S. Nat. Mus. xlvii.
PGA (LO Lye MPL CEL OSUUC areie nds ni waewSeatou iene ned PANAMA.
Sabormania pia Strand, Arch. Naturg. 78. A. Hft. 12, p. 80
Leese C—O OCLEOCETCiaaase esas evenosesy SPANISH GUINEA.

Q*

a

ON VISIBLE AND INVISIBLE CHARACTERS IN SILKWORMS. 133

6. First Report on the Inheritance of Visible and Invisible
Characters in Silkworms. By Miss Mauve L. CiEc-
HorN, F.Z.8., F.L.S., F.E.S.

[Received December 3, 1917: Read March 9, 1918. ]

The mulberry silkworm races of Bengal are all with the excep-
tion of one (Bombya textor) multivoltine, but their cocoons are
not so good as those of the European or Japanese races, and
therefore when the question of reviving the Indian silk industry
arose, the following suggestions were put forward, viz.: that
endeavours should be made to obtain an improved multivoltine
race by crossing an indigenous variety with a European one, and
that our knowledge of hybridization according to Mendel’s law
should be utilised to help in the process.

With these suggestions in my mind I obtained from Italy, in
December 1910, some European silkworm seed (eggs of the
Bombyx mort) and some Nistri (Bombyx croest) seed cocoons from
the Berhampore Government Nursery.

The European seed was of the Italian-Japanese Hybrid, and
was the first generation of a cross between the univoltine (pro-
ducing one brood a year) yellow Italian (Indigeno giallo) male,
and the univoltine white Japanese female—pure yellow Italian
seed not having been available at the time. This Italian-Japanese
Hybrid is very hardy, being a first cross, and so I started my
cross-breeding experiments with it.

The cocoons of the multivoltine Nistri silkworm weighed (with
the chrysalis removed) from 1 grain to 1°6 grains, whilst the
cocoons of the Italian-Japanese Hybrid (being of a univoltine race
and therefore bigger) weighed from about 2°5 grains to 4’6 grains.
My aim in my experiments with these two varieties of silkworms
has been (1) to make a multivoltine race (because though its
cocoons are smaller than those of the univoltine, yet they are
compensated for by the numerous broods produced during the
year), producing cocoons of about 4 grains in weight, and (2) to
see how far the good qualities of the univoltine varieties can be:
combined with the multivoltine character.

I made two series of experiments :—

A. Across between the multivoltine Nistri 9 and the univoltine
Italian-Japanese Hybrid ¢.

B. A cross between the univoltine Italian-Japanese Hybrid 9
and multivoltine Nistri ¢.

Experiment A was made with individuals selected usually from
three or four families in each generation, except in one of the
experiments in F,, which was made with a whole family.

Experiment B was made with the complete family in each
generation, with the exception of F,, when only a few worms
were reared out of four layings.

The layings produced in F, by Experiment A and _ those

134 MISS MAUDE L. CLEGHORN ON THE INHERITANCE OF

produced by Experiment B were entirely different from each
other, for they resembled the maternal parent in each
case, all the layings of the Nistri 9 and Ital.-Jap. ¢ being
multivoltine, and all those of the Ital.-Jap. 9 and Nistri
$ proving univoltine.

Result of the Experiment as regards the Size of the Cocoon.

Eaperiment A.—The parent. cocoons of the Ital.-Jap. o
and Nistri Q cross weighed 2°9 and 1°5 grains respectively. The
cocoons of the first generation of this cross resembled those of the
Nistri (7. e. the maternal parent) more than those of the Ital.-
Jap. parent, being rounded at both ends and very thick but
of the loose texture of the Nistri, the firm texture of the Ital.-
Jap. being entirely unrepresented. They were uniform in
size and shape, but varied in weight from 2°5 to 4:1 grains.

In the second and the immediately succeeding generations, the
cocoons were not so uniform in size and shape. Many were large
and rather pointed at the ends, whilst they were all thinner and
firmer than those of F,, and I found that in the earlier g generations
moths which gave complete multivoltine layings had nearly
always cut out from cocoons which were about 3 grains or less in
weight. I did not make use of these 3-grain cocoons for rearing
purposes, even though they were far better than the original
multivoltine Nistri cocoon, but I carefully selected the best out
of the most multivoltine layings of cocoons weighing from about
3°5 grains to 5 grains in weight. The cocoons in all the generations
of the cross were far superior to the original Nistri cocoon, and in
many of the generations they are also superior to that of the
Ital.-Jap. Hybrid.

Up to F, many of the cocoons were 4 grains in weight, in F,,
F,, F,, many were over 4°D grains, while some weighed 5 grains
and over. In F, and I’, there were no 4-grain cocoons, and on
the whole the cocoons. of these two generations were very poor
compared with those of the preceding generations, but they were
‘nevertheless still superior to the original Nistri cocoon. As the
layings from which these cocoons were produced were nearly
entirely multivoltine, the cocoons appeared to be also becoming
more multivoltine in character. The rearings in I, from which
F, hatched were all entirely multivoltine, while the I, silkworm
were better again and seriposited cocoons, nearly all of which
were over 3°5 grams, many over 4 grains, and some nearly 5 grains
in weight (with chrysalis and outer fluff removed). The layings
of the moths from these cocoons were also entirely multivoltine,
which showed that it is possible to obtain a 4- or 5-grain cocoon
from entirely multivoltine layings in F, after a cross without any
recrossing. Care was, however, always taken to have the parent
moths as distantly related as possible. (Table 1.)

In this experiment I found that, after the direct influence of
the fresh cross seemed to have disappeared, every third eee ation

produced the best cocoons, for the cocoons of F,, F,, F,, F,,

VISIBLE AND INVISIBLE CHARACTERS IN SILKWORMS. 135

TABLE 1, SHOWING THE DESCENT OF COCOONS UP TO THE NINTH
GENERATION, WITH DATES OF ELEVAGES.

Ital.-Jap. Hyb. 3 y, Nistri ? iaiaehr ane mina oe a ee eau ans ONE
No. 1, 2°9 grs. No. Af8 1°5 ors.

No. 229 No. 15g No. 269 No. 13g . ... March 28 ,,
37 grs. “38ers. 33 grs. “*2°8 grs.
No. 262 x NG: 3389) cntitiedienc’e Api 28s 4
4°2 gs. 37 grs.
|
No. 43 ? No, 213, No. 209 i ieiveecise UNO w Goss

41 grs. Sl grs. “*5 grs.

No.59¢ . No.15? No.69 y. No. 4d... duly 8 ,
3°8 grs. 43 grs. 4°8 grs. 5 grs.

No. 52¢ ,No.19 No. 259 /No.18g No.159No.50¢ No.52g... Aug. 30 ,,

4 ors, 38ers. 44 ers. “S4ors- BG grs. “32ers. 4 ers.
No. 102 No.52 No.1é No. 179 x Norls)c.. Oct. 10. ,,
2°2 grs. 2°8 grs. 3°6 grs. 3'6 gers. 3°6 grs.
No.41g . No.559 No.619 . No.27g = No.41¢ ............. Nov. 30 ,,
2°5 grs. 2°4 ors, 1°9 grs. 2°9 grs. 2°5 grs.
No. 40? / No. 378 No. 23 , No. 209 sinler eelatvassatenintea a ceviaucen Ee De suedl Oe
3 grs. 28ers. 22ers. “24ers,
No.115 No.113 No.119 No.221 No.216 o.cecccccceseneeeee. Mar. 22,

37 ors. 48ers. 4legrs. 4ers. 3'3 grs.

were much superior to those of the intervening generations.
Even those of IF, seriposited during the rains in July were
much better than those of F, , (April) and F,, seriposited in
October. As the raiyats in India grow about three or four crops
of cocoons a year (only rearing a few to keep the breed alive
when they are short of leaf) it would be to their advantage to
arrange that the crops reared are of every third generation.

The cocoons obtained from this multivoltine race are much
superior to the Nistri; those of F, were valued at about 9% francs
per kilo in the Milan market, which is close up to the price of
good Italian cocoons, and the cocoons of F.,, which were not
near as good as those of F,, were valued at about 7 francs per
kilo. The correct rendement could not be obtained, as the 175
cocoons sent to be tested were too few, 1 lb. weight of cocoons
being necessary.

P,

1a

F,

Fs

Fy

136 MISS MAUDE L. CLEGHORN ON THE INHERITANCE OF

Experiment B.—Cross between univoltine Ital.-Jap. Hybrid
© and multivoltine Nistri ¢.

This was carried out for four generations only and then
discontinued, as the cocoons were not as good as those of the
reciprocal cross, rarely averaging a weight of 3°5 grains.

Results of Hxperiment as regards the Multivoltine and Univoltine

character of the Silkworm Moths.
Leperiment A.

TABLE 2, ILLUSTRATING THE DESCENT OF THE UNIVOLTINE CHARACTER IN

THE CROSS BETWEEN ITAL.-JAP. 5 AND NistRI @.

Multivoltine Nistri @ X Univoltine Ital.-Jap. f... 0.00.0... ccc cee eee

Layings all multivoltine...............
Many more?s than 's.

22 layings univoltine. 30 layings almost univoltine. 6 half multivoltine ......

More ¢s than @s. ;

6 layings univoltine. 16 about half univoltine. 2 multivoltine ..,.........

More 9sthan és.

A few layings univoltine. One 75/y multivoltine. Many partly multivoltine ...

More gs than 9s.

One 75 °/y multivoltine. 2 half multivoltine. 165 partly multivoltine ........ :

és and 9s about equal.

All about half multivoltine ...............

Nearly all about half multivoltine. A few about 75°/p multivoltine...........

Nearly all completely multivoltine
All multivoltine

Allnulneolene

P

F3

Is

F;

Fs

Fy

VISIBLE AND INVISIBLE CHARACTERS IN SILKWORMS. 137

From the results obtained it will be seen that all the eggs in
the layings of the multivoltine Nistri 9 when crossed with the
univoltine Ital.-Jap. ¢ hatched, 7¢.e., the layings were all
multivoltine, whilst the univoltine paternal parent had no visible
effect on the character of the layings. This showed that the
multivoltine character was dominant in the 9, but recessive mn
the ¢. Toyama, the Japanese authority on silkworms, also
found that in crossing pure breeds the first cross always resembled
the maternal parent.

T usually found that the completely multivoltine layings did-

not give the most multivoltine results as might be expected, but
the half multivoltine laying produced the least number of
univoltine layings in the long run.

In the Nistri 2 and Ital.-Jap. ¢ cross I discarded the
multivoltine layings which appeared in the earlier generations,
and mostly selected from the half, or partly multivoltine layings,
tillin F, when most of the layings were almost entirely multi-
voltine. Some of the families have been entirely multivoltine
since F’,, others have had a few partly univoltine layings, but in
none of the generations since F, have any of the layings been
entirely univoltine.

Toyama does not give any detailed account of the results he
obtained in the brood characters. It would have been interesting
and useful, as the inheritance of these invisible characters appears
to be complicated. He selected from multivoltine forms, but
does not say if they were entirely or partly multivoltine. He
states as follows :—‘'Those forms raised from the first cross do
not remain true to the parents in subsequent generations. Even
when we selected multivoltine forms for five generations we failed
to get any constant multivoltine breed.”

In his.interesting pamphlet ‘Sulla riproduzione degli Incroci”
Dott. Quajat, when referring to his experiments with bivoltine
and univoltine races, states that ‘nelle successive riproduzioni,
bivoltinismo tende a diminuire, ed alcune volte anzi a scomparire
completamente. Sara oria interessante constatare se le ovature
ottenute univoltine, abbiano allo stato lateute il bivoltinismo, e
se questo si potra manifestare in seguito a nuovi incroci o
spontaneamente.”

I found that from F, the layings began to show a great
tendency to become completely multivoltine, and in those of the
F,, moths, from which F,, worms hatched, only one laying out of
the whole generation was half univoltine, all the others were
completely multivoltine.

To account for the 22 layings in F, appearing univoltine, the
univoltine character must have dominated in the maternal parents
of these layings. When a female F, was crossed with a ‘“ pure”
multivoltine Nistri ¢ only 3 eggs out of 250 hatched, and from
the results obtained, in the reciprocal cross, it was found that if
the maternal parent was a pure univoltine none of the eggs
hatched for about a year, and if she was a pure multivoltine all

138 MISS MAUDE L. CLEGHORN ON THE INHERITANCE OF

the eggs were multivoltine. As this female of F, had hatched
from a completely multivoltine laying, it might naturally be
’ expected that, when the paternal parent was pure multivoltine all
the eggs laid would be multivoltine, but as only a few eggs
hatched it showed that the maternal parent was dominant in the
univoltine character, and that the dominance of the univoltine
character was inherited by the F, 2 from the paternal parent in
which it was not a dominant character.

It is clear that the female influences the reappearance of the
character in a dominant form, and shows that the descent is of a
sex-limited inheritance.

In my experiments I found that the univoltine or multivoltine
character of the maternal parent showed itself in the layings and
not that of the paternal, for the character of the paternal parent
always appeared to be masked or suppressed.

Besides the difficulties a sex-limited inheritance presents the
univoltine and multivoltine characters are not visible in the moths
but only in their layings, for moths which outwardly resemble the
univoltine parent may have a multivoltine laying and vice versa.

When all the moths of a generation are bred inter se the
character of only the maternal parents can be determined by the
eges laid; but to prove that the males and females of each
generation are either homozygous or heterozygous, dominant or
recessive, they have each to be bred with pure univoltines and
multivoltines. So to find out the exact composition of all the
moths in each generation would require a multitude of experiments,
and [I could not spare many moths from F,, for I knew that in
F, only a very small percentage of eggs would hatch.

“The layings in F, which were laid by the F, moths give a clue
to the character of some of the parent moths in F,. For the
maternal parents of the six univoltine layings (Table 2) must
have been dominant in the univoltine character, and those of the
two multivoltine layings, in the multivoltine Shae a! but the
sixteen 50 per cent. univoltine layings point to the maternal
parents being heterozygous.

Heperimént B.

From the results obtained in the Ital.-Jap. 2 and Nistri ¢
cross (Table 3) it will be seen that although all the layings in F,
were univoltine, yet there were some multivoltine, and partly
multivoltine layings in F,, I, and F,; and, again, though the
moths of F, were reared from one of ‘the multivoltine layings
of F., yet 37 layings of the F,, moths were univoltine.

Forty- -seven moths from an I, family of the Ital.-Jap. 2 and
Nistri ¢ cross were tested with pure multivoltine Nistri moths
and the results obtained were as follows :-—

(1) Out of 21 layings of the F, 2s and pure multivoltine
Nistri gs 19 layings were entirely univoltine and 2 entirely
multivoltine.

VISIBLE AND INVISIBLE CHARACTERS IN SILKWORMS. 139

(2) Out of 26 layings of pure multivoltine Nistri Qsand F, ds.
all the eggs hatched, thus all being multivoltine.

These results give a clue to the gametic composition of the F,
moths, for they show :—
(1) that 19 F, 2s were dominant in the univoltine character
and 2 in the multivoltine character. |
(2) that all the F, ds which may have been univoltine were
recessive in the univoltine character.

TABLE 3, ILLUSTRATING DESCENT OF THE UNIVOLTINE AND MULTIVOLTINE
CHARACTER IN THE CROSS BETWEEN THE UNIVOLTINE ITAL.-JaAp. 2
AND MULTIVOLTINE Nistri ¢.

Univoltine Ital.-Jap: 9 X Multivoltine Nistri f......c0000., et a P,

Vayines QE wnivoweine ooo. veccas sci caviesstseantescncsss ses EY
11 2 moths 21 ¢ moths.

!
7 layings univoltine. 2 multivoltine. 1 half multivoltine............... Fe
62 Gs. 34 bs.

37 layings univoltine. 2 almest univoltine. 2 multivoltine. 2 almost multivoltine. Fy,

34 9s. 47 os.

14 layings univoltine. 7 almost univoltine. 4 almost multivoltine ......... FF,
(Discontinued.)

Unequal Sex-ratios.

I had noticed that in the earlier generations of the Nistri ?
and Ital.-Jap. ¢ cross (which were reared a year previous to that
of the Ital.-Jap. 2 and Nistri ¢ cross), the number of males
and females seemed very unequal, and that in one generation
males predominated, and in another, females. However, these
remarkable fluctuations gradually decreased in the latter
generations.

In my second series of experiments, which were with the Ital.-
Jap. 2 and Nistri ¢. I noted the exact number of males and
females in each generation, and found that in this reciprocal cross
the sex-ratios were also very unequal, but just the reverse in the
character of the predominating sex, for in F, of the Nistri 2
cross there were many more females than males, while in F, of
the Ital.-Jap. @ cross there were more males than females.

The unequal sex-ratios could be accounted for by supposing

140 MISS MAUDE L. CLEGHORN ON THE INHERITANCE OF

that in some of the generations the univoltine females do not
hatch, and in others the univoltine males, but in F, when all the
eggs hatched the sex-ratios were also unequal.

: Method of Rearing of Worms.

To ensure choosing distantly related moths for rearing from,
I reared the silkworms of each laying separately, and the cocoons
were also kept separate by keeping the silkworms after their fourth
moult in rounded trays with a double inner circle of plaited strips of
bamboo, about an inch and a half from the outer edge, forming
a space into which the silkworms readily crawl to sereposit their
cocoons. When the cocoons are removed from the trays, they
are placed in rows on large sheets of white paper on which is
noted the number of the laying from which they were reared.

Kach cocoon is covered over with a small earthen cup, the
common Indian chilum. A day or two before the moths come out,
the cocoon is cut open to remove chrysalis and weighed. .

The weight is noted and the chrysalis put back. When the
moths emerge, the pierced cocoons are removed from under the
earthen covers and placed within the opening at the top of the
covers for reference, and this also to prevent the moths from
getting out. By referring to the number of the layings and to the
numbers of the two parent cocoons (which is also noted on
the paper), the choosing of the distantly related moths for
rearing is simplified.

If I had been rearing on a large scale I should have kept four
or six families separate for seven or eight generations, then bred
them together, and from these again four or six families would
have been separated out.

In these experiments the silkworms have always been fed most
during the night, as I have noticed that, in the wild state, the
larve of moths mostly feed up to 9 or 10 a.m., and then restart
feeding at about 5 or 6 p.m. Even young silkworms, which
I kept on a small potted mulberry plant under observation, hardly
ate at all during the day, though surrounded by fresh leaf on all
sides. I find that silkworms always eat most voraciously between
10 p.m. and 4 a.m., so it is unnatural for them to be forced to
feed all day, which is usually the case by the native rearers, and
may well be one of the causes of their degeneracy.

They should have a rest of at least seven or eight hours during
the day, and be fed every three or four hours during the night,
starting late in the afternoon at about 6 p.m. If there is qn
abundant supply of leaf, they might with advantage be fed every
two hours. Newly hatched worms should always be fed every
two hours between 5 p.m. and 10 a.m. The worms if fed well in
the last stage after the fourth moult will give good cocoons, even
if not very ‘well fed in the earlier stages, but it is most essential
that they be well fed throughout the larval stage for them to be
vigorous and healthy.

VISIBLE AND INVISIBLE CHARACTERS IN SILKWORMS. 141

One of the advantages gained by the nocturnal feeding of
worms is that the leaf can be gathered at sundown, and so keeps
fresher, and does not fade as readily as when plucked in the
morning. There is also very little chance of attacks from the
dreaded silkworm fly, which is always on the alert during the
brightest and hottest hours of the day.

I have found that the leaf from male trees of Morus Indica
gives better results in feeding silkworms than that of the female
trees, The diccious character of the mulberry is not, as far as I
know, known to sericulturists, for in most handbooks on the
subject they advise rearers to select trees which bear little or no
fruit. The inconspicuousness of the catkins and pseudo-spikes of
the mulberry has evidently led to the difference between the
trees being overlooked by all but botanists. As the flowers are
inconspicuous they are anemophilous, and the trees, when in
flower in February, may be seen giving off pollen in little puffs
like smoke. This is best observed when there is no breeze.

Additional Observations made during Haperiments.

The inheritance of the visible colour character of the cocoons is
clearly Mendelian. |

The parent cocoons of the Ital.-Jap. Hyb. were of the yellow
Italian ¢ and white Japanese 9.

The colour of the yellow Italian cocoon is of a deep pinkish
yellow (carneo-giallo or flesh colour), which sometimes varies from
a deep orange to almost white, but never of the vivid yellow of
the Indian cocoon. All the cocoons of F, of this hybrid were of
a pale flesh colour, none were white like that of the Japanese
parent. A character which dominates after a eross is made was
described by Mendel as dominant, and the character which
seemed to have disappeared he called recessive. So the flesh
colour of a cocoon is a dominant character, and the white
a recessive. All observed results in the study of heredity point
to the dominance of a character being due to the presence of that
character, and the recessive to the absence of the dominant
character.

The flesh colour of the Italian is due to the presence of the
flesh cqlour and to the absence of the vivid yellow colour of the
Nistri. While the white colour of the Japanese cocoon is due to
the absence of both the pinkish yellow and bright yellow colours
—the example of the inheritance of eye-colour in man might
make this clearer. Here the dominance of brown eyes can be
traced to the presence of a brown pigment ; and the recessiveness of
blue eyes to the absence of the brown pigment. Ali human eyes
(except those of Albinos) have a layer of deep purple pigment on
the inner surface of the iris, but in brown, hazel, green, and grey
eyes there is also a layer of brown on the outer surtace of the iris,
and it is this brown layer which entirely (if abundant) or partly
conceals the purple layer. In clear grey and clear deep and pale

149 MISS MAUDE L. CLEGHORN ON THE INHERITANCE OF

blue eyes the brown pigment is absent and the purple pigment
shows through the tissue forming the iris, and makes it appear
of a clear deep blue, when the tissue through which it is seen is
very delicate—and of a clear grey or pale blue when the tissues
are more or less coarse. So, to trace the dominance of brown
eyes the real nature of the various kinds of light eyes must be
carefully made out; it then becomes clear that the presence of the
brown pigment in brown eyes makes brown eyes dominant to blue
eyes in which it is absent. If one parent has very dark brown
eyes and the other clear blue eyes, all the children will have dark
eyes, which sometimes include hazel, green or grey, but none will
have clear grey or blue eyes. The reason for there being no
blue-eyed children is apparent, for all the children inherit a
factor, or unit character for pigment from the dark-eyed parent,
but a factor lacking in pigment from the blue-eyed parent—and
the presence of this pigment in all the children makes brown eyes
dominant to blue.

To return to the colour of the cocoons the simplest explanation
will be found in the interaction of two simple Mendelian
characters. These two characters are flesh colour I’, and yellow
colour Y, and the two pairs of unit-characters involved are—

1. Flesh colour F, Absence of flesh colour f.
2. Yellow colour Y. Absence of yellow colour y.

The parent moths, then, have the following constitution :—

(1) Yellow Italian possessing the flesh colour and lacking the
yellow colour of the Nistri HS ene Hie? FFyy
(2) Japanese white lacking both characters... 2» NES

(3) Nistri lacking the flesh colour and possessing the yellow
colour ae ie a ey. bee Sle Vit

The actual parents in the experiment were :—

P, the Ital.-Jap. Hyb. an F, of a cross between the Yellow
Italian and White Japanese.

P, the Nistri.

The Yellow Italian has a deep pinkish yellow (flesh coloured)
cocoon. It inherits two factors, one for the flesh colour F, and
one for the absence of the yellow colour y, from each parent, and

so consists of the union of two similar pairs of factors Fy and iy:

The Yellow Italian is therefore pure (homozygous) as regards the
colour, for the germ-cells (gametes) by the union of which it was
formed, each carried the same kinds of factors Fy and Fy. So
the gametic composition of the Yellow Italian for the colour of its
cocoon is represented as F'Fyy in Table 4, Diagrams (1) and (2).
As the whiteness of the Japanese cocoon is due to the absenee
of both the flesh colour of the Italian, and the yellow of the

VISIBLE AND INVISIBLE CHARACTERS IN SILKWORMS. 143

Nistri, the gametic composition is represented in small letters as.
ffyy (Table 4). There was no colour in either of its parents for
it to inherit, for the gametes, from the union of which it was
formed, bore the factors fy and fy. Table 4, Diagram (1).

The cocoons of the Ital.-Jap. Hyb. are a pale flesh colour as
they inherited the flesh colour from the Yellow Italian parent
only, for the gamete from the White Japanese parent bore colour
factors lacking in both the flesh and yellow colours. The colour
factors inherited from the Yellow Italian parent were Fy
and from the white Japanese parent fy. The Ital.-Jap. Hyb. is
therefore not pure as regards the colour of its cocoon, for the two
gametes, from the union of which it was formed, were unlike and
did not carry pairs of similar factors. It is a hybrid (heterozygote),
the gametic composition being Ffyy, and it will give gametes Fy
and fy. When the Nistri (YYff), which has a deep yellow
coloured cocoon, was crossed with the Ital.-Jap. Hyb. all the
cocoons of I’, of this cross were of a bright yellow colour, but not
quite the deep yellow of the Nistri. So the bright yellow of the
Nistri was dominant to the pale flesh colour of the Ital.-Jap. Hyb.,
for the deeper yellow entirely masked the pale pinkish yellow
of the flesh colour even when it was present (Table 4). The
gametic composition of F, of the Nistri 9 and Ital.-Jap. ¢ cross
was YyFf and Yyff, and it therefore contained two classes,
both numerically equally represented. Table 4, Diagram (1).
All the cocoons were yellow, but not quite uniform in tin.

The gametes given off by these two classes were :—
YyFf giving YF, Yi, Py, yf.
Yyff ee bre 3

In F, there were a few deep yellow cocoons like those of the
Nistri, a good many bright yellow like those of I, a few pinkish
yellow and a very few white out of 52 cocoons. The exact
numbers were—-13 deep yellow, 29 yellow, 6 flesh coloured, and 4
white out of 52 cocoons.

The gametes for the colour character in I, were YF, Yf, Fy,
and yf, and'their union at random would give, in F,, cocoons of
the following gametic compositions—YYFF, YYFf, YYff, Yy FF,
Yylf, Yyff, yy FY, yyl'f, and yyff, which resulted in cocoons of
various shades with just a few white ones.

This is just what occurred, and it will be seen that the
proportions of the various colours obtained out of 52 cocoons
run the proportion calculated very close, for according to the
analysis in ‘Table 4 there should be 12 deep yellow, 24 yellow,
5 flesh coloured, and 7 white out of every 48 cocoons. ‘Table 4,
Diagram (2). There was a difficulty, however, in distinguishing
between the deep yellow and the bright yellow from the
intermediate forms which occurred,

In F, cocoons I found that in some cases when both the parent
cocoons were deep yellow, all the cocoons produced by the offspring

144 MISS MAUDE L. CLEGHORN ON THE INHERITANCE OF

were deep yellow too; in other cases, two deepish yellow F,
~ cocoons produced some flesh-coloured and even white ones.

I also found that flesh-coloured parent cocoons would, in some
cases, give only flesh-coloured ones, and in others flesh-coloured
and white.

Table 2 shows that among all the vellonae in I’, there are two
kinds—the pure (homozygous) dominants YYff and YYFF and
the hybrid (heterozygous) dominants Yyff, YyFF, and YyFf. So
when YYff and YYff are chosen as the parents, all the offspring
will produce deep yellow cocoons, but if YyFF and YyFf are
chosen, about a quarter of the cocoons will be flesh-coloured, and
if the light yellow Yyff and Yyff are chosen as the parents, about

a quarter of the cocoons will be white.

“In his experiments with the white Japanese and yellow
Siamese, Toyama obtained from the F, yellows some which
produce only yellows, and 2 yellow cocoons which gave 221 yellow
and 77 pinkish yellow, and 2 yellows which produce 254 yellow
and 77 white cocoons. These results obtained by him give almost
the exact proportions (75 per cent. and 25 per cent.) I have
obtained in my analysis of the colour factors, Diagrams (2)
and (3) Table 4.

Among the flesh-coloured cocoons they are two kinds—the
homozygous FFyy and the heterozygous I'fyy. Parent cocoons
which are both IFyy will produce all flesh-coloured ones, but
two Ffyy parent cocoons will give a few white ones. Table 4,
Diagram (3). Toyama appears to have obtained both the pure
and impure flesh-coloured cocoons in his experiments. For,
referring to them, he states ‘the pale-pinkish-yellow form pro-
duces some uniform (producing only pale-pinkish-yellow) and
some mixed (the white 25 per cent. and the pale-pinkish-yellow
75 per cent.) offspring in each succeeding generation.’

The pure white cocoon bred inter se always gave white cocoons.
The yellowish white cocoons. The yellowish-white ones, how-
ever, mostly gave a small percentage of yellow cocoons.

In the gametes of the pure white cocoons there is no colour
factor, so when the parent cocoons are both pure white the
offspring inherit only colourless factors and will all be white.
Table 4, Diagram (3).

The white recessives are also easily distinguished from the
dominant yellows, and as they are pure (homozygous) in the
colour character, a race with white cocoons can be easily
made from white cocoons which appear in any of the generations.
Toyama also found that “every white form from its first pro-
duction remains true to itself.” |

The recessive character always breeds true in whatever genera-
tion it occurs, but as it never is present in the I’, after a cross,
the F, tndigaduails must always be reared, as the recessives make
their first appearance in F’,.

On the whole the results of these experiments, excepting that

145

VISIBLE AND INVISIBLE CHARACTERS IN SILKWORMS.

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10

Proc. Zoou. Soc.—1918, No. X.

146 ON VISIBLE AND INVISIBLE CHARACTERS IN SILKWORMS.

of the visible colour character, show great complications. The
univoltine layings which appeared in the generations after F, m
the multivoltine Nistri 2 and univoltine Ital.-Jap. ¢ cross, and
the multivoltine layings which appeared in the reciprocal cross,
show that the maternal parents are dominant in the wnivoltine
and multivoltine character respectively, and that these characters
were inherited from the paternal grand-parents in which they were
dominant characters. So these recessive characters in males
appear to become dominant when inherited by the females.
Hither the female sex is in some way closely associated with the
dominance of the multivoltine and univoltine characters, or there
issome factor present in the male which makes the univoltine and
multivoltine characters lie latent, but does not hinder them from
being handed down to the offspring, the females of which may show,
in a dominant form, the latent character of their paternal parent.
The inheritance of the invisible univoltine and multivoltine
character does not appear to be quite Mendelian; however, it
may be that the sex-limited descent affects the inheritance, and —
there 1s really no failure in the segregation of the unit characters

BIBLIOGRAPHY.

Bulletin of the College of Agriculture.—Tokyo imperial Univer-
sity, Japan, 1906, Vol. VII. “Studies on the Hybridology of
Insects’: K. Toyama. :

“Sulla Riproduzione degli Incroci e su alcuni caratteri ereditari
che presenta la Sericaria Mori in relazione alle leggi di Mendel” :
EK. Quajat.

‘¢ Mendel’s Principles of Heredity ”: W. Bateson.

‘“ Breeding and the Mendelian Discovery”: A. D. Darbishire.

ON CETACEA STRANDED ON THE BRITISH COASTS. 147

7. Notes on Cetacea stranded on the British Coasts during
1913-1917. By Sipney F. Harmer, S8c.D., F.R.S.,
F.Z.8., Keeper of Zovlogy in the British ‘Aiesoun

(Natural History) *.

[Received March 19, 1918: Read March 19, 1918. }

The stranding of specimens of Cetacea on various parts of the
British Coasts is no new thing, but in the majority of cases the
evidence derivable from these occurrences has not been sufficiently
used. About six years ago the Trustees of the British Museum
decided to examine this evidence more systematically ; and
arrangement was accordingly made with the Board of Trade
and with the Admiralty, by which instructions were issued to
Receivers of Wreck and members of H.M. Coastguard that the
stranding of specimens of Whales should be reported by telegram
to the Museum. These orders were given during 1912, and the
system had become fully operative by the beginning of 1913.
‘The notes here recorded give a summary of the results thus
obtained.

The telegrams received are based on leaflets which were
distributed on behalf of the Museum, calling attention to some
of the more obvious characters by which the species of Cetacea
can be distinguished. They have generally been sufficient to
give some preliminary idea of what the stranded animal was
likely to be; and further evidence has been obtained, wherever
possible, by means of correspondence and by securing the
specimen in question, or some part of it, when this could be “done.
A written Report, on a form requesting answers to certain
specified questions has generally proved very instructive,
particularly in cases where the answers have been supplemented
by sketches or photographs.

The species of Whalebone Whales can usually be distinguished
by the characters of their whalebone or baleen; and a special
effort has accordingly been made, in each such case, to obtain a
blade of baleen. In the case of the Toothed W hales, the receipt
of a lower jaw is often sufficient for specific deter mination ; but
in a number of instances the entire head and flippers have been
secured, or even the entire animal or its skeleton. The records
may accordingly claim to have been based on satisfactory
evidence, in the majority of instances in which a positive result
has been recorded.

During the five years under review, fifteen species out of some
twenty generally recognized as British have been recorded. The
species which have not at present been reported are the Atlantic
Right Whale or Nordkaper (Balena glacialis Bonn.), the Blue
Whale or Sibbald’s Rorqual (Lalenoptera musculus L.), the

* Published by permission of the Trustees of the British Museum.

LOF

148 DR. SIDNEY F. HARMER ON CETACEA

Humpback (Megaptera nodosa Bonn.), the White Whale
(Delphinapterus leweas Pall.), and the Narwhal (Jonodon
monoceros lL.) The following observations refer in the first
instance to specimens actually recorded as having been stranded
during the five years in question (Harmer, 1914-1918); though
the statements of previous observers are noticed to some extent.
It may be remarked that the number of records of the larger
Cetacea in the neighbourhood of the British coasts has been
largely increased during recent years by the results of the
Whaling Companies which have operated at certain Stations
im Scotland and Ireland. Information on this subject is given,
for the Scotch Stations, by Thompson (1912, p. 393), and for
the Irish Stations in a paper by Lillie (1910), amd 4 in the Reports
to the British Association by Burfield (1913) and Hamilton
(1915, 1916); as well as in other memoirs cited by those authors.

MYSTACOCETI or WHALEBONE WHALES.

(1) Lesser RorQuat (balenoptera acutorostrata Lacep.).

This has proved to be the species of Whalebone Whale most
commonly stranded on our coasts; and twelve specimens have
been definitely recorded (including two from 1911 and 1912
respectively), besides one or two others which probably belonged
to the same species. All have been found during the summer or
early autumn, from June to October; and from two distinet
regions of ie coast :—(a) the Eastern coast of Scotland and
E: rgland, from Caithness to Yorkshire; (6) the South-Western
district, including the North coast of Cornwall, the Welsh coast
and the South-West of Ireland.

The Lesser Rorqual reaches. a length of about 33 feet, and
it can be recognised by the broad white band which runs across
the outer side of its pectoral limb or flipper and by the colour of
its baleen, which is entirely white or yellow, with hairy fringes
of the same colour. The baleen-blades may, however, have a
rosy tinge at their base, owing to the presence of vascular papillee
containing blood.

Although the British Museum possesses but little material
bearing on this point, I believe that there isa material difference
in the thickness of the hairs of the baleen-blades between young
and fully adult specimens. Thus I find that in the baleen of a
Lesser Rorqual, stranded at Perranporth, Cornwall, on June 5,
1916, the hairs are much finer than in an individual of the same
species stranded at Ulrome, Yorkshire, on Oct. 21, 1915. In
both instances the determination of the species is confirmed by
other evidence. ‘Thus in the specimen from Perranporth the -
pectoral fin was described as having a white band on its outer
surface—a marked specifie character of the Lesser Rorqual ;
and this evidence was completed by a sketch of the part in ques-
tion. In the animal from Ulrome the baleen was described as

STRANDED ON THE BRITISH COASTS. 149

being all of one colour, white or yellow—another distinguishing
feature of the same species. The Perranporth Whale was only
18 feet long, while that from Ulrome measured 33 feet. The
much finer character of the baleen-hairs of the former specimen
was thus probably due to immaturity.

This species is known to occur off the Norwegian coast at all
times in the year, and it feeds to a large extent on fish.

(2) Rupotpui’s Rorquat or Ser WHALE (Lalenoptera borealis.
Less.).

Jn my Annual Reports on Stranded Cetacea (1914-1918) I have
recorded the following four specimens as belonging to this
species :—

1914, Feb. 28.—-Derrynane, Co. Kerry, 60 ft.

» sept. 21.—Crosskirk, Caithness, 43 ft.

» Nov. 17.—John o’ Groats, Caithness, 47 ft.
1917, Oct. 13.—Annet, Scilly Islands, 45 ft.

These determinations were made mainly on the evidence of
single blades of baleen; and J have to admit that on a re-
examination of these specimens, and taking into consideration
the other evidence available, I have come to the conclusion that
they were probably incorrect in three of the four cases. The
mistakes (if they were wrong conclusions) arose principally from
the comparison of young specimens with old ones.

There can be little doubt that the specimen from Crosskirk
was correctly determined. Its baleen-fringes are composed of
extremely fine, flexible hairs, which have the curly, wool-like
texture which has been noticed by other writers on Rudolphi’s
Rorqual. In this respect they agree precisely with a series of
baleen-plates, 1912.6.25.1 in the British Museum collection,
obtained from one of the Shetland Whaling Companies and
undoubtedly correctly determined. In the other three specimens
which were referred to BL. borealis, the baleen-hairs, although
relatively fine, particularly in the individuals from John 0’ Groats
and the Scilly Islands respectively, were straight and not curly.
The fact that they are finer than those of the large blades of a
fully grown Common Rorqual is probably due to immaturity *,
and perhaps to the small size of the blades, which may have been
taken from near one end of the baleen-series. I am accordingly
led to the conclusion that the three individuals in question were
Common Rorquals; and some further evidence in support of this
conclusion is forthcoming.

It was pointed out by G. O. Sars (1881, pp. 3, 5) that the
Common Rorqual is characterized by a very remarkable asymmetry
in the coloration of the head, and that this feature is invariable
in both sexes and at all ages, at any rate in the Northern

* See the remarks on this subject under the preceding species.

150 DR. SIDNEY F. HARMER ON CETACEA

specimens which were examined by this observer. On the left
side of the head, the upper jaw and the lower lip are dark in
colour, like the dorsal surtace generally. On the right side, the
anterior end of the upper jaw, the greater part of the lower jaw
and a number of the anterior baleen-blades are white; these
blades, as Sars points out, being much like those of the Lesser
Rorqual.

In the specimen from John o’ Groats the baleen-plates were
described as being blue on one side, and white for a distance of
two and a half feet from the front end of the beak on the other
side, the remaining plates of the same side being blue. It 1s
fortunate that two photographs of this individual were supplied,
taken from the two sides. In these, the upper part of the head and
the lower jaw appear dark on the left side, while no difference
in the colour of the baleen is apparent anywhere in the series.
On the right side, the front part of the upper jaw, the lower lip
and the baleen-plates of the anterior end of the series are white ;
and these characters make the reference of the specimen to
BL. physalus practically certain.

In the individual from the Scilly Islands the evidence is less
complete, though the baleen in the fore part of the mouth was
described as having been white, “‘ getting gradually darker till it
becomes black.” In the photographs received of this individual (see
p. 13 of my Report on the specimens stranded during 1917), the
left side is not visible. On the right side, however, the lower
lip at least 1s white, and apparently part of the upper region of
the head. ‘he baleen is not visible. The blade received is a
small one, probably from near one end of the series. It agrees
so closely with that of the individual from John o’ Groats, in the
fineness and straight character of the hairs, that I feel little
doubt that it belongs to the same species; while it may further
be remarked that its coloration agrees better with that of the
baleen in the Common Rorqual than with that in Rudolphi’s
Rorqual. ‘This specimen, too, thus seems to have been an
immature L. physalus. |

The evidence with regard to the specimen from Derrynane was
very inadequate; but I am inclined to refer this one also to
B. physalus on the evidence of a single incomplete baleen-blade,
in which the hairs are not curly, but straight and somewhat
coarser than in the other two specimens, in correlation with its
greater size. In my original record of this specimen I remarked,
on the assumption that it was a Rudolphi’s Rorqual, that the
length (60 ft.) recorded was probably too great; but if the
determination then made was incorrect there is no reason to
suppose that the size of the animal was overestimated.

If the foregoing corrections are justifiable, the only record of
B. borealis, in these observations, is the specimen from Crosskirk,
Sept. 1914. According to Collett (1912, p. 597), this species,
which feeds principally on pelagic Crustacea, only reaches the
Norwegian coast during the summer. It becomes from 45 to 50

STRANDED ON THE BRITISH COASTS. deeok

feet long when adult, and its baleen is usually black in colour,
though some of the blades are partly white, with white hairy
fringes of the fine, silky texture and curly character noted above.
An elaborate account of this species has recently been given by
Andrews (1916); while the memoir of Allen (1916, p. 234) may
also be consulted for its coloration and other characters. The
maximum length of B. borealis is said to be about 53 ft.

(3) Common Rorquat (Laleenopterayphysalus L.).

Including three of the individuals which were originally
referred to B. borealis, as above explained, this species is
represented by eight records: namely, North Devon (Feb., 1913),
Kerry (Feb., 1914), Northumberland (May, 1915), Caithness
(July, 1913), Donegal (Aug., 1913), North Kent (Oct., 1914),
Scilly Islands (Oct., 1917), and Caithness (Nov., 1914).

The Common Rorqual usually reaches a length of 65 to 70 feet
in Northern waters. Its baleen-blades are for the most part
slate-coloured, with darker and lighter longitudinal streaks; but,
as pointed out in the account of the preceding species, a number
of them at the anterior end of the right series are white. Their
hairy fringes are yellowish in colour. The food consists partly of
fish and partly of pelagic Crustacea, the species showing a
preference for the latter form of diet (Collett, 1912, pp. 581, 582).
Like the Lesser Rorqual, it occurs off the Norwegian coast at all
times in the year.

B. physalus is known to be very variable in colour; and to such
an extent that various forms of the species have been distinguished
by the Norwegian whalers. Tor information on this subject see
Cocks (1884, p. 458; 1887, p. 215), True (1904, p. 119), and Allen
(1916, p. 181). The specimen here recorded from the Scilly
Islands appeared to have a good deal of white on the dorsal
surface, a type of coloration which is unusual for this species.
But I have recently been informed by Mr. King, who took the
photographs published on p. 13 of my Report for 1917, that in
his opinion much of the white colour observable was due to the
loss of the skin in patches and the consequent exposure of the
underlying blubber.

ODONTOCET!I or TOOTHED WHALES.
Fam. PHYSETERIDA,
Subfam. PHYSETERINA.

(4) Sperm WHALE (Physeter catodon L.).

Three records, from E. Caithness (May, 1917), Galway (Sept.,
1916), and Inverness (Dec., 1913).

The Sperm Whale, which occurs principally in the warmer
seas, is remarkable for the striking difference in size between the

152 DR. SIDNEY F. HARMER ON CETACEA

males and the females; the males reaching a length of at least
60 feet, and the females probably not much more than half that
length. Its most obvious character is the possession of a long
series of extremely large teeth, about 20-25 in number, as much
as 83 in. in length and 34 or even 3? in. in basal diameter, on ~
each side of a narrow lower jaw ; the upper jaw being edentulous
or with vestigial teeth of irregular form. The maxillary teeth of
the Sperm Whale are figured by Owen (1840-1845, pl. 89. figs. 3,
4), and by Sir William Turner (:912, pl. ix.), according to whom
they may be as many as 15 in number, and may reach a length of
80 mm. (p. 74). Owen (p. 354) gave the number as 8 on each
side. Other striking features of the species are the enormous
truncated head extending some distance in front of the tip of the
lower jaw, and the position of the blow-hole on the left side, at
the anterior end of the head.

Many records of the occurrence of the Sperm Whale in the
British seas have previously been published, a large proportion of
them having been on the Scotch coast (cf. Turner, 1871, 1872,
1904). Part of the skull of a large Sperm Whale, stranded in
1582 at Caister, on the Norfolk coast, may be seen in the Church
of St. Nicholas at Great Yarmouth. Mr. A. B. Van Deinse has
just published a paper (1918) on 37 specimens recorded on the
Dutch coast during the period 1531 to 1788. It is a remarkable
fact that, with very rare exceptions, the Sperm Whales recorded
in European waters are of large size and are therefore presumably
males. It is believed by the Whalers that these are roving
individuals which have been driven away from the herds by the
competition of other males (cf. Thompson, 1912, p. 397); the
species being polygamous. Although two of the. records here
given conform to the general rule, the third, from Galway, is of
special interest ; having been a young individual, with uncut
teeth, of only 18 feet in length. The condition of the dentition
shows that this individual was a “sucker”; and that an adult
female must have been somewhere in the neighbourhood, although
its presence was not recorded.

The Sperm Whale feeds largely on Cuttlefish, but partly at any
rate on fish.

Subfam. ZIPHIINA.

The Ziphioid Whales are distinguished from the Physeterine
by a further reduction of the dentition; the functional teeth
being commonly a single pair, or more rarely two pairs, in the
lower jaw. Other vestigial teeth have, however, been recorded
in both jaws in all the three genera known in British seas. The
functional teeth generally remain beneath the gum in young —
individuals and in the females even when adult; but they pierce
the gum in males sooner or later, sometimes only when fully
adult. In Hyperoodon and Ziphius there are, moreover, strongly
marked differences in cranial characters between the adults of the
two sexes,

STRANDED ON THE BRITISH COASTS. 153

(5) Borrte-xoseD WHALE (/Lyperoodon rostratus Mull.).

Fourteen records, mostly from the Northern coast of Scotland,
along the entire Eastern coast of Scotland and England, and in

the South of England as far west as Somerset.

' The Bottle-nosed Whale has long been hunted in Northern
seas (cf. Collett, 1906), and its movements are better known
than those of many other species. According to Hjort (1912,
p- 649, chart on p. 650) it is present in considerable numbers in
the Norwegian Sea during April—~July; extending as far North as
Lat. 76° in June. In September, when it is migrating South, the
Faroe Islanders get their last Bottle-noses. The localities where
it is principally found are on the Western side of the Gulf
Stream water, in the transition-belt between the Arctic and
Atlantic currents. The individuals of this species are said to
follow the 800-fathom line during their migration.

; It has been pointed out by several authorities (cf. Turner,
(1886, p. 45) that Bottle-nosed Whales are most commonly
stranded on the British coasts in the autumn, when on their
Southward migration; the individuals recorded being either
young specimens of either sex or adult females, often accompanied
by a calf. ‘The occurrence of the species, as shown by the British
Museum records, is in general agreement with this statement,
except that the sex of the majority of the specimens is not
definitely known and there are no records of females with calves.
There is one record for each of the months July (1917) and
August (1916), both from Scotland ; four for each of the months
September and October; and one from the South of England
(Somerset) in November, 1914. The only others of which
evidence has been obtained are two in March (Northumberland,
1914; Caithness, 1915), and one near Nairn, in May, 1914.

A remarkable alteration is known to occur in the male Bottle-
nosed Whale, with increasing age, both in external form and in
cranial characters. The changes in the skull are principally due
to the immense increase in the height and thickness of the
maxillary crests which are so characteristic a feature of «this
species, the adult males having formerly been described as a
distinct species, HZ. latifrons Gray. With this increase in the
maxillary crests is associated a corresponding change in the profile
of the head ; the “‘ forehead” becoming more and more prominent
until in old age its anterior outline is quite vertical. The old
males appear to have a migration-route which is further from the
land than that of the adult females ; with the result that they are
hardly ever recorded on the British coasts (Turner, 1886, p. 45).

According to Southwell (1883, p. 480) the young Bottle-nose is
black, the colour becoming lighter with age; old specimens being
almost yellow, with a greyish white ventral surface, while
the beak and front of the head are quite white. The oldest
males may become white all over (Millais, 1906, p. 298).

This species is provided with a pair, or sometimes two pairs, of

154 DR. SIDNEY F. HARMER ON CETACEA

large teeth, which may reach a length of 40 mm. (Collett, 1906,
p. 11), at the extreme front end of the lower jaw. According to
most authorities these teeth remain concealed throughout life in
females, but they pierce the gum in the oldest males, and are then
commonly provided with a tuft of the Stalked Barnacle,
Conchoderma auritum. Vestigial teeth, in addition to these,
have frequently been described in both jaws (figured by Turner,
1912, p. 82); and traces of them have been found in one or two
of the individuals here recorded. The male in this species reaches
a length of at least 30 feet, while the female is said not to grow
much beyond 24 or 25 feet (Munsterhjelm, 1915, p. 9). Collett
(1906) states that the Bottle-nosed Whale is believed to be able to
remain under water at least two hours, and that, although it feeds
principally on Cephalopods, it also eats Herring or Cod, or even
pelagic Crustacea. Newly-born young have been observed in
June, while other females possess a small foetus at this season.

(6) Cuvier’s WHALE (Ziphius cavirostris Cuv.) *.

Before these records were instituted, this species was known
as British on the evidence of a single skull, obtained by Sir
William Turner from the Shetland Islands. One of the most
interesting results of the system inaugurated by the British
Museum has been the demonstration that Cuvier’s Whale is
not the extreme rarity it had been supposed to be. ‘Three
individuals have been definitely determined, from Cork (Feb.,
1913), North Cornwall (June, 1916), and Wexford (July, 1915).
It is by no means unlikely that some of the older records of
*‘ Bottle-nosed Whales” may have belonged to this species.
Two of these specimens have already been recorded by me in a
communication to this Society (Harmer, 1915, p. 559).

True (1919, p. 54) has given reasons for believing that the
adults of the two sexes of this species are distinguished by marked
differences in cranial characters and in the teeth. In adult males,
according to this authority, the mesirostral ossification is enor-
mously developed and there is a deep “ prenarial basin” in thé
skull. The teeth, of which one pair are present at the front end
of the lower jaw, as in Hyperoodon, are fusiform and reach a
diameter of 25 to 30 mm. In adult females, the mesirostral
ossification is only slightly developed and a prenarial basin is not
formed. The teeth are slender, with a diameter of 10 to 14 mm.

Although the skulls of the specimens here recorded are not yet
all available for study, they appear to confirm True’s statements.
The largest individual (Co. Cork) measured 26 feet in length,

* [In my Annual Report for 1917, specimen No. 7, recorded on June 9 from
Co. Clare, was described as a Ziphius. ‘The examination of its skeleton, which has
just been cleaned, proves that it belongs to a species of Mesoplodon, having its two
teeth at the extreme anterior end of the lower jaw. It is proposed to publish
a further account of this highly interesting specimen in due course.—S. F. H.,
July 16, 1918. |

STRANDED ON THE BRITISH COASTS. 155

and its cranial characters agree with those regarded by True as
belonging to adult females. he teeth are slender, with a
diameter of about 13 mm., and were completely concealed beneath
the gun. Although the sex was not ascertained from observations
made on the entire specimen, there can be little doubt that
the animal was an adult female. he Cornish specimen was
incomplete, the part recorded measuring 15 feet in length. Its
teeth were uncut, and it was presumably. a young male or female.
The other specimen, 18 ft. 2 in. long, was definitely known to
be of the male sex. Its teeth were large and massive, with a
diameter of 35-37 mim., and they projected beyond the gum. It
thus appears probable that the teeth remain uncut in the female
Cuvier’s Whale throughout life, unless they become external in
old age, while in the male they are probably cut relatively
early.

The coloration of this species appears to be variable. Certain
individuals have been recorded in which the upper part of the
head was white; and the Wexford specimen possessed this type
of coloration. In other cases the colour has been described as
dark above and light below.

(7) SowmrBy’s WuAur (JMesoplodon bidens Sowb.).

This rare species is represented by three records, from Inver-
ness (Aug., 1915), Lincolnshire (Sept., 1916), and Wexford (Sept.,
1914), respectively.

The male Sowerby’s Whale is provided with a pair of large
triangular teeth im ‘the lower jaw, at about the middle of the
length of the mouth, on either side. Females have a pair of
similar teeth in the same position, but, so far as is known, always
beneath the gum. The three specimens here recorded had their
teeth concealed by the gum, and were presumably females. The
Wexford individual was, however, definitely known to be of this
sex. The Skegness specimen was reported to have been 18 feet in
length, which is unusually long for this species; and it was white
below—a type of coloration which has been reported by other
observers, although Sowerby’s Whale is often completely black. A
list containing 33 records of this species has been given by
Kiikenthal (1914, pp. 98, 99).

The Ziphioid Whales are said to feed principally on Cuttlefish ;
and it is not impossible that the reduction of their teeth may be
associated with this diet. Piscivorous animals, such as the
Delphinide and Crocodiles, seem to require a number of sharp
teeth adapted for holding their prey. In the Ziphioids, teeth of
this character are obviously not required ; and it may be suggested
that the absence of teeth is an advantage to them, since the
suckers of the Cuttlefish would probably attach themselves to
such convenient pegs, were these present, and the operation of
swallowing the prey might thus be rendered more difficult. It
must not be forgotten, however, that the Sperm Whale, which is

156 DR. SIDNEY F. HARMER ON CETACEA

also ‘ teuthophagous, ” has a specially strong series 0 teeth in its
lower jaw.

(8) KILLER or GRAMPUS (Orcinus orca L.).

A single record, May 1916, from the Solway Firth. The
specimen, a male, was unfortunately incomplete, but the part
remaining was 25 ft. 6 in. long; and the animal must have been
at least 30 feet long when alive. The species is very common on
the Norwegian coast (Hjort, 1902, p. 120).

It has been pointed out by Liitken (1887, p. 367) that a very
remarkable alteration in the proportions of the fins takes place
during growth in male Killers. While in young males, and in
females throughout life, the fins are relatively small and weak, in
the males all the fins —pectoral, dorsal and caudal—become
disproportionately larger as the animal grows older. The
difference between the small pectoral fins of the young male and
those of the old male is described by this author as being “ per-.
fectly astonishing.” These alterations in the fins have more
recently been described and figured by Grieg (1906, p. 9; see
especially his figs. 2-6, showing alterations in form and size of
the dorsal, caudal, and pectoral fins).

The specimen from the Solway Firth was an ececen illus-
tration of these statements. The dorsal fin was about 5 ft. 6 in.
in height, while the pectoral limbs were 6 ft. 8 in. long and
3 ft. 7 in. broad ; thus greatly exceeding in absolute size those of
a large Sperm Whale. While the pectoral limbs of young
animals and of females generally measure about one ninth of
the total length of the animal, those of the old males are as
much as one fifth of the entire length. Assuming the correctness
of this statement of Liitken’s, the total length of the old male
here recorded would have been well over 31 ft. It seems
probable that the female of this species hardly exceeds half the
length of the largest males. The failure to recognize the
occurrence of these changes in the males has resulted in the
introduction of more than one specific name for the Killer.

Grieg (1906) has given an account of a school of about 47
Killers which were observed on the Norwegian coast at the
middle of January, 1904. Four of the females were ascertained to
be pregnant, and others were accompanied by a calf. The young
of 2°5 m. in length were regarded as not more than 2 months
old, and those measuring 3:5 m. as being probably 1 year old.
Birth and pairing are believed by Grieg to take place during the
later months of the year.

O. orca may be easily.recognized by its very large teeth, which
have a basal diameter of as much as 17 or even 1} in. and are
10 to 13 in number on each side of each jaw, as well as by its
strikingly marked black and white (or yellowish) coloration.
A white patch behind the eye is characteristic, and the white
area of the ventral surface, although variable in extent, typically

STRANDED ON THE BRITISH COASTS. 157

sends up a lateral extension backwards on each side, behind the
region of the dorsal fin. ;

(9) BuackrisH or Proor-WHALE (Globicephala melena Traill).

Three records in the five years: Hampshire, 1913; Northum-
berland, 1914; and Cork, 1915; all in March; besides a school
of about 50 individuals which were stranded at Penzance on
July 1, 1911.

The Pilot-Whale reaches a length of at least 26 ft. and is
usually completely black. It possesses about 10 large teeth,
about j7; in. in basal diameter, on each side, in the front half of
each jaw. The “forehead” is enormously swollen, and the
pectoral limbs are very long and narrow, reaching a length of
about 4 ft.6 in. It is well known to associate in large schools,
which are hunted in the Orkneys, the Faroe Islands, and
elsewhere. As many as 2000-3000 individuals have been driven
ashore at one time on the coast of Norway (Hjert, 1902, p. 119);
and a record of 1000 specimens observed at the Lofoten Islands,
on Sept. 4, 1890, has been given by Grieg (1897, p. 8).

(10) Risso’s GRAMPUS (Grampus griseus Cuv.).

Four records: Jersey, Aug , 1913; 5S. Devon, Aug., 1917; and
S. Devon, Nov., 1913; the last record consisting of two
individuals, presumably mother and calf, although the sexes
were not definitely ascertained. It will be observed that all
these records belong to the 8.W. extremity of the British Islands ;
and there is little doubt that this is a more Southern species
which only just reaches our seas.

Rissos Grampus is recognizable by its blunt head, without
beak; and by the considerable reduction of the number of
its teeth. Of these, none are present in the upper jaw, while the
lower jaw possesses 4 or 5, of considerable size, with a basal
diameter of as much as #? in., at the front extremity. Adult
individuals reach a length of about 12 ft. Flowers Memoir
(1874) may be consulted for information regarding this species.

(11) Borrnie-noseD Doupnin (Tursiops truncatus Mont.).

Ten records: one in February (Scilly Islands, 1915), one in
May (Merioneth, 1916), and the remainder during the period
June-August. With the exception of one individual (Essex,
1914), the determination of which was not quite certain, all were
from the Southern coast of England, or from the Welsh or
Lancashire coast.

The Bottle-nosed Dolphin reaches a length of about 12 ft.,
and possesses about 25 teeth on each side of each jaw. As in
some other Dolphins, about two of these, at the frout end of the
series, are very small and usually remain concealed beneath the
gum; while the remainder are relatively large, with a basal

158 DR. SIDNEY F. HARMER ON CETACEA

diameter of as much as ;4 in. As indicated by its name, the

head has a well-marked beak, about 3 in. long.

(12) Wairre-BeakeD DowpHin (Lagenorhynchus albirostris Gray).

Eight records: one from the North coast of Ireland (March,
1917), one from the Island of Islay (Oct., 1913), and the remainder
from the Eastern coast, from Caithness to Lincolnshire—to which
may be added three specimens stranded in Suffolk and Kent in
February, 1918. Although the two species are known to overlap
in their distribution, it 1s noteworthy that in this series of
observations the distribution of the White-beaked Dolphin
and of the Bottle-nosed Dolphin have not overlapped and are
complementary to one another.

The White-beaked Dolphin is characterized, as indicated by its
popular name, by possessing a well-marked short beak, which is
white in colour. It reaches a length of rather more than 9 ft.
Its teeth are about 25 on each side of each jaw, but are distinctly
smaller than those of the Bottle-nosed Dolphin, their basal

° 1 fe { pS Yin 1
diameter being about 16-16 in.

(13) Wuarre-sipED DoLpHIn (Lagenorhynchus aucutus Gray).

Three records: Fair Island, between the Orkneys and the
Shetlands (March, 1913); Lincolnshire (May, 1917); Co. Mayo
(June, 1916).

This species, which is said to be one of the commonest of the
Cetacea off the Norwegian coast, where it may occur in schools of
as many as 1000 individuals (cf. Hjort, 1902, p. 118), may be
regarded as a boreal species which does not often occur in our
seas. Most of the specimens previously found have been from
the Orkneys and other parts of Scotland; and one or two have
been noticed from Ireland. So far as I am aware, the Lincoln-
shire specimen here recorded is the first to have been obtained on
the English coast.

The White-sided Dolphin has a characteristic longitudinal
whitish area on each side, in the middle and posterior half of its
body. Its teeth are smaller (basal diameter about 53; in.) and
more numerous (about 30 to 35 visible during life on each side of
each jaw) than in the White-beaked Dolphin, from which it
differs in certain other respects, the following of which may be
noticed. ‘The beak, which resembles that of JZ. albirostris,
is black. ‘lhe pectoral fins are faleate, with a very convex lower
border; those of the other species being blunter and broader,
and less convex below. ‘These fins, moreover, originate from the
white part of the body, being connected with the black part of
the head by a narrow dark streak; while in JL. albirostris the
black of the dorsal surface extends as far as the base of the
flipper (cf. Liitken, 1887, pp. 377, 386, 395). LZ. acutus is said
to reach a length of about 12 feet (Hjort, 1902, p. 117).

STRANDED ON THE BRITISH COASTS. 159

(14) Common Dotpuin (Delphinus delphis .).

About 20 records, one or two of which have been not quite
certain, though probable. Three individuals have been recorded
from the Northern part of Iveland (Mayo, Donegal); two from
the North-Eastern coast of Scotland (Inverness, Kincardine) ;
one, somewhat doubtful, from Kast Angha (Suffolk); and the
remainder, either from the English Channel, from Kent to the
Scilly Islands, or from the entrance to St. George’s Channel, on
both sides, from Cork (1918) to Wexford on the Irish side, and
on the Welsh coast on the opposite side. None have been
recorded in the North Sea area, from Forfar to Norfolk ; and
none on the West coast of Scotland; which, however, has
provided a curiously small number of records during the whole
of these observations. The distribution thus indicated is in
agreement with the supposition that this is an oceanic species
which is frequently stranded on the more exposed parts of the
coast-lines, but comparatively seldom makes its way into the
North Sea. The three records from the S. coast of Ireland
(including one for 1918) were obtained in February, that from
Inverness in April, and the remainder from August to eo

The Common Dolphin reaches a length of about 7 ft. 6 in.
and is distinguishable by its very long ae and numerous er
teeth, of relatively small size, wi ith a basal diameter of about
.in. The teeth are more numerous than in any of the other

6 op : :
species here considered, being about 45 in each half of each jaw.

(15) Common Porpotisse (Phocena phocena M..).

Numerous records, indicating, as generally supposed, that this
is the commonest species in British waters. Of those which were
certainly determined, the great majority were recorded from the
Kast coast of England, and most of them during the period May
to August. Evidence has been obtained in support of the belief
that the Common Porpoise gives birth to its young in the early
summer, and that the length at birth is from 2 ft. to 2 ft. 6 in.
According to Prof. Meek (1918, p. 197), the occurrence of
Porpoises near the coast during July and August may be
regarded as an inshore migration for parturition and pairing.

This species differs from all others found on the British coasts
in the form of its teeth, which instead of being conical, as in the
majority of the species, are broadened at the free end ese
shaped”), although the exact form of the broadened part
is variable. The teeth undergo a considerable amount. of
thickening as growth proceeds, and the broadened blade is not
infrequently worn away in some of the teeth. About 25,
or rather more, are present on each side of each jaw; though
usually two, at the anterior end of the series, remain small and
are concealed below the gum. The length of the adult is about
5 ft. 6 in.—and this is distinctly the smallest of the British
Cetacea. ‘Ihe head is not provided with a beak.

160 DR. SIDNEY F. HARMER ON CETACEA

MEMOIRS CITED.

ALLEN, G, M., 1916.—‘“'The Whalebone Whales of New England.”
Mem. Boston Soc. Nat. Hist. viii. No. 2, p. 105.

Awnpbrews, R. C., 1916.—** Monographs of the Pacific Cetacea,” II.
“The Sei Whale (Balenoptera borealis Lesson).” Mem.
Amer. Mus. Nat. Hist. (n.s.) i. part vi. p. 289.

BurFig.p, 8. T., 1913.—‘ Belmullet Whaling Station.” Rep.
82nd Meeting (Dundee, 1912) Brit. Assoc. p. 145.

Cocks, A. H., 1884.—“The Finwhale Fishery on the Coast of

_ Finmark.” The Zoologist, (3) viii. p. 455.

Cocks, A. H., 1887.—‘“‘The Finwhale Fishery of 1886 on the
Lapland Coast.” The Zoologist, (3) xi. p. 207.

Coutuerr, R., 1906.—‘* Nogle Meddelelsen om Naebhvalen
(Hyperoodon), og Hvidfisken (Delphinapterus).” Bergens
Mus. Aarbog, 1906, No. 6.

Cotuert, R., 1912.—‘* Norges Pattedyr.” Kristiania, pp. 543-.

Fiower, [Sir] W. H., 1874.—“On Risso’s Dolphin, Grampus

_ griseus (Cuv.).” Trans. Zool. Soe. viii. p. 1.

GrieG, J. A., 1897.—* Nogle cetologiske notiser.” Bergens Mus.
Aarbog, 1897, No. 6.

GrizG, J. A., 1906.—‘‘ Nogle notiser fra et spekhuggersteng
ved Bildgstr@mmen i januar 1904.” Bergens Mus. Aarbog,
1906, No. 2

Hamiuron, J. K., 1915.—* Belmullet Whaling Station.” Rep.
84th Meeting (Australia, 1914) Brit. Assoc. p. 125.

Hamitton, J. E., 1916.—‘“ Belmullet Whaling Station.” Rep.
85th Meeting (Manchester, 1915) Brit. Assoc. p. 124.

Harmer, 8. }'.— “ Report(s] on Cetacea stranded on the British
Coasts during [1913—-1917].” Published by the Trustees of
the British Museum.

1914. No. 1, Cetacea stranded during 1913.

191500 No, 2am a a.) Gea
1916) “No, 3° i Oa
1987) ENG ee & MOTE:
1918. No. 5, iio pony,

Harmer, 8S. F., 1915.—On Specimens of Cuvier’s Whale
(Ziphius cavirostris) from the Irish Coast.” Proc. Zool.
Soc. 1915, p. 559.

Hyort, J., 1902.—* Fiskeri og Hvalfangst i det Nordlige Norge.”
Aarsberetn. vedk. Norges Fiskerier for 1902, i Hefte, Bergen
(udgivet af Norges Fiskeristyrelse).

KiKENrHAL, W., 1914.—“ Untersuchungen an Walen (Zweiter
Teil),” VI., “ Zur Kenntnis des Mesoplodon bidens (Sow.).”
Jen. Zeitschr. li. p. 93.

Lituiz, D. G., 1910.—‘ Observations on the Anatomy and
General Biology of some Members of the Larger Cetacea.”
Proc. Zool. Soc. 1910, p. 769.

Lirxen, C, F., 1887.—“ Kritiske Studier over nogle Tandhvaler
af ‘Slaegterne Tursiops, Orca og Lagenorhynchus.” Vid.

STRANDED ON THE BRITISH COASTS. 161
Selsk. Skr. (6), naturvid. og math. Afd., Bd. iv. No. 6
(Copenhagen), p. 337. .

Meer, A., 1918.—‘‘The Reproductive Organs of Cetacea.” Journ.
of Anat. lil. p. 186.

Miuuais, J. G., 1906.—‘* The Mammals of Great Britain and
Treland,” vol. 111., London (Longmans, Green & Co.).

MounsteruwJetmM, L., 1915.—“ Anteckningar om Hyperoodon
rostratus ( Mill.) gjorda under en ishavsresa sommaren 1910.”
Troms@ Museums Aarshefter, xxxvii. 1914, p. 1.

Murray, [Sir] J., and Hsorr, J., 1912.—“'The Depths of the
Ocean.”? London (Macmillan & Co.).

Owen, [Sir] R., 1840-1845.—* Odontography,”’

Sars, G. O., 1881.—‘ Fortsatte Bidrag til Kundskaben om vore
Bardehvaler.” Forh. Vid.-Selsk. Christiania, Aar 1880,
INo.. 12.

SoutHwELL, T., 1883.-—‘‘On the Beaked or Bottle-nose Whale
(HTyperoodon rostratus). Trans. Norfolk and Norwich Nat.
Soe. ii. p. 476.

Tuompson, D’A. W., 1912.—‘‘ Whales, Seals and Sea-Serpents,”’
in ‘Science of the Sea.” Edited for the Challenger Society -
by G. H. Fowler, ch. xii, pp. 383-402, London (John
Mutray).

True, F. W., 1904.—‘*The Whalebone Whales of the Western
North Atlantic compared with those occurring in European
Waters, with some Observations on the Species of the North
Pacific.’ Smithson. Contr. xxxiil. p. 1.

True, F. W., 1910.—‘‘ An Account of the Beaked Whales of the
Family Ziphiide in the Collection of the United States
National Museum, with Remarks-on some Specimens in
other American Museums.”’ U.S. Nat. Mus., Bull. 73.

Turner, [Sir] W., 1871.—‘“ On the Capture of a Sperm Whale
on the Coast of Argyleshire, with a Notice of other Specimens
caught on the Coast of Scotland.” Proc. Roy. Soc. Edinb,
vil. p. 365.

Turner, [Sir] W., 1872.—“ Additional Notes on the Occurrence
of the Sperm-Whale in the Scottish Seas.’ Proc. Roy. Soe.
Edinb. vii. p. 632.

Turner, [Sir] W., 1886.—“On the Occurrence of the Bottle-
Nosed or Beaked Whale (Hyperoodon rostratus) in the
Scottish Seas, with Observations on its External Characters.”
‘Proc. R. Phys. Soc. Edinb. ix. 1885-1888, p. 25.

Turner, [Sir] W., 1904.—“ The Occurrence of the Sperm Whale
or Cachalot in the Shetland Seas.” Ann. Scott. Nat. Hist.
1904, p. 4.

Turner, [Sir] W., 1912.—‘‘The Marine Mammals in the
Anatomical Museum of the University of Edinburgh.”
London (Macmillan & Co.).

Van Deinszg, A. B., 1918.—‘ Over de Potvisschen in Nederland
gestrand tusschen de Jaren 1531-1788.” Zool. Meded. R.
Mus. Nat. Hist. Leiden, Deel iv., Aff. 1, p. 22.

Proc, Zoou. Soc.—1918, No. XI. 11

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ON THE VARIATION OF THE PIT-VIPER. 163

8. On the Variation of the Pit-Viper, Lachesis atrox.
By Miss Joan B. Procter, F.Z.S.

[Received March 19, 1918: Read April 9, 1918. |
(Text-figures 1-5.)

INDEX...
Page

Variation and Atiology (Evolution, heredity, etc.). Variation
in Lachesis atrox ; derivation of markings.................. 167-180

Systematic (New names or changed names, or Vacs of
I. atrox ; revisions, discussions, or elucidations affecting
any part of the system, to be indexed under the names)... 163-167, 180

The distinction of the forms, whether they be regarded as
species or as varieties, which cluster around the tropical American
Pit-Viper, Lachesis atrow L., and of which the principal are
L. lanceolatus Lacepéde, L. yararaca Wied, and L. jararacussu
Lacerda, is a subject which needs renewed investigation, especially
in view of Dr. Vital Brazil’s recent publications*, in which he
claims specific rank not only for L. lanceolatus but also for
L. jararacussu, which has generally been regarded as a mere
colour variety.

Mr. G. A. Boulenger, who has kindly helped me with advice in
my study of reptiles, suggested to me that I should take up this
investigation, and he has given me not only access to the collection
in the British Museum, of which he is in char ge, but the benefit
of his experience, especially as regards the bibliography of the
subject. My best thanks are due to him for these favours.

IT. HISTORICAL.

The number of. names which Mr. Boulenger has quoted under
the synonymies of LZ. atrow and L. lanceolatus is very great, but
for my present purpose it will be sufficient to discuss only the
principal, such as have had more general currency, or which are
accompanied by figures enabling me to form an opinion on the
forms for which they are intended.

The earliest name is that of Coluber atrow Linnzeus, Mus. Ad.
Frid. (1754) pl. xii. fig. 2. The specimens are described as having—
ventrals 200 and 196, subcaudals 70 and 67 pairs, and scales with
“carima elevata.” However, Mr. L.G. Andersson, Cat. Lin. type-
spec. Sn. (Bih. Sv. Vet.-Ak. Hand]. xxiv. iv. No. 6) p. 19, has
been able to supplement this definition, as he has had access to
the type-specimens in the Stockholm Museum, and he finds that
the keels on the scales “are low and extend nearly to the tip
of the scales,” thus agreeing with Mr. Boulenger’s L. lanceolatus,

* La Défense contre l’Ophidisme. Sao Paulo, 1911, 8vo, 2nd edition, 1914.
1 a

164 MISS JOAN B. PROCTER ON THE

not with his LZ. atrow. He counts 204 and 200 ventrals (v.),
67 and 70 subeaudals (c.). Both descriptions appear to corre-
spond with two specimens in the British Museum collection, from
Martinique (Cat. Snakes, ii. p. 536, spec. 7 & s).

Coluber lanceolatus Lacepéde, Hist. des Serp. (1789) p. 121, is
based on specimens in the Paris Museum: 228, 225 v., 61, 59 ec.
Habitat Martinique and perhaps Dominica and Cayenne. Figure
worthless, probably from a bleached specimen. Duméril and
Bibron (Erp. Gén. vii. 1854, p. 1505), under Bothrops lanceolatus,
do not give us any particulars concerning the type-specimen,
which they must have examined, but the fact that the ‘“ Vipére
jaune de la Martinique,” also found in St. Lucia and Dominica, is
the form intended. There can be no question as to the appli-
cation of the name lanceolatus, the Fer-de-lance, which must be
regarded as a strict synonym of ZL. atrow of Linnzeus according to
Andersson.

Cophias jararaca Wied, Abbild. Nat. Bras. (1825), from
EK. Brazil, 193-201 v., 59-68 ce. ‘The description and figure
indicate a snake similar to LZ. atrow, but with markings consisting
of dark brown, darker-edged transverse bands, narrower on the
back than on the sides, such as is figured by Jan, in Icon. Oph.
47me livr. pl. 111., and corresponding to several examples in the
British Museum collection from Rio Janeiro and Para (Cat. Sn.
iil. p. 537, spec. 7, p. 539, spec. v). This variety must be the
prevalent form in the Province of Bahia, as the following
descriptions and figures of Wagler in Spix’s Serp. Bras. (1824)
are evidently referable to it. Bothrops megera, p. 50, pl. xix.:
195 v., 53 ¢ This figure shows dark olive upper parts with
dark, darker-edged transverse bands and immaculate ventrals.
B. fur ia, p. 52; pl. xx.: 201 v., 65:¢.. A uniformly coloumed
specimen which ey be taken to have lost its markings.
B. leucostigma, p. 53, pl. xxi. fig. 1: ¢v.,66c¢. Brownish, with
darker transverse bands: ventrals powdered with grey. SB. tessel-
latws, p. 54, pl. xxi. fig. 2: 190 v., ¢c. _ Dorsal markings as in
B. lewcostiyma, ventrals checkered with brown. 3B. teniatus,
p. 55, pl. xxi. fig. 8. Colours lighter than the above, transverse
bands always double.

Bothrops jararacussu Lacerda, Leg. sur le ven. des Serp. (1884) p. 9.
From the province of Rio de Janeiro. An adequate description
of the striking markings of the Jararacussu is given (black above
with yellow markings, longitudinal on the h 50, and obliquely
pronged forks on the sides, yellow beneath spotted with black).
ae to the B. atrox figured in Jan, Icon. Oph. (47me livr.
pl. 1. fig. 3), and to two specimens in the British Museum
eee from Sao Paulo, presented by Dr. Brazil.

Having thus indicated the sources to which I have referred in
order to fix the exact meaning of the principal names which have
had currency in the past, I will review the various opinions which
have been expressed in the principal works on Ophidia.

VARIATION OF THE PIT-VIPER. {65

Schlegel (Phys. serp. (1837) pp. 932, 535, 537) recognises three
species :—1l. Trigonocephalus jararaca, from Sao Paulo, sta.
Catharina I., Sta. Cruz de la Sierra: 172-204 v., 44-62 ¢. From
the description, this snake is the true Jararaca of Wied.
2. 7. atrox: 190,196 v., 60, 68 c. Scales surmounted by a strong
keel with tubercular tendency, i in which respect it does not agree
with the Linnean type specimen. Habitat N. Brazil, Dutch ‘and
French Guianas. 3. 7’. lanceolatus: 271, 220 v., 60, 60 c. Mar-
tinique and St. Lucia. Scales surmounted by a simple keel.
Colour of ventrals a clear greenish yellow. The description
corresponds to the type-specimen of L. atrox as described by
Andersson. Schlegel maintains that 7’. danceolatus has a higher
number of ventrals: than the two preceding, but the number 271
is probably due to some mistake.

Dumeril and Bibron (Erp. Gen. (1854) pp. 1505, 1507, 1509)
describe three species :— Bothrops atrox, B. lanceolatus, B. jararaca.
The only differentiation made between the first two in the
“Tableau Synoptique des Espéces du Genre Bothrops” (p. 1504), is
that the ventrals of BL. atrow are spotted, and those of L. lanceo-
latus unicoloured. In describing 4. atvou, however, they mention
that the scales are move strongly keeled. 4. jararaca is distin-
guished from the preceding by a relatively obtuse canthus rostralis
and larger scales on the snout. ‘The description corresponds with
that of the Jararaca of Wied.

B. atrox. 29-32 scales. Habitat Dutch and French Guianas.

B. lanceolatus. 271, 240, 220 v., 68, 64, 60 c¢c.; from Marti-
nique, Dominica, St. Lucia.

B. jararaca. 172, 195, 204 v., 44, 62, 65 ¢.: from Sao Paulo,
Sta. Catharina I., never in Surinam.

The number of ventrals given by Dumeéril and Bibron do
not seem very reliable, and 271 for 2B. lanceolatus is probably
copied from Schlegeél’s statement, which there is good reason to
question.

Jan, Icon. Oph. 47me livr. Pl.i.: Bothrops lanceolatus (Merr.).
Antilles? Geneva Museum. The specimen figured has 29 rows
of scales. Markings irregular, but similar to those of two spe-
cimens in the British Museum collection from Martinique (Cat.
Sn. p. 536, spec. 7, s).—Pl.u.: B. atrow L., var. dirus Jan, from
Buenos Ayres. Turin Museum. 25-27 rows of scales, with keels
similar to those figured for 4. lanceolatus. The markings repre-
sented in fig. 3 nnn. are matched by two specimens in the
British Museum collection, from Sao Paulo (presented by
Dr. Brazil), and, as before mentioned, agreeing with those of the
Jarvaracussu of Lacerda.—PIl. il. fig. 4: var. tesselatus Neuw.
Milan Museum. Another form of dorsal markings, similar to a
specimen in the British Museum collection, from St. Lucia (Cat.
Sn. p. 536, spec. v).—PIl. ii: B. jararaca Neuw. Freyburg
Museum. An excellent figure of the typical Jararaca; 25 rows
of scales.

166 MISS JOAN B. PROCTER ON THE

A. E. Brown, in a paper in the Proc. Acad. Philad. 1893
(pp. 435-6), describes three species, but adds that he doubts their
right to more than a varietal distinction :—1. Bothrops atrox,
described from two specimens from British Guiana; 196, 195 v.,
42, 59.c¢., 27, 27 rows of scales, which are broad, with a -high
short keel not extending to the tip. Abdomen unspotted.
2. B. lanceolatus, described from two specimens from Martinique ;
199, 200 v., 69, 69 c., 25, 25 sc. Scales longer than in B. atrox,
and bearing a simple keel to the tip. 3. SB. jararaca, described
from one specimen from Brazil; 206 v., 57 c., 27 se.; besides the
anntlated markings, Mr. Brown notes that the canthus rostralis
is less sharp and the scales on the snout are larger than in the
preceding species, a fact which was also observed by Duméril
and Bibron.

G. A. Boulenger, Cat. Sn. vol. ii. 1896, pp. 535-537.

Lachesis atrox. 161-216 v., 47-73 c.
L. lanceolatus. 180-240 v., 46-70 ¢.

Mr. Boulenger finds that the only distinctive character between
these species 1s that of scale-structure. In the former the scales
are ‘‘strongly keeled, the keels on the posterior part of the back
very high, swollen in the middle, and much shorter than the
scale.” In the latter the scales are merely “sharply keeled,” the
keel extending nearly to their extremity. He also describes the
ventrals of LZ. lanceolatus as yellowish, uniform, or powdered, or
spotted with brown, and in this respect many of the specimens
in the British Museum collection certainly disprove the state-
ment made by other authors, that the ventrals of this snake are
constantly immaculate. He adds that these species may have to
be united, as some specimens of L. lanceolatus ‘approach L. atrox
in the swelling of the scales at the base of the keels, and are thus
intermediate between the two species.” Cophias jararaca and.
Bothrops jararacussu are placed in the synonymy of L. lanceo-
latus.

V. Brazil, Def. contre l’Ophidisme (2nd ed. 1914), pp. 78, 84, 88.

L. lanceolatus. 195-200 v., 50-53 ¢. Tropical America.

L. atrow, 202 v.,55 ¢. Tropical America, less abundant than
the above in the southern states of Brazil.

L. jararacugu. 170-176 v. Brazil (Sao Paulo and Rio).

In describing the first two species Dr. Brazil maintains that
one of the main points of difference between these snakes lies in
the ‘‘systéme de coloration” of L. atrox, of which he says: “ La
coloration du fond, sur lequel se dessinent des figures pareilles a
celles constatées dans l’espéce précédente (L. lanceolatus) est d’un
gris rougeatre, parfois un ton gris cendré. Cette combinaison de
couleurs donne un aspect velouté a Vanimal, ce qui permet de le
reconnaitre a premiere vue.’ The ground-colour of L. lanceolatus
is described as ‘‘ vert trés foncé, gris et quelquefois jaunatre,”
and the abdomen “vert foncé, parsemé de taches jaunes,” whilst

4

VARIATION OF THE PI'T-VIPER. 167

that of ZL. atrow is of a “trés beau jaune clair marqueté sur les
bords de noir ou de gris foneé” *. |

In examining a large number of these snakes I have found the
ground-colour to be equally variable in both; the ventrals may
be yellow, yellow checkered with black, or of that greenish colour
produced by a powdering of black over yellow, in either L. atrox
or L. lanceolatus. It was therefore not. quite clear to me why
Dr. Brazil considers the general aspect and coloration such striking
points, or how the latter should produce a velvety appearance in
L. atrox. Mr. Boulenger has looked into this matter with me,
using specimens of L. atrow and L. lanceolatus named and sent to
him by Dr. Brazil himself. There certainly is a difference in the
appearance of these specimens, but this is not due in any way
to the markings, which are strikingly similar. The specimen
marked “ atrow” has, however, the very prominently keeled scales
mentioned before, and Mr. Boulenger has pointed out to me that
the “velvety” appearance of this snake is entirely due to the
consequently greater breaking up of light upon its dorsal surface.
Dr. Brazil’s Z. atrow is therefore the same as Mr. Boulenger’s.

L.. jararacussu.—Dr. Brazil maintains the specific rank assigned
to the Jararacussu by Lacerda: first, on account of its striking
black and yellow markings; secondly, because of the comparatively
low number of its ventrals; and, thirdly, because of its much
more triangular head. With regard to the first point, there are
specimens in the British Museum collection which can be graded
to form a transition series from the typical ZL. atrow to the
Jararacussu, a specimen from W. Ecuador being exactly inter-
mediate between the two types of markings. With regard to the
second point, the ventrals of the Jararacussu are as a rule fewer
than those of Z. atrox. Dr. Brazil mentions 170-176, but two
specimens received from him have 180 and 184. 180-240 ventrals
is, however, the range given for L. lanceolatus in Mr. Boulenger’s
Catalogue of Snakes, and, as I have failed to detect any structural
differences in the shape of the head or in the scaling, I think
that the Jararacussu cannot be regarded as more than a colour
variety of L. atrow L.

Il. Form Aanp LEPIDosIS.

As I think I have shown in the annexed table that there is no
correspondence between variations in markings and the number of
ventrals and caudals, I will describe the differences in form and
lepidosis which have led to the distinction of species.

The most important is that of the two types of scaling found
in LZ. atrov. It has been generally accepted that the high short

* The coloured plates which accompany Dr. V. Brazil’s descriptions do not convey
these supposed differences, and it is well to point out that but little care has been
bestowed on the rendering of the markings. I think a L. lanceolatus with only
about 10 markings on the body, as figured, to be an impossibility.

168 MISS JOAN B. PROCLER ON THE

keel is proper to L. aérov and the low long keel to LZ. lanceolatus.
Dr, Andersson has shown, however, that the latter form is really
the Coluber atrow of Linneus. so that if the former is maintained
as a distinct species on account of its scale-structure, it will be
necessary to alter the name which has usually been given to it.
I suggest that of ZL. affinis Gray, as the specimens of Bothrops
affinis (Catalogue of Suakes (1849), p. 7, specimens Z & 0), which I
have examined, answer the definition, and this appears to be the
earliest name which can be applied to it.

The dorsal scale of the typical Z. atrox L. is of a long narrow
diamond shape, usually about twice as long as broad, and bearing
a simple keel extending to the apical pits in the tip of the scale.
That of ZL. affinis is broader in proportion, more rounded, and
sometimes truncate behind; it bears a short keel, supported upon
an extremely convex area, leaving only a narrow margin of flat
scale. This convexity of the scales is so pronounced in some speci-
mens that they can be distinguished from the preceding form by
touch alone. ‘The scales of ahen specimens, however, are inter-
mediate in type; they may be of the narrow form with the long
keel distinctly swollen at the base, or, while maintaining the long
keel of atrox, they may be both broad and convex as in affinis.
Other individuals present scales of both the extreme types. On
examining the middle third of the body in one of these, I find that
the median dorsal scales are of the high short type, and the lateral
dorsals of the long low type. Further examination shows that,
in every case, the anterior part of the snake is of the atrox
type and the posterior of the affinis type. ‘The evolution of the
scale-structure is thus clearly shown. ‘The long low-keeled scale
gradually broadens, whilst its keel swells along its base and
shortens, until it becomes completely transformed into the affinis
type. This transformation is, as described above, completed upon
the median dorsal scales sooner than upon the laterals.

As regards the snout of Z. jararaca, its shape is very slightly
different from that of Z. atrow, in that the canthus rostralis is
somewhat more obtuse, and the scales of the upper surface are
slightly larger than those on the vertex.

These characters are not very distinct, as several specimens of
the typical ZL. atroaw present snouts of a similar form; in fact,
one speeimen (Cat. Sn. 11. p. 536, spec. v) is indistinguishable
from ZL. jararaca (spec. g, Cat. Col. Sn. [1858] p. 226) in this
respect. Duméril & Bibron, and A. E. Brown, both mention
this point as one of the distinguishing features between the
two forms; and Schlegel (Phys. Serp.), in his figure (pl. xix.
fig. 1) of the dorsal view of the head of the Jararaca, represents
the difference between the scales on the snout and those on the
rest of the head as very considerable. This is certainly a point
of variation which J cannot consider proper to the Jararaca
alone, having found no fewer than six young specimens of
L. atrox with this characteristic.

The different types of form and lepidosis described above are

VARIATION OF 'THE PIT-VIPER. 169

therefore so inconstant, and so completely connected by inter-
mediate forms, that they cannot in themselves be considered
sufficient for the distinction of species.

111. Sysrem or MARKINGS,

Before discussing the evolution of markings and their individual
variation, I will outline the general system of arrangement of
markings common to all colour varieties of LZ. atrox. Asa working
hypothesis on this subject, Mr. Boulenger drew my attention to
Dr. Zenneck’s paper ‘“ Die Zeichnuung der Boiden ”*, and I have
found that his theory—that the dorsal and lateral markings of
Boids are made up of four paired longitudinal series -—is equally
applicable in the case of ZL. atrox. Of these, only three are
usually present, but in some specimens the fourth is present upon
the head. It is the constant relationship between the spots
of the series of one side of the snake which forms the regular
pattern, but those of one side are quite independent of those
on the other, thus accounting for the asymmetrical appearance
of the markings on the median dorsal line, which is sometimes
very marked ¢.

When present on the head, the first or dorsal series commences
as a streak, or a few broken spots. Occasionally it forms a
A-shaped marking with its fellow. On the body this series is
usually the broadest and most marked, and, in individuals
where it tends to disappear, it is always the last to go. The
dorso-lateral or second series, when present on the head, may
either consist of an oblique narrow streak or of broken spots of
this streak, originating above the ocular shield, or of a cress-bar or
blotch in front of the eyes. This second series is rather unstable
on the body, for though it may be of equal development to the
dorsal, it is more often hardly discernible and certain spots
frequently become confluent with their neighbours of the first
series. In some forms the markings of these two paired series
form a single dark triangular area on the head, but are more
often irregular and indistinct.

The lateral series is always present on the head of those snakes
exhibiting dorsal markings. It forms a broad black streak from
the posterior border of the eye to the commissure of the jaws,
usually passing through the sixth and seventh labials. Rays
proceeding from it may be present on the other upper labials, or
may take the form of isolated spots. The post-ocular streak 1s
always separated from the superciliary streak by a narrow area
of the lighter ground-colour, and is sometimes also outlined in

* Tiibing en Zool. Arb. iii. (1898).

+ These four series are termed Dorsal (D.), Dorso-lateral (D.L.), Lateral (L.); and
Ventro-lateral (V.L.) by Mr. Boulenger, ‘Snakes of Europe’ (1913), p. 30. This
lettering is followed in my text-fig. 1.

t Mr. Boulenger has drawn attention to this bi-lateral asymmetry in his op. cit.
p. 33, and he mentions a specimen of Lachesis alternatus with 24 markings on the
left side and 27 on the right.

170 MISS JOAN B. PROCTER ON THE

cream-colour, which brings it out very sharply. On the body this
series 1s well defined. ‘

The fourth or latero-ventral series is never present on the body,
and not always upon the head. It is evidently a most primitive
character, which has disappeared in many specimens. In the

Text-figure 1.

most marked form this series only consists of three or four black
spots on the lower labials, and one larger spot which has become
confluent with the post-ocular streak. This latter spot is the
last of the series to disappear, and is entirely the cause of the
apparent broadening and lengthening of the post-ocular streak in’

VARIATION OF THE PIT-VIPER. 171

many snakes. This series must not be confounded with the outer
ventral series, which commences on the chin-shields and is usually
distinctly marked throughout the body.

The markings develop upon regular lines, and the degree of
this dévelopment varies on uorenit regions of the body. That
on the anterior end of the body is the most primitive, as I shall
endeavour to show later on, but markings immediately behind
the head are always irregular.

In Lachesis atrox the spots of the series on each side of the
body form themselves into distinct alternating groups, which I
shall call A and B. Group B is less stable than A, and frequently
disappears altogether, either temporarily in the evolution of the
markings or permanently.

Text-figure 2.
Group A. Group B.

On one snake the relative development of groups A and B. is
not always constant, but it is always alike on both sides of
the body.

Since group A is usually the best developed, it is this group.
which I have counted in giving the numbers of markings in the
table of statistics. This number is not the same for the two sides
of many snakes—another reason for the faulted appearance in
the general dorsal pattern. |

The dorsal markings of L. atrow are therefore made up on each
side of three longitudinal series of spots, which in turn form
themselves into alternating groups A and B. Each of these
consists of six primitive spots, which, according to their presence,
absence, or confluence, determine the variety of marking pro-
duced. In order to follow their evolution and variation, I have
numbered those of each group in order of their stability (see
text-fig. 2), so that each may be referred to by a definite
formula.

The ground-colour appears to be formed by the varying pro-
portions of a black (or dark brown) and a yellow pigment, the
black being the more superficial. When the shade is greyish,
there is a smaller proportion of yellow pigment. The usual brown
tints are composed of a very fine powdering of the dark pigment:
over the yellow. Quite apart from the regular markings, the scaleg_

172 MISS JOAN B. PROCTER ON THE

often each show a delicate arrangement of black pigment standing
out from the ground-colour and having the same pattern effect
as the fine dendritic marks seen in many rocks. This arrange-
ment is often exactly repeated from one scale to another. When
this style of markings is present on the clear yellow of the
ventral shields, it produces a dark green effect, merging, when -
seen ata distance, into the ground-colour.

IV. INDIVIDUAL VARIATION AND Evouutrion oF MARKINGS.

Individual Variation.

As a starting-point in the study of the markings and colour
varieties of L. atrox, it was suggested to me by Mr. Boulenger
that I should make an examination of a female and her 26 young
from Trinidad (received from Mr. Urich) in order to ascertain
the amount of individual variation among specimens unquestion-
ably pertaining to one form only.

The young snakes range from 160 to 300 mm. in length, and
are more brightly marked than their mother. Their ground-
colour is a soft shade of brown, except the tip of the tail, which
is a pale yellowish. Particulars as to the number of ventrals
and subcaudals etc. will be found in the annexed table. In the
majority of cases, the pattern on each side is made up in the
following way, as shown on text-fig. 3 :—

Group A consists of the six primitive spots arranged in a
pyramid (see text-fig. 2, p.171). <A large triangular or rather
trapezoid spot (A 1, dorsal series) forms the apex, and rests
upon two small rhombic spots (A 5, 6, dorso-lateral series), which
are sometimes confluent with it. These rest upon three similar
spots, the outer ones being somewhat the larger (A 2, 4, 3, lateral
series), This triangular marking is margined in cream-colour,
and encloses an area darker than the surrounding ground-colour,
a state of things which seems to suggest a centripetal aggregation
of pigment. Group A, though equal in size to group B, is in
every case darker and more marked.

Group B is also triangular in form, but the base coincides with
the mid-dorsal line, instead of the apex as in group A. The said
base is composed of three oval spots (B 5, 1, 6, dorsal series), the
middle one being much the largest. Three more small oval spots
enter the remainder of this triangle, the largest at the apex
(B 3, 4, dorso-lateral, B 2, lateral). (Text-fig. 3, A.)

When the markings of the two sides of the snake correspond
exactly, the general dorsal design is a single string of light-
spotted ovals on a dark ground; this is formed by the paired
groups B having their bases applied together. They appear oval
rather than diamond-shape, on account of the curved border of
the ground-colour of these groups. If the markings should alter-
nate with each other, a light wavy band on a dark ground is the
general result. Usually, however, a single specimen exhibits

VARIATION OF THE PIT-VIPER. 173

many different combinations of these forms, owing to the different
number of markings on the two sides. In most specimens mark-
ings of one or more series are present on the head, and in several
the latero-ventral series is very marked.

Text-figure 3.

Many specimens do not possess all the primitive spots described,
especially upon the posterior parti of the body, where the pattern
becomes modified. In the first stage of this modification A 5
and 6 become confluent with A 1, which tends to divide into two
triangular spots; the spots of group B, with the exception of
Bland 2, merge into the background (text-fig. 3, c). Some
specimens are of the more primitive type anteriorly only, whilst
others have the markings of the greater part of the body of the
slightly modified type, and become still further altered on their

174 MISS JOAN B. PROCTER ON THE

posterior parts. -One specimen, however (see Table, Trinidad 1 A),
is. inclined towards the Jararaca type of marking. Anteriorly
A lis divided, each spot being confluent with A 5 and 6; A 4 is
absent, and of group B only an occasional B 1 is present (text-
fig. 3,D). At about the middle of the snake the twin spots
A 1,1, have become some distance apart, and are in sufficiently
close proximity to A 2 and 3 to give the appearance at a distance
of paired transverse bands, or, since the intermediate space is
darker than the surrounding ground-colour, of a single dark-
edged band; these bands, which may pair with those of the
opposite side or not, are very little more than their own width
apart.

In all specimens the pattern tends to break up into a spotted,
type just before the tail, all the spots being more or less equal in
size and distribution. It consists of A 1, 1, 4, 2, 3, and B 1,
3, 4, and 2.

In the markings of the mother, A 1, 2, 3, 5, and 6 are con-
fluent, forming a somewhat truncate chevron, in the middle of
which A 4 is sometimes situated. Group B is made up of the
six primitive spots, all except B 1, 2, however, being indistinct
(text-fig. 3, B). On the whole, the individual variation shown by
these 27 specimens is very slight ; the most apparent differences
between them are due not to variation of the actual markings,
but to the inconstant relationship of those of the two sides of
the snake.

I have since examined another mother and young, from Anda-
goya, Colombia (received from Dr. Spurrell), and in this case
there is no variation at all among the young.

The markings of both families are of the most primitive type
occurring in ZL. atrow, ail other forms being evolved from it, as
will be shown presently.

Evolution.

In the course of evolution, these primitive markings become
modified in the following way, which is the same for all forms of
L. atrox.

A 1 tends to divide transversely to the axis of the body, and
A5 and 6 become confluent with it. B5 and 6 merge into the
ground-colour, A 4 and B3 and 4 disappear (this stage has
already been described in the variations of the young snakes),
A 1 then divides completely, but is still distinct from A 2 and 3,
A 4 disappears, and the whole of group B tends to merge into
the- background. The spots Al, 1 then become a considerable
_ distance: apart, narrow, and in close proximity with A 2 and 3,
which, since the interspace is darker than in group B, thus
produce the effect of a dark, darker-bordered, transverse band.
Except for an occasional Bl, this group is pale and quite
indistinct. At this point the bands, which may or may not pair
with: those of. the opposite side, are slightly more than their own.

VARIATION OF THE PIT-VIPER. 175

width apart. A 1,1 and A 2, 3 now become confluent, forming
the dark unbroken marginal lines of the transverse band which
constitutes group A. (text-fig. 4, A). These lines may be nearer
together on the back than on the sides, and are sometimes much
serrated. The bands, 7. e. groups A, gradually become further
apart from each other, with the result that the markings are
slightly fewer in number than in the primitive type. For the
present I shall call this annulated form of marking Type Il. It
constitutes a distinct branch in the direct evolution of markings
(text-fig. 3,.a). Type III. evolves directly from the primitive

Text-figure 4.

arrangement, in exactly the same way as Type II. up to the point
where A 1 is completely divided, but distinct from A 2 and 3.
A 1, 1 are, however, squarish, and form a square group with A 2
and 3, which they equal in size. In the course of evolution A 4,
B 2, 3, and 4 reappear (text-fig. 4, B), and are also all of equal
size. In this type A 4 belongs to the dorso-lateral series, being
situated immediately above its position in the lateral series. in
Type I. Inthe evolution from Type I. to Type III. the series
equalise, the dorsal narrowing and the dorso-lateral broadening.:
The general pattern is formed, therefore, of three series of equal

176 MISS JOAN B. PROCTER ON THE

and alternating spots (text-fig. 4, c). Group A can only be
distinguished from B’by the fact that these spots are more square
than those of the latter. In this type the ground-colour is not
usually darker at A, as is the case in the preceding types.

Text-figuxe 5.

Further development of this type is shown on a specimen from
St. Lucia (Cat. p. 536, spec. v). On this snake both the dorso-
lateral and lateral series are absent, or but faintly marked, the
markings thus consisting of two squarish spots—A 1, 1—followed
by one roundish spot—B 1—all of which are for the most part
confluent with those of the opposite side.

VARIATION OF THE PIT-VIPER. Lig

The fourth type—Z. jararacussu (text-fig. 5)—constitutes the
third branch in the evolution of the markings. A 1, 5, 2,6,and 3
become confluent, forming a subcrescentic marking, in the middle
of which A 4 may be present or not. All the primitive spots of
group B unite together to form a dark triangular area, which in
the course of evolution becomes confluent with group A—. é.,
the apex of A with the base of B—along the dorsal line. At this
point the markings darken, and the usual brownish, greyish, or
olive ground-colour becomes a bright clear yellow. Finally, more
and more black pigment becomes deposited at A and B, until the
markings have usurped the place of the ground-colour, even the
crescentic marking (A) being partially filled in with black. The
general effect of Type IV. is therefore a black snake, with yellow
forked markings issuing transversely from the yellow ventrals,
which are checkered with black. The dorsal surface of the head
is completely black, owing to the development and confluence
between the streaks of the dorsal and dorso-lateral series. The
post-ocular streak is separated fom this area by a narrow stripe
of the clear yellow ground-colour, which continues slightly below
the commissure of the jaws.

The most striking feature about this type of marking is the
clear yellow of the interspaces in the dorsal pattern; in other
types this is only found on the ventral surface of the snake, the
dorsal ground-colour always exhibiting a superficial powdering of
dark pigment. It therefore seems probable that in the course
of evolution this pigment has all aggregated in the spots of
Groups A and B, and, as more and more becomes deposited, these
areas further encroached upon the yellow ground-colour.

V. PHYLOGENETIC RELATIONS BETWEEN THE TYPES.

In tracing the evolution of markings from one type to another,
I have assumed that the anterior parts represent the primitive
pattern, as in lizards and many mammals (ex. Lacerta muralis,
Equus quagga). It might appear, however, that in LZ. atrow this
is not the case, the more regularly spotted type, characteristic of
the West Indies representing the original pattern, from which the
clustered, annulated, and forked markings of C. and 8. American
forms were derived. On careful consideration I cannot adopt
this view for the following reasons :—1. The manner in which
the pattern of the Jararacussu develops from Type I. into the
complicated design of Type IV., and develops on each individual
posteriorly until the tail is a uniform black, cannot be looked upon
as primitive, and if this elaborate pattern, become simplified,
is a further development of the clustered type of marking, the
other forms must also be more advanced. 2. I hold the greater
number of spots to be the more primitive (and for this assumption
there is much correlative evidence among other snakes and
lizards). The clustered pattern is therefore less advanced than
the more evenly spotted, for in the former groups A and B

Proc. Zoot. Soc,— i918, No, XII. 12

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Taste oF PARTICULARS RELATING TO SOME SpeciMENS or LZ. ATROX,

Viz. : Colour variety and form of marking. Form of scales, at the anterior (u), middle (m), posterior (p), of the body ; either

of the affinis (af), intermediate (in), ov atroa (at) types; number of ventrals (v), pai

of subcaudals (c), and dorsal

* Mr. Boulenger tells me that the locality of this specimen is not to be trusted, as it forms part of the old collection, and bears no
register number. It was entered as Craspedocephalus lanceolatus in the appendix of Gray’s Cat. Col. Sn. (1858) p. 226 (spec. g)
and was referred to Lachesis lanceolatus by Mr. Boulenger. ee

scales (s). Number of markings on the right (r) and left (1) sides of the snake, Locality.
————— ae aes ae fxs as = eS
Scales. Markings. |
Colour Variety. Form of Number of Number of Locality.
low ore Com a ie =)
a m p eS € a oa 1
Var. affinis | . a ae
Almost black ..... af af aul 27 OU SeU emit:
Indi orconsarn in it 225 27 22 2 St. Lucia.
Primiti 5 at at 1833 24 | Atoyac, Mexico.
ve | fe
. af af 213 Di | British Honduras
i af 202 23 Malaga
F j ie |) 23 ‘equame
(2 and yg.) at af 218 28 Trinidad.
Printitive at out 198 27 a
” a af af 190 2 b!
Very primitiy in aft 198 27 c
Primitive .. af af | 193 27 d
as af af | 191 27 | e
te af af | 192 27 f
e in af | 192 27. &
ca af nf | 199 25 h
i af nf | 197 27 i
BS af af | 195 7 j
3) Soro Sauscoer | af af-|} 191 25 k
Tendency to var. jararaca . af af 216 25 1
Primitive peach af af ; 200 25 m |
.s af af 199 Py/ me {fr
x af af | 191 25 o
ns in af ; 196 2a p
a af af | 193 27 q
x) af at 188 26 | ty
4 af af 189 27 s
D af af 194 27 | t
Very primitive af af 189 2’ | u
Primitive 4 é af af 197 25 | v
Tendency to f. typica, primitive in in 194 25 | w
Primitive . : in af 212 27 x
in af 192 24 y
Very primitive (deformed specimen).. at af 164 27 vA
Primitive fs af af 200 24 ‘Trinidad. 2
Very primitive af af 200 27 | ”» 3
= o
Primitive af af. af | 181 67 19 21 | Berbice. 1
x af af af 195 60 19 19 4
a ae af 209 ns is a8 ios de Cacuta, Colombia. |
a a a 90 5 19 ‘ayaria. |
Tendency to var. jararaca at at in 175 61 16 16 W. Ecuador.
” mn ay at at at ae Ss | Bs 18 Sao Paulo, 1
at at in 5 5 9 18 2
af laf af || 185 66 | 19 21 3 3
af af af | 195 63 18 16 4 4
Intermediate between affinis and jararaca af af af 196 55 25. | 16 17 Berbice.
Var. jararaca, |
at at at 208 60 24 16 17 Martinique ®,
Be af af at 185 63 97 19 18 Rosario de Cacuta.
UGS B ROSS cccrooeacie-eneeccenis erebereetanauecececced|| fle at at 200 P 24 | anterior unmarked} Rio Janeiro.
i at at at 197 60 25 16 16 Brazil.
Intermediate between var. affinis, jararaca, and af at af 186 63 25 17 18 Berbice. 1
ENP U RECO) eernmenies uariner Bee copenate ae af af af 189 64 24 14 16 7 2
| |
Forma typica. Z
Indistinct .... at at af 228 62 30 22 21 Guadeloupe. |
at at ae nae of pS BS Be Martinique. 1 |
; in in ct 5 2: . 2
Dorsal Series only at at at 204. 69 26 19? 22? | St. Lucia.
Modified af af af 192 60 23 obscure Venezuela.
Indistinct ... af af af | 179 69 24 Fe Berbice. 1 |
ie af af af 178 = «6B 19 19 ir 2 |
ndistinct ... af af af 194 71025 16 16 Pampa del Sacramento.
Modified in in in 195 70 25 obscure Pozuzo, Peru.
odified ._. af af af 196 56 24 18 16 Sarayacnu.
Intermediate between var. is and j af af at 200 65 25 7 18 W. Ecuador. 1 |
en var. affinis and Jararacussu.}| Af BF AE | ie BB 25 19 18 2 |
Var, fararacussn ...cessesevecsecerseeeee ged ape an af ay) GB 20 20 | Sao Paulo. 1 |
at in af 184 53 25 19 19 ms 2 |

180 MISS JOAN B. PROCTER ON THE |

consist of 6 spots.each, whilst in the latter group A consists of 5
and B of 4. Besides these two points, I attach importance to
the fact that the eight primary series of spots never extend
beyond the head.

As we have already seen, Types II., III., and IV. are all
directly evolved from Type I.; but their relative positions have
yet to be discussed. Of the stages of development evinced
between each of these three and the primitive type, many show a
tendency for the markings to break up into the spotted type
(Type III.) just before the tail. Nearly all specimens with mark-
ings ranging from Types I. to II. show this peculiarity, also all
forms intermediate between Types I. and III. In the series
from Types I. to IV. this is the case in intermediate forms, but
Type IV. itself maintains its direct line of development, the
black markings encroaching more and more upon the yellow
ground-colour towards the posterior per of the snake, the end of
the tail being black.

It will therefore be seen that phylogenetically, the markings
of Type III. are not to be regarded as so primitive as those of
Type Il., and that Type IV. fee the most modified.

In the next chapter, on classification, varieties will be based
on these types of markings in conjunction with their geographical
distribution.

Having traced the markings of ZL. atrov from the most
primitive onwards, it is of some interest to try to trace these
markings back to the hypothetical four-paired streaks of primitive
snakes.

The first stage towards this would be the equal development of
group B with group A, and the commencement of the fourth or
lower labial series on the body. The series would then become
more equalised, until a pattern were reached of 4 rows of equal
and alternating spots—independent from those on the oppo-
site side—which would be but the result of the breaking up
of the primitive streaks. But Z. atrov is very far ahead of this
prototype, and such cases as might appear at first to be an
approximation to it, prove on careful consideration to be
secondary, an instance of deceptive apparent reversion.

VI. CLASSIFICATION.

It will be seen from the foregoing that no very definite
boundaries can be traced between the varieties’ of L. atrox,
whether based on structure or on coloration, so complete are the
links connecting them. I can, however, recognize four principal
forms based chiefly on markings, more or less 1n conjunction
with geographical distribution, which I think afford on the whole
a more satisfactory basis for classification than the characters of
scale-structure, etc.

The first of these forms is ZL. atror, var. affinis, which ex-
hibits the more primitive type of marking (Type I.), viz., the

VARIATION OF THE PIT-VIPER. 181

characteristic chevron, or triangle, which, however, varies slightly
according to localities.

The most primitive type of marking is not found in the
W. Indies proper, or south of Colombia and the Guianas, but
prevails north as far as Mexico, and appears to be constant in
Trinidad. Specimens of this form have scales with the high short
keel. In the W. Indies proper a less primitive variation occurs,
in which the elemental spots are more confluent and the pattern
less distinct; the scales may either be of the short high-keeled
or long low-keeled type. As this variety extends southwards
the markings, though still undoubtedly of Type I., frequently
approach those of the Jararaca, or Jararacussu in type, and the
scaling may conform to either type. Thus the ZL. atrox and
L. lanceolatus of Dr. Vital Brazil, which are marked simply with
dark triangles on the sides, undoubtedly approach the Jararaca
of Wied in pattern. A pattern of markings intermediate between
the primitive. type of this variety and that of the Jararacussu
occurs in W. Ecuador. JZ. atrox, var. affinis, is the most widely
ranging of the four varieties (Mexico to Southern Brazil and
Peru). All forms of it are marked with triangles or chevrons,
with their apex turned towards the mid-dorsal line. Group A is
always darker than B, and encloses an area darker than the sur-
rounding ground-colour. The ventrals may be uniform cream-
colour or yellowish, or blotched or speckled with black. In the
young the elemental spots, when present, are more distinct, a
certain amount of confluence or fading taking place with age.

Form 2, L. atroa, var. jararaca, has been considered a distinct
Species since the time of Wied, but as it does not differ from the
typical Z. atrow in any appreciable way, save in the dorsal mark-
ings, I consider it to bea variety only. The Jararaca inhabits
Brazil, and is especially prevalent in Bahia, Sao Paulo, and
Rio. It is recognized by its annulated form of markings, pre-
viously described as Type IT., consisting of dark, darker-edged
bands, narrower on the mid-dorsal line than on the sides. These
bands are more than their own width apart, the interspaces being
lighter and unspotted. The ventrals may be uniform, spotted, or
powdered with black.

Figures of the Jararaca are given in Wied, Abbild. Nat. Bras.,
under the name of Cophias atrox; Wagler in Spix, Serp. Bras.
pls. xix., xxi., under the names of Bothrops megera, B. leuco-
stigma, B. tessellatus, and B. teniatus; and in Jan, Icon. Oph.
xlvii. pl. 11. under B. atrozx.

A typical specimen of this variety could not be confounded with
the preceding, but in the British Museum collection there are
some young specimens from Berbice (Cat. Sn. p. 539, spec. e—x),
which are exactly intermediate between these three varieties.

Form 3, the typical Z. atrox (ZL. lanceolatus) occurs principally
in the West Indies, which is the acknowledged home of the
Fer-de-lance, but also in the Guianas, Venezuela, and curiously
in Peru, in a somewhat modified form. The markings of this

182 ON THE VARIATION OF THE PIT-VIPER.

variety are of the spotted type, described as Type III., consisting
of three longitudinal series of subequal and alternating dark spots,
more or less uniformly distributed on a lighter ground-colour ;
the scales are usually of the low, long-keeled type. In the
modified form from Venezuela, Berbice, and Peru the spots of
group A are darker than those of group B, the whole pattern
thus having a less even appearance. In some cases the markings
are very indistinct, only the principal spots of group A (A 1, 1,
2, 3) being discernible. The scales may be of either type.

Form 4, LZ. atrow, var. jararacussi, is the distinct colour variety,
which Lacerda regarded as a valid species in 1884, and which is
still maintaind as such by Dr. V. Brazil. There is, however, no
difference between the Jararacussu and JZ. atrow except that of
colour and markings.

The Jararacussu inhabits Brazil, Sao Paulo, and Rio, and is
distinguished from the preceding forms by its peculiar markings
described above as 'l'ype IV., and the fact that the interspaces in
the black areas are bright yellow instead of the brown, grey, or
olive shades of all the other varieties. The pattern appears to
consist of transverse yellow forks issuing from the yellow ventrals,
which are blotched with black. The tail is black. The head of
the Jararacussu is extremely characteristic, the dorsal surface
being of a uniform black, from which the black post-ocular
streaks are separated by a narrow streak of yellow, continued
obliquely from the yellow throat. A coloured plate of this variety
is given in Dr. Brazil’s ‘ Défense contre l’Ophidisme’ (2nd edition),
pl. 14, under the name of Lachesis jararacussu. It is also
represented in Jan’s [conographie, 47me livr., pl. il. fig. 3, as
Bothrops atrow.

a

ON DEATHS IN THE SOCIETY’S GARDENS, 183

9. Report on Deaths of Animals in the Gardens in 1917.
By J. A. Murray, M.D., B.Sc., F.Z.S., Director,
Imperial Cancer Research Fund, Pathologist to the
Society.

[Received April 23, 1918: Read April 23, 1918. |

The accompanying tabular statement of the numbers of deaths
of mammals, birds, and reptiles (including amphibia and fishes)
in 1917, shows only minor differences from the figures recorded
in 1916 by Professor Plimmer or from the average numbers in
the previous 5 years. In 1917 rather less than 25 per cent. of
the animals dying had been less than 6 months in the Gardens,

TABLE I,
MamMats. | BirRDs. REPTILES.
613 1940 446 In Gardens, 1.1.17.
215 745 368 Admitted in 1917.
828 2685 814. | TOTAL.

GS" oe oo. 2. 62 | Under 6 months. 7.
167 eee ad 61 ae 80 yas Over 6 months. Died.
284 | 20°6 17°4 Per cent. of total.
283 23°4 286 | 4 in 1916.

27°0 | 23'3 31°2 i in 1911-15.

As has been pointed out by Dr.Chalmers Mitchell, little statistical
value attaches to such tabular statements of combined mortality
of species with widely different liability to disease and duration
of life. The only conclusion to be drawn is that there has been
freedom from severe epidemics during the year.

Table II. summarises the results recorded under the more
important causes of death for the chief mammalian orders, and
for mammals, birds, and reptiles generally. ‘These have been
compiled from the careful records made by Professor Plimmer as
regards the first 7 months of the year. There was an interval
of nearly two months before I entered upon the duties of
pathologist, and the incomplete data recorded in this interregnum

184

DR. J. A. MURRAY ON

eo is

Ou

General Diseases.
Tuberculosis 4.06 see ae
Mycosis . ee eRe ites ae HON
Septicgomias yi. yo ds.ccetensnabane
AN DSCOSSae heater ose cores
Peritonitis .. Bea ratty Soe
Cestodesies.. . tae eee
Filariasis

Ascarides ei i eae 2
Heemogregarines ...... ...........

. Diseases of Respiratory

Organs.
A Cel ectasiseat castcacsssseunatiaas
PVCUIMONIA: © 255. .cucnens wore
Pleurisy ..

. Diseases of Heart.

Degeneration of Heart-muscle.
V alvular disease

. Diseases of Liver.

Hepatitis

. Diseases of Alimentary y Tract.

Gastro-enteritis

Gastric ulceration ............
Enteritis

Gastritis .. DA Ay eae
Tntestinal obstr uction VARA Aarne
Intussusception Dee oe
Prolapse of Rectum 225 3..2609. <0:

. Diseases of Urinary and

Generative Organs.
Nephritis sicko eeeh same
Salipimeitis 2. Ailes. ces

. Various.

Carcinoma... ethane
SVARCOMIA Caran trey yet tn ree emeae
Heomormliages is nhs sccadaee
Injuries discovered BEST:
mortem Sas

Starvation and malnutrition! A
Killed by companions, es &e.
Killed by order .

Not diagnosed ee oneal

Primates.

O7

—_

bb re OD

ll el

|
|

a]

@mwre

ore aS | Carnivora, |

9 |

TABLE IT.
MamMALs.
ue ee
| 2 Sele
~~ x ee ee
ela|+=|°e
laa] oe | @ | a)
° By) ers lcs
Se \> eS
288
HI
"Oe aii
Bevel
| |
asf 1 ||
26 9 | 2. |
a oi |
|
1 fl Pe
OO coh all
1 |
2 |
|
2 3
1 |
38
13 |

—:

ee:

| Marsupialia.

Total. |

Dore Sw

a

Brrps. | REPTILEs. |

46 Ao |

R) oes
9

:

I |

D) 2
a

197 ‘| 22

Sea

Seeel

88 4

7 1

2

i |

| 4

5

5

|

| a |

liar kean ae

10+ || 1

ig +) 1

81 78

are partly responsible for swelling the totals under the heading

‘not diagnosed ” in Table IT.

Protozoan Parasites.—In addition to the three reptilian deaths
in Table IT. ascribed to Hemogregarines, these parasites have
been observed in a number of snakes dying from other causes.

DEATHS IN THE SOCIETY’S GARDENS. 185

They have revealed an apparently definite dimorphism of the
schizonts in the circulating blood, a long slender form which
produces great deformation of the red cell, the haemoglobin being
retained, and a short stout form which produces much. less
deformity but exhausts the hemoglobin very rapidly. The
significance of the two forms and their relation to the schizogony
in the liver-cells remain to be determined by further study, for
which a considerable material has been preserved.

Malignant new Growths.—In two mammals death was ascribed
to carcinoma and in two birds to sarcoma. Of these one case of
squamous-cell carcinoma of the fauces and palate of a Dingo,
probably arising from the epithelial covering of the tonsil, has
been examined personally. In addition a new growth was found
on examination of a Leopard which died with bronchiectatic
cavities in the lungs. The growth in this case proved to be an
adeno-carcinoma lying on the ventral surface of the trachea in
the thorax. The tissue of origin could not be determined.

(Two further cases have been observed in 1918: a carcinoma
of the liver in a Marsh-Buck and a teratoma of the testis in a
Golden Eagle.)

Comparative Pathology of the Thyroid.—The special value of
the material occurring in the Prosectorium consists in the
opportunity offered for a comparative study of pathological and
physiological problems which are of general interest, and special
attention has been directed to the thyroid gland. The thyroid
occurs throughout the whole vertebrate series and, with the
exception of the cyclostome fishes, presents the same histological
picture in all. From studies in man and mammals in various
pathological conditions an important réle has been ascribed to it
in the reaction of the body toa variety of intoxications. The
results of physiological experiment indicate other important
functions both in health and disease. It was therefore of interest
to note the contrast in the appearance of the gland in cold- and
warm-blooded animals respectively in severe infections. In
warm-blooded animals dving under. these conditions ane
congestion of the whole gland is practically constant. Nothing
of the kind has been Sneountered in the reptiles examined:
although a large proportion presented severe septicaeemic con-
ditions after death. This result is unfavourable to the view that
the thyroid plays the part of a neutraliser of toxic substances in
the body. It is in much better harmony with the view that the
changes in the thyroid in these conditions are the expression of
its participation in the heat-regulating mechanism of the body.
In poikilothermic animals one would expect these changes to be
absent.

Effect of Diet on the Thyroid.— Although attention has been
drawn to the effect of a meat diet on the thy roid, no observations
are recorded on the results of the natural experiment pr esented
by the occurrence of meat-eating and plant-eating groups in birds
and mammals. The analysis of the observations in birds presents

186 ON DEATHS IN THE SOCIETY’S GARDENS.

many difficulties, and will be undertaken later. In mammals,
however, the comparison is much easier, and the preliminary
survey which has been carried out shows that in Carnivora the
thyroid is from two to three times as large as in Ungulata of
corresponding body-weight. The key to the problem is probably
furnished by Marine and Lenhart’s work on thyroid enlargement
in trout kept in hatcheries. These authors have shown that the
goitrous condition, which so frequently supervenes, is due to the
diet of horseflesh with which the fish are fed and that it can be
obviated and even cured by supplying a ration of sea-fish. Their
investigations have shown that the constituent deficient in the
horseflesh diet is iodine. ‘The necessity for fixing the inadequate
amounts present in flesh is met by a hypertrophy of the gland.
When adequate amounts of iodine are supplied either in the form
of sea-fish or soluble iodides, the enlargement subsides.

THE SECRETARY ON ADDITIONS TO THE MENAGERIE, 187

EXHIBITIONS AND NOTICES.
February 5th, 1918.

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

The Secrerary read the following Report on the Additions
made to the Society’s Menagerie during the months of November
and December, 1917 :—

NOVEMBER.

The registered additions to the Society’s Menagerie during the
month of November were 38 in number. Of these 28 were
acquired by presentation and 10 were received on deposit.

The number of departures during the same period, by death
and removals, was 88.

Amongst the additions special attention may be directed
OS
1 Brindled Gnu (Gorgon taurinus) 2, from South Africa,
deposited on November 21st. ,

1 Wilson’s Bird of Paradise (Schlegelia wilsonz), from Waigiou,
and 2 Himalayan Goldfinches (Carduelis caniceps), presented by
Alfred Ezra, F.Z.S.

1 Lesser Vasa Parrot (Coracopsis nigra), from Madagascar,
presented by The Marquess of Tavistock, F.Z.5.

DECEMBER.

The registered additions to the Society’s Menagerie during the
month of December were 17 in number. Of these 7 were
acquired by presentation and 10 were received on deposit.

The number of departures during the same period, by death
and removals, was 81.

Amongst the additions special attention may be directed
tou

1 Grey Seal (/Halicherus grypus), from the North Sea, presented
by 'T. Witherwick on December 24th.

1 Kuropean Flamingo (Phaenicopterus roseus), from S. Kurope ;
2 South-American Flamingoes (Ph. chilensis), from S. America ;
2 Black-necked Swans (Cygnus melanocephalus) and 2 Coscoroba
Swans (Coscoroba coscoroba), from 8. America: deposited on
December 31st.

*

Mr. D. Sera Sirs, F.Z.S., Curator of Birds, exhibited and
remarked on a series of lantern-slides made from photographs

188 MR. T. E. WHITEHEAD ON THE WILD DINGO.

of Reptiles taken in the Gardens, directing special attention
to those showing feeding habits of the Black Cribo (Owyrhopus
clelia), which attacks and devours poisonous snakes.

The Wild Dingo (Canis familiaris, var.).

The Secrerary read the following letter from Mr. Thomas E.
Whitehead :—

“During a recent visit to England, I spent a delightful day at
the Zoological Gardens, but was rather surprised at the animals
exhibited as Dingos. Now, Sir, I spent a great many years as a
stockman and drover in the back country of South Australia,
New South Wales, Queensland, and Northern Territory, besides
travelling and prospecting over a large part of Western Australia
during the past 23 years, but I have never met with a Dingo
anything like those I saw at the London Zoo.

‘““T have enquired from my friends if they could tell me what
part of Australia they could have come from, and I am told that
many years ago a Mastiff got away with the Dingos about Mount
Kosciusko and crossed the breed in that district. So probably
that is where your Dingos came from. They are too tall, too
long, too thick muzzled, and too short-haired to be true Dingos.
The true Dingo resembles the fox a great deal, but is larger, his
nose is pointed, forehead fairly broad, has longish hair, and a tail
almost as bushy as the fox. There are two colours, according to
the colour of the country he inhabits. Those in the Tableland
and Mountainous districts are usually of a sandy brown colour,
while those in the yellow sand-hill country are usually sandy
yellow, and have longer hair than those on the Tablelands. I
once saw a pack of about a dozen of the darker kind in the fore-
noon on the south side of Lake Yandandaninna, and towards
evening I saw a pack of the lighter colour on the north side of
the lake ; they had come for a drink from the sandhills to the
northward.

‘‘Dingos never bark, neither do they learn to bark when
tamed. The Bushman can always distinguish between the howl
of a Dingo and that of a Domestic Dog. Like the fox, he is
very partial to poultry, and is a terror to poultry-owners in the
new townships. He will kill sheep by the dozen just for the
fun of doing so. If hungry, he will rip a hole in the flank and
eat his fill of the liver and leave the rest. I once heard sheep
rushing on a bright moonlight night. On going to investigate I
saw our Overseers prize Kelpie rounding up about 300 sheep,
while in the centre was a Dingo slut enjoying herself by killing
sheep wholesale. I fired my rifle at the Kelpie but missed, then
fired at the slut, and next morning tracked her up by blood
marks and found her nearly dead, so quickly despatched her.

ON A NEW SPECIES OF THE GOLDEN MOLE. 189

She had killed 11 sheep and had bitten many more, some of
them dying during the next few days.
‘“THos. EK. WHITEHEAD,
262 Bulwer Street,
North Perth,
Western Australia.”

On a New and a Rare Species of the Golden Mole
(Bematiscus).

Dr. R. Broom, C.M.Z.S., exhibited specimens of two rare
South-African Golden Moles, and made the following re-
marks :—

Hitherto the giant moles from the eastern part of Cape
Colony have been referred to Lematiscus trevelyani, but it 1s
quite manifest that there are at least two species. ‘The specimen
which I make the type of this new species was sent to me by
Miss Ivy Lesch from St. Cuthberts, [solo, and as it differs very
markedly trom Bematiscus trevelyani of the Pirie Bush or any of
the hitherto described species, [ have much pleasure in naming it
after the discoverer, to whose interest in natural history we also
owe Lematiscus transvaalensis.

Bematiscus lesche is a moderately large mole, probably about
180 mm. in length, but as I have only the dried skin, it is
impossible to be quite sure of the length. It is manifestly con-
siderably smaller than 2B. trevelyant. The fur is of a fairly
uniform dark brown—lighter on the abdomen. On the lips the
short fur is somewhat reddish brown. On the upper side of the
head the fur is also rather hghter than further back, owing to
the short under-fur being pale reddish brown. On the back and
greater part of the body the fur is about 18 mm. in lengti
dark slaty very fine under-fur with coarse-tipped hairs whose tips
of 7 mm. overlap the under-fur. The tips have the last 2 o1
3mm. dark brown and the more proximal § light brown.

The following are the measurements of aie aa aws of the front
foot :—I1st 2°5 mm., 2nd 6 mmn., 3rd 11 mm., 4th 1:5 mm.

The following are the principal measurements of the skull —

mm.
Greatest lemoth..nccij,.ssaks naceee Oe 35°6
Creatests widths ..: ett. cces eee 21:4
Gueatest hetoht <i. sc.css.ncsanceasden.- LZ
Width between orbits.................. (a5)
DentalSeries: icsisels ca voeeienecedden 14
Miglaie SOMES). iacsdeahsreaces sts kes. es 8
Width across the premolars ......... 10°8
Width between last premolars ...... 4:6

The skull has the large bulla for the malleus well projecting
as in B. villosa and B. transvaalensis, and the affinities are much
closer with these two species than with B. trevelyani. The mole

190 THE SECRETARY ON ADDITIONS TO THE MENAGERIE.

from Maritzburg described by Dobson, and doubtfully referred by
him to B. villosa is, I think, undoubtedly a distinct species, and
may be called Bematiscus dobsoni.

To the genus Lematiscus there would thus be referred five
known species—JS. villosa and &. dobsoni from Natal, B. trans-
vaalensis from the Transvaal, and Bb. trevelyant and B. lesche
from Eastern Cape Colony.

The second exhibit is a specimen of the rare Golden Mole—-
Chrysochloris sclateri. Originally discovered in the Nieuwveld,
north of Beaufort West, and named in honour of Mr. W. L.
Sclater, it was next discovered at Movija in Basutoland, 350 miles
from the original locality. In an unexplored country a species
occurring at two such remote localites would not excite any
wonder, but in South Africa our researches on the fauna have
gone so far that we can say not only that the species has never
been found in the intermediate area, but that no Golden Moles
inhabit the northern part of Cape Colony. Six years ago I
discovered tracks of Chrysochloris on Compass Berg, and I have
since been fortunate, through the kindness of Miss K. Jansen, of
Wilgebosch, New Bethesda, in getting a specimen which proves
to be also C. sclaterit. This discovery brings the known localities
130 miles nearer, and suggests that the species may originally
have extended to Basutoland by the Winterberg and Stormberg.
Possibly it may yet occur in these localities. The black mole of
the south coast, which I named C. duthiec, is an extremely closely
allied species. It occurs at Knysna, and possibly extends east-
wards to Port Elizabeth. C. sclatera with its 40 teeth is probably
near the ancestral form from which C. hottentota and allied forms
with 36 teeth are derived.

February 19th, 1918.

Dr. A. Smurrn Woopwarb, F.R.S., Vice-President,
in the Chair.

The Secrerary read the following Report on the Additions
made to the Society's Menagerie during the month of January
1918 :—

The registered additions to the Society’s Menagerie during the
month of January were 26 in number. Of these 16 were
acquired by presentation, 8 were received on deposit, and 2 by
purchase.

The number of departures during the same period, by death
and removals, was 80.

Amongst the additions special attention may be directed to:—

1 White-nosed Coati (Vasua narica), from Tampico, presented
by H. C. Banbury on January 19th.

AN AFRICAN CIVET ATTACKING HUMAN BEINGS, 191

_ 1 Otter (Lutra lutra), from Hampton Court, presented by
H. Tagg on January 18th.

2 Straw-necked Ibises (Carphibis spinicollis), from Australia ;
1 Scarlet Ibis (Hudocitmus ruber), from Para; 1 Himalayan
Monaul (Lophophorus impeyanus) $, from Himalayas; 1 Peacock
Pheasant (Polyplectrim chinquis) 3, from Barmah; 2 Swinhoe’s
Pheasants (Genneus swinhow) 3 9, from Formosa, presented by

Major The Hon. Waldorf Astor, F.Z.8.

Dr. Smrruw Woopwarb, F.R.S., V.P.Z.8., exhibited a copy of
an incised drawing of a hunted deer, pierced by arrows, made by
Paleolithic man in the cave of La Pena, San Roman de Candamo,
Asturias, Spain. It was lately published by Dr. Hernandez-
Pacheco in no. 17 of the memoirs of the Spanish commission on
prehistoric investigations.

An African Cwet attacking Human Beings.

The following letter, communicated by Professor Pouuron,
F.R.S , F.Z.8., was read from Captain G. D. Hale Carpenter,
M.D., giving an account of a case which had come under his
personal observation in which an African Civet attacked human
beings.

“On the night of Dec. 10-llth, 1917, an Indian and an
African, employées of the railway, were sleeping in the verandah
of the station building when the latter was awakened by a bite
on his toe, and found to his alarm what he thought was a young
leopard—it was a very dark night and without a moon. It
viciously attacked the two men, but they managed to catch it
and throw it down a well, as they had no stick or other weapons
handy: they both came to hospital in the morning to have their
wounds dressed—the African had a contused and punctured
wound on the ball of the great toe: the forearm of the Indian
was superficially lacerated. When they brought up the body of
the culprit, which I had decided in my mind would prove to be a
Serval cat, as I thought it unlikely the two men would without
weapons have overpowered a Leopard cub. But to my astonish-
ment the draggled carcase was that of a rather small, old, Civet !

‘The men said there had been two of them.

‘“¢T should think undoubted instances of unprovoked assault by
a Civet on mankind must be rare.

““G. D, Hate Carpenvrer, M.D.,
Capt. Uganda Med. Service.”

“On the Central Railway of ex-German Kast Africa, :
17 miles W. of Tabora,
December 12, 1917.”

192 A CHEHTODONT FISH WITH ARABIC CHARACTERS,

Professor EK. W. MacBrips, M.A., D.Sc., F.R.S., F.Z.8., gave
an account, illustrated by lantern-slides, of bis recent investi-
gations into the development of the Sea- Urchin (£chinocardiwm
cordatiune).

March 5th, 1918.

Dr. A. SmitrH Woopwarb, F.R.S., Vice-President,
in the Chair.

Mr. D. Sera Smita, F.Z.S., exhibited skins of the Hoatzin
(Opisthocomus cristatus), and described the habits and distribu-
tion of the species, illustrating his remarks with lantern-slides.

A Chetodont Fish with Markings resembling Arabic
Characters.

My. C. Tare Reean, M.A., F.R.S., F.Z.S., exhibited photo-
graphs of an Indo-Pacific Chetodont Fish (Holacanthus sema-
circulatus Cuv. et Val.). Two of these had been sent to him
by Major H. R. Cartwright, Commandant of Police, Zanzibar ;

Text-figure 1].

they were of a specimen that had been sold in the fish-market
for a penny; the man who bought it was going to eat it and cut
off the tail and threw it on the ground; another man picked it
up and called out that it had writing on it, and, indeed, on one
side of the caudal fin was written in old Arabic characters
‘ Laillaha Ialah” (There is no God but Allah) and on the
other side “Shani-Allah ” (A warning sent from Allah). The

THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 193

news caused great excitement in the market place; the fish
changed hands at rapidly increasing prices until 5000 rupees was
offered ; the fish was regarded as sacred and Major Cartwright,
who put it in formalin for the owner and had it photographed,
thought it might become the object of pilgrimages.

The other photographs exhibited by Mr. Regan were of ex-
amples of this species in the collection of the British Museum
(Natural History), and illustrated the changes in coloration that
take place during growth. In young specimens the ground-
colour is almost black, the body is crossed by a few curved white
stripes, and the posterior half of the caudal fin isclear. In larger
fish 3 or 4 inches long the general ground-colour is paler, but is
_ dark-spotted ; the stripes have increased in number by the addi-
tion of narrow ones between the original ones, and still narrower
ones between these, so that there may be as many as 24 stripes
instead of the original 6; also the posterior part of the tail is
now darked and is crossed by 3 pale stripes, of which the posterior
2 may be discontinuous or connected or replaced by Jongitudinal
bars ; it is these which may simulate Arabic characters. This
stage was described as Holacanthus alternans Cuv. & Val., and
from now onwards the ground-colour becomes paler and the dark
spots better defined, the pale stripes disappear anteriorly, and
posteriorly still increase in number but break up into spots and
vermiculations, so that a fish of seven inches long has quite a
different appearance, and indeed was described by Bleeker as
a distinct species, H. lepidolepis.

March 19th, 1918.

Dr. A. Smita Woopwarp, 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 February
1918 :—

' The registered additions to the Society’s Menagerie during
the month of February were 19 in number. Of these 14 were
acquired by presentation, and 5 were received on deposit.

The number of departures during the same period, by death
and removals, was 71.

Amongst the additions special attention may be directed
LO" ——

2 Stanley Cranes (Tetrapteryx paradisea), from South Africa,
and one Black-necked Crowned Crane (Balearica pavonina),
from West Africa, presented by W. H. St. Quintin, F.Z.S., on
February 21st.

1 American Bison (Dison americanus), from N. America,
deposited on February 15th.

Proc. Zoou. Soc.—1918, No. XIII. C3:

194 GON FOSSIL ROSTRAL TEETH OF SAW-FISHES.

April 9th, 1918.

Dr. A. Suita Woopwarp, F.B.S., Vice-President,
in the Chair.

The Sfcrerary read the following Report on the Additions
made to the Society's Menagerie during the month of March
1918 ;— %

The registered additions to the Society’s Menagerie during the
month of March were 31 in number. Of these 23 were acquired
by presentation, 6 were received on deposit, 1 by purchase, and
1 in exchange.

The number of departures during the same period, by death
and removals, was 93.

Amongst the additions special attention may be directed
UO
1 Jungle-Cat (felis chaus) 2, from India, purchased on
March 9th.

2 Hog-Deer (Axis porcinus), $ 2, from India, deposited on
March Sth.

1 King Penguin (Aptenodytes patagonica), from South Georgia,
received in exchange on March 19th.

Miss L. E. Currsman, Assistant Curator of Insects, exhibited
specimens of an Hast-African homopterous insect, /éyreea nigro-
cincta, sent by the Curator of the Museum at Nairobe.

Special interest is attached to these insects by reason of their
habit of clustering in colonies on a plant-stem In imitation of the
inflorescence. There is a coloured form and a green form of
the adult insect, and when the latter are found clustering on the
apex of the flower-spike, they bear an extraordinary resemblance
to unopened buds.

An interesting account of a similar species was published by
Dr. Gregory in his book ‘The Great Rift Valley,’ with a sketch
of the insects. They have also been figured in a paper read by
Mr. Hinde before the Entomological Society. in 1902.

Dr. A. SmrtrH Woopwarp, V.P.Z.S., exhibited fossil rostral
teeth of Hopristis and Pristis, and referred to the progressive
changes in the rostral teeth of the Pristide, or saw-fishes, during
geological time. In the Cretaceous genus, Sclerorhynchus, these
teeth only differ from the ordinary shagreen in their enlargement
and the elongation of their apical portion. Their pulp-cavity is
restricted to the basal half, and the distal portion is traversed by
several very irregular large vascular canals, mainly longitudinal
in direction. In the genus Hopristis, not known later than the

' ON FUR SALES IN THE UNITED STATES. 195

Eocene period, the teeth are still loosely ranged along the edge of
the rostrum, not in deep sockets ; but their structure is: newly
that of Pristzs, the vascular canals being only much more irregular
and not yet clearly the centres of well-marked hexagonal prisms.
Pristis itself, with the rostral teeth in deep sockets, and with the
well-known prismatic structure, ranges from the Eocene period
onwards.

Mr. G. A. Bouuencer, F.R.S., exhibited the head of a Hydro-
cyon goliath Blgr., from the Congo, a fish attaining a length of
four feet, the largest and most formidable representative of a
genus of the family Characinide inhabiting the principal rivers
of Africa. The Nile species is known as the Kelp-el-Bahr, or
River-dog, of the Arabs, and a third species, common in the
Zambesi and Limpopo systems, is called Tiger-fish by the
colonists.

The object of this exhibition was to show the enormous shark-
like teeth to which special interest attaches owing to a recent
publication by Dr. Eastman, who has pointed out their close
similarity to the fossils known as Onchosaurus Gervais (originally
referred to the Mosasaurs), Jschyrhiza Leidy, and Gigantichthys
Dames, which appears to indicate the existence of the Characinide
as far back as the Upper Cretaceous, a range in time which
Mr. Boulenger had predicted as probable thirteen years ago, and
whieh is of special importance in explaining the present distri-
bution of this family *.

April 23rd, 1918.

A. Ezra, Esq., Vice-President, in the Chair.

Mr. D. Serx Smirn, Curator of Birds, exhibited and made
remarks on a Zulu Head-dress made of the plumes of the male
Long-tailed Whydah, Chere progne.

The SECRETARY called attention to an advertisement that had
recently appeared in the London Press, announcing Fur Sales by
Public Auction about to take place in the United States. The
sales in question were only examples of what took place annually
in London and other important commercial centres. The numbers
advertised were smaller than usual, no doubt on account of
the War, but they included very large quantities of animals the

* Of. C. R. Ac. Sci. Paris, clxvi. 1918, p. 197.

196 ON SPECIMENS AFFECTED WITH RICKETS.

extinction of which could not be far distant, unless measures
were carried out to protect them. Inthe opinion of the speaker,
which was confirmed by the Meeting, there was urgent need for
drastic measures to protect Mammals. The protection of Birds
appealed to popular sentiment, and was zealously advocated by
many influential organisations. The danger that threatened
Mammals was even greater, and, on account of their higher
intelligence and more sensitive nervous organisation, the cruelty
involved in the methods of hunting, trapping, and killing them
was incomparably greater.

Professor Woop-Jones, F.Z.S., Honorary Acting Prosector,
exhibited and made remarks on specimens from the Prosectorium
illustrating the effects of Rickets. He also exhibited a set of
anatomical preparations useful for teaching purposes, made from
material obtained from the Society’s Collection.

INDEX.

1918.—Pages 1-196.

[New names in clarendon type.

Systematic references in italics.

(z.8..) indicates additions to the Society’s Menagerie. |

Agama sanguinolenta, 14.
Alamosa bipunctella, 65.
piperatella, 65,
Anerastia ablepta, 88.
acrophea, 88.
anemopsis, 88.
argosticha, 88.
baliora, 88.
ephestiella, 88.
—— erasmia, 88.
—— korbi, 79.
—— pleurochorda, 88.
xiphimela, 88.
Aptenodytes patagonica (z. 8. L.), 194.
Atascosa quadricolorella, 104.
Aurora longipalpella, 106.
AVES:
Head-dress made of plumes of Chere
progne, 195,
Axis porcinus (z. s. .), 194.

Balenoptera acutorostrata, 148.
borealis, 149.

physalus, 151.

Balearica pavonina (z. s. L.), 193.
Bandera binotella, 89.

carneella, 90.

Proc. Zoou. Soc.—1918, No. XIV.

_ Bandera cupidinella, 90.

perlepidella, 90.
subluteella, 90.

—— virginella, 90.
Baptotropa, gen. n., 116.

——— tricolorella, 116.

| Barberia affinitella, 131.

_ Bematiscus leschze : exhibited, 189.
| Biafra concinnella, 115.

rhodinella, 115.

Bison americanus (z. s. L.), 198.
Boiga trigonatwm, 16.

Cabnia myronella, 131.
Calamotropa, gen. n., 91.
pulverivena, sp.n., 91.
Calera punctilimbella, 59,
albicostella, 62.
Canis familiaris, communication, 188.
Carduelis caniceps (z. 8. L.), 187.
Carphibis spinicollis (z. 8. u.), 191.
| Ceara albifasciata, sp, n., 63.
ee discinotella, 63.

Chere progne: exhibited, 195.

Chortonceca, gen. n., 80.

—— eurysticha, 81.

-—- leucocraspia, sp.n., 81.
14

x1

Chortoneca minimella, 81.

minoralis, 81.

Chrysochloris selateri: exhibited,
190.

Cenochroa californiella, 59.

iibella, 58.

—— inspergella, 59.

-—— monomacula, 59.

Coenotropa, gen, n., 89.

—— limitella, sp. n., 89.

Commotria albinervella, sp. n.,
Ee

—— arrhabdella, sp. n., 108.

—— castaneipars, sp. n., 112.

—— crassiscapella, 110.

—— enervella, sp.n., 112.

—— erythrograpta, sp. n., 109.

— invenustella, 108. ‘

—— laticostella, 108.

—- mesiella, sp. n., 108.

—— miosticta, sp. n., 110.

—— neurias, sp. n., 109.

oberthuri, 108.

opacella, 108.

phlebicella, sp. n., 112.

—— phoenicias, sp. n., 110.

phycttella, 109.

—— propheella, sp. n., 112.

—— rhodochroa, sp.n., 111.

——-- rhodoneura, sp. n., 110.

—— rosella, sp. n., 111.

roseopennella, 108.

—— rufidelineata, sp. n., 110.

——— rufimedia, sp. n., 109.

—— tripartella, sp. n., 109.

-—— venosella, sp. n., I11.

Coracopsis nigra (z. 8. L.), 187.

Coscoroba cescoroba (3.8. L.), 187,

Critonia hilgerti, 121.

holorhoda, 120.

-—— leucopleura, sp. n., 120.

ochracealis, 121.

—— pheoneura, sp. n., 119.

promelena, 120.

—— purpureotincta, 120.

. —— rhodessa, 121.

roseistrigella, 120.

sarcoglauca, 121.

INDEX.

Critonia sarcoida, sp. n., 121.
—— subconcinnella, 120.
Cygnus melanocephalus (uz. 8. L.), 187.

Delphinus delphis, 159.
Dembea venulosella, 64.
Discofrontia normella, 119.

Echinoecardium cordatum: lantern-
slide, 192.

Ematheudes crassinotella, 115.

euchlytella, 115.

— lentistrigalis, 114.

—— paleatella, 114.

pseudopunctella, 114.

punctella, 114.

straminella, 114.

——-- tunesiella, 114.

varicella, 115.

—— vitellinella, 115.

Emmalocera actinoleuca, sp. n.,
127.

albicostalis, 1381.

—— anerastica, 129.

aurifusella, 128.

—— bifidella, 128.

—— carnatella, 130,

costella, 130.

cremoricosta, 130.

—— depressella, 128.

—— diversella, 130.

—— endopyrella, sp. n., 127.

— eremochroa, sp.n., 130.

erythrinella, 130.

laminella, 129.

latilimbella, 129.

— leucocincta, 128.

longiramella, 127.

— lucidicostella, 129.

monochromella, 130.

ornatella, 127.

——- polychroella, sp. n., 128.
pulverealis, 127. ;

radiatella, 127.

sanguifusalis, 128.

— strigicostella, 128,

INDEX.

Emmatlocera subfasciatella, 181.

tricoloralis, 130.

—— umbricostella, 129.

umbrivittella, 131.

varregatella, 129.

Epidauria chionocraspis, sp. n.,
2. |

discella, 92.

—— perfasciella, sp. n., 92.
—— pheniceella, 92.

—— strigosa, 92. |
—— subcostella, sp. n., 93.

—-- transversariella, 92.

Eremias intermedia, 14.

-— (Mesalina) guttulata, 15.

—— velox velox, 14.

Eryx miliaris, 17.

Ethiotropa, gen. n., 116.

— pyromerella, sp. n., 116.
Eruovoey.

Insecta: [Exhibition of specimens |
of an East-African homopterous |
insect, 194.

Eudocimus ruber (z. s. u.), 191. |
Eumeces schneideri, 15.
scutatus, 15.

Felis chaus (z. s. u.), 194.
Fondunkia translucidella, 131.
Fossifrontia leuconewrella, 107.

Genneus swinholi (z. s. u.), 191.
GEOGRAPHICAL:
Insecta: An _ East-African homo-
pterous insect from Nairobi, 194.
Mammatta: On Cetacea stranded on
the British Coasts, 147.
Pisces: Head of Hydrocyon goliath
from the Congo, 195.

Reptitia:. Reptiles from the River |

Tajan, 11.
Globicephala melena, 157.
Gorgon taurinus (z. s. L.), 187.
Grampus griseus, 157.
Gymnodactylus caspius, 14.
—— microlepis, sp. n. (Pl. I. fig. 1), |

AN

x1ll

Halicherus grypus (%. 8. 1.), 187.

Holacanthus semicireulatus (Fig. 1),
photograph, 192.

Hosidia ochrineurella, 64.

Hydrocyon goliath ; exhibited, 195.

Hyperoodon rostratus, 153.

Hypsotropa acnidias, 77.

adumbratella, 73.

— albivenalis, 70.
— approximella, 77.

— hipartella, 78.

—— biscrensis, sp. n., 69.
— castella, 78.

— chionorhabda, sp. n., 70.
—-- contrastella, 68.

—— cremoricosta, sp. n., 7+.
—— diaphza, sp. n., 75.

—— dyseimata, 7d.

—— endorhoda, sp. n., 76.
— euryzona, 75.

— fuscostrigella, 68.

—- fusifasciata, sp. n., 71.
—— graptophlebia, sp. n., 76.
— heterocerella, 68.

—— icasmopis, 7d,

— ichorella, 72.

—— illectalis, 69.

—— infumatella, 71.

— laropis, 78.

—— laterculella, 74.

—— leucocraspis, sp. n., 77.
— leucophlebiella, 77.

—— limbella, 71.

—-~— luteicostella, 70.
monostidza, sp. n., 76.
neurica, 78.

niphopleura, 75.
niveicosta, sp n., 74.
ochricostella, sp. n., 70.
ocraceella, sp. n., 69.
—— papuasella, 78.

—- paucipunctella, 72.

— periphea, sp. n., 74.

—— pervittella, sp. n., 68.
—— phyrdes, 738.

polyactinia, 76.

—— polystictella, sp. n., 77.
— punctinervella, sp. n., 72.

Riv

Hypsotropa purpurella, sp.n.,71.
—— pusillella, 73.

—- psamathella, 76.

— — guadripunctella, 76.

—— ramulosella, 7 6.
rhodochroeila, sp. n., 69.
rhodosticha, 77.
rosescens, sp. n., 79.
sabuletella, 73.

sceletella, "72.

semirosella, 69.
solipunctella, 72.
stereosticha, 75.
subcostella, sp. n., 68.
syriacella, 72..
tenutcostella, 69.
tenuinervella, 73.
tripartalis, sp. n., 74.
unipunctella, 71.
vertheiimsteini, 72.
vulneratella, 72.
zophopleura, 71,

INsncra.

On the Pyralids, subfamily Hypso-
tropine : syetematic, 55; On visible
and characters in Silk-

03: On
Kast-African homopterous insect,
194.

Ityrea nigrocincta: exhibited, 194.

invisible

worms: structure, an

Lagenorhynchus aculus, 158.

—— albirostris, 158. _

Laurentia albivenella, s). n., 90.
——- inclarella, 91.
Leotropa, gen. n., 64.

—— papuanensis, sp. n., 65.

-~—— phoenicias, sp. n., 64.

—— Sarcina, sp. n., 64.
Lophophorus impeyanus (z. s. u.), 191.
Lutra lutra (z.s. u.), 191.

Mabuia sepiemteniata, 1d.

INDEX.

MAMMALIA.

Skull of Rana tigrina: structure, 1:
External Characters of the Lemurs
and of Zarsius: structure, 19; On
Oetacea stranded on the British
Coasts; structure, 147; Communi-
cation of letter on the Wild Dingo,
188; On a new species of the
Golden Mole: structure, 189; On
the rare Golden Mole, 190: Copy
of drawing of a hunted Deer, 191;
On an African Civet attacking
Human Beings, 191: On Fur Sales
in the United States, 195.

Martia arizonella, 118.
Megalophota, gen. v., 117.
—— leonella, sp. n., 118.
Menuthia nanellu, 6).
Mesodiphletia crassivenia, 62.
—— deliquella, 62.
-—— ochraceella, sp. n., 62.
-——- rosellu, 02.

stricticostella, 62.
Mesoplodon bidens, 155.
Metacrateria, gen. n., 79.
—— metallactis, 79.

——- miasticta, sp. n., 79.
—— yerirrorella, sp. n., 79.
neh

Monoctenocera brachiella, 122.

pulverulella,

—— leucania, 122.

Morpnouocy. See Structure.

Nasua narica (z. 8. u.), 190.

Nutri tessellata, 17.

Navasota chionophlebia, sp. n.,
66. :

-_— discipunctella, sp. n., 67.

-—— hemapheella, sp. n., 66.

——— hebetella, 66.

—— leuconeurella, sp. n., 66.

—— myriolecta, 67.

—— persectella, sp. n., 66.

—— syriggia, sp. n., 67.

Oligodon hamptoni, sp. u. (Fig. 1),

9.

INDEX. xV

Ollia honoponerella, 104.

parvella, 104.

santaritela, LOA.

Orcinus orca, 156.

Osacia lineolella, 123.

Oxyrhopus cleelia: lantern-slide, 188.

PATH 0.40 G Y.

Animals in the Society's Gardens,
183.

Patna brunneicostella, sp. n., 117.
eboricostella, 117.

—-- venatella, sp. n., 117.
Pectinigeria pamponerella, 104.
violodis, 104.

Peoria albidel/a, 78.

Phocena phocena, 159.
Phenicopterus chilensis (z. s. L.), 187.
—— roseus (%.s.u.), 187.

Physeter catodon, 151.

Piscus.

On the development of the Sea-
Urchin, 192; A Cheetodont Fish
with Arabic characters, 192; On
fossil rostral teeth of Eopristis and
Pristis, 194; Exhibition of head
of a Hydrocyon goliath, 195.

Polyocha achromatella, sp. n.,

126.

—— cinerella, 125.

detritella, 126.

flagrantella, 125.

—-— foucarti, 126.

—— fuscicostella, sp. n., 126.
gensanalis, 125.
leucopleurella, 124.

—— neuropterella, 126.

—— plinthochroa, sp. n., 124.
—- rhodesia, 131.

—~—- sanguinariella, 125.

stipella, 125.

—— strigivenella, 125.

venosa, 125,

vesculella, 125.
Polyplectrum chinquis (z. 8. u.), 191.
Powjadia cyttarella, 104.

—— leuconeura, 104.

Powjadia pimella, 104.
Prinanerastia incarnatella, 80.
—— lactealis, 80.

— lotella, 80.

nitidicostella, 80.
Prophtasia amplichea, sp. n., 1095.
—— bistriatella, 106.

—— epiteuxis, sp. n., 15.
-—_— glaucophza, sp. n., 105.
-—— platycerella, 104.

—— pyrostrota, sp. n., 106.
—-— sphalmatella, sp. n., 105.

Ragonotia dotalis, 124.
Raphimetopus, gen. n., 78.
—— ablutella, 78.

—— spinifrontella, 78.
REPTILIA !

On a new Snake of the Genus
Oligodon : systematic, 9; Reptiles
from the River Tajan: systematic,
11; On the variation of the Pit-
Viper: structure, 163; Lantern-
slides made from photographs of
Reptiles, 188.

Rhinaphe apotomella, 82.
—— approximella, sp. n., 82.
biseriella, 85.
brunneovittella, 85.

castanealis, 838.

—— celsella, 83.

-—— conspersella, 86.

ella, 86.

—— enervella, 8&4.

—— endonephele, sp. n., 87.
—-— flavescentella, 84.

-—- furvimacula, sp. n., 87.

haploschema, 84.
hemirhodella, 85.

-_— holophea, 8°.

—-— ignetincta, sp. n., 87.
—— infumella, 86.

-—— lateriticlla, sp. n., 86.
leucoteniella, 8d.

——- lotricella, 85.

—— mictochroella, 87.

——- neesimella, 83.

XV1 INDEX.

Rhinaphe nigricostalis, 83. Saluria inficita, 96.

—— pailidicosta, 84. | —— insignificella, sp. n., 103.

-—— pheostrotella, sp. n, 85. | —— interpunctella, sp. n., 103.
plinthina, 83. ont lentistrigella, sp. n., 97.

—— sangirensis, sp. n., 85. | —— macreila, 100.

~-— seeholdi, 87. | —- maculivittella, 102.

-—— separatella, 84. magnesiella, 103.

-—— signicollis, 87. | ~—— mesomelanella, sp. n., 103.

——_— stictella, 84. i | eee minutella, 95.
syssema, 86. | -— museella, 28.

ne - taliella, 86. | —— neotomella, 101.

— vectiferella, 88. | —— neuricella, sp. n., 98.
venelia, 83. — nigritella, 99.

— venilineella, sp. n., 86. —— nilgiriensis, sp. n., 95.

—— virginella, 84. --— ochridorsella, 97.

Rhodochrysa superbella, 89. —— opificelia, 101.

—— ostreella, 102.
—— paranensis, 98.
Sabormania pia, 131. — paucigraphella, 94.

Saborina forcipella, 122.

pectigerella, 102.

——— papuacola, sp. n., 123. —— pleurostichu, 98.
vicina, 128. | -— proleucella, sp. n., 99.
Saluria albivenella, 99. | —— psammatellia, sp. n., 94.
—— anchridis, 98. | pulverosa, 102.
-—— arcticostella, 100. | — rhodopheza, sp. n., 100.
ardiferella, 99. | rosella, 97.
hreviculella, 101. | -— rufella, sp. n., 101.
—— callirhoda, 103. _ — — semirosella, sip, 1) one
cancelliella, 102. | —— sepicostella, 95.
——— carnescens, sp. n., 94, | —— spurcella, 96,
—-— claricostella, 95. | —— stictella, sp. n., 96.
——— ctenucha, 94. | ——- stictophora, sp. n., 96.
—— desertella, sp. n., 97. | —-— subcarnella, 100,
devylderi, 100. | -—- subcostella, sp. n., 100.
—- dichroella, 101. | ——- tenuicosta, sp. n., 98.
—— distictella, sp.n., 101. | —— tetradella, 102.
ensiferella, 102. it see tripartetla, 103.
erodelia, 93. —— varicosella, 99.
flammella, 94. | —— verecundella, 97.
——- flavicosta, sp. n., 96. | Schenectadia merilesella, 131
—— flavipurpurella, sp.n.,98. = Schlegelia wilsoni (2. s. u.), 187.
—— floscella, 96. Siboga albimediella, sp.n., 113.
—- furvella, 100. — dialeucella, sp. n., 113.
—— gemmatella, 100. —— falsella, 113.
— glareosella, 95. —— zeavora, sp. n., 113.
~—- grammivena, sp. n., 99. Statina bifasciella, 60.
—~—— hemiphealis, 94. — cashmiralis, 60.
— holochroa, 102. —— gaudiella, 60.

INDEX.

Statina punctilineella, 60.
rhodobaphella, 60.
roseotinctella, 60.
—— rosinella, sp. n., 60.
STRUCTURE.
Insecta: On visible and
characters in Silkworms, 133.
Mammnatia: Skull of Rana tzgrina, 1;
External characters of the Lernurs
and of Tuarsius, 19; On Cetacea
stranded on the British Coasts,
147; On a new species of the
Golden Mole, 189.
ReptiutraA: On the variation of the
Pit-Viper, 163.
Sudania subcostella, 88.

invisible

Tampa dimediatella, 63.
Taphrometopon lineolatum, 16.

XVil

‘Tetrapteryx paradisea (z. 8. L.), 193.
Tinerastia discipunctella, 61.
Jissirella, 61.

Tursiops truncatus, 157.

VARIATION:

RepritiA: On the variation of the
Pit-Viper, 163.

Pisces: A Chetodont Fish with
Arabic characters; On fossil ros-
tral teeth of Eopristis and Pristis,
194.

Vipera lebitina, 15.

Zamenis diadema, 16.
rhodorahchis, 16.
Ziphius cavirostris, 154,

PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEBYT STREET,

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PROCEEDINGS

OF THE
GENERAL MEBTINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY

OF LONDON.

1918, pp. 197-310,

wita 2 Puates and 30 TEx'r-FIGURES.

PRINTED FOR THE SOCIETY,
SOLD AT ITS HOUSE IN REGENT’S PARK,
LONDON:

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

Lalas

OF THE

COUNCIL AND OFFICERS

OF THE

ZOOLOGICAL SOCIETY OF LONDON.

1918.

Patron.
His Masesty Tuer Kina.

COUNCIL.
His Grace Tore Duke or Beprorp, K.G., F.R.S., President.

Tue Hon. Cecrz Barine, M.A.)

Ricuarp H. Burnes,
M.A.

Lr.-Cot. S. Monckton Copet-
MAN, M.D., F.R.S.

Esq.,

CHARLES DrumMonD, Ksq.,
Treasurer.
ALFRED Ezra, Esq., Vice-

President.
Capr. Haroup 8. FEerRGuson.
Cart. Huan §. GLADSTONE,
M.A.
SipNeEY F. Harmer, Esq., M.A.,
Sc.D., F.R.S., Vice-President.
Pror. James P. Hit, D.8c.,
WitiiAM Huntsman, Esq.
Sir Epmunp G. Lover, Br.,
Vice-President.

Pror. Ernest W. MacBripe,
D.Se., LL.D., F.R.S., Vice
President.

Con. Sir A. Henry McManon,

G.C.M.G., K.C.I.E.,G.C.V.O.,
Case.

P. Cuatmers Mircney, Esq.,
C.B.H., M.A., DiSe:, Tales
E.R.S., Secretary.

ApvpriAN D. W. Potiocg, Esa.

THE LorD QUEENBOROUGH.

THe MARQUESS OF SLIGO, F.S.A.,
Vice-President.

AvuBYN Trevor-Batryg, KsqQ.,
M.A.

AntHony H. WINGFIELD, Esq.

A. Smira Woopwarp, Ksq.,
LL.D., F.R.S., Vice-Pre-

sident.

PRINCIPAL OFFICERS.
P. Cuatmers Mircuett, C.B.E., M.A., D.Sc., LL.D., F.RB.S.,

Secretary.

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

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

Lieut. Epwarp G. BouLencer, Curator of Reptiles. ;

Prof. H. Maxwexu Lerroy, Curator of Insects.

JoHN Barrow, Accountant.

W. H. Coun, Chief Clerk.

LIST OF CONTENTS.

1918, pp. 197-310,

EXHIBITIONS AND NOTICKS.

Mr. R. I. Pocock, F.R.S., F.Z.S., Curator of Mammals.
Exhibition of, and remarks upon, a series of the
molar tecul OF MIO VHANUS.9...5..cc;hecessssoalgsete ores seen

Mr. E. Heron-Auien, F.L.S., F.R.M.S., F.Z.S. Lantern-

exhibition of Arenaceous Foraminifera...............+6%

The Secretary. Report on the Additions to the Society’s
Menagerie during the month of April, 1918............

Mr. C. Tate Reean, M.A., F.R.S., F.Z.S. Account, illus-
trated by lantern-slides, of the Freshwater Fishes of
(TES Or 03 WN 0 eae ee an Pe Rea Pe ne rer ene re Ae

The Secrerary. Report on the Additions: to the Society’s
Menagerie during the month of May, 1918 ............

Lt.-Col. S. Monckton Coprman, M.D., F.R.S., F.Z.S.
Exhibition of, and remarks upon, a colony of Burrow-
mas bees on Primrose: Pill: 2... ccasenactaesseceteytsecne et

The Secretary. Report on the Additions to the Society’s
Menagerie during the months of June, July, August,
dase puenmmle4r LONG.“ ai.c jesse vedas cee cen ey cienie ood ae

Mr. R. I. Pococx, F.R.S8., F.Z.8., Curator of Mammals.
Exhibition, on behalf of Mr. EK. Gerrard, of the skin
of an abnormally coloured Red Deer.............. aes

Prof. H. M. Lerroy, F.Z.S., Hon. Curator of Insects.
Account, illustrated by lantern-slides, of Wheat
Weeeval- jm Australia ii nis eeccetee cic cae cdeedeamesed ses ee,

Page

303

303

303

303

304

304

306

307

iv

Mr. D. Seru-Smitn, F.Z.8., Curator of Birds. Exhibition
of a mounted specimen of a hybrid Cockatoo .........

Dr. R. T. Lereer, M.D., D.Sc., F.Z.S. Lantern-exhibition
on Diagnosis of Helminth Infections.....................

Dr. R. T. Lerrer, M.D., D.Sc., F.Z.S. Demonstration on
the “new” Rabbit disease . 2. sscc0..:ceresseee ee eee

Prof. H. M. Lerroy, F.Z.8., Hon. Curator of Insects.
Exhibition of a series of lantern-slides from photo-
graphs taken in the Zoological Gardens, Sydney,
Ni Si. Wales jester cic atc cae cee oes d onto nenaiietne oie eee eee

The Secrerary. Report on the Additions to the Society’s
Menagerie during the month of October, 1918.........

Miss K. Lanver, F.Z.8. Description and exhibition of the
method of preparing skeletons by the use of trypsin...

Mr. E. Harscuex. Description and exhibition of investi-
gations into the forms assumed by drops and vortices
of gelatin in various coagulants ........:is--s50s4er eee

Prof. F. Woop-Jonus, F.Z.8. Exhibition of a cast and set
of Rontgen-ray photographs taken from a Chimpanzee
which had died from pulmonary tuberculosis .........

PAPERS.

10. Comparison between the Lower Jaws of the Cynodont
Reptiles Gomphognathus and Cynognathus. By Dvr.
Branistav Perronizvics. (Text-figures 1-8.).........

11. On a new Genus of Extinct Muscardine Rodent from
the Balearic Islands. By Dosxoruma M. A. Bars,
Hon. M.B.0.U. (Plate I. and Text-figures 1 & 2.)...

12. A Case of Hermaphroditism in a Lizard, Lacerta

viridis. By Nort Tayiur, B.Sc. (Lond.). (From

the Zoological Department, University of London,
University College.) (Text-figures 1-3.)

eeceeoesreoeersereoose

Page

308

308

308

309

309

310

310

310

i

209

Vv

Page
13. On two new Elasmobranch Fishes (Crossorhinus juras-
sicus, sp. nov., and Protospinax annectans, gen. et sp.
nov.) from the Upper Jurassic Lithographic Stone
of Bavaria. By Arruur 8 .1ra Woopwarp, LL.D.,
HBS ateVecZe, EA CO Le oo intudausgaa ten ieed cans 231
14, The Function of Pathological States in Evolution. By
MORLEY AINQBERIS 5: t60 2 scss00e02 00s vot ec eect oe sae enrasniaens 237

15. Notes on the Beavers at Leonardslee, 1916-1918. By
Sir EpMunD G. Lopmr, Bart., Vice-President Z.S. .... 255

16. On the Madagascar Frogs of the Genus Mantidactylus
Blgr. By G. A. Boutmenesr, F.R.S., F.Z.8. ......... 257
17, Ciliary Action in the Internal Cavities of the
Ctenophore Pleurobrachia pileus Fabr. By JAmEs
F. Gemuiut, M.A., M.D., D.Sc., F.Z.8. (Text-
ATU eSey lie rges Ntararesai2,.'o she Seale Serato Se Dswe ihe coahs eels ve secon 263

18. On Seymouria, the most primitive known reptile. By
D. M. 8. Watson, M.Sc., Capt, R.A.F., Lecturer
in Vertebrate Paleontology, University College,
Hondons.2:(Rext-figures 11D.) cavin sce scses cause eee nes 267

Alphabetical List of Contributors .......0+00..c0escsererseseeeees vii

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CONTRIBUTORS,

With References to the several Articles contributed by euch.

(1918, pp. 197-310.)

Page

Bats, Miss DorornEa M. A., Hon. M.B.O.U.

On a new Genus of Extinct Muscardine Rodent from

the Balearic Islands. (Plate I. and Text-figures 1 & 2.)... 209

Bouuenaer, G. A., F.R.S., F.Z.S.
On the Madagascar Frogs of the Genus Mantidactylus

BUI ee a Bat ee etree Hh vamucte herein nilotiews: Diubove aces waee tele 257

CopEMAN, Lt.-Col. 8. Monckton, M.D., F.R.S., F.Z.S.

Exhibition of, and remarks upon, a colony of Burrow-

mie. bees on brimarose till iy. oo s iweseescccs.etssh cela eeachoce 304

Gemini, James F., M.A., M.D., D.Sc., F.Z.S.
Ciliary Action in the Internal Cavities of the Cteno-

phore Pleurobrachia pilews Fabr. (Text-figures 1 & 2.)... 2

Harscuer, E.

Description and exhibition of investigations into the
forms assumed by drops and vortices of gelatin in various

coagulants Bg EOE tet fe eae OR OE ae TRO ena

Herron- Auten, E., F.L.S., F.R.M.S., F.Z.S.

Lantern-exhibition of Arenaceous Foraminifera......... 303

vill
Lanper, Miss K., F.Z.S.

Description and exhibition of the method of preparing

skeletons by the’ use of trypsin. ..-..2.0..2..00..5.... eee

Lerroy, Prof. H. M., F.Z.S., Hon. Curator of Insects.
Account, illustrated by lantern-slides, of Wheat Weevil
In. Australias? 3) o.aeds ak ate occa. cee ae ee

Exhibition of a series of lantern-slides from photo-
graphs taken in the Zoological Gardens, Sydney, N. 8.
Wales a5 cdi teh cad Basis cans cess aren eee

Lerrer, Dr. R. T., F.Z.S.

Lantern-exhibition on Diagnosis of Helminth Infec-

Lover, Sir Epmunp G., Bart., Vice-President Z.S,
Notes on the Beavers at Leonardslee, 1916-1918 ......

MitcHELtL, P. Cuatmers, C.B.E., M.A., LL.D., F.R.S.,
F.Z.8., Secretary to the Society.

Report on the Additions to the Society’s Menagerie
during the month-of April, 1918< 2... 522.0...<1 cae
Report on the Additions to the Society’s Menagerie
during the month of May, 1918 ....0..0...... ee
Report on the Additions to the Society’s Menagerie
during the months of June, July, August, and September,
POV S Gee ec ne eaoeee. Fe oO eee One ee er 6 Sh:

Report on the Additions to the Society’s Menagerie |

during the month of October; V91S .... 2.2.3... ....- omen cer

Perronigvics, Dr. BRANISLAV.
Comparison between the Lower Jaws of the Cyno-
dont Reptiles Gomphognathus and Cynognathus. (Text-
FiGures EBs) | oe ssn. cee cence ase Macs Ooh sere eee Saree eee

Page

310

307

309

308
308

259

303

304

306

309

ix

Pocock, R. I., F.R.S., F.Z.S8., Curator of Mammals.

Exhibition of a series of the molar teeth of Elephants.
Exhibition, on behalf of Mr. EK. Gerrard, of the skin

of an abnormally coloured Red Deer ........................

Recan, C. Tats, M.A., F.R.S., F.Z.S8.

Account, illustrated by lantern-slides, of the Fresh-

water Fishes of Great Britain ............... eee Oe

Ropers, Morey.

The Function of Pathological States in Evolution......

Sera-Smiru, D., F.Z.8., Curator of Birds.
Exhibition of a mounted specimen of a hybrid

eal EAC LOO Gare ee eC ar eee a

TayLER, Nort, B.Sc. (Lond.).

A Case of Hermaphroditism im a Lizard, Lacerta

endian bext-neures’ 1-3.) 1. 0a. isn. ac ehs oevkree enna vers

Watson, D. M.S8., M.Sc., Capt. R.A.F.

On Seymouria, the most prunitive known reptile.
(lesctsmcmecnt — WOi Sahar ce eats, te thine atl oko awe ee

W oov-JoNness, Prof. F., F ZS.
Exhibition of a cast and set of Rontgen-ray photo-
graphs taken from a Chimpanzee which had died from

MUMMON ay UM DETCUlOGIG neers Gear egos seeeanwee oe eta g team aes

Woopwarp, Dr. A. Smirn, F.R.S., V.P.Z.S.

On two new Elasmobranch Fishes (Crossorhinus juras-
sicus, sp. nov., and Protospinax anneclans, gen: et sp. nov.)
from the Upper Jurassic Lithographic Stone of Bavavria.
Giga Voml a) ear str fh ens seh G totus iedan os ev okaae nl anne tes

Proc. Zoou. Soc.—1918. b

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310

PROCEEDINGS
OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
or Tite

ZOOLOGICAL SOCIETY OF LONDON.

PAPERS.

10. Comparison between the Lower Jaws of the Cynodont
Reptiles Gomphognathus and Cynognathus. By Dr.
BRANISLAV PETRONIEVICS *.

{Received March 6, 1918: Read May 7, 1918.]
(Text-figures 1—8.)

Being occupied at the end of last year (1917), when in London,
with the problem of the double articulation of the lower jaw,
which is to be expected in the direct reptilian ancestors of the
Mammals, I was led to examine the lower jaws in the specimens
of the Theriodont Reptiles that are preserved in the Natural
History Museum. One of the skulls in question, that described
by Seeley in 1895 as one of the two specimens of Gomphognathus
from Lady Frere (8S. Africa), struck me on account of the strange
appearance of its lower jaw. On comparing carefully this latter
with the lower jaw in Cynognathus crateronotus, described by
Seeley in 1895, IL have arrived at the conclusion that the difference
between them is a Very considerable one, and that in the lower
jaw of Gomphognathus we have the greatest known development
of the dentary bone in any theriodont reptile, a development
representing the initial state of the double articulation, if not the
actual double articulation itself.

The main points of difference in the lower jaws of Gompho-
gnathus and Cynognathus are the following :—

1. In Gomphognathus the articular bones are placed inside and
laterally at some distance from the dentary bone, whilst in

* Communicated by Dr. C. W: Anprews, F.R.S., F.ZS.
Proc. Zoou. Soc.—1918, No. XV. 15

198 DR. B. PETRONIEVICS ON THE LOWER JAWS OF THE

Cynognathus they are attached closely to the dentary bone
(comp. the right ramus of the lower jaw of Gomphognathus
in text-fig. 1 with the right ramus of Cynognathus in text-fig. 7).

Text-tigure 1.

art. srt gu.

4
a/\
artsrt-dent.; | art

oO ‘

Gomphognathus : inferior surface of skull and mandibles. 3.

ang.; angular.

ang.dm.; angular lamina. ¥
ang.n.; angular notch.

aré.; articular.

art.srf.dent.? ; articular surface of dentary.
art.srf.qu.; artieular surface of quadrate.
dent.; dentary.

dent.n.; dental notch.

prart.; prearticular.

sang. ;, supra-angular.

spl.; splenial.

st.?; stapes.

2. In Gomphognathus the posterior end of the dentary bone
goes almost as far back as the articular bones (comp. text-figs, 2

CYNODONT REPTILES GOMPHOGNATHUS AND CYNOGNATHUS, 199

and 4), whilst in Cynognathus the articular goes farther back
than the dentary * (and it does this more externally than internally
—comp. text-fig. 5 and text-fig. 6, in which the length of these
portions of the dentary bone is the same).

3. In Gomphognathus the posterior edge of the dentary bone
covers externally the articular bones almost entirely (comp. text-
figs. 2and 4), whilst in Cynognathus the hind part of the articular
bones is clearly visible and considerably external to the dentary
bone (text-fig. 5).

Text-figure 2,

prang-

Gomphognathus: right ramus of lower jaw, external view. X 5.
Shaded part covered with matrix,

dent.; dentary.

in.con.dent.; incipient condyle of deutary.
pr.cor.; coronoid process of dentary.
pr.ang.; processus angularis.

In connexion with the first point we must add that Seeley’s
contention (comp. Seeley (1), 1895, p. 26), that the articular bones
of Gomphognathus were ‘somewhat displaced, being drawn away
laterally from the edge of the dentary bones,” must be wholly
rejected, because the lateral distance of these bones from the
dentaries is exactly the same on both sides (comp. text-tig. 1),
and the bony connexion of the almost horizontally situated
reflected angular lamina, which is well preserved on the left side,
is such (comp. text-fig. 1) that no doubt about the naturalness of
its position can be entertained. The angular notch limited by
this lamina is a large one, and comparatively large, also, is the
notch lying iu front of it, between the dentary on the one side,
and angular and splenial on the other (comp. text-fig. 1; the hind
part of the splenial is preserved only on the right side).
In Cynognathus the reflected angular lamina, whose position is
almost a vertical one, is preserved only partially (comp. text-fig. 6

* In Cynognathus platyceps the hind end of the dentary is more prolonged back-
wards than in Cynognathus crateronotus, but it did not reach the squamosal.
Comp. Seeley (2), 1895, fig. 29, p. 185, and Broom, 1904, text-fig. 100, p. 496.

200 DR. B. PETRONIEVICS ON THE LOWER JAWS OF THE

and text-fig. 7), but it must have been relatively smaller than
in Gomphognathus (comp. text-fig. 6).

In Gomphognathus there 1s a bony connexion between the
articular bones and the dentary (whose thickness could not be
established, but which probably is not a considerable one), this
bony connexion, forming the roof of the angular notch, being
well preserved and visible on the left side (comp. text-fig. 1). The

Text-figure 3.

Gomphognathus : right ramus of lower jaw, internal view. X 9.

Shaded part covered with matrix.

ang.; angular.

art.; articular.

er.?; coronold.

dent.; dentary.

in.con.dent.; incipient condyle of dentary.
prart.; prearticular.

spl.; splenial.

Text-figure 4.

Gomphognathus : left ramus of lower jaw, external view. X 2.

ang.; angular.

dent.; dentary.

in.con.dent. ; incipient condyle of dentary.
pr.ang.; processus angularis.

sSang.; supra-angular.

sutures between the articular bones are not to be traced in their
whole length in Gomphognathus (comp. text-figs. 3 and 4), while
they are clearly visible in Cynognathus (comp. text-figs. 5, 6,

CYNODONT REPTILES GOMPHOGNATHUS AND CYNOGNATHUS. 201

and 7): it is probable that the mutual limits of these bones are
in both eases the same. The suture between the articular and
prearticular is obliterated in both cases.

As to the second point. The ecoronoid process of the left
dentary in Gomphognathus is only partially preserved (text-fig. 4),

Text-figure 5.

Cynognathus crateronotus: right ramus of lower jaw, external view. X 4.
Shaded part covered with matrix.

ang.; angular.
art.; articular.
dent.; dentary.
sang.; supra-angular,

Text-figure 6.

7 \ cr prart. oat

‘ ca

ang. ' angn. .
analm

Cynognathus crateronotus: right ramus of lower jaw, internal view. XX 4,
Shaded part covered with matrix.

ang.; angular. dent. ; dentary,
ang.im.; angular lamina. prart. ; pre-articular.
ang.n.; angular notch. . sang.; Surangular.
art.; articular. spl.: splenial.

cr. 3 coronoid.

whilst the right dentary shows this process in its entirety, and
although the upper part of it is covered with matrix, its upper
edge is clearly visible from above (comp. text-fig. 2). As the hind

202 DR. B. PETRONIEVICS ON THE LOWER JAWS OF THE

end of this edge lies in direct prolongation of the lower part of
the posterior edge of the coronoid process uncovered by the new

Text-figure 7.

are.

Cynognathus crateronotus : right ramus of lower jaw,
inferior surface. X 2.

ang.; angular. dent.; deutary.
ang.dm.; angular lamina. prart.; pre-articular.
ang.n.; angular notch. sang.; surangular.

art.; articular.

preparation, so the naturalness of this posterior edge and conse-
queutly of the prominent backward condylar prolongation of the
right dentary (text-fig. 2) cannot possibly be doubted. As this

*

CYNODONT REPTILES GOMPHOGNATHUS AND CYNOGNATHUS. 203

condylar prolongation is damaged beneath and at its posterior end,
whose transverse section shows a triangular shape, the articular
surface of the condyle cannot be certainly established. On the
left side (text-fig. 4) only a part of the prominent backward condylar
prolongation is preserved, so that on the outer side of the left
jaw the hind end of the articular bones seems to go farther back
than the hind end of the dentary, while on the right side the
reverse seems to take place. But the hind end of the articular
bones on the right side is evidently damaged, so that it is not
impossible that ie dentary reached as far back as the articular
bones, and that a double articulation really did take place in
Gomphognathus *.

The possibility of the double articulation in Gomphognathus
follows also from the fact of two different articular surfaces
preserved in our specimen on the left side (comp. text-fig. 8).
A comparison with the corresponding region in the type skull of
Diademodon described by Watson (comp. fig. 3 in Watson 1911)
shows (comp. also fig. 8 in Watson 1911 representing the quad-
rate in Gomphognathus polyphagus) that only the triangular
hollow on the front face of the squamosal, in which the missing
quadrate was received, is preserved in our specimen on the left
side together with the two notches which received the two pro-
cesses of the quadrate (these notches being filled with matrix).
As there is in our specimen on the left of these notches and
in a more forward position another flat surface on the squa-
mosal, which lies in the same direction as the hind end of
the dentary (comp. text-figs. 1 and 8), it is quite possible that
into this flat surface was received the hind end of the condylar
prolongation of the dentary. Indeed, as the articular end in
our specimen did not reach the squamosal on the external side
of the dentary as in Gomphognathus kannemeyert (comp. fig. 1 in
Seeley (1) 1895, p. 5, and especially fig. 2 in Broom, 1904,
pl. xxxv.), or as in C2 ynognathus (comp. fig. 8 in Seeley (2) 1895,
p. 81, and fig. 1 in Broom, 1904, pl. xxxv.), so the flat surface in
question could be reached in our specimen only by the hind end
of the dentary (supposing that it was reached by the lower jaw
at all).

The third point is a corollary of the second. As the posterior
edge of the left dentary of Gompho gnathus, extending from the
angle up to the condylar prolongation, is almost undamaged
(comp. text- fig. 4), whilst the corresponding posterior edge of the

right dentary is not inconsiderably damaged, the difference in the
extent of the covering of the articular bones by the dentary from
outside in Gomphognathus and Cynognathus is strikingly shown

* The length of the two dentaries of Gomphognathus, as they are preserved, is
almost the same (149 mm.), when measured from the hind point of their symphysis.
But a certain asymmetry of them is not improbable, because when we compare the
length of the upper edge of the left dentary with the corresponding line in the
coronoid process of the right dentary, we find the first to be 68 and the second
66 mm.

204 DR. B. PETRONIEVICS ON THE LOWER JAWS OF THE

when text-fig. 4 is compared with text-fig. 5. So that, although the
hind end of the dentary in Cynognathus goes farther back on the
inner than on the outer side, it is clear from this comparison that it
did not reach the squamosal and that the possibility of an incipient
condyle of the dentary in Cynognathus is excluded.

Text-figure 8.

arts. dent. 4

Gomphognathus: articular surface of the left squamosal. Nat. size.

art.; articular,

art.sf.dent.?; probable articular surface of dentary.
art.srf qu.; articular sur‘ace of left quadrate.

dent.; dentary.

The inner Side of the jaws shows the same bones in_ both
exses (comp. text-fig. 3 with text-fig. 6). The comparison shows
that the splenial goes farther back in Gomphognathus than in
Cynognathus.

‘The essential difference above mentioned that exists in respect
to the relation of the articular bones and the hind end of the
dentary to the squamosal between the Gomphognathus-skull of our
specimen and the other specimens known under the names of
Gomphognathus and Diademodon (a general survey of which has
been, given by Watson, 1911, p. 397 seq.) suggests the well-
founded supposition that our specimen is generically different
from all the other specimens of the same kind, which are all
essentially similar in the above point. Beyond that, a comparison
between the lower jaw of Gomphogaathus kannemeyeri (figured
by Seeley (1) 1895, tig. 2, p. 8) and of Diademodon mastacus
(figured by Broom, 1905, fig. 1, pl. x.) shows that they are similar
in shape, while the shape of the lower jaw of our specimen
(comp. text-fig. 2), especially the shape of its coronoid process, 1s very
different. So that I agree with Watson in his identification of
Gomphoguathus and Diademodon for all the other specimens

—— ~~

a

4

CYNODONT REPTILES GOMPHOGNATHUS AND CYNOGNATHUS. 205

except the one in question, which alone has to retain the old
generic name of Gomphognathus ™.

Having finished the comparison between the lower Jaws of
Gomphognathus and Cynognathus, | will add here some general
remarks concerning the origin of the mandibular articulation and
the origin of the Ossicula auditus in Mammals, inasmuch as these
two very closely connected problems receive new light from our
comparison.

That the mandibular articulation in the most primitive mammals
(the hypothetical order Promammaha of Gregory—comp. its
definition in Gregory, 1908, p. 164) was a double one, our
Gomphognathus-skull puts this point beyond question, as it shows
that in. this advanced Theriodont the dentary almost if not
actually reached the squamosal. That the dentary articulation
was situated in the same plane with the articular articulation, and
not in front, is also a point put beyond question. ‘That the two
articulations have worked closely. together and practically as one
articulation, is also very probable (on these three points in
primitive Mammals comp. Gregory, 1908, pp. 135-138).

Our Gomphognathus-skull shows also the way in which the
cotylus and condylus of the dentary articulation in Mammals
have probably arisen. As the articular surface of the dentary
situated on the squamosal is a smooth plane surface in our
specimen, so we have to suppose that also the corresponding
surface on the condylar prolongation of the dentary was a smooth
plane surface. Consequently we may suppose that this latter
surface was a mechanical result of the lateral movements of the
lower jaw, through which the hind end of the dentary (7. e. the
condylar prolongation of its ascending process) was brought in
contact with the squamosal. The primitive condition of the
deutary articulation in Mammals would, according to this
supposition, be a simple syndesmosis, as we find such a syndes-
mosis secondarily in Zatusia hybrida among living forms. The
ditferent forms of the mandibular articulation in higher Mammals
would then be considered as further mechanical results of jaw-
movements according to the mechanical theory of normal articu-
lations of Fick t, Tornier t, and Cope §.

Still greater is the importance of our specimen in respect to

s
* Between the skulls of Cynognathus on the one side and the skulls of Diade-
modon (and perhaps also of Gomphognathus) on the other, I find an essential
difference in a separate ossification that exists in front of the epipterygoid and below
the postfrontal bone in the British Museum specimen R 3587 of Diademodon-
skull (described by Watson, 1911, who has not recognized its separate nature), an
ossification that J suppose to be an incipient orbitosphenoid. The front edge of
the epipterygoid bone in Cynognathus crateronotus is damaged, but the under
surtace of the postfrontal bone is so smooth, that a corresponding separate orbito-
spheénoidal ossitication is a very improbable one (comp. fig. 5 in Watson, 1911, p. 300,
with fig. 6, p. 76, in Seeley (2), 1895).
Pas Comp. R. Fick, ‘“‘ Ueber die Form der Gelenkflachen ” in ‘ Archiv fiir Anatomie
und Physiologie,’ 1890, p. 391.
~ Comp. J. Tornier, ‘Das Entstehen der Gelenkformen” in ‘ Archiv fiir Ent-
wickelungsmechanik,’ 1897, pp. 124-158, 224-268, 307-346.
~ § Comp. H. Cope, ‘ Primary Factors of Organic Evolution,’ 1904, p. 288 seq.

206 DR. B. PETRONIEVICS ON THE LOWER JAWS OF THE

the question of the origin of the Ossicula auditus in Mammals.
Our specimen shows the beginning of the separation of the
articular bones from the dentary, a state of things that the
classical theory of the origin of Mammalian ossicula auditus
necessarily presupposes. As we have seen, the bony connexion
between the articular bones and the dentary is a relatively much
reduced one, especially in the hind part (comp. text-fig. 1). The fact
that the quadr ate has been lost in our specimen on ‘the left side,
where it might have been preserved, shows that it was probably
only insutticiently fixed in the corresponding groove of the
squamosal. The movable condition of the quadrate, that the
classical theory presupposes, is here, as it seems, also in its
beginning. And finally we find in our specimen on the left side
a displaced bone (comp. text-fig. 1, st¢.), that is, according to
another specimen, to be considered as the bone connecting the
region of the fenestra vestibuli with the quadrate, consequently
as the stapes or the columella’ (on this bone comp. Seeley (1
1895, p. 25, Broom 1904, p. 491, who considers it in Cynognathus
platyceps wrongly as a ‘tympanic ring, and especially Watson,
1911, p. 324).

From the above it follows that we have in Gomphognathus
realized all the elements that constitute the morphologically
initial state in the evolution of the mammalian ossicula auditus,
and we have only to suppose that in the descendants of the cor-
responding relatives of Gomphognathus (the direct ancestors of
Mammals) this state of things increased in the direction of a
further separation of the stapes—quadrate-articular bone chain
from the squamosal and dentary bones, in order to reach the final
state, when in true Mammals the former were transformed into
the ossicula auditus. Into the question how this transformation
took place, and into the corresponding question of the homologies,
I cannot enter here (comp. the Literature, Nos. 7-11). I mention
only that the tympanic membrane of the Mammals is most pro-
bably a neomorph (and I agree in this point with Gaupp—comp.
Gaupp, 1911, p. 641 seq. and p. 659, while in manv other con-
clusions reached by Gaupp—comp. p. 633 and p. 656—I must
disagree), and that the manubrium mallei in Mammals cannot be
homologized with the extra-columella of the Reptiles, as eueslay
dloes (comp. Kingsley, 1900, p. 232 seq.).

In finishing this paper I desire to express my thanks to
Dr. Smith Woodward and Dr. Andrews, of the British Museum
(Natural History), for the loan of the new preparations (executed
by Mr. Hall), and to Dr. Andrews for some valuable help.

LITERATURE.

1. H. G. Sretey.—Researches on the Structure, Organization,
and Classification of the Fossil Reptilia. Part LX. Section 4.
On the Gomphodontia. In ‘ Philosophical Transactions
Royal Society,’ 1895, vol. 186, B, pp. 1-58 (On Gompho-
gnath us, pp. 8-31).

CYNODONT REPTILES GOMPHOGNATHUS AND CYNOGNATHUS. 207

2. H. G. Seetey.-—Researches, etc., Part LX. Section 5. On
the Skeleton in new Cynodontia from the Karoo Rocks. In
‘Philosophical Transactions Royal Society,’ 1895, vol. 186, B,
pp. 99-148.

3. R. Broom.—On the Structure of the Theriodont Mandible
and on its Mode of Articulation with the Skull. In ‘ Pro-
ceedings of the Zoological Society of London,’ 1904, vol. i.,
pp. 490-498.

. R. Broom.—On some Points in the Anatomy of the Theriodont
Reptile Diademodon. In ‘ Proceedings of the Zoological
Society of London,’ 1905, vol. 1., pp. 96-102.

5. D. M.S. Warson.-—The Skull of Diademodon, with Notes on
those of some other Cynodonts. In ‘ Annals and Magazine
of Natural History,’ vol. viii., 1911, pp. 293-330.

D. M. 8. Warson.—On some Reptilian Lower Jaws. In
‘Annals and Magazine of Natural History,’ vol. x., 1912,
pp. 573-587,

“4. R. Broom.—On the Fate of the Quadrate in Mammals. In

‘Annals and Magazine of Natural History,’ vol. vi., 1890,
pp. 409-411.

AS

=

8. J. S. Kryestey.—The Ossicula auditus. In ‘Tufts College
Studies,’ No. 6, 1900, pp. 203-274.
9. H: Gapnow —The Evolution of the Auditory Ossicles. In

‘Anatomischer Anzeiger,’ Bd. xix., 1901, pp. 896-411.

10. W. Kk. Grecory.—The Orders of Mammals. In ‘ Bulletin of
the American Museum of Natural History,’ vol. xxvii.,
1910, chp. I: “The Origin of Mammals and the Problem
of the Ossicula auditus,”’ pp. 113-143 (comp. also Literature
on “Evolution of the Mammalian Ossicula auditus,”
pp. 484-5).

11. G. E. Gaupp.—Beitriige zur Kenntniss des Unterkiefers der
Wirbeltiere. IIT. Das Problem der Entstehung eines
“ secundiren ” Kiefergelenkes bei den Saugern. In

‘Anatomischer Anzeiger,’ Bd. xxxix., 1911, pp. 609-666.

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uy * t

P. Z.S. 1918, Miss BATE, eral:

aggersamosseene

J. M. Woodward, det pate & Danielsson, Ltd.

HYPNOMYS, gen. nov. et LE\THIA.

A NEW GENUS OF EXTINCT MUSCARDINE RODENT. 209

11. On a new Genus of Extinct Muscardine Rodent from
the Balearic Islands. By Dorotuea M. A. BATE *
Fon. MB.O.U.

| Received April 9, 1918; Read May 7, 1918.)
(Plate I.t and Text-figs. 1, 2.)

INDEX.
SYSTEMATIC: Pare
LT ONOMIYS, COMM! xo) ceseac veghteis cpus stehiavks ae cbeaPaasceseane” CLO
EEA RONCII SUS MAD, MNS), 1h aaa See ice od notes come tawt oui tose LO
LE OPP HOUS eS Delis | ste cintaulomeene ocsian cect oh ssiee es yonuite LO

During a second visit to Mallorca in 1910 in search of ossiferous
deposits a few rodent remains were obtained; the following year
further researches were carried out in Menorca by means of a
grant from the Trustees of the Percy Sladen Memorial Fund, and
here similar remains were found to be somewhat more plentiful,
occurring in several fissures in the Miocene limestone. A de-
scription of the deposits from which the collection was obtained
has already been publishedy. A first cursory examination
of the specimens led me to suppose that they represented large
forms of Hliomys§ or Leithia||. Since then a number of spe-
cimens from Menorca have been developed from the hard matrix
in which they were embedded, and all have been carefully
examined, with the result that it is found that they cannot be
included in any genus with which I have been able to compare
them. The examples from the two islands differ considerably in
size, those from Menorca being the larger, and they are probably
specifically distinct: in this Konnection itis inter esting to remember
that, after the examination of a very large quantity of material. a
SGailar variation in size was found tee obtain in Myotr agus ¥.
This seems to point to a longer period of isolation in Menorca,
the most easterly and probably the first of the group to be
separated from the mainland.

All the remains obtained are now in the collection of the
British Museum (Natural History). No complete skulls and
only a few limb-bones were procured, but it will be seen from
the descriptions of the specimens given below that the Balearic
genus should undoubtedly be included in the Muscardinide.
The. specimens to be described are intermediate in size be-
tween the largest recent forms and the extinct Lezthia from
Malta, and aor a number of points of resemblance both to
the recent Hliomys and to the Maltese Leithia, but at the same
time differ to an equal extent from both these genera. The

* Communicated by Dr. A. Sm1tH WoopwarpD, F.R.S., F.Z.S.

+ For explanation of the Plate see p. 222.

+ Geol. Mag. [6] vol. 1. 1914, pp. 337-45.

§ Lbid. p. 100.

|| Proce. Zool. Soc. 1916. p. 424.

| See Andrews, Phil. Trans. Roy. Soc. ser. B, vol. 206, 1915, p. 301.

210 MISS DOROTHEA BATE ON A NEW GENUS

range of species in neighbouring islands is always of interest,
which may perhaps make it worth mentioning here the present-
day distribution of Hliomys in the Balearics. Mr. Oldfield
Thomas has described 4. gymnesicus* from Menorca, a species
smaller than, though similar in coloration to, Z. quercinus, which,
in spite of this, he believes to be most nearly allied to the Southern
forms. In an earlier paper f the same author mentions that this
animal is also well known in Mallorca but is said not to occur in
Ibiza. Unfortunately I have not seen one from the larger island,
but in the British Museum there are now several specimens from
the small island of Formentera to the south-west of Ibiza, which
prove to belong to the large form FZ. lusttanicus which is found
on the neighbouring mainland of Spain.

Remains of Muscardinide are known only from European
deposits, and occur from Miocene times. Hliomys itself is repre-
sented by #. pomelt in the Lower Miocene of Allier, and by
EL. hamadryas in the Middle Miocene of La Grive St. Alban and
Steinheim. Both species are of small size, and from the mandible
do not show any close affinity to the remains which form the
subject of this note, than which they would both appear to be
already more highly specialised. In the British Museum there
are two mandibular rami supposed to represent an Hliomys
from the Pleistocene of Malta (B.M. 49342c and 49351)¢. These
are considerably more robust than in any recent member of the
genus, and are found to resemble in shape the Mallorcan speci-
mens and to be only slightly inferior in size, although the length
of the alveolar area is considerably less. Unfortunately none of
the cheek teeth are preserved, and the formation of the roots can-
not always be satisfactorily deduced “from an examination of the
alveolar cavities, though in one of these specimens (B.M. 49351)
it seems that in the first and second molars the posterior roots
had become confluent as in the genus now to be described. These
specimens seem worth mentioning as suggesting a possible exten-
sion of the known distribution of the Balearic dormouse. So far
as | am aware, no other rodent remains have been described from
the Balearics, although so long ago as 1855 De la Marmora
wrote § that he had observed indications of an ossiferous breccia
in the hill of Belver near Palma, in which he had seen a bone
which seemed to belong to a Lagomys or to a rabbit.

The following description of the most important specimens
obtained may be taken as applying to the genus so far as known
at present. It is proposed that the genus be known as

HyYPnomys, gen. nov.

Skull, mandible, and limb-bones as in Lliomys but more robust ;
interorbital region wide and anterior portions of frontals greatly

* Ann. Mag. Nat. Hist. ser. 7, vol. x1. 1903, p. 494.

+ Proc. Zool. Soc. 1901, p. 41.

+ Cat. Foss. Mamm. Brit. Mus. Pt. 1. p. 225 (1885) (as Myowxus sp.).
§ Mem. Accad. Se. Torino (ser. 1), vol. xxxvil. 1885, p. 59.

OF EXTINCT MUSCARDINE RODENT. 211

expanded. The infraorbital foramen opens anteriorly and the
outer wall of the infraorbital canal is very robust with a wide
base. The anterior palatine foramina penetrate for some distance
the palatal plate of the maxille which forms the greater portion
of the ee The angle of the mandible is perforated. Dental
formula i. +, pm. 1m. #, molariform premolars and molar crowns
sub- atindrated in shape with low transverse ridges. Upper molars
with one large wide internal and two smaller external roots. In
the first and second lower molars the two posterior roots may be
confluent for the greater part of, or for their eatire length; the
last molar has two anterior and one large posterior root. Tibia
and fibula joined.

Skull.—The skull is represented only by some few fragmentary
specimens none of which show the posterior portion, that is to say
behind the frontals. An isolated and imperfect auditory bulla
(B.M. M11671j was obtained which agrees in form and in the shape
and position of the meatus with the corresponding bone in Dyromys.
So far as available specimens show, the skull in general resembles
and may be compared with that of Eliomys, which for our present
purpose will be taken in a broad sense and to include Dyromys.
Even the smaller examples (those from Mallorca) are larger and
comparatively, as well as actually, more robust than EF. lusitanicus,
which is the largest of the recent species of that genus. Viewed
from above, the nasals are seen to be practically flat and about
the same width throughout their length. The interorbital
portion of the frontals is wide and smoothly flattened and is
defined by sharp lateral edges; a measurement taken at the
narrowest point is nearly half the antero-posterior length of the
nasals in Hypnomys, whereas in /. lusitanicus it is not quite a
third. Anteriorly the frontals expand rapidly by a smooth
swelling and attain to a great width—about twice that of the
intererbital region—where they are joined by the nasals and
the nasal processes of the maxilla. This expansion is caused by
the very large size of the olfactory cavities (Pl. I. fig. 14). There
is a fine but distinct ridge on the dorsal aspect of the maxilla at
the upper root of the zygoma from where the skull narrows
rapidly to form the snout. The premaxille are robust, slightly
inflated dorso-anteriorly as in Hliomys, and deep dorso-ventrally,
the rostrum being stouter than in ZL. lusitanicus. The infra-
orbital region is not very well preserved in any of the Balearic
specimens, but sufficient is‘shown by two or three examples to
demonstrate that in this respect Hypnomys shows no close
resemblance whatever to either a typical Murine such as Rattus
or to typical Sciurines such as Sezwrus or Xerus. Though showing
more general resemblance, this portion of the skull in the Balearic
genus still differs somewhat from that of any other of the
Muscardinide with which I have been able to compare it; in
some respects 1t seems to agree with that of Leithia, though
unfortunately all the anterior portions of skulls of this last in

Fo? MISS DOROTHEA BATE ON A NEW GENUS

the British Museum are either very imperfect or else considerably
crushed and distorted. The infraorbital foramen opens anteriorly
and, as well as can be seen in tne type specimen of Hypnomys
mahonensis (Pl. I. fig. 2), it has a greatest length of about 3 min.
and occupies a median position horace the two roots of the
zygomatic process of the maxilla. The lachrymal foramen shows
an extension of similar length, but ovoid in form, in front of the
infraorbital foramen much as in Muscardinus, and without the
lower lateral expansion seen in Hliomys. The outer wall of the
infraorbital canal is very robust, broadly concave anteriorly, with
a distinct outstanding point in front of its base, which is wide
owing to the hinder border being produced outwards from above
the first molar.

The anterior palatine foramina start a short distance behind
the incisors and penetrate the palatal plate of the maxille for
some distance, their extent being comparable with that obtaining
in EHliomys. The palate (Pl. L. fig. 1) is not completely preserved
in any of the specimens, but can be seen to be wide and gently
coneave and chiefly composed of the palatal plate of the maxille,
which extends posteriorly about as far as the hinder border of
the second molar. The palatal plate of the palatines is hardly
shown but can certainly bave formed only an insignificant
portion of the palate.

Text-figure 1.

Left mandibular rami of :—

A. Leithia melitensis (B.M. 49342 D),
B. Hypnomys morpheus (B.M. M 11697),
C. Hliomys lusitanicus (recent specimen). X 15.

Mandible.—In general plan the mandible (text-fig. 1, B) is
essentially as in Hliomys (text-fig. 1, C), from which it yet differs
strikingly, although this difference is easier to observe than to
describe. That of Eliomys may be said to have an aspect of

.

OF EXTINCT MUSCARDINE RUDENT, 2b

attenuation, 1ts posterior portion particularly being very shght
and the cheek-teeth closely crowded together. This applies also
to the skull, though perhaps to a lesser degree. The mandible of
the Balearic genus is altogether more robust, and gives the im-
pression of being possibly less highly specialised. The angle of
the jaw is perforated, and also resembles in shape that of Eliomys,
but compared with this last the coronoid process originates
further forward and rises more abruptly; it tapers to a slender
point but is comparatively shorter and the space between it and
the condyle less deepiy excavated: this last also applies to the
region between the condyle and the highest point of the angle,
causing the hinder portion of the jaw to be more solid owing to
this comparatively greater extent of bone. The articulating
surface of the condyle is strongly marked. The symphysial
region and that between it and the cheek-teeth are likewise
robust. The incisor extends considerably behind and rises above
the cheek-teeth row, causing a marked protuberance on the out-
side of the jaw at the base of the coronoid process, and the
inferior dental foramen occupies a correspondingly high and
backward position.

Teeth.—The dental formula is the same as that of other of the
Muscardinide, that is to say, i.4, pm. +, m. 3. The incisors are
of medium size, with the anterior band of enamel smooth and
stained the characteristic orange-yellow colour. The upper
incisor originates above and just in front of the premolar; its
inner surface (Pl. I. fig. 10) is flat and the outer gently rounded,
this being also the case in the corresponding tooth of the lower
jaw. Both ave considerably compressed laterally with the antero-
posterior thickness much greater than the lateral width, this being
nearly double in the lower teeth. -Their transverse sections (PI. I.
figs. 11 & 12) are more or less elliptical in shape, thus differing
from those for instance of Hliomys, Glis, and Leithia, in which the
section forms practically an isosceles triangle with the anterior
face as the shorter base (PI. I. fig. 13). There is naturally a
corresponding difference in the shape of the worn surfaces of the
teeth. As Mr. Hinton has pointed out to me, these differences in
the form and proportions of the incisors are of importance as
indicating probable differences in the modes of life in these
various genera.

It is interesting to find that an examination of the microscopic
structure of the enamel in the incisors of Hypnomys seems to
bear out the conclusions independently arrived at from a general
study of its remains. Mr. Thornton Carter very kindly under-
took the task of making this examination, and allows me to
include his report of his investigations, which is as follows :—

“The structure of the enamel of the incisors in the specimens
from Menorca and Mallorca is identical with that of Leihia
melitensis.

“The ‘pattern’ is distinctive and presents characters which
would seem to place it between Sciuride and Myoxide. In

Proc. Zoou. Soc.—1918, No. XVI. 16

s)

Hyg MISS DOROTHEA BATE ON A NEW GENUS

longitudinal sections the enamel rods (or fibres, as Sir John
Tomes designates them in his classical memoir) leave the surface
of the dentine with a slight curve and then proceed outwards,
slightly flexuous, at an angle of about 70° with the dentinal sur-
face for about half the thickness of the enamel, where they bend
sharply and abruptly to proceed in a straight course to the
surface.

“There is appearance of serrations of the margins such as is
usually seen in the Dormice.

“Thus the course of the rods is as in Myoxide, whilst their
form is as in Sciuride.

‘‘In transverse section alternate groups of rods cross one
another at an angle of about 120°, and when half-way through
turn sharply outwards and run parallel out to the enamel surface
which they reach at right angles, the pattern assumed resembling,
though not identical with, that seen in a transverse section of the
incisor of Hliomys.”

While the palate is almost uniform in width, the rows of cheek-
teeth appear curved owing to their inner borders being practically
in a straight line, so that all the differences in the sizes of the
teeth, especially that of the premolar, affect only the outer
border of the rows, which thus have a curved outline. The plane
of wear of the premolar is almost horizontal, while that of the
molars lies at a considerable angle, the inner border being the
higher; this character will be referred to again later. Some
scarcely worn upper molars are slightly concave and their enamel
ridges incline towards being cuspidate, but in most of the speciniens
obtained the worn surfaces of the crowns are nearly flat. The
molariform upper premolar is bluntly triangular in outline and
the molars subquadrate in shape, the second being very little
larger and the third not much smaller than the first. In the
above characters these teeth somewhat resemble those of Leithia,
those of Hliomys differing markedly in the concavity and greater
comparative width of their crowns.

In the Balearic molars the ridges are low, and in those of aged
individuals, of which a number were obtained, these become
almost entirely worn away. In little-worn specimens (PI. I.
fig. 1) it can be seen that in each of the upper cheek-teeth the
crown is crossed by four complete and one incomplete transverse
ridges, with the addition of one or more further incomplete
ridges in the second and third molars. The second complete
ridge in the first and second molars runs from the inner border
in an anterior direction for more than half its distance before
turning to reach the outer margin. The two posterior complete
ridges run more or less parallel with the hinder border of the
teeth, and it is in the space left between the complete ridges that
the incomplete ones occur. The inner edge of the crown surface
almost invariably remains unbroken, while the outer border is cut
by the valleys between the ridges. In much worn teeth there
is found a confluent valley alongside the inner enamel border of

OF EXTINCT MU. CARDINE RODENT. DAES)

the tooth for the whole of its distance; this condition of wear is
also seen for instance in Leithia, Hliomys, and Xerus. Apart
from size, the upper molars of Lezthia chiefly differ from those of
Hypnomys in their transverse ridges being considerably higher
and more sharply defined, and in the complete ones in the first
and second molars originating at the postero-internal border of
the crown as already described and figured by Mr. Lydekker *.

Of the lower cheek-teeth the premolar is the smallest and is
obtusely triangular in shape; the third molar is somewhat larger
with a rounded posterior margin. although the crown has a
tendency to become squarer in outline when much worn. The
first and second molars are the largest of the series, almost equal
in size and in shape quadrate, or with the antero-posterior length
slightly greater than the width. The number of ridges appears
to be much the same as in the upper cheek-teeth, though they
become broken up earlier and into a greater number of incomplete
ridges. In a moderately worn specimen (B.M. M 11674, Pl. I.
fig. 6) the first and second molars seem to have three complete
ridges, two distinct incomplete ones running half-way across the
crown from opposite sides and a shorter one coming in from the
antero-external corner, while the antero-internal area is broken
up into several rather indistinct loops. The third molar shows
four complete ridges and two small accessory ones. In the case of
aged individuals the confluent worn surface is of course along the
external border of the crown, it being on the internal side in the
upper cheek-teeth. In less worn specimens this external enamel
border is interrupted at two points, and the enamel between and
on either side of these channels, owing to its being slightly folded
over, is at this stage of wear more massive than in other parts of
the ridges,

The number, position, and conformation of the roots of the
cheek-teeth in rodents form a subject of considerable interest and
importance, and it is one that has received a good deal of atten-
tion from investigators, notably Schlosser t and, later, Dr. Forsyth
Major{. The specimens under discussion are no exception in
this respect, for their molar roots are unlike those of any Pleisto-
cene or recent genus with which I have been able to compare
them with the single exception of Leithia, and that only in the
case of the upper molars. It may be mentioned that I have not
found this character previously noted in the descriptions of the
Maltese genus. The alveolar formula can be most easily realised
by reference to the accompanying text-figure (text-fig. 2), but at
the same time it should be borne in mind that such a formula
alone is of very little value for purpoess of comparison. The
present examples are a case in point, as, for instance, the first and
second upper molars of Hliomys and Glis are three-rooted, as in

* Proc. Zool. Soc. 1895, pp. 861-3, text-fig. 1.
Te Paleontographica,’ Band xxxi. 1885.
t See, for instance, the following: “ On Fossil Dormice,” Geol. Mag. Nov. 1899,
p. 492, and “On some Miocene Squirrels,” Proc. Zool. Soc. 1893, pp. 179-214. :

Ge

216 MISS DOROTHEA BATE ON A NEW GENUS

Hypnomys and Leithia, but an examination of the roots them-
selves show them to be essentially different.

In Hypnomys the upper premolar has three long and rather
stout roots, which terminate in blunt rounded ends (B.M. M 11658,
Pl). I. fig. 3). The anterior one takes a forward direction and is
the shortest of the three, the other two project in a parallel
direction from the hinder border of the tooth and may be con-
fluent for a varying distance, but apparently not for more than
half their length. This description is based on examples of
Hypnomys mahonensis, but the alveolus in a Mallorcan specimen
indicates a similar formation. Unfortunately there has been no
opportunity of examining any milk-teeth in the case of either
this genus or Leithia.

Text-figure 2.

A. io lgeamest

e. 5
s re
iS ei ° fem hr e jLim
@
mS gM! aa ES | oe
@ @ @
ee ch et
=)
Sey hs ee 38 4

Ualvedi. L.alveoh. U alveoli. L.alveoli

Diagram of alveoli of cheek-teeth of A. Hypnomys and B. Leithia.

The upper molars each have three roots, one large internal and
two small external ones, a number commonly found in many
rodents. ‘The distinctive feature of the Balearic teeth lies in the
construction of the large internal roots: in Hliomys, Glis, and
also in other forms such as Xerws and Sciurus, which have the
same alveolar formula, these are single, rather flattened conical
roots projecting from the centre of the border of the crown which
they support. In Hypnomys the internal root is quite different,
this being clearly portrayed in the accompanying illustrations.
Fig. 3 (Pl. I.) shows the latero-internal view of the first and
second molars of H. mahonensis: it will be seen that there isa
slight constriction at the base of the crown where the enamel
ends, and from there the confluent root continues practically of
the same size for its whole length, which is a little less than twice
the antero-posterior length of the crown-surface. In the figured
specimen the first molar measures 4 mm. from the crown-surface
to the tip of the root, while the antero-posterior length of the
crown is very little more than 2mm. In the second molar the
root is very slightly shorter and not rounded at its apex, but this
uray perhaps be partly due to damage in developing the specimen.
The internal root of the third molar is exposed in the type

OF EXTINCT MUSCARDINE RODENT. yA le

specimen of //. mahonensis (B.M. M 11657), in which it is seen
to be similar to that of the other two. In some specimens there
is a slight groove along the centre, indicating the compound
nature of the roots ; the ends are closed in all examples examined.
The slight intero-external width is well shown in the section
(B.M. M 11661, Pl. I. fig. 4) and in the posterior view of the
second upper molar (B.M. M 11660, Pl. I. fig. 5), in which it can
also be seen that there is a certain amount of curvature in the
direction of the compound roots. They are considerably longer
than the external roots, although, owing to the slope of the
crowns, they do not penetrate much deeper into the alveolar
cavity.

As already mentioned, the upper molar roots of Leithia are
sunilar to those of Hypnomys, the only other genus, among many
examined, showing a closely similar condition of root structure is
L'rechomys. In the British Museum there are two fragments
of the upper jaw of 7. platyceps (B.M. M 1627) from the Phos-
phorites of Caylux, containing the premolar and first molar, the
roots of the latter being as in Hypnomys except that the broad
inner root is perhaps not quite so wide compared with the crown.
It must be mentioned that in 7rechomys the roots of the premolar
are the same as in the molars, which is of course not the case
in either Leithia or the Balearic genus. The molar roots of
Theridomys are somewhat similar but are accompanied by much
more hypsodont crowns.

The roots of the lower cheek-teeth of Zypnomys (B.M. M 11678,
~Pl. I. fig. 7) show a very considerable amount of variation both
as regards their length and conformation, The premolar may
perhaps be said to have two roots, but these are confluent for
the greater part of their considerable length, diverging at a vary-
ing but never very great distance from their apices, the anteiior
of which is slightly the longer of the two. The upper, posterior
portion of the root is wide and flattened, being very evidently the
result of the fusion of two roots. In some specimens (as in B.M.
M 11678) there are clearly three confluent roots.

In the first and second molars the two anterior roots are
usually long, equal in size, and separate though not very widely
divergent. In the posterior roots the length and antero-posterior
thickness are about the same asin the anterior pair, so that
viewed laterally little or no difference is observable (Pl. I. fig. 7).
But viewed from behind they are seen to be confluent transversely
for either the whole or three-fourths of their length (PI. I.
figs. 8, 9, BM. M 11675-6); in the latter case the stout terminal
portions of the roots may diverge considerably. The resemblance
to Hypnomys seen in the roots of the upper molars of Leithia
does not hold good for those of the lower jaw, for in the
Maltese genus the first and second molars each have four roots,
entirely separate from each other for their entire length. In
Trechomys, however, the lower molar roots seem to agree with
those of the Balearic genus with the possible exception of the

218 MISS DOROTHEA BATE ON A NEW GENUS

third molar. Schlosser gives the number of roots for this tooth
as three, but judging from the alveolus of a specimen of 7’. platyceps
in the British Museum (M1627) it appears that there may some-
times be two large, transverse roots only.

The last lower molar-in Hypnomys has three roots, two anterior
slender ones and a large, backwardly projecting posterior one.
This condition is very similar to that obtaining in the correspond-
ing tooth of Leithia and the recent Hliomys, though in the latter
there is a more marked constriction immediately below the crown.

Limb-bones.—Very little of the skeleton was obtained, but a
few femora, a tibia, and fragmentary portions of humeri and
other bones, chiefly from a Mallorcan cave-deposit, show characters
which also indicate that the genus should be included in the
Muscardinide. The ulna is very similar to that of Hliomys, as is
also the femur, except that the third trochanter seems to be
shehtly less prominent and less sharply separated from the great
trochanter. As contrasted with the Hystricide, Sciuride, and
Anomalurids, which have the tibia and fibula free, a charac-
teristic of the Muride and Muscardinide is the joining of these
two bones for a third or more of their distal portions. It may be
worth noting that in Rattus (Kpimys), for instance, the fibula
at its distal extremity is once more separated though closely
adpressed to the tibia. On the other hand, in Hliomys quercinus
the tibia and fibula continue completely ankylosed at their distal
ends, and it is with this latter that the Balearic specimens agree,
as Leithia likewise does in this respect. ‘These bones are joined
for more than a third of their total length in the three last-
mentioned genera. In Hypnomys the tibia isa straighter bone
with a less well-developed cnemial crest than in Hliomys.

’

The above are the chief characters of all the Balearic specimens,
and it will only be necessary to add a brief note and a few
measurements in the two species from the different islands.

The species from Menorca may be named

Hypnomys MAHONENSIS, Sp. N.,

and regarded as the type species of the genus, distinguished from
the Mallorean species by its greater size and different habitat.
Further differences would probably be apparent were a larger
amount of material available. All the specimens were obtained
from fissures in the Miocene limestone, and were in one or two
instances associated with remains of J/yotragus and Testudo
gymnesicus. Some imperfect lacertilian Jaws also occurred, and
these have been very kindly examined by Mr. G. A. Boulenger,
¥.R.S.: they prove to be those of a Lacerta of the viridis- ocellata
group and a species of Chalcides. Although representatives of
both these occur commonly in the Mediterranean region, neither
ave found in Mallorca or Menorca at the present day.

Owing to the fragmentary state of the specimens not many

OF EXTINCT MUSCARDINE RODENT. 219

measurements can be given, but the following are afew. The
length of the nasals is about 18 mm. in the single example in
which these bones are completely preserved, and the interorbital
width of the frontals is 8mm. The base of the outer wall
of the infraorbital canal measures 6 mm., and the incisive
foramina are approximately 8 mm. in length. In the type
specimen (PI. I. fig. 1) the complete row of left upper cheek-teeth
is almost 10 mm. in length, this measurement being 14 mm. in a
corresponding example of Leithia, and barely 7 mm. in one of
Eliomys lusttanicus. In H. mahonensis the width of the palate
between the first molars is not quite 6 mm., in Leithia it is little
more, being 6°5 mm.

The length of the lower cheek-teeth series is about 10 mm.;
as already mentioned the length of their roots is very variable:
in one specimen having teeth with moderately worn crowns
(B.M. M 11673, Pl. I. fig. 7) the first molar has a crown-width
of 2°5 mm. with a root-length (measured from’ the crown-surface
to root-tip) of almost 5mm. In the third molar in the same
specimen the greatest length of root is 3 mm., the large posterior
root attaining an antero-posterior width of 1:5 mm., while the
antero-posterior length of the crown is 2°5 mm. In an example
of a left lower second molar (B.M. M 11676, PI. I. fig. 9) with a.
less worn crown than the above the crown-width is 2 mm., and
the greatest root-length 3 mm.

It is proposed that the species from Mallorca be known as

HyYPNOMYS MORPHEUS, Sp. n.,

characterised by its smaller size and different habitat from
HT, mahonensis. The few specimens of jaws and limb-bones by
which it is represented were obtained from cave-deposits in
Mallorca, and were found associated with remains of MJyotragus,
and in one case with a few mandibular rami and limb-bones of
Apodemus, which still occurs plentifully in the island. In size it
appears to agree with the larger forms of Glis. The type speci-
mens (B.M. M 11695), consisting of the anterior portion of the
skull with the incisors and right molars and two mandibular
rami, are believed to have been associated and fell apart on being
separated from the earthy matrix in which they were preserved.
The base of the outer wall of the infraorbital canal measures
5 mm., and the antero-posterior length of the molar row is 6 mm.
The length of the lower cheek-teeth series is 8mm. In the
upper incisor (B.M. M 11696) shown in fig. 10 (PI. I.) the antero-
posterior width is 2°5 mm. while the thickness is 15 mm.; ina
specimen of the lower incisor the antero-posterior width is 2 mm.,
with a thickness of barely more than 1 mm.

The greatest length of a right tibia, wanting its proximal
epiphysis, is 41 mm., the distal 18 mm. of which are joined to the
fibula. A femur has a total length of 55 mm., that of a right
ulna being 36 mm.

220 MISS DOROTHEA BATE ON A NEW GENUS

Summary and Conclusions.

The above description may be very briefly summarised as follows.
Ilypnomys 1s represented by two species known by remains from
the two largest islands of the Balearic group and is considei ed
to belong to the Muscardinide, though seeming to be a very
distinct form not closely related to any other at present known.
Its dental and osteological characters so far observed not only
seem to show that //ypnomys belongs to the Muscardinide but
also appear to afford further support to the opinion, now very
generally held, that this group cannot be included in either the
so-called Sciuromorpha or Myomorpha and lending additional
weight to the argument against employing these terms. This
seems to hold good also for the modern pee tees which
have been briefly described by Mr. Muiller* as “.. . mostly
arboreal animals with habits and aspect somewhat intermediate
between mice and squirrels. aa

The Balearic genus may be compared with the recent Lliomys
and the extinct Leithia trom the Pleistocene of Malta. It agrees
with the former in general plan of skull, lower jaw, and limb-
bones, and it may be noted that there is also a perforation in the
angle of the lower jaw. It differs in a number of points, among
which the following may be cited :—In the pattern of the molar
crowns, which are very slightly coneave and lack the marginal
cusps, both distinctive characteristics of Hliomys. In Hypnomys
the worn surfaces of the molars are subquadrate in shape, whereas
in the recent genus the width is distinctly greater than the
antero-posterior length; the alveolar formula differs in the two
genera, and the large inner root in the upper molars of Zypnomys
is quite distinctive. This root-structure was evidently attained
by a process of simply filling up the space between two parallel
roots, whereas the single contcal root seen in the Sciuride and
others would seem to be arrived at by the drawing together of
the apices of the two roots with a similar fillmg up of the
intervening space. It would be interesting could one arrive at
a reasonable hypothesis to account for this root-formation in
ITypnomys and Leithia, for presumably it must have been acquired
to meet a special stress or strain consequent on some peculiarity
of diet or mode of life.

Hypnomys agrees with Leithia in its squared molars and in
the large, confluent and single inner roots of the upper molars,
but differences are seen in the former having a perforation in the
angle of the jaw, which is also of a different shape (see text-fig. 1),
in the crown pattern of the molars, much lower enamel ridges,
and in the roots of the lower molars.

It has also been noted that in the roots of both upper and
lower molars, but noé premolars, Hypnomys shows a striking
resemblance to Trechomys. However, it is not for a moment

* “Cat. Mammals of West. Europe, London 1912, p. 549.

OF EXTINCT MUSCARDINE RODENT. Dil

suggested that this similarity, which is apparently unaccompanied
by other points of resemblance, indicates any close affinity. On
the contrary, it would seem that these three genera each exhibit
in this character what would have been a stage, and that a very
early one, in the history of the evolution from the low-crowned
four-rooted towards the hypsodont and rootless or semi-rootless
molar. The cheek-teeth of Zheridomys also seem to suggest the
possibility of this result being attained without the necessity of
including a stage having a large conical inner root in the upper
molars accompanied by a somewhat triangular-shaped crown, as
seen in some of the Sciuride (Xerus for instance) and which
Dr. Forsyth Major has fully described in his invaluable paper on
the Miocene Squirrels *.

Hypnomys is an interesting addition to the extinct fauna
of the Balearics so far known by Myotragus and Testudo
gymnesicus, both forms totally distinct from the present-day
fauna of the islands. Owing to the small amount of material
available, the question as to the precise age of these Balearic
deposits and their contents seems a difficult, if not impossible,
one to answer at present. Lately this matter has been the
subject of some interesting remarks by Prof. M. Boulet, who
seems inclined to consider the deposits of earlier age than
Pleistocene. Seeming to support this view are the absence of
human remains and the character of the chief remains found, 2. ¢.
Myotragus, Hypnoniys, anda giant Testudo. The occurrence of
remains of this last in the limestone fissures of Menorca certainly
suggests that there must have been great changes of climate and
vegetation, and that there have been considerable alterations in
the land surface is indicated by extensive stretches of sheets of
stalagmite now exposed on the weathered surface of the Miocene
limestone in both Mallorca and Menorca.

On the other hand, it should be realised that not much weight
should be placed on the absence of human remains owing to the
fact that not a single one of the ossiferous deposits was found to
be in an undisturbed condition, all the upper layers in which such
remains might be expected to occur having been entirely destroyed.
Another point to be borne in mind is that in the Mediterranean
region several species now extinct have been proved to have per-
sisted until the time of man’s occupancy of these particular areas,
The well-known Prolagus, remains of which were found in
Neolithic settlements in Corsica by Dr. Forsyth Major, is a case
in point. Another somewhat similar example is that of a small
deer, Anaglochis, whose remains occur abundantly in some of the
cave-deposits of Crete, while a number of its antlers were found
by Sir Arthur Evans in a shrine in the famous Minoan Palace
of Knossos.

I should hke to take this opportunity of expressing my grateful
thanks to the Trustees of the Percy Sladen Memorial Fund for

* Proc. Zool. Soc. 1893, pp. 179-214.
+ ‘L’Authropologie,’ tome xxviii. (1917) pp. 160-3.

229. A NEW GENUS OF EXTINCT MUSCARDINE RODENT.

enabling me to visit Menorca. Also to Dr. A. Smith Woodward,
F.R.S., for continued kindness in giving me every facility for
working in the Geological Department of the British Museum
(Nat. Hist.), to Mr. M. A. C. Hinton for kind advice and for
looking over some of my material, and to Mr. J. Thornton Carter
for his valuable investigations into the enamel structure in the
teeth of Hypnomys and other genera.

EXPLANATION OF PLATE I.

Fig.1. Hypnomys mahonensis. Type-specimen (B.M. M 11657). Palate showing
the left premolar and molars and right M!. & 4,

Das Do. Side view of same specimen showing anteorbital region. XX 3.

3. Do. Lateral view of right upper premolar and Ist and 2nd molars,
showing the large confluent roots of the latter (B.M. M 11688).
s:

4. Do. Transverse section of left M! (B.M.M 11661). x 4.

5. Do. Posterior view of right M2 (B.M. M 11660). x 4.

6. Do Crown view of right lower molars (B.M. M 11674). x 4.

ae Do. Lateral inner view of right lower cheek-tecth (B.M. M 11673).

x 3. :
8. Do. Right My, (B.M. M 11675) showing posterior roots. X 4.
9. Do. Left Mg, (B.M. M 11676) showing posterior roots. X 4.
10. H. morpheus. Right upper incisor (B.M. M 11696), inner view. X 3.
11. H. mahonensis. ‘Transverse section of upper incisor (B.M. M11662). x 4.

12. Do. Transverse section of lower incisor (B.M. M 11677). x 4,
13. Leithia melitensis. Transverse section of upper incisor (B.M. M 49345).
x 4.

14. H. mahonensis. Dorsal view of interorbital region of skull (B.M. M 11659).
xX 2.

bo
oo

ON HERMAPHRODITISM IN A LIZARD. 2

12. A Case of Hermaphroditism in a Lizard, Lacerta viridis.
By Nort Tayuer, B.Se. (Lond.).* (From the Zoo-
logical Department, University of London, University
College.)

[Received April 23, 1918: Read May 28, 1918.]
(‘Text-figures 1-3.)

The specimen, the urogenital system of which is described in
the following pages, was “placed i in my hands. by Prof. J. P. Hill,
F.R.S. It turned up in the course of class-work in the Senior
Laboratory in the College, and its abnormal condition being
observed, it was fixed in corrosive sublimate and preserved for
detailed examination. The specimen presents certain features
of interest which, it is believed, are worthy of being placed
on record.

I wish to express my thanks to Prof. Hill for his advice and
assistance in the preparation of these pages.

I. DESCRIPTIVE.
a. General Morphology.

When this specimen came into my hands, it was in a partly
dissected condition, the greater portion of the abdominal viscera
having been removed.

The Fat Bodies were present and well developed, the right
being rather larger than the left. They are not represented,
however, in text-fig. 1, which gives a general view of the
urogenital system, since in situ they obscured the more anterior
portions of the oviducts.

The testes (text-fig. 1, &.7. and Z.7'.) were well developed and
suspended in the folds of the longitudinal mesorchia (J/es.).
The right was placed somewhat more anteriorly and was rather
larger than the left, the dimensions of the former being about
‘9 cm. Xx *4 cm., and of the latter -65 cm. x °3 cm.

Both gonads seem normal in shape apart from the remarkable
stalked outgrowths on each (Ov.). These on section were found
to contain ova, and the gonads may therefore properly he
designated ovotestes. The right gonad, it will be seen, possessed
two of these spherical ovarian appendages, each joined to the
dorso-lateral border of the testicular portion of the organ by a
well-marked stalk; the more anterior was further sub. divided
into two by a median constriction.

The left gonad also possessed two outgrowths, the surface of
the larger being subdivided into five or six hemispherical pro-
jections. The ep:didymes were well developed (L.Hp., R./p.)

* Communicated by Prof. J. P. Hii, D.Sc., F.R.S., F.Z.S.

224 MR. NOEL TAYLER ON

ext-figure 1.

KA By 8.

--LAg.
s--d,aort
B.Lg 2 Fe P log
Lv. toes x
“Sp.
‘R M.D--- Z--L.M.D.
R.T.---
Lb sh
(@) i a 4 Ov:
<--Mes'
Eff.Ren Z--L.Ep.
Mes:
REpss
Zi,
R.L= =
Rt
R.K=--
=-BL.
oy,

RCop sere =2 --see- bap.
Lacerta viridis: dissection of a hermaphrodite animal.
Explanation of the lettering.

B.Lg. Broad Ligament. Mes. Mesorchium.
if Bladder. Ov. Ovary.
d.aort. Doreal Aorta. R.Cop. Right Copulatory Organ.
L.Cop. Left Copulatory Organ. R.Ep. Right Epididymis.
L.Ep. Left Epididymis. Tt Kes Right Kidney.
L.Fl. Left Funnel. Teal: Muscular Band.
LMD. Weft Oviduct. R.M.D. Right Oviduct.
Ing. Lung. R.T. Right Testis.
LT. Left Testis. Rt. Rectum.
Lv. Liver. Sp. Spleen.

HERMAPHRODITISM IN A LIZARD. - D225

and situated in the broad ligament parallel to but separate from
the testes. They were not attached, as is normally the case, to
the inner border of the testes, and vasa efferentia were conse-
quently absent. Posteriorly, the epididymes passed into the
vasa deferentia, and these, having united with the ureters, opened
into the posterior division of the cloaca by the urogenital papille.

The two kidneys were apparently normal, each consisting of
an anterior and posterior lobe (in the figure, only the right
kidney, &.4., is indicated by reference letters).

The copulatory organs (2. Cop., L. Cop.) were present as in
the normal male lizard.

The oviducts (R.ILD., £.M.D.) were developed for about
a third of their lengths. Each opened into the body-cavity by a
well-developed funnel (only the left, Z./J., is indicated by
reference letters in the figure), and behind that was continued
into the duct, the plaited glandular walls of which are seen in
the outer border of the broad ligament (B.Lg.). The right
oviduct attained the greater complexity as in the normal female.

Passing down from the posterior tip of the ccelomic funnel on
each side, on the extreme outer border of the broad hgament
was a narrow but well defined ribbon-like muscular band which
continued right back on each side to the cloaca. Similar bands,
which were at first taken for the oviducts, were referred to by
Howes (5) as the round ligaments, and he thus describes them
in his specimen: ‘ From this (the oviducal aperture) there passed
back a ribbon-shaped muscular band which skirted the free edge
of the broad ligament, remindful of the round ligament of
mammalian anatomy. This structure was wholly absent on the
side destitute of an oviducal vestige, as indeed it is in the normal
male. Its development is correlative with that of the oviduct.”

b. Histology.

Transverse sections were made of the gonads and stained with
hematoxylin and eosin. The main body of each gonad was found
to consist of normal testicular tissue, 2.e. seminiferous tubules
with an interstitial stroma, the lining cells being in active
mitotic division (text-fig. 2, s.¢.).

The stalked outgrowths were found to consist of ovarian tissue,
the bulb-like extremities containing large and’ fully-grown ova.
A section of the gonad through the nuclear plane of the most
anterior ovum is somewhat diagrammatically represented in
text-fig. 2.

The large yolk-laden oyum (ov.) is, it will be seen, surrounded
by the relatively thin follicular epithelium (/oll. ep.), outside
this is a fibrous layer continuous with that sheathing the
testicular portion of the gonad. In the stalk region all the
normal histological elements of the ovary are represented.
Outside is a layer of cubical epithelial cells continuous with
the peritoneal epithelium, while the main mass of the stalk is
formed of a loose stroma of connective tissue, contained in which

226 MR. NOEL TAYLER ON

are numbers of young follicles of various ages, the youngest
being nearest to the testis. The connective-tissue body of the
stalk is confluent with the tunica albuginea of the testicular
portion of the gonad, which is somewhat thickened in the region
of junction.

The sections of the oviducts revealed in their anterior portions
a typical structure. They are lined with a well developed
ciliated and glandular epithelium, the lumen of the duct being
filled with the coagulated secretion of the gland-cells. Sperma-
tozoa were not detected.

Text-figure 2.

Lacerta viridis, hermaphrodite animal: sections through gonad.

ov.; ovum.
foll. ep.; follicular epithelium.
s.t.; testicular stroma.

The appearance of the epididymis was quite normal, but no
spermatozoa were found in the lumen of the vas deferens, and
examination of the sections failed to reveal the presence of
vasa efferentia through which the spermatozoa normally pass
from gonad to epididymis.

HERMAPILRODITISM IN A LIZARD. DAgATE

In the dissected specimen two or three fine tubule-like filaments
were seen to pass from the posterior ends of the oviducts to
the anterior extremities of the corresponding epididymes; these
were best marked on the left-hand side (see text-fig. 1). These
filaments were carefully examined in the sections. ‘The epi-
thelial lining of the oviduct was found to end posteriorly in
a cul-de-sac, as did that of the epididymis anteriorly. The
“filaments” in section appeared as actual tubular passages
devoid of any epithelial lining and running in the substance of
the broad ligament. They commenced at the extreme anterior
tip of the epididymis, and while two of them ended blindly, the
third ran up as far as the posterior extremity of the oviduct, its
lumen being continuous with the spaces in the mesentery in
which were contained the glandular portion of the oviduct.

Text-figure 3,

hec. fol.

Lacerta viridis, hermaphrodite animal: secti on through kidney.

kid.; kidney tissue.
foll.ep.; follicular epithelium.
thee. foll.; theca folliculi.

The morphological interpretation of these spaces would seem
a problem of some difficulty. Being devoid of epithelial lining
and making no connection with the lumen of the epididymis, one
would hardly seem justified in regarding them as rete tubules
proper, while the close relationship of one of them with the
oviduct also seems anomalous, on this view.

The condition which Howes (5) describes in one of his
specimens seems of some interest in this connection. ‘In one
instance,” he says, ‘I discovered an interesting modification of
the condition recorded by Leydig. There was buried up in the

228 MR. NOEL TAYLER ON

peritoneum in a line with the head of the epididymis a delicate
filament which, while it answered in every respect to the
rudiment described by him, instead of ending abruptly and
blindly became suddenly enlarged, opening into the body-cavity
by a wide-mouthed funnel-shaped extremity, identical with that
of the oviduct, and lined by a ciliated epithelium.”

This without doubt seems the description of a vestigial oviduct,
yet, posteriorly, according to his figure, it appears to arise from
the anterior tip of the epididymis. Unfortunately he does not
seem to have investigated the precise nature of its relations to
this organ.

The sections through the kidneys reveal the presence on the
dorsal portion of one of them of an embedded mass of almost fully
grown ova. ‘Text-fig. 3 is a semi-diagrammatic representation
of one of the sections in which five ova occur. It will be seen
that the mass of ova lies actually embedded within the kidney
tissue (kid.). Hach ovum is surrounded by a layer of follicular
cells (foll. ep.), while externally to this and separating it from
the kidney substatice is a thin fibrous layer (thec. foll.) presum-
ably representing the theca folliculi.

II. Discussion.

No instance of complete hermaphroditism in the Lacertilia
seems yet to have been put upon record, though cases of the
more or less complete developmeut of the Miillerian Ducts in
adult male lizards have been described.

Leydig in 1872 (7) described the persistence in the males of
Lacerta agilis of the Miillerian Ducts as small blind and con-
voluted tubules, while Braun in 1877 (2) noted the development
of rudimentary Miillerian Ducts in the young male of the same
species, making no mention, however, of its presence in the
adult form.

In 1887 Howes (5) published a brief but important paper,
“On the vestigial structures of the reproductive apparatus in
the male of the Green Lizard” (Lacerta viridis). One of his
specimens was a male lizard in which both the oviducts were all
but fully developed, while in another the oviduct was fully
developed on one side.

In thirteen out of twenty-five specimens examined certain
segments of the oviduct were well developed, the other portions
being only represented as delicate filaments, thus giving a series
of conditions analogous to those described by Matthews for the
male toad.

In 1893, Hill (4) published an account of the persistence of
vestigial Miillerian Ducts in the full-grown male of an Australian
lizard, Amphibolurus muricatus ; while two yeais later, in 1895,
Jaquet (6) described the presence of Miullerian Ducts identical
with those of the normal female in an adult individual of
Lacerta ayilis. |

HERMAPHRODITISM IN A LIZARD. 229

All the above mentioned cases are, it will be noted, pre-
dominantly male, indeed as far as their gonads are concerned
wholly male, for in no case is any reference made to the
presence of ova or ovarian tissue,

In this respect the specimen described in the present paper
stands in striking contrast to previously described cases; more-
over, while it is distinguished by the presence of well-marked
ovotestes, it must have been physiologically sterile.

Among lower forms the occurrence of well-developed Miillerian
Ducts seems often to be accompanied by the existence of an
ovotestis. Cases of this kind have keen described by Marshall
in the Frog (8), but Fantham (3) seems to have been the first to
record a case of true hermaphroditism in the Reptilia. The
specimen of Testudo greca described by him possessed well-
developed oviducts, the lumen of each being continuous through-
out. Of the two gonads the right was a typical testis, on the
ventral surface of the left however was a ‘conspicuous yellow
egg.” On section another was found devoloping in its proximity.
while “‘a few groups of bodies resembling developing ‘ ovarian
ova’ were seen scattered in separate groups (follicles) among
otherwise testicular tissue, more especially near the periphery of
the anterior portion of the gonad.” Epididymes, vasa efferentia
and vasa deferentia were present as in normal specimens, the
former being rather large.

It seems a point worthy of note that the development of the
oviducts in the cases referred to above, viz., those of Howes,
Hill, and Jaquet, is much more complete than in the subject of
the present paper; in all these three cases the oviducts were
developed throughout their whole length and opened into the
cloaca, yet in none of these cases is any mention made of the
presence of ova or ovarian tissue. In the present specimen, on
the contrary, numerous ova occur, though only the anterior
thirds of the two oviducts are fully developed.

Literature referred to.

1. Bourng, A. G.—“‘ On certain abnormalities in the Common

“ Frog; i. he occurrence of an ovotestis.” Q.J. M.S. xxiv.

2. Braun.— Das urinogenital Syst. der embheinnischen Rep-
tilien.” Arbeit. aus dem zool.-zocotomisch. Instit. Wiirzburg,
vol. iv., 1877-8.

3. Fanruam., H. B.—‘ On Hermaphroditism and Vestigial
Structures in the Reproductive Organs of Zestudo graca.”
Ann. Mag. Nat. Hist. xvi. 1905, p. 120.

4. Hii, J. P.—‘ Note on the presence of Vestigial Miillerian
Ducts in a full-grown male lizard (Amphibolurus muricatus).”
Proc. Linn. Soc. N.S.W. vol. viii. (Series 2nd), Sept. 27th,
1893.

Proc. Zoou. Soc.—1918, No. X VII. Ag

230 ON HERMAPHRODITISM IN A LIZARD.

5. Howes, G. B.—‘ On the Vestigial Structures of the Repro-
ductive Apparatus in the Male of the Green Lizard.” Journ.
Anat. & Phys. xxi. pp. 185-9, 1887.

6. Jaquet, M., (1895).—‘“‘ Note sur un cas d’hermaphroditisme
incomplet observé chez le Lacerta ayilis.” Biblogr. Anat,
(Paris—Nancy) iii. p. 267. :

7. Leypie.— Die in Deutschland lebenden Arten der Saurier.”
Tubingen, 1872.

8. Marsuaut, A. Mitnes.—‘‘ On certain abnormalities of the
Reproductive Apparatus of the Frog.” Journ. Anat. &
Phys. xvii. pp. 121-144.

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ON TWO NEW ELASMOBRANCH FISHES. 231

13. On two new Elasmobranch Fishes (Crossorhinus juras-
sicus, sp. nov., and Protospinax annectans, gen. et sp.
nov.) from the Upper Jurassic Lithographic Stone of
Bavaria. By Arruur SmitH Woopwarp, LL.D.,
HOO Vader Zia:

[Received May 23,1918: Read June 11, 1918. ]
(Plate I.)

Most of the modern groups of Elasmobranch fishes seem to
have arisen during the Cretaceous period, but some are of still
older date, and a few interesting types are represented by well-
preserved fossils in the Upper Jurassic lithographic stone of
Bavaria, Wiirtemberg, and France. Two remarkable new
examples of these early forerunners of the existing fauna have
lately been identified in the British Museum, one apparently
indistinguishable from an existing genus, Crossorhinus (ov Orecto-
lobus), the other evidently of a new genus and family closely
related to the Spinacidee.

Family CROSSORHINIDAK,

CROSSORHINUS JURASSICUS, sp.n. (PI. I. fig. 1.)

Specific Characters.—Head gently rounded in front; length of
head and trunk about equal to that of the tail, Three pairs of
fringing dermal lappets, all undivided, the first extending along
the sides of the front half of the head, the next pair diminutive,
and the third pair largest, extending along the sides of the
branchial region. Pectoral fins rounded, relatively large, ex-
tending nearly as far back as the origin of the pelvic fins, which
are also rounded and about two-thirds as wide as the pectorals.
Dorsal fins rather small and apparently nearly equal in size; the
first dorsal arising opposite the hinder limit of the pelvic fins, the
second ending in advance of the much smaller anal fin, which is
close to the lower lobe of the caudal. Body and fins covered with
very fine shagreen, of which some granules between the pectoral
fins have a fluted sculpture.

Description of Type Specimen.—The fossil, which is shown of
the natural size in Pl. I. fig. 1, 1s exposed in its anterior half
from below, in its caudal half from the side. The snout is short
and bluntly rounded, and the rami of the jaws are vaguely seen,
meeting in an acute angle at the symphysis, where there are
remains of a cluster of very slender, smooth, pointed teeth. The
branchial region is relatively long, but the branchial arches are even
more obscured by the crushed shagreen than‘the jaws. A single

i aa

Vee DR. A. SMITH WOODWARD ON

pair of large dermal lappets (IIL), widest in front and gradually
narrowing backwards, extends along thé whole length of the bran-
chial region, Another pair of minute simple lappets (II) is well
seen just in front of this, and ‘the mode of staining of the fossil
suggests that there is a long and narrow fringe of skin (1) along
each side of the rostral region. The vertebral centra are much
constricted and smooth, but where broken they seem to exhibit
traces of some secondary calcification round the primitive double-
cone. Their arches are not distinguishable, but the comparative
shortness of the centra in the front part of the caudal region
evidently results from diplospondyly. The large pectoral fins are
remarkably rounded, slightly longer than wide, and the stout
basal cartilages do not extend more than half-way towards the
distal margin.» The long unjointed proximal radial cartilages are
well seen in the left pectoral. The tapering ascending parts of
the pectoral arch are crushed backwards. The pelvic fins, which
are much longer than wide, are supported in their basal half by
very stout radial cartilages ranged along the basipterygium, which
is not produced into claspers. ‘The individual represented is
therefore female. The two dorsal fins are crushed downwards to
the left side of the fossil, and seem to have been nearly equal in
size; but the parts projecting beyond the edge of the tail
probably represent only their apical halves. The first dorsal,
which is rather fragmentary, arises just behind the end of the
pelvic fins, while the second must have been completely in advance
of the anal. The anal fin is relatively small, short, and rounded,
and close to the lower lobe of the much extended caudal fin. The
lower lobe of the latter is clearly notched near its distal end.
The head, trunk, and fins are completely covered with very fine
shagreen. Most of the granules appear to be flat and smooth,
but some are pointed, and a few on the back of the trunk between
the pectoral fins are both pointed and slightly enlarged and
coarsely fluted.

Affinities —So far as preserved, there is nothing in the fossil
thus described to separate it from the existing genus Crossorhinus,
but it is distinguished from all known species* by the simplicity
‘ of the dermal lappets fringing the head, and by the relatively
large size of the pectoral fins.

Family PROTOSPINACIDA, nov.

Body depressed, but base of pectoral fins not produced forwards.
Vertebral centra well calcified (probably tectospondylic). Radial
cartilages of paired fins not extending to the margin; two dorsal
fins on the tail, each with an anterior spine; anal fin present.

* Compare C. Tate Regan, P. Z. S. 1908. pp. 354-357, pl. xi. fig. 2, pl. xii. fig. 2;
also Ann. & Mag. Nat. Hist. [8] vol. iii. (1909), p. 529. J. Douglas Ogilby &
A. R. MeCulloch, Journ. Roy. Soc. N.S. Wales, vol. xlii. (1908), pp. 269-280, pl. xlii.,:
pl. xiii. fig. 1. For skeleton see also W. A. Haswell, Proc. Linn. Soc. N.S. Wales,
vol. ix. (1884), pp. 92-98, pl. 1. figs. 6-8, pl. ii. fig. 13.

TWO NEW EDASMOBRANCH FISHES. 233

Genus PROTOSPINAX, nov.

Snout short and obtusely rounded. Teeth small, compressed
to a sharp edge. Pectoral fins extending as far backwards as the
pelvic pair; dorsal fin-spines large, laterally compressed, and
smooth, the first inserted opposite the pelvic fins; anal fin very
small, close to the elongate-ovoid caudal, which is not notched.
Shagreen dense and fine, none enlarged; lateral line supported
by a series of calcified ringlets.

PROTOSPINAX ANNECTANS, sp. n. (PI. I. figs. 2, 3.)

Specific Characters.—Attaining a length of about a metre.
Length of cranium slightly less than one-fifth, length of caudal
fin about one-sixth of the total length. ‘Teeth smooth and
lozenge-shaped, their sharp-edged crown sometimes with a
prominent middle point. Antero-posterior measurement of
pectoral fin about equal to the length of the cranium, and nearly
twice as great as the length of the pelvic basipterygium. Dorsal
fins about equal in size, the first arising slightly in advance of the
middle of the fish.

Description of Type Specimen.—The se which is shown of
one-sixth the natural size in Pl. I, fig. 2, is very fragmentary,
but there are definite points of contact between the pieces of
rock in which it is contained, and most of it is preserved in
counterpart, so that its general shape and proportions are recog-
nisable. ‘The head and trunk are seen directly from above,
while the greater part of the tail is exposed in side-view. The
edges of the head and fins are sharply outlined by fine dense
shagreen, while the distinctness of part of the margin of the
caudal region is due to fossilised muscle. The cranium is well
calcified in the usual small tessere, and evidently not much
distorted. Its postorbital part is about as broad as long, and the
postorbital processes are small and slender. There is very little
constriction between the orbits, which are completely within the
hinder half of the cranium. The olfactory capsules form relatively
large rounded lateral prominences in the middle of the cranium.
The rostral part is short and wide, not tapering but nearly
truncated in front, and remarkable for the large size of the
elongated anterior fontanelle which extends backwards between
the olfactory capsules. There is no indication of a posterior
fontanelle in the cranial roof. ‘The jaws are not seen, but there
appears to be a vague trace of the mandibular articulation on the
right side well behind the occiput. All the vertebra are crushed
and broken, but they show much secondary calcification round
the primary double-cone, and this seems to have been in concentric
lamine (on the tectospondylic plan). As in many other fossil
Elasmobranchs from the lithographic stone, the body-muscles are
well preserved; and it is clear that while in the abdominal region
each myotome corresponds with one vertebral centrum, in the
anterior part of the caudal region each myotome comprises two

Dok DR. A. SMITH WOODWARD ON

vertebral centra. There is thus the common diplospondyly. The
vertebral arches are scarcely seen, except within the caudal fin,
‘which is displayed in direct side-view. Here the hemals are
apparently stouter and less inclined backwards than the neurals.
The pectoral arch is only imperfectly shown, but the right
pectoral fin is complete. It is relatively large, and the supporting
cartilages extend only about half-way from its insertion towards
the distal margin. The three basals are distinct, the propterygium
being comparatively small and narrow, the triangular meso-
pterygium about as wide as long, and the metapterygium longer
than wide but very little pr oduced backwards. ‘The radial
cartilages, which are not much longer than the basals, are rather
sparsely arranged and do not clearly exhibit any transverse
articulations. About 12 are arranged along both the mesoptery-
gium and the metapterygium. Faint striations are seen in part
of the fin-membrane, but there are no distinct remains of dermal

rays. The pectoral fins extend as far backwards as the pelvic
fins, which are much smaller. The pelvic basipterygium is long and
gradually tapering, and bears at least 17 radial cartilages, which
(like those of the pectoral) are not closely pressed together and do
not show any transverse articulations. The cartilages occupy only
half of the total expanse of the fin. As they are imperfect behind,
the sex of the individual is uncertain. Of the median fins, one large
dorsal, bordered in front by the remains of a smooth, laterally
compressed spine (d'), arises just behind the origin of the pelvic
fins; but it is very imperfectly preserved. The impression of the
fin-membrane shows some fine striations, which may perhaps
denote strengthening dermal rays. As the tail of the fish is
relatively long, this is doubtless the first. dorsal fin, but the fossil
is too fragmentary to exhibit the second dorsal. The caudal fin,
which is displayed in direct side-view and only incomplete at the
upper extremity, is long and ovoid, with the lower lobe the larger.
The membrane here again shows faintly some fine striation. Just
in advance of its lower lobe, a small deep and narrow fin is shown
(a.), with most of the outline defined by oxide of manganese.
Though its separation from the caudal is a little obscured by the
rough fracture of the rock, it is almost certainly distinct and may
be regarded as an anal fin. Fine shagreen covers the whole of
the trunk and fins. Near the margins it is especially smooth and
dense, but on parts of the trunk the granules are rather stellate.
On the trunk in front of the pectoral fins, and again on the
tail just behind the pelvic fins, it is interesting to notice that the
course of the lateral line is marked by a close series of incomplete
ringlets (fig. 2a), as in Chimeroids and in the extinct dog-fish,
Mesitera.

Young Specimen.—A second specimen in the British Museum
(No. 37014), from the same formation and locality, only 30 cm.
in length, evidently represents a young individual of the same
species. The cranium and vertebral column are in undisturbed
series, with the two dorsal fin-spines in their natural position on .

TWO NEW ELASMOBRANCIHI FISHES. 230

the tail; but the parts of the paired fins are scattered, and only
fragments remain. The specimen, however, is of special impor-
tance, because displaced portions of both jaws with groups of the
teeth are also preserved. The characteristic large anterior fon-
tanelle in the cranium is well displayed as in the type specimen.
The teeth (fig. 3) are relatively small and closely arranged, several
series evidently functional at one time. Their exact shape is
difficult to determine, but they seem to be transversely elongated
rhomboids, with a low crown, which is smooth, compressed antero-
posteriorly to a sharp edge, and sometimes rising in the middle to
a little cusp. Many of the vertebral centra clearly exhibit the
secondary calcification round the primary double-cone. <A frag-
ment of a pelvic fin seems to denote a male individual, and a row
of slightly enlarged, pointed shagreen-granules may have belonged
to the clasper. The two dorsal fin-spines (fig. 3a, d', d*), though
fractured, are shown to be nearly similar in size and shape, and
their length equals about one-quarter of the distance between
their insertions. ‘The spine of the first dorsal is supported not
only by a short triangular cartilage behind, but also by a larger
and more extended cartilage in front. Traces of the fine stellate
shagreen are seen on the rostrum.

Affinities. —The new genus and species now described evidently
represent a family closely related to the Spinacide, but still re-
taining the anal fn and a less specialised dentition. Protospinax
is indeed a generalised type such as might be expected among
Jurassic Elasmobranchs when the Batoide were beginning to be
differentiated. The Batoids themselves were first represented by
the Rhinobatide. and it is interesting to notice that one member
of this family (Belemnobatis) contemporary with Protospinax had
a spine in front of each of its two dorsal fins.

EXPLANATION OF PLATE I.

Fig.l. Crossorhinus jurassicus, sp.'n.; nearly complete fish, nat. size—Litho-
graphic Stone; Kichstadt, Bavaria. J, II, LIT. the three paired dermal
lappets. [British Museum no. P. 11211.]

2. Protospinax annectans, gen. et sp. n.; fragmentary fish, one-sixth nat.
size.—Ibid. a. anal fin; d!. spine of first dorsal fin. [British Museum
no. P. 8778. |

2a. Ditto: portion of lateral line of same specimen ene ged four times to show
supporting ringlets,

3. Ditto; group of teeth enlarged ten times.—Ibid. [British Museum
no. 87014. ]

3a. Ditto: portion of tail of same specimen, showing dorsal fin-spines (d}, d?),
nat. size.

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ON’ PATHOLOGICAL STATES IN EVOLUTION. Dat

14. The Function of Pathological States in Evolution.
By Moruey Rosertrs*.

[Received May 7, 1918: Read June 11, 1918.)

That dissatisfaction with much orthodox biological opinion is
growing can hardly be denied. Not a little of this feeling 1s
due to the fact that what is often given as explanation cannot
be resolved into factors capable of appreciation, and, possibly, ot
measurement, by the intellect. The theory has to be accepted
as more or less a matter of faith. Where there is a general
tendency to rely on authority, speculation is discouraged, for
orthodoxy everywhere rests on the native conservatism of man,
and even the revolutionary is capable at last of fatigue... As a
result, tentative hypotheses offered by the great leaders tend to
become objects of faith, and among their less enterprising
followers there arises a more or less fervent conviction that,
however unsatisfactory they appear now, they will presently
become demonstration. Thus the theory of the germ-plasm,
even in its later modified form, seems heid too dogmatically by
many: the ‘nature’ of inherited living matter accounts for
every organ as it appears; while all changes are due to obscure
variations of an advantageous kind which give the survivors in
the struggle a better chance. On analysis, such opinions do not
seem truly scientific, for the “nature” of the germ-plasm can
barely be distinguished from the directing entelechy of Driesch,
and if the Weismannian cloud of ids and biophors is now
somewhat condensed, the magic determinant still remains in a
concealed vitalism which is exactly analogous, as regards the
organism, with pantheism as regards the universe. Nor, if we
are told with certainty that altered characteristics are not trans-
mitted, is the theory of small advantageous variations much more
satisfactory, if we know neither how they come, nor how they
are inherited. To say so much must not be regarded as treating
with disrespect its great author, without whom we might still be
wandering in the barren field of teleology.

To regard these theories as hasty and, perhaps, unsound
explanations 1s not to accept without scrutiny the theory
of the transmission of acquired, or modified, characteristics.
Though this is a view that can be defended on the physico-
chemical grounds of catalysts which are measurable determinants
of a really scientific order, experiments to prove the fact must
take a very long time, and we are compelled to rely on other
methods of proof. That the experiments of Tower and Kam-
merer, for instance, suggest the transmission of modifications
cannot be denied. Such as oppose the general view that the
environment has thus an inheritable moulding influence on the
organism, seem to reply that these are only rare and doubtful

* Communicated by the SECRETARY,

238 MR. MORLEY ROBERTS ON THE FUNCTION OF

cases, Whereas the theory of inherited advantageous variations,
whether continuous or discontinuous, can be made responsible
for the whole of thetphenomena. As the conclusion is gradually
being strengthened that large variations of a Mendelian character
deal with other characteristics than those which are racial, all
who rely on inherited spontaneous variations are forced back on
the Darwinian view that small variations can gradually, if of an
advantageous kind, convert one species into two or more, and
that all living characteristics, or organs themselves, are due to
such a cumulative effect. It is, of course, inferred and definitely
stated by eee that any variation in the least degree injurious
would inevitably be destroyed. It is this statement I propose to
examine, and for the purpose of such an enguny, it must be
clearly understood what is meant by the word ‘disadvantageous’

or injurious.

At first sight nothing seems clearer. Why should we doubt
that any functional or organic failure is a handicap in the
biological race? By functional trouble of which the cause is
not obvious we mean some hindrance, which may be recovered
from, to normal or physiological action. It is due to factors
which, for the most part, are unknown. We do not doubt that
there is a failure somewhere, which, as regards certain cells,
might be called organic, but often we carers do more than guess
where the actual failure occurs. In that advanced disorder of
function which has visible lesions and destruction or irremediable
alteration of the individual parts of the machine there is un-
doubted organic disease. Can anything seem more certain than
the conclusion that any organism which fails in the established
functions of its species is as a fact severely handicapped, that
the variation is disadvantageous and cannot possibly be trans-
mitted either directly or by survival? There are, however, some
reasons for believing that this inference is inaccurate and that
the function of disease in evolution is of much greater import-
ance than that of mere elimination. But pathology has very
naturally been neglected as a study by biologists. On the views
generally held, it has seemed sutficient to recognize that disease
destroyed organisms which obviously left oftspring, if it left
them at all, that were handicapped even more heavily ‘than their
parents. It has been understood that their elimination was only
a matter of time and that neither their virtues nor their failures
could influence the race.

If there is one thing more than another which has struck me
when attempting to study these questions, it is that too many
men of science appear to believe that any serious investigation
of other branches than their own is for them a waste of time.
The physiologist ignores the pathologist, who in his turn is far
too likely to fix his eyes on morbid phenomena which cannot be
properly appreciated save by those with a knowledge not only
of normal function but of the general physiology which underlies
it. The same can be said of most workers in science, but in no

‘PATHOLOGICAL STATES IN EVOLUTION. 239

case is it more likely to occur than in that of the biologist, who,
by the very name and nature of his task, should inelude in his
apparatus a considerable knowledge of everything which deals
with the organic, and even inorganic, world. Science, however,
is kept in more or less water-tight compartments, and it seems
left to the mathematician to hold the opinion that his own
branch of learning has, somehow or another, deep relations with
all things, including life itself. Even by him it does not seem
to have been pointed out that in things living and non-living
certain principles of construction rule alike. However much
they were wedded to mechanico-physical explanations, biologists
have assuredly often ignored the fact that any organism is con-
struction, and knowing little of the laws of construction have
ignored basal facts familiar to every architect or even every
artisan. It was reserved for Wolff, in formulating his law of
bone-growth and reaction to stress, to propound a principle more
far- reaching than he recognized, when he showed that living
bone, reacting to normal or abnormal stimulation, can be proved
to develop in accordance with the principles of engineering and
architecture. ‘This iaw may, I feel assured, be extended to every
living tissue, and in such an extension will be found the key to
many phenomena still awaiting explanation.

To one who holds this view, the work lately done by Starling
on the “ Law of the Heart,” which shows that the force with
which the heart contracts is directly proportional to the length of
the muscular fibres at the end of the preceding diastole, is by
no means surprising. It is indeed on a par with the conclusions
of Wolff as regards bone, and might, I believe, have been deduced
from it or from the form I suggest, provided it is understood
that each varying tissue has its own acquired typical reaction.

If, then, it can be shown that disease has had.a profound effect
upon the evolution of all organisms, and that analogous results
are found in every kind of human constructive effort in such
numbers as to suggest as a law that all great variational develop-
ments result not from the happy-go-lucky aggregation of smail
advantageous variation or frcm discontinuous variation, whether
of a Mendelian character or not, but rather from partial failure
and repair, we seem to be in sight of a general principle of pro-
found importance. If this principle proves sound, it is obvious
that immense labour has been spent by biologists endeavouring
to explain life without seeking help from other workers. Thoug h
they may show some general knowledge of the cell, and even
special knowledge of the reproductive cells, I find few who appear
to have studied general embryology, to speak only of one branch
of physiology. On the other hand, many physiologists and patho-
logists have done good work in some branches of ev olutionary
theory. Bland- Sutton, i in his fruitful little bock ‘Kvolution and
Disease,’ pointed out that “Pathology is only a department of
Biology, and it is important to bear this in mind in studying
disease.” It is true that he went little further than to show that

240 MR. MORLEY ROBERTS ON THE FUNCTION OF

what is pathological in one organism may be physiological in
another and that many diseases are reversions, that is, failure in
normal growth. Yet this greatly needed to be shown, and it
is not to be expected of a great pathologist and surgeon, and
perhaps the less the greater he is in his own branches of work,
that he should-attempt tasks from which many of the biologists
themselves seemed to shrink. Claude Bernard made similar
remarks as to pathology. It is to be regretted that a stumbling
block was placed in the path of progress by Darwin’s hopeless
dictum as to the explanation of variation, just as another was by
Huxley when he declared consciousness an insoluble problem.
In every science great discoverers have too often delayed progress
as much by authoritative unsound opinion as they have advanced
it. Every Bible is first a book of revolution and then a refuge
for reaction. Yet no man can possibly know all he should know
for the purposes of his own work. ‘This fact affords the only
justification for those, who cannot pretend to profound knowledge
in any special line, attempting to solve problems which by their

nature are beyond the specialist. They may have been able to
grasp in a measure the general conclusions of each science and
by a happy, perhaps accidental, combination, show at least part
of the forest to those more particularly oceupied with the trees
themselves or the flora of the undergrowth.

It is remarkable that hitherto no one seems to have made the
observation that reaction to an actual, or threatened, breakdown
is one of the basal laws of all construction and organization.
Yet none can read engineering without observing that all
development has followed such lines. As new stresses are
introduced failure is threatened and steps are taken to obviate
disaster. What is a patch in one engine becomes organic in the
next. Since waste of energy can be looked on as pathological,
we observe the reaction in the engineer against such failures, as
the atmospheric engine is succeeded by improved forms ending
in the quadruple expansion engine. Many other instances could
be adduced in general or special engineering evolution, but
the best illustration of the facts which need elucidation can
perhaps be found in Gothic architecture. If such a demon-
stration of this general principle can be made it will go far to
obviate the objection, very likely to be made, that what occurs
in human construction has no relevance to the living organism,
especially if it can be suggested forcibly that human intelligence
is in itself a reaction, fal that the law obtains in developments
of all kinds. That trial and error are at the base of evolution is
indeed implied in the current teaching as to variation, and its
extension to intellectual processes will surprise no worker who
has had to deal experimentally with the unknown. We may
expect, but never know where to look for, failure till we see it.
When it is seen we can do our best, as reacting agents, to remedy
it. Having said so much, and leaving aside the wider implica-
tions of such views, we may turn to such a problem of construction

PATHOLOGICAL STATES IN EVOLUTION, vat

as the evolution of a cathedral, in the hope that it may throw a
light on other than architectural puzzles: merely observing, on
the way, that no general principle yet discovered is confined in
its application to one branch of knowledge. Having once found
it, our task is to employ it as a weapon of further analysis.

It is more or less a commonplace that function creates
structure, however Lamarckian that may sound; and in the case
of architecture of a religious order the function which constructs
is public worship. In fine climates the necessary structure is
often a roofless temple. In tropical climates a flat roof may be
needed as a protection against the sun. In temperate climates a
walled enclosure is insuitticient and a flat roofed structure cannot
keep out rain effectually or bear heavy snow. Thus arose the
pointed or sloping roof. But it has been said that ‘Gothic
architecture is not a style. Itis a fight.” The arch is a mighty
warrior. It gives and receives thrusts. ‘The sloping roof partakes
of the same nature. Need created it and the nature of materials
and the positional energy we call gravity caused thrusts which
endangered the simple walls of the building, walls at first meant
to endure nothing but flat roofs probably covered with brush or
the like material. To build stronger walls might have occurred
to the primitive architect, but as the danger was immediate, he
probably at once shored those in existence, and then built others
at a right angle to act as buttresses. In the meantime the
worshippers increased in numbers, and it is indulging in no
flight of fancy to suppose the later builder saw that if the new
external walls were roofed over and doorways cut into the main
building, there would be an immediate increase of space by the
creation of chapels. Such a series of embryonic additional walled
spaces, with further doorways in them leading to each other,
obviously gave him the aisles. ‘The flying buttresses which are
such a feature in great Gothic architecture had, I can only
suppose, a like origin. They were originally buttress walls carried
up to the roof. At some period a genius, already acquainted
with arcuated structure, saw that if the inside of these walls was
cut away they would still take a heavy thrust and lighten the
rest of the building. If, however, on being converted into such
slender stone shores they showed signs of yielding, what could be
easier than to pile some of the mater ial taken away on the base
of the flying arch and thus create the beginning of the pinnacle ¢
Though an architect might develop such a rough statement, he
would be the first to admit that it represents in few words much
of the evolution of a church; that is, he would own the structure
sprang from need, and that each new need caused a constructional
failure which, when strengthened and corrected, was the cause of
further structure. He would further tell us that all good orna-
ment is organic; that ib springs naturally from the work already
done, being in its origin just the little more needed to give a
margin of safety, though on it later are exercised the esthetic
faculties of man, which are e again a response to the need of full

242 MR. MORLEY ROBERTS ON THE FUNCTION OF

satisfaction for the instinct of workmanship. Human ornament
is in fact strongly homologous, if we may use that word here,
with the beauty of very energetic birds, who carry out by virtue
of their free energy the extension of structures and colours
already existing in their less brilliant forms. ‘hat, however, is
by the way. ‘The main fact we are concerned with is that the
structure as a whole evolved through trial and error, through
failure and repair, through a threatened structure to a more
complete and adequate one for increased function. In a word,
the great origin of structure was failure after failure duly com-
pensated for. Is there any reason for believing that variation
in the structure of ving organisms follows exactly the same
principle¢ Are we entitled to say that the mammal, for in-
stance, with all its complexity, is the result of infinite ages of
functional failure or disease which was met by processes of repair
and reaction? In a word, can we speak of the evolutionary
value of disease, of impaired function, of disadvantageous varia-
tions ¢ It seems possible to do so, if what is true of one structure
is roughly true of another.

It may seem absurd to speak of the value of disadvantageous
variation, but it is no more absurd than to imply that all
variation is advantageous because it 1s perpetuated. What is
useful at one period may be harmful at another, and embryolo-
gists thoroughly understand that developments useful in feetal
or larval hfe may open up many dangers for the adult. ‘The
real point to be considered is whether organisms as species do
not vary and run great, even largely destructive, risks by an
increased pressure of function which, in the few that finally
react or whose descendants react to such stress, results at last
in structure that is advantageous as altered. The given variation
in itself may be a failure of what was normal function in the
species, and we should therefore as pathologists or physiologists
speak of it as. disease, but if the few that recover become a new
species, a mended race, it is no longer disease. After many
generations it may be truly advantageous to individuals. Have
such processes occurred in the evolution of organisms as they
undoubtedly have in the arts and social progress, where we often
observe political failure or organization result in ad hoc reaction
which leads to a changed social form? J have no doubt that
they do, and many organs in mammals, to speak only of them,
show it. It is, in fact, a universal principle. As beaveis patch
up a dam when it yields or threatens to give way, so tissues,
organs, and societies react to threatened disaster. In no tissue
is this clearer than in bone. It is true that Wolff's law only
deals directly with mechanical stresses, since it runs—-‘‘ every
change in the form and position of the bones or their function is
accompanied by certain definite changes in their internal archi-
tecture and by equally definite secondary alterations ef their
external conformation in accordance with mathematical law;”
but I hope to show reasons for concluding that such a law may

PATHOLOGICAL STATES IN EVOLUTION. 943

be stated in more general terms and applied to every tissue and
organ, provided we add that the more complex the tissue or the
organ the greater the liability of failure, and that each tissue
reacts in a typical way.

It is unnecessary to go into details of osteogenesis and mor-
phology, It has been recognized by eugineers that the head of
the femur is formed exactly in accordance with mechanical law.
Had any of them been required to design a structure fit for
undergoing the stresses borne by the femur in its development
and after-life, he would have sketched a figure extremely like it,
not only in its general shape, but in the trabeculee which support
the bone in every direction where extra stresses are applied by
normal function, The important point to note is the fact that
femoral developement follows stress in individual development,
from which we must draw the conclusion that it followed stress
during evolution, not that its value for complex function was
gradually increased by chance or “spontaneous” variation, unless
we attribute to “spontaneous” a meaning which Darwin never
gave it, seeing that he denied knowing how variation arose. All
the variations were definite responses, and it is easy to infer that
before response became rapid and easy every kind of disaster
and disablement must have occurred to those subjected to re-
action-provoking stresses. The very process of adaptation (and
on these lines “adaptation” is no longer a mystic word) implies
long periods of disordered function and poor structural response
even in those who survived after repair. But now bone is so
plastic and fluent that when it is grafted the osteoblasts and
osteoclasts shape it according to the form of the main bone of
which it becomes a part.

When we speak of repair it may be noted that the treatises
on this subject are strictly limited in their purview. ‘They mostly
follow Hunter, a vitally important figure in the history of
pathology and indeed of all medical science, who, however,
lacked the apparatus of knowledge now at every one’s disposal.
We learn a great deal about the repair of wounds and fractures :
of .the functions of the fibroblasts or of the wandering cells
of the blood-stream, and are told, lately, much of regeneration,
but of the evolutionary value of organized exudations we hear
nothing. Nor has it been suggested that it is to this and
analogous processes that much new structure is due. © That this
is so is strikingly apparent, as I shall attempt to show, in many
organs of a highly specialized type. In no structure, perhaps, is
the process so clearly seen as in the mammalian heart, which is
a perfect museum of evolutionary failures and dislocations, com-
pensated for by an extraordinary complication of patched-up
tissues and organized exudations in which, perhaps, one tissue
takes on the functions of another and some evolutionary rem-
nants long survive without function. J was, indeed, first led to
take this generai view of the variational value of pathological
conditions by observing that the heart, when laid open from any

244 MR. MORLEY ROBERYS ON THE FUNCTION OF

aspect, powerfully suggested an organized or cured aneurism.
Many must have made the same observation, even if they have
not come to similar conclusions. The anatomist and pathologist
perhaps know their subjects too well and are necessarily greatly
dominated by current theory. The general adaptation of the
heart to the work it performs may well delight the anatomist as
he studies its machinery. His main business is not evolution.
The pathologist on the other hand, observing its many failures,
is scarcely likely to discern that by failure. itself may come
eventual perfection, and while the physiologist considers its
functions rather than its apparatus. he studies it as it 1s, not as
it was. In each case the observer may not see the forest for the
trees. Yet when we look at the partially repaired aneurism with
its fibrous growths and turn to the opened heart, the essential
relationship of the chorde tendine, for all their definite functions,
to the rude fibres of an aneurism is obvious. Is such a likeness
an accident of evolution and pathology, or are we to consider the
heart as much an organized dilatation sac of the whole fused
circulatory canal as the cured aneurism is of a part of 16? It is
in embryology that we seek for confirmation of what is suggested
by anatomy. But even anatomy alone offers powerful proof of
the view that. the heart, as we know it, is the latest result of
repeated failures of the circulatory canal under strain and of the
repairs effected by the stressed tissues in their response to
changed and abnormal stimuli, just as bone alters under its
particular stresses. During embryological life there is found in
the heart a small patch of non-functioning muscle in the anterior
segment of the mitral valve. Its presence is intelligible if we
consider it a relic of a disrupted and repaired organ. The
muscles of the heart are obviously homologous with those of
the arteries. Yet they have become striated although they are,
of course, still involuntary. Non-striated muscle is the earliest
in evolution. It seems that the increased functioning of the
cardiac muscle has converted it into its striated form so that it
resembles skeletal muscles, which are much more active than
non-striated muscle. The whole histology of cardiae muscle
probably represents the result of great strains. Structures such
as the disks or bands of Ebarth are found nowhere else and may
be the result of peculiar stress. There are even portions of
muscle which no longer perform muscular functions. Their
fibres do not contract but serve instead to conduct stimuli as
if they were nervous tissue. All tissue is conductive, but the
bundle of His, with its Purkinje fibres, which carries the impulse
from the sino-auricular or Keith-Flack node to the ventricle,
transmits messages at ten or twelve times the normal muscular
rate. When it fails there is heart block. In the embryo the
valves arise from the cardiac walls and are composed of muscular
tissue which by the action of fibroblasts gradually become non-
muscular. This must have been originally a pathological process.
It is a reversion, a degeneration made use of. We observe

PATHOLOGICAL STATES IN EVOLUTION. 245

analogous, or shall I say homologous, results in the hypertrophied
heart. The normal male heart weighs about eleven ounces. In
some cases of aortic stenosis it may weigh over thirty ounces.
In such hypertrophied muscle are often found fibrous tissues
which probably represent the connective tissue of muscular fibres
which have atrophied from overstrain. The chorde tendinee of
the mitral valve are less muscular and more fibrous than the
same attachments of the tricuspid. This adaptation difference
lessens strain on the thinner right ventricle. It has, indeed,
remained thinner on that account. In the reptile with a function-
ing foramen the valves are purely mechanical, as pressure is
relieved by the patent orifice. The fossa ovalis in the mammal
is a remnant of the early communication between the auricles.
In a large number of normal hearts there is a small valvular
passage yet remaining in the left margin of the fossa. None of
these phenomena seem capable of explanation as the result of
spontaneous variations arising from some theoretic instability of
the organism. ‘To argue that they are 1s to give biologic mystics
a chance. It appears obvious from all these facts taken together
that cardiac evolution has been a series of caused variations due
to increased and varying stresses which acted not only as a
moulding force on the shape and musculature of the heart but
on all its appendages. In the muscle of the ventricular walls
with its extraordinary complexity of layers and interlaced fibres
lies powerful evidence of such reactions. In both ventricles
there are seven muscular layers, while in the arteries there
seems but one. In the left ventricle these layers are obviously
thicker and stronger than in the less stressed right cavity. But
how did the ventricular cavities acquire more layers than the
arteries¢ There is obvious reason for believing that stress can
be responded to by increase of muscle fibre during evolution.
In the gravid uterus the smooth fibres of the wall increase to
eleven times their normal length and are from two to five times
as broad. There may be new fibres in it. I doubt if any one
knows. But in evolution new fibres are undoubtedly found. In
the arteries, the fibres of non-striated muscle in the tunica media
are for the most part circular, but they appear to have more or
less longitudinal branches which interlock with like branches of
the neighbouring fibres. One of the most prominent features of
an individual aneurism is the thinning out, and sometimes the
disappearance, of the tunica media. The muscle fibres in such
cases are completely broken down, and ifthe aneurism is repaired
in individuals the work is done mostly by an increase of the
connective-tissue elements. The process 1s said by some to be a
reparatory endarteritis, in which the tissues of the adventitia
proliferate actively. But the evolutionary process has obviously
taken the path of increase and reactive proliferation of the
muscular elements of the media.

It is often observed that the aneurism which displays sufficient
reacting power to the stresses of the blood stream accumulates

Proc. Zoou. Soc.—1918, No. X VITI. 18

246 MR. MORLEY ROBERTS ON THE FUNCTION OF

blood clots in layers, and it is therefore all the more interesting
to note that in the embryo the columne carnee and chorde
appear to rise from a spongy network which at an early age fills
the primary ventricle. Such origin is strongly suggestive of
some process analogous to blood clotting or to an irritative
reaction of the embryonic ventricular wall. The evolutionary
dilatation sacs which I suggest were originally pathological can
be seen in the embryo during the process of their formation.
The path laid down by pathology is trodder by physiology. It
follows that during evolution there must have been an immense
destruction of organisms whose circulating canals did not react
and numbers which retained their unaltered “specific” characters.
The same process goes on to-day. Though many die of cardiac
disease, it may be that much youthful functional trouble and
even more serious adult disorders are even now remoulding the
heart. No organ is perfect: if it does not degenerate it pro-
gresses. ‘Though such processes are ‘ disease,” it by no means
follows that they will be destructive, any more than that the
functional incapacity of the tricuspid valves in athletes, which
probably precedes what is known as ‘‘second wind,” is anything
now but a cardiac safety-valve.

As we learn more of the heart and its latent capacities we
may perhaps say with the late Dr. H. G. Sutton, “we trust
nature too little, to say the least of it.” But there are, of course,
ereat difficulties to overcome before we can hope to understand
how the cardiac musculature has altered and may still be
changing by the addition of new fibres. As yet, little is known
of myogenesis. Like a neurone, a muscle cell seems to last a
life-time, and though both may degenerate or die, neither pro-
liferates after the early period of development. But whatever
their histogenesis, new fibres do appear in evolution. Harvey
did not refuse to believe in the validity of his own conclusions
because he lived before Leeuwenhoek. With considerable hesi-
tation I venture to suggest that morphogenetic stress is at its
height during fcetal development. The child in utero has not,
perhaps, the calm and happy life commonly attributed to it. On
the contrary, it probably leads a strenuous existence, and if it
inherits a new weakness this is shown just where and when new
stresses find plastic embryonic tissues to respond to them. If
such a speculation is sound it accounts for many phenomena.
But in any case, whatever the machinery of inheritance and
evolutionary repair, it 1s certain that new fibres arise where they
are needed.

If such views in any way represent the biological history of
the heart, it is obvious that many of the opinions of variation
usually held are without foundation. Every variation is definitely
caused; it is in no sense accidental or spontaneous; it may not
be even at once advantageous to the individual: on the contrary,
it may be a severe handicap which puts greater general stress on

PATHOLOGICAL STATES IN EVOLUTION, 247

all who experience it, though such stresses fall short of those
which cause death. Variations of this order may only be
advantageous to the whole species as a continuing race. They
inay destroy, and doubtless have destroyed, individuals without
number at an earlier age than the usual life-period of the
unvaried type. We may possibly imagine a part of humanity,
now responding to stresses which make the heart do more work
and fail earlier, displaying such energy during their shorter
life as to displace those with a normal cardiac mechanism which
survives to the average age of man. It is to be inferred from
these considerations that the structure of an organism is not a
congeries of minute fortuitous advantageous variations, nor the
gradual massing of details in an orthogenetic line, nor the result
of large discontinuous variations due to chromosomatic in-
heritance, but a complex of definite reactions to definite stresses.
The true theory of living structure is that its growth is neither
casual nor foreseen, but that is what we may call, in political
language, the ‘‘ opportunism ” of the organism as a whole. Every
advance is a forced, even a desperate, experiment. Life, like a
hypothesis or a dain, is built up by stopping leaks.

The evolution of the stomach seems to have followed the
lines suggested for cardiac development. From the physiological
point of view, a straight intestinal tube which becomes dilated
cannot be considered anything but pathological. It has failed
under the stresses imposed on it, but the organism which
reacted turned a weak dilatation sac into a strong perma-
nent food pouch. The results to the reacting organisms were
many. The ingested food became temporarily static, was more
thoroughly dealt with, and the organism was not continually
feeding. Its whole available energy was not devoted to nutrition:
it had time at its disposal and could develop other functions
leading to further structures. That the human stomach is such
an organized failure is suggested forcibly by the musculature. In
the small intestine this is composed of two layers of fibres,
circular and longitudinal. In the stomach it is made of three
sets, an inmost layer of oblique fibres being added. This oblique
layer is obviously a later growth and, as would be expected on
the lines laid down as to disaster and repair; its strongest fibres
are found just where they are wanted, that is, supporting the
greater curvature or dilatation of the stomach. This later
layer is naturally less well developed than the longitudinal and
cireular fibres. Other oblique fibres are formed about the pylorus
where they form the sphincter. I suggest that these oblique
muscle fibres arose as points of strain, under intense stimulation.
The dilated pouch has weacted in accordance with mechanical
law, just as the heart did with its more complex arrangement of
oblique fibres woven into a structure capable of giving in the
left ventricle a thrust of over fifty pounds. The reacting organism
is no fool of a mechanic either in its bones or its muscles, and

LSs

248 MR. MORLEY ROBERTS ON THE FUNCTION OF

these phenomena are additional reasons for extending Wolff’s
law to all tissues. If protoplasm did not so react there would be
no problems to solve.

Such views on the mammalian gastric apparatus are so
obviously supported by the embryology of the organ that there
is no need to go into details beyond noticing that in the fourth
week there comes the first dorsal bulging in the fore-gut. Of
the curiously shaped fundus, Keith has remarked that it is in
origin like the cecum, but I do not think it has been suggested
that its form has possibly been moulded by the presence of the
large air bubble so often seen in X-ray photographs. It is an
elastic air-reaction pouch just as the whole stomach itself is a
food-reaction pouch. It began to give way there, but the process
was stayed. So far as I am aware, it is not provided with
obvious oblique fibres. Further investigation may find them.

Tf evolution is still proceeding, is it absurd to suggest that
the common disorder known as dilated stomach may be a patho-
logical process actually in the process of becoming physiological ?
According to some physicians, few modern stomachs do not
suffer at times from an amount of dilatation which is patho-
logical; 7@. e., their gastric musculature fails to react correctly.
The stomach may yet be such a functioning dilatation pouch as
to enable the human race to do with no more than one meal a
day or even less. Our descendants will have all the more time
for work. This by no means implies that’ the empty stomach
should be any larger than it is now in healthy subjects. Before
the invention of X-rays the gastric apparatus was always
pictured in text-books as usually seen on the post-mortem table.
The dead stomach was shown as the portrait of the live one:
the weakened pouch of the sick man as that of the live and
healthy subject. But nowadays it is known that such extreme
dilatation is natural only when a large meal has been taken.
When the healthy stomach is empty it contracts so that it
nearly resumes its ancient cylindrical character and is of a size
not much greater than that of the small intestine. With further
development it might hold still more and yet react in the same
way. The suggestion that functional failure or disease which
becomes organic and destructive in many, may, in reacting and
surviving organisms, alter their outlook on life and all their
activities, seems to me powerfully reinforced by these considera-
tions. The disadvantageous variation does work, and finally
improves the race.

It can even be shown that disadvantageous variations actually
become permanent racial characters. We may consider hernias.
tm the prone position of most animals, hernial sacs, now known
not to be essentially pathological until they are forced open by
mechanical stresses or relaxed by organic weakness, are not a
great source of danger. They may even be considered as an
additional means of securing the peritoneum to its connective
tissue. During the processes of evolution, however, a mammalian

PATHOLOGICAL STATES IN EVOLUTION. 249

hernia seems to have occurred almost universally and to have
established itself as normally physiological. The twnica vaginalis
propria of the testis is actually part of the original peritoneal sac,
as can be seen in the embryo. During feetal life it is separated
from the parent tissue. In whatever sense we now call such a
change physiological it seems impossible to regard it as originally
anything but pathological. Is it too startling to declare that it
is an evolutionary sloughed tissue such as is often seen in
strangulated hernias ? i certainly do not know how we can
describe the scrotum as anything else than the coverings of an
evolutionary hernial sac which is not only of no advantage but a
positive danger to most male animals. In some, the pigs for
instance, the testicles do not descend into an external pouch but
are supported and protected by the normal skin tissues, not by a
thinned and delicate integument of later development like the
scrotum, a tissue still scantily supplied with the non-striated
muscular fibres which might have reinforced it and are perhaps
now developing slowly. When we consider the rarity of mus-
cular fibres in human skin tissues in comparison with those of
animals, their greater frequency in the scrotum and perineum
suggests that they are a reaction product. They act in the
dartos, or deeper layers of the scrotal dermis, at right angles to
the ruge and are something of a support. The pink colour of
this structure is due to the presence of these muscular fibres.
They are not connected in any way with the cremaster muscle
and therefore not affected by the cremasteric reflex. In no sense
can the descent of the testes be called advantageous. It causes
a weak spot, recognized as such by men and animals. The
Japanese wrestlers are trained from youth to return the testes
into the inguinal canals. If the translation of the testis from
a safe place to an exposed one has had any good results they
have been indirect and only discoverable, though not yet
discovered, over long periods during which the change must
have been disastrous to many. ‘To argue that they were advan-
tageous to begin with is to destroy the authority of reason.

It may seem an undue extension of the view that pathology
has played an immense part in evolution if it is suggested that
it was upon pathological conditions that the very existence of
the Metazoa depended. There can be no doubt that they origi-
nated from some protozoon by a failure of normal physiological
fission. We see here how theories of disease may be modified
according to the point of view taken. From that, shall I say, of
a protozoan Hippocrates or Hunter nothing can be more obvious
than that a failure of mitosis would be a calamity, the birth of a
monster, of Siamese twins, among the normally constituted uni-
cellular organisms. It is still in the processes of reproduction
that we find the strongest evidence of the part played by disease.

When considering such problems in this light 1t seems some-
what difficult to account for the satisfaction of many with the
theory of small cumulative advantageous variations. What ground

250 MR. MORLEY ROBERYS ON 'THE FUNCTION OF

is there for imagining such machinery could result in a complex
series of adaptations such as the uterus and what we may call its
habits and customs in dealing with the embryo from the entrance
of the ovum till birth? Even those who adapt to their own
ideas some theory of large discontinuous variation will, in the
end, be compelled to attribute the uterine growth and functions
to a mystic power or virtue in the original germ. They may
follow some philosophers and “unpack” powers out of a con-
jurer’s bag without telling us how they got there. Yet if we
regard the uterus as the result of tissue reactions under abnormal
stimuli, being guided in research by the processes seen every day
in disease, che variations, whether small or large, continuous or
discontinuous, assume an aspect neither fanciful nor mystical,
and vur need for biological faith is reduced to a decent scientific
minimum.

The fact that the embryo acts upon the maternal organism as
a parasite against which the mother has to be protected,
commonly recognized, but I have not seen the obvious conclusion
dvawn that the whole history of the mammal must have been
due originally to a pathological accident in some one or more of
their ancestors. The mammalian animal still lays eggs, but they
are not extruded. When such retention first took place, it must
have been due to an accidental pathological delay of the travelling
ovum, owing perhaps to catarrh of the tube. Even now the
mother has to be rendered immune to the products of the offspring.
Many of the phenomena of early gestation are those of immuni-
zation, in many cases a very slow process, as 1s shown in human
beings by vomiting and malaise. It has, moreover, not been
clearly or generally recognized except by pathologists that the
very methods by which the ovum attaches itself to the uterine
wall are, so far as the hostess is concerned, actually pathological
and bordering on the malignant. Yet they have resulted in a
series of protective reactions which save the parent and permit
the growth of the parasite. The method by which the ovum
becomes partially buried in the tissues is obviously of a destruc-
tive kind and curiously analogous to the malignant processes
seen in chorion-epithelioma, Bland-Sutton remarks, ‘This disease
is instructive because the erosive action of the trophoblast is the
physiological type of the invasiveness so characteristic of many

varieties of cancer.” It may, 1 think, be added that it is the
balance established by reaction which makes the trophoblastic
action physiological.

That the influence of the ovum on the undeveloped tube must
have been of an exceedingly dangerous character is now seen in
tubal pregnancies during which the chorionie villi frequently
penetrate the wall of the tube, which does not react as powerfully
as the uterus. Such a process in the uterus, which is itself a tubal
dilatation, is now normal because these villi, the earlier nutrition
roots or organs of the parasite, are prevented from injuring the

PATHOLOGICAL STATES IN EVOLUTION, 251

uterine wall irrevocably by the transformation of the reactive
uterine decidua and the chorionic villi and the allantois of the
foetus into the combined temporary organ known as the placenta.
It may be noted that the non-placental mammals are less
exposed to the destructive and toxic effects of their offspring
as they are. born at an earlier stage than in the case of the
deciduate mammals. The marsupial foetus is about half an inch
in length when transferred to the milk-pouch. It is impossible
to look at the placenta without recognizing that it is what we
may call a compromise growth, one which serves the embryo
without destroying the parent hostess. That all mammals are
not yet fully armed against any morbid alteration of function
in the penetrating chorionic villi is seen, as suggested above, in
chorion-epithelioma, where the energy of the villi trophoblasts
leads to a malignant overgrowth of the epithelial elements,
which the maternal tissues fail to inhibit. The hydatid mole,
which does not as a rule become malignant, is a case where such
inhibition has been sufficient. All these cases, malignant or
benign, support the view held by many that malignancy always
depends on the failure of some tissue inhibition, and that if
bacilli play any part in the drama it is that of helping to upset
the balance between tissues which are fundamentally hostile
though they exist normally in symbiosis. These phenomena
regarded as a whole establish on a firm foundation the
view that the uterus and its reactions during gestation are
definite protective processes or variations springing originally
from a purely pathological accident in some ancestors of the
mammalans. However complex the embryology of the uterus
and its appendages, the broad facts are compatible with this
view, which is strengthened by the later parasitic history of the
offspring after birth. ‘The mamme appear to be a compromise
between the needs of the infant and the protection of the
mother: they originated in sore or tender spots on the epithelium
most exposed to the assaults of the parasite. The growth of the
nipple is a complex variation depending on the mechanical action
of sucking with a reaction proliferation of the epithelial elements
of the sweat and sebaceous glands and an increased blood-supply
as special maternal protections against oral infection. It seems
to me that few stronger instances can be found of the fact that
the development of many organs, if not all of them, is the result
of direct reactions or adaptations which are in the nature of
repair to tissues otherwise likely to suffer disastrously.

It is large macroscopic results of this order which enable us to
reason about other finer reactions, and even help us to link to the
general process those of a microscopic and_ ultra-microscopic
character which we class under “immunity.” Such phenomena
are reactions under stress which, by the provocation of catalysts,
influence life. Much of human character, even, is similar re-
action, perfect or imperfect, to the infections to which the race

x

D2, MR. MORLEY ROBERTS ON THE FUNCTION OF

has been and still is exposed. Thus psychology itself must at
last be classed as the result of physical reactions, a conclu-
sion fully in accord with the work’ of Pavloff on conditioned
reflexes.

If any further illustration of the conclusions so far suggested
1s necessary, it may be found in the growth of the mesentery.
It has often been pointed out that the embryonic processes by
which it is formed are histologically those of plastic organized
exudations. The attachments of the whole tube do not come
about at the same period of fcetal development, and it seems of
significance when we note that the mesentery of the small gut has
an oblique attachment, to the posterior abdominal wall from the
duodenum to the right iliac fossa, only found in animals which
have assumed the upright posture. This comes into existence as
late as the fourth or fifth month of fcetal development. Before
this band was formed there must have been a great series of
disasters, for even now the last part of the mesentery to become
attached to the abdominal wall, that is, the angle between the
ileum and ascending colon, sometimes remains free. A volvulus
may easily form there by rotation of the ileo-colic loop. The
whole history suggests a series of lymph effusions, caused by
pathological states, some of which were sorted out by the lethal
process of natural selection, the remainder surviving and leaving
offspring with the hability to or ganize the effusion in the safe
way. The pathology of those cases in which what are known as
Lane’s Kinks can be found is obviously of a similar character.
The stasis of the affected bowel causes lymph effusion and the
formation of a band which is morphologically homologous with
the early mesentery.

After reviewing phenomena such as these, the conclusion
seems inevitable that single small favourable variations have
not done the whole work of evolution. They may play their
part as correlated changes, but they then take their place in a
series of which the causes can be recognized. In combination
with reasonable views of use and disuse and of increased or
decreased blood-supply they may, perhaps, be held to explain
such phenomena as the delicate co-aptation of some cardiac
valves. Their place in the explanation of the phenomena of
mimicry seems obvious. But though they may help us to com-
prehend how tissues become finished structures if they are

- combined with the results of functional energy, they yield no

hint as to great or decisive developments and the mechanism
involved in them. If the reasons adduced for the thesis laid
down carry any weight, it is obvious that many, if not most, of
the really decisive variations in all internal structure depended
and still depend, not on variations which can be called favourable
but on those that for the major portion of the organisms involved
are directly disastrous: not on variations which are small but
on those which are big enough to be appreciable as the cause of

PATHOLOGICAL STATES IN EVOLUTION. 253

immense functional and structural results; not on changes which
can in any sense be called spontaneous, by which we may suppose
is meant those no cause can be assigned to, but on variations,
which, though they occurred ages ago, were obviously due to the
very same causes that the pathologist can demonstrate to be
working at the present day. Only such organisms as respond by
direct reactions in a manner that finally turns out to be useful,
or at the very least compatible with life and reproduction, are
able to survive. The whole of growth and development thus
becomes largely a function of effective morphogenetic repair to
organic failure and disease.

Though this is not the place to deal at length with the vexed
question of transmission of modifications, it may be remarked
that the foregoing arguments seem to imply that such alterations
as a matter of fact are inherited. i think some progress can be
made if we simply assume provisionally that organisms do tend
to repeat themselves and that it 1s wnlikeness rather than likeness
which requires explanation. We know that gross unlikeness is
almost always due to a lack or excess of some internal secretion,
hormone, or enzyme, and from this it may be inferred that
likeness is due to such catalytic machinery coming over in the
zygote, and to each differentiation producing anew its own
peculiar products which stimulate or inhibit further growth and
differentiation. Some time ago I was struck by a remark of
Starling’s that each new organism seemed a fresh “creation.”
He gave this up on account of the difficulty he found in the
‘time element” of the problem, but I venture to think he was
right in his surmise. There is a growing body of opinion in
support of this view, as the names of Cunningham, McBride,
Dendy, and Bourne seem to bear witness. We must certainly
take into account these regulators of metabolism, and if we
accept the view that hypo-thyroidism determines cretinism, or,
in the adult, myxedema; that hyper-thyroidism is the direct
cause of the phenomena seen in Graves’ disease, just as hyper-
or hypo-pituitarism causes giantism ov infantilism in children
while a later overgrowth of the gland causes acromegaly, I see
no difficulty in accepting the hypothesis that growth is deter-
mined, 7@. e. stimulated or finally inhibited, by non-living catalysts
or secretions not necessarily confined to the endocrine organs.
In this way a bridge may perhaps be built between the orthodox
Weismannian and the Lamarckian. Growth and character are
caused by determinants, but these are not part of the cytoplasm
itself, they are the machinery by and through which living
matter acts. The organism is not built up by special protoplasm
or by entelechies or by any mysterious élan créatif. It arises
from the definite influence of definite catalysts originating, in an
orderly sequence, as the organs become differentiated, while the
individual is as a whole exposed in an infinite progression to
the internal and external stimuli of a like but slowly changing

254 ON PATHOLOGICAL STATES IN EVOLUTION.

environment to which it reacts. The factors which did the
work are working now.

To recapitulate the tentative conclusions arrived at, it may be
suggested that—

1. Mechanical reaction to stress is a general law of all tissues.

2. Morbid conditions in many cases give rise to repair which
sbi physiological.

. Such repairs lead to new functions, new stresses, a
mor bid states and further repair.

4, These factors are some of the main causes of specific and
generic differences.

5. In all probability transmission of changes caused in the
way indicated does take place by a morphogenetic reply in utero
to increased functional stresses.

6. As it is a narrow view to assume that pathology in all cases
tends to death, the study of pathology and general physiology
should be part of the preparation of the biologist.

NOTES ON THE BEAVERS AT LEONARDSLEE. 25d

15. Notes on the Beavers at Leonardslee, 1916-1918.
By Sir Epmunp G. Lopsr, Bart., Vice-President Z.8

(Received June 24, 1918; Read October 22, 1918. ]

From the books on Natural History we have been given to
understand that Beavers breed only once in the year, and that
the young ones are born between the end of April and the
beginning of June, after a period of gestation which is believed
to last about fourteen weeks.

In a book called ‘In the Beaver World,’ by E. A. Mills, it is
stated that the number in a litter varies from one to eight, and
that the eyes of the young ones are open from the beginning,
and in less than two weeks they appear in the water accompanied
by their mother.

It is difficult to give an opinion as. to how old the young
Beavers are when we first see them at Leonardslee. ‘They are
then about the size of rabbits, and we have supposed them to
have been born six weeks or two months before, but we feel we
have very little evidence to go on. I have always noticed the
young ones of this size swimming about alone, the mother taking
no notice of them.

In January 1916 a pair af Canadian Beavers were received here.
(I will cali these No. 1 and No. 2.)

On Dec. 11, 1916, a young beaver was seen, about the size ofa
rapoit: (1 will call this one No. 4.)

On July 10, 1917, three young ones were seen; these again
were about the same size as No. 4 when it was first seen. (I will
call these three Nos. 6, 7, and 8.)

Some time in August 1917 two young beavers were seen on
the bank together. One was considerably larger than the other.
The smaller one was recognized as No. 4, and we have to come to
the conclusion that the other must have been born in the spring
of 1916, soon after the arrival of the pair Nos. | and 2. (1 will
call this one No. 3.)

On June 15, 1918, a young beaver was seen for the first time.
Although only seen so lately, it is clear from its size that it
must have been born some months ago, perhaps in December
1917 or January 1918. (I call this one No. 5.)

At the end of 1917 we had noticed that the old female seemed
heavy in young, and were rather disappointed not to have seen
any signs of a litter, but it seems that she had one after all.

On June 18, 1918, a very small beaver was seen (I call this
one No. 9). It was not larger than a big rat. The little one
was obviously out before the authorized time, for the mother (not
the old female) went after it, and taking hold of a piece of its skin
swam back with it to the mouth of the burrow, which is under
water, and, letting go with her mouth, pushed it with her paws.

I think this young mother must be No. 3, which I suppose was

256 NOTES ON THE BEAVERS AT LEONARDSLEE.

born in May or June 1916, therefore she was only just two years
old when this little one was born.

It wiil be interesting to see how long it will take the young
one to grow to “ rabbit” size.

Now that there are two females bearing young ones, it will be
difficult or impossible to make any exact observations.

It is certain that the old female had young ones twice in one
year: in October 1916 (seen December 1916) and in May 1917
(seen July 1917), and it is probable that she had young ones on
the following dates :—

April 1916,
October 1916,
May TOL,
December 1917.

For some periods Beavers will show up continually in the day-
light, and then will come an interval when they are seldom on
view, or only one or two.

When the pair of Canadian Beavers first arrived in January
1916, I saw only one at a time for several months, until at last
I was afraid that one must have died ; but, just as I had come to
this conclusion, I saw the two together on the bank.

To get accurate statistics is not so easy as might be imagined,
but I think that the notes I have made are not far from the
truth.

MADAGASCAR FROGS OF 'rHE GENUS MANTIDACTYLUS. 257

16. On the Madagascar Frogs of the Genus A/antidactylus
Blgr. By G. A. Boutencer, F.R.S., F.Z.S.

[Received September 18, 1918: Read October 22, 1918. ]

(Published by permission of the Trustees of the British Museum.)

INDEX.

GEOGRAPHICAL: Page
Meadavascan MrOgSyisitysocs st sac tengindheaece aedoce tiene oshvleeeipyece 2OF
SysTEMATIC:
Mantidactylus, Synopsis of Species .°................cse ee 258
ESA MOUONTNICOMO Ly SP, Ws. sagndseeu sve eerste ninadetenies penpiasecre LOO
Aglyptodactylus, g.n., for Limnodytes madagascariensis

Among the many peculiar features of the herpetological fauna
of Madagascar is the fact of the genus Nana, so numerous in
species in Continental Africa and the Indo-Malayan Region,
having only two representatives: /. (Zomopterna) labrosa Cope,
allied to the South African 2. natalensis A. Smith, and &. (Pty-
chadena) mascareniensis D. & B., distributed over the greater
part of Africa, the Seychelles and the Mascarenes included.

Most of the Madagascar frogs originally referred to Rana or
Limnodytes (Lylorana) have proved to be distinguished by the
presence of an intercalated bone between the penultimate and
distal phalanges of the fingers and toes and have been referred to
an autochtonous genus, Wantidactylus*. In the species grouped
by me under this genus, the swellings or discs in which the
fingers and toes terminate bear on the lower surface a ring-shaped
groove, defining a circular or transversely elliptic area, thus afford-
ing a further distinctive character by which to recognise them
among those species of Rana in which digital discs are likewise
present.

One species, Limnodytes madagascariensis A. Dum. (2. ingui-
nalis Gthr.), which I had left in the genus Rana, has been shown
by the late Dr. F. Mocquard to be also provided with the inter-
calary phalanx and therefore referred by him to Mantidactylus ;
but as in this species the small digital terminal expansions are
devoid of the groove to which I now draw attention, I consider
it to be entitled to generic distinction, under the new name of
Aglyptodactylus. In this A. madagascariensis, the omosternum
is forked at the base, as in Mantidactylus, the nasal bones are
small, oblique, and separated from each other as well as from the
frontoparietals, and the terminal phalanges are obtuse; there are
no femoral glands.

We are now acquainted with 22 species of Mantidactylus, to
which a twenty-third is here added. A key to the identification

* Ann. & Mag. N. H. (6) xv. 1895, p. 450.

258 MR. G. A. BOULENGER ON MADAGASCAR

of the species was drawn up by Mocquard in 1909*, but as this
key does not seem to me to work well and as the arrangement
therein followed does not at all express the natural affinities, I
have prepared a synopsis in which I have endeavoured to make
good these deficiencies,.so far as it is possible to do so in a linear
sequence.

Synopsis of the Species of Mantidactylus.

I. Glandular dorso-lateral fold, if present, not confluent with the supratemporal
fold.

A. Discs of fingers very small, usually smaller than those of the toes; snout
rounded or very obtusely pointed, not or but feebly projecting beyond
the mouth; loreal region oblique; belly perfectly smooth, or very feebly
granulate behind.

1. Toes entirely or nearly entirely webbed; head broader than long; back
granulate.

Tympanum hidden or small, very indistinct, and

distant from the eye; tibio-tarsal articulation

not reaching beyond the eye; heels meeting or

not, when the hmb is folded at right angles to

the body ; tibia 23 to 3 times as long as broad,

24 to 22 times in length from snout to vent...... M. guttulatus Blgr. 1881 7.
Tympanum distinct, small and distant from the

eye; tibio-tarsal articulation reaching eye or tip

OF SNOUt . fo. .c6sccces eds sos teseed asesdeseasssece ween all, grandidiert Mocquards Iss.
Tympanum distinct, {diameter of eye; tibio-tarsal
articulation reaching posterior border of eye...... M. inaudax Peracca, 1893.

2. Toes $ to } webbed; head as long as broad; tympanum very distinct, $ to
# diameter of eye.

Series of vomerine teeth nearly equidistant from

each other and from the choanz; tibio-tarsal

articulation reaching eye; heels meeting; tibia

22 to 3 times as long as broad, 24+ to 24 times

in length from snout to vent; back smooth...... M. alutus Peracca, 1893.
Series of vomerine teeth much nearer the choane

than each other; tibio-tarsal articulation reach-

ing tip of snout, or between eyeand tip of snout ;

heels strongly overlapping; tibia 33 to 4 times

as long as broad, 12 to 2 times in length from

snout to vent; back with glandular longitudinal

FOLDS A 8 E dd nee ioc. deren sete pe tneteeaseecsson, DL betsileanusmenemleeone

B. Discs of fingers small, as large as or larger than those of the toes; belly
smooth or granulate only on the sides and behind.

1. Head as long as broad or a little broader than long; snout rounded or
obtusely pointed; toes at least 4 webbed.

a. Tibio-tarsal articulation reaching tympanum or posterior border of eye ;
heels not overlapping; tibia 23 to 3 times as long as broad, 2 to
22 times in length from snout to vent; first and second fingers equal;
loreal region oblique; back smooth or with indistinct flat warts; belly
smooth.
Tympanum 3% to 2 diameter of eye; toes # webbed. M. curtus Blgr. 1882.
Tympanum 3 to $ diameter of eye; toes 4 to 9
WEDDED. fo.iccsscu non fas ceuuudnes oot ohare: wuedeedas dette to ae OLPOTUS D loma 18S Oo:

* N. Arch. Mus. (5) 1. p. 55.
+ Includes MW. piger Mocquard, 1900.
t Includes W. multiplicatus Boettg. 1913.

FROGS OF THE GENUS MANTIDACTYLUS. 259

b. Tibio-tarsal articulation reaching eye or between eye and nostril; toes
2 to 2 webbed ; tympanum # to once diameter of eye.
’ Loreal region oblique; tympanum not more than
14 times its distance from the eye; tibia 25 to
3 times as long as broad; inner metatarsal
tubercle + to 4 length of inner toe; back with or
without small ane warts ; belly perfectly
smooth.. . M. ambohimitombi, sp. n.
Loreal region ‘oblique ; tympanum 2 to 3 times its
distance from the eye; tibia 3 to 3$ times as
long as broad; inner metatarsal faperele <4 to;
length of inner toe; back with elongate warts
or glandular folds; belly paints on the sides
and bebind . : Liseesssscssesesesseee M, ulcerosus Boettg. 1880.
Loreal region ‘nearly. vertical ; “inner metatarsal
tuberele 4 % length of inner toe; two glandular
folds along the back . Baas . M. bellyi Mocquard, 1895.
c. Tibio- ‘areal, aficbelation: réaclane’ tbe:
yond tip of snout; toes 4 webbed ;
tympanum a little ‘smaller than the
eye; three glandular folds along the
back ; belly smooth ..................... WL. opiparis Peracca, 1893.

3. Head a little longer than broad; snout pointed, strongly projecting beyond
the mouth; tibio-tarsal articulation reaching anterior border of eye or
nostril; toes not more than + webbed; belly smooth.

Tympanum larger than’ the eye; first finger
shorter than secoud; two glandular folds along

the back’ ....... M. erumnalis Peracca, 1893.
Tympanum : about + diameter of eye; first finger a as

long as or slightly shorter than second ; “back | Hew. 1913.

with large se lands” 2.2 .ss..i le cectesesd ceetesede serene DE. glandulosus 3 Meth. &

C. Discs of fingers rather large, at raat nearly twice as broad as the penultimate
joint, as lar ge as or larger than those of the toes.

1. Belly perfectly smooth; tibio-tarsal articulation reaching eye; heels meet-
ing; tibia 3 times as long as broad, 2 to 24 times in length from snout to
vent; toes entirely or nearly entirely webbed; first finger much shorter
than second ; loreal region nearly vertical.

Head longer than broad; snout pointed, strongly
projecting beyond the mouth ; eo ae 2
diameter of eye ....... ois
Head as long as broad ; snout rounded or : obtusely
pointed, moderately projecting ; a ae to
% diameter of eye .......... vecssseseeeeee ME, cowaniit Bigr. 1882.

2. Belly granulate belind. anid on the sides only; heels overlapping; tibia
34 to 4 times as long as broad; toes # to nearly entirely webbed; inner
metatarsal tubercle a tO: length of inner toe; first finger a little shorter
than second; tympanum 4 to # diameter of eye; loreal region nearly
vertical.

Tibio-tarsal articulation reaching eye, nostril, or

tip of snout; tibia 14 to 2 times in length from

snout to vent; tongue usually with a conical

papilla in the middle of the anterior third ...... M. lugubris A. Dum. 1853*.
Tibio-tarsal articulation reaching tip of snout or

pees tibia 13 times in length from snout to

(E00 IRON Reno ree ; nee M. flavicrus Blgr. 1889.
3. Belly peamulate Meals: strony OV serlappiie:

a. Tibio-tarsal articulation reaching eye or tip of snout; tibia 33 to
45 times as long as broad, 14 to 2 times in length from snout to vent ;
tympanum 3} to = diameter of eye; loreal region nearly vertical.
Toes 3 webbed; ipner metatarsal tubercle small,
feebly prominent, + length of inner toe; a narrow
dorso-lateral glandular fold ..............0.....0.0.0.. M. granulatus Boettg. 1884.

M. majori Blgy. 1896.

* Includes M. femoralis Blgr., 1882, and ambreensis Mocquard, 1895.

260 MR. G. A. BOULENGER ON MADAGASCAR

Toes 4 webbed; inner metatarsal tubercle strong
and prominent, compressed, } to } length of
inner toe; no dorso-lateral fold ..................... MM. redimitus Blgr. 1889.
6. Tibio-tarsal articulation reaching beyond tip of snout; tibia 4% to
5 times as long as broad, 15 to 13 times in length from snout to vent ;
loreal region oblique.
Toes nearly entirely webbed; inner metatarsal
tubercle 4 length of inner toe; tympanum 4% to
3 diameter of eye; a pair of inwardly curved
glandular folds on the anterior third of the back,
trom the upper eyelids; heel with a dermal pro-
CESS OF SPUY 1.0... beesecsecseccng cee stecsssss ess cenerssssace DL, Lutens Meth<&Jlewsmleies
Toes % to 3 webbed ; inner metatarsal tubercle $ to
4 length of inner toe; tympanum 3 to 2 diameter
of eye; a curved glandular fold on each side
from the upper eyelid to between the shoulders,
followed by a straight fold ....................0... ML pliciferus Blgr, 1882.
Toes 4 webbed; inner metatarsal tubercle } to 2
length of inner toe; tympanum # to # diameter
of eye; upper parts rough with prominent glan-
dular folds and tubercles; heel with a dermal
PTOCESS OF SPUY * 2222.6 .2ceece. oan anos eeoense-una: hoeeeee Ue eSDEr DIA alemae

II. Glandular dorso-lateral fold extending from
behind the eve to the hip; loreal region ver-
tical; tympanum # to once diameter of eve;
discs of fingers and toes rather large; tibio-
tarsal articulation reaching tip of snout or a
little beyond ; tibia 45 to 5 times as long as
broad, 13 to 12 times in length from snout to
vent; toes 3 webbed ; belly granulate behind. I. albofrenatus F, Mill. 1892*,

Nothing is known of the development and lJarvee of these frogs,
but thé eggs are remarkably large, measuring 5 mm. in diameter
in M. guttulatus (¢ 120 mm. from snout to vent), 3 mm. in

M. lugubris (2 50 mm.), 2° mm. in WM, betsileanus (2 33 mm.).

MANTIDACTYLUS AMBOHIMITOMBI, sp. n.

Vomerine teeth in short transverse or oblique series behind the
level of the choane, equidistant from the latter and from each
other. Head a little broader than long; snout rounded, feebly
projecting beyond the mouth, with indistinct canthus and very
oblique, concave loreal region; nostril equidistant from the eye
and from the end of the snout ; interorbital region as broad as or
a little narrower than the upper eyelid ; tympanum distinct, 2 to
3 the diameter of the eye, 1 to 13} times its distance from the
latter. Fingers moderately long, first and second equal or first a
little the longer, the discs small, not very much larger than the
well-developed subarticular tubercles. Toes moderately long,
¢ webbed, the discs about as large as those of the fingers; no
tarsal fold; inner metatarsal tubercle oval, moderately promi-
nent, ¢ to $ the length of the inner toe; no outer tubercle.
Tibio-tarsal articulation reaching the eye or between the eye and
the nostril; tibia 23 to 3 times as long as broad, 14 to 2 times in

* M. frenatus Boettg., 1913, is probably identical with this species, although the
hind limb is longer and the discs of the fingers and toes are described as very small.

2

FROGS OF THE GENUS MANTIDACTYLUS. 261

length from snout to vent, as long as or a little shorter than the
foot. Skin finely granulate above, with or without elongate flat
warts on the sides of the body; a strong, curved glandular fold
from the eye to the shoulder ; lower parts smooth; femoral gland
more or less distinct, with a single pit, or absent. Brown above,
spotted or marbled with darker, often with a large dark trian-
gular spot between the eyes; a dark canthal streak and a tem-
poral band, usually light-edged beneath ; a yellow vertebra! streak
sometimes present; limbs with more or less distinct dark cross-
bands; hinder side of thighs usually dark brown, with small
yellow spots. White beneath, uniform or mottled with brown,
or nearly entirely brown. Male without vocal sacs.

Nasal bones rather large, narrowly separated from each other
and from the frontoparietals.

From snout to vent 65 millim.

Several specimens from the Ambohimitombo Forest, Mada-
gascar, from the collection of Dr. Forsyth Major, 1896.

Proc. Zoo. Soc.—-1919, No. XIX. 19

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CILIARY ACTION IN PLEUROBRACHIA PILEUS. 263

17. Ciliary Action in the Internal Cavities of the Ctenophore
Pleurobrachia pileus Fabr. By Jamus F. Gremnitt,
MAS MD: D.Se., F238.

[ Received October 2, 1918: Read November 5, 1918. ;
(Text-figures 1 & 2.)

During life ciliation is active throughout the internal cavities
of Pleurobrachia; and the latter are wide enough to allow the
divection of the ciliary action on their different surfaces to be
made out from the motion of particles suspended in the con-
tained fluid, e.g. small oil globules, alimentary particles, and
debris.

The ‘‘ circulation” * is an extremely orderly one and meets the
physiological need for continuous regulated change through the
whole of the internal cavities.

On the whole the circulation inside the funnel system goes
on independently of that within the stomodzumt. So far as I
could make out, except under the influence of peristaltic action,
only slight interchange of fluid between the stomodeum and the

funnel takes place.

I. Stomodewm (text-figs. 1 & 2).

Round the margin of the mouth there is a very narrow band
best seen in yonng specimens, the cilia of which strike into
the mouth-cavity. Up the middle of each lateral wall of the
stomodeum, and continued on the thickenings in this region, and
to the infundibular opening, there is a track with aboralward
ciliation. Over the rest of the lateral wall of the stomodeum the
ciliation is oralwards with a slant towards the sagittal angles.
Along the sagittal angles from the opening of the funnel to that
of the mouth the ciliation is stronger and in the oralward

direction.

Il. Funnel and Canal System (text-fig. 2).

We may best follow the circulation here by beginning in the
floor (oral wall) of the funnel at points on opposits sides of the
opening from stomodeum into funnel. It will be remembered
that the aboralward currents up the middle of the sides of the
stomodzum lead to these points. Working transversely outwards

* Reference may be made to the following recent papers on ciliation :—
Carlgren,O. Biol. Centralbl. xxv. 1905, pp. 8308-322 (Actinians, Madreporarians).
Orton, J. H. Journ. Mar. Biol. Assoc. U. K. ix. 1912, pp. 144-478 (Ascidians,
Molluscs).

Orton, J. H. Ibid. x. 1913, pp. 19-49 (Amphiowus, Ascidians, Molluscs).
Gemmill, J. F. Proc. Zool. Soc. Lond. 1915, pp. 1-19 (Starfish).
Widmark, E.M.P. Zs. Allg. Phys. Jena, xv. 1913, pp. 33-48 (Aurelia aurita).

+ In this paper the whole of the cavity between mouth and funnel-opening is

called stomodgeum.
19* |

264 DR. J. F. GEMMILL ON CILIARY ACTION IN THE

Text-figure 1.
Mn.0

Diagram of lateral wall of stomodzeum of Plewrobrachia, showing direction of
ciliary currents.
M.O., mouth-opening ; F.O., funnel-opening.
(For explanation see text.)

Text-figure 2.

Diagram of internal cavities of Plewrobrachia (transverse or infundibular plane in
plane of paper). ‘The arrow-heads in the walls of the cavities point in the
direction of the ciliary currents.

1. Stomodzeum. 2. Funnel. 3. Paragastric canal. | 4. Tentacular canal.
5. Perradial canal. G. Interradial canal. 7. A sub-sagittal canal. 8. A sub-
transverse canal. 9. Adradial canal.

INTERNAL CAVITIES OF PLEUROBRACHIA PILEUS. 265

from either of the points in question we find the current strongly
outwards towards the equator, in the floor of the funnel, but soon
meet the opening into the paragastric canal. The axial wall of
this canal carries cilia which strike oralwards, while on its abaxial
wall the cilia strike in the aboral direction. The canal is thus
bathed mesially by an in-going and laterally by an out-going cur-
rent (text-fig. 2,3). Exactly the same thing holds good for the
ciliation and currents within the tentacle-canals (text-fig. 2, 4).
On the floor (oral wall) of the perradial, interradial, and adradial
canals the ciliation is outwards, 7. e., towards the entrances into
the meridional canals.

The adjacent halves of neighbouring sub-sagittal canals are
elliated along the whole length of their axial walls in the oral-
ward direction. This holds good also for the adjacent halves
of neighbouring sub-transverse canals. The areas of oralward
ciliation thus correspond with the distribution described for the
female gonads at the sides of the canals.. The axial walls of the
remaining halves of all the meridional canals (cf. distribution of
the male gonads) are ciliated in the aboralward direction.

Along the roof (aboral wall) of the advadial, interradial, and
perradial canals the ciliation is inwards, 7. ¢., towards the main
cavity of the funnel. ‘The walls of the aboral extension of the
funnel have an aboralward ciliation, while an oralward reflux
takes place down the middle of this extension into the main
cavity of the funnel. )

Here mixing occurs and currents are caught up anew into the
paragastric, tentacle, and other canals.

The arrangement above described proved constant for a con-
siderable number of large and small Plewrobrachia examined.
The tissues are transparent enough to allow the examination to
be made in undissected specimens with the help of a binocular
microscope. Fine carmine grains can be used to supplement the
evidence of the particles floating in the gastrovascular fluids.
Mixed with sea-water the powdered carmine will occasionally be
ingested into the stomodeeum by natural peristaltic action, or it
may be injected into this cavity with the help of a pipette.
After a short interval the carmine is expelled from the mouth,
but meantime in successful cases sufficient particles to be visible
have found their nan into the funnel and the cavities leading
therefrom.

The specimens were obtained last June at the Muillport
Biological Station, and the work was done partly there and
partly at Glasgow University, the cost of obtaining material
being met out a a grant from the Carnegie Scottish Univer sities
Trust. :

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CAPT. D. M. S. WATSON ON SEYMOURIA, 267

18. On Seymouria, the most primitive known reptile. By
D. M. S. Watson, M.Sc., Capt. R.A.F., Lecturer in

Vertebrate Paleontology, University College, London.
[Received September 9, 1918: Read November 19, 1918. ]
(Text-figuires 1-15.)

The reptile Seymouria bayloriensis was originally described
from two imperfect skulls, one in connection with a few vertebre
and the shoulder-girdle, by Prof. Broili of Munich, whose excel-
lent description made us well acquainted with the structure of
the upper and lateral surfaces of the greater part of the skull, and
gave general information about the palate, occiput, and anterior
axial skeleton.

In 1910 Williston described as Desmospondylus anomalus a
collection of vertebre, a humerus, femur, and some other bones,
which he subsequently recognised as belonging to a young Sey-
mouria. In 1911 the same author described a_ beautifully
complete skeleton, publishing a restoration to whose accuracy I
am glad to be able to bear testimony. At the same time he
suggested that the skull described by Cope as Conodectes favosus
was really Seymouria, a determination which is undoubtedly
correct. As Cope’s description of Conodectes is quite insufficient
for recognition of the skull, and Seymouria is a name universally
known, I propose to regard Conodectes as a nomen nudum and
relegate it to the synonymy.

In 1914, v. Huene published figures of the type skull of
Conodectes, but added nothing to our knowledge; and in 1915
I gave a short description of such knowledge of the otic region
as could be obtained from the rather badly preserved skull of
Conodectes.

In January 1914, through the kindness of Prof. Broili, [ was
able to make a careful examination and drawings of the type
material of Seymouria in Munich, which at that time, owing to
a new and more complete preparation, showed many features of
the structure of the palate and occiput which were not referred
to in the original description.

In 1915 I was so fortunate as to collect some Seymouria
material in ‘Texas. The most important of my specimens was
found weathered out on the side of a small hillock about 20 yards
away from the Cradock bone-bed quarry. I at first supposed it
to be a single individual, but have sttbsequently found that two
are represented. The better individual is represented by the
pelvis, both hind legs, fifteen presacral, the sacral and the caudal
_vertebree in a connected series, and many ribs; these bones are

connected by matrix and are all articulated. Almost certainly
belonging to this individual are the atlas and axis and three

268 CAPT, D. M. S. WATSON ON SEYMOURIA,

succeeding vertebree, with the incomplete right scapula, coracoid,
and clavicle attached by matrix, the occipital region and part of
the right side of the skull and lower jaw, the lower end of a
radius and ulna, and a metacarpus. ‘Lhe associated skeleton is
represented by a sacral and nine presacral vertebre, an incom-
plete femur and ischium.

Another specimen collected on West Coffee Creek consists of
numerous fragmentary bones of a young individual washed
perfectly clean. I also obtained certain isolated bones.

This material and the new preparation of that at Munich
allow me to add materially to our knowledge of the structure of
this most interesting form, perhaps the most perfect annectant
type known to us.

Skull.

Profs. Broili and Williston have given a satisfactory account
of the upper and lateral surfaces of the skull. The Munich
skulls and that of my skeleton give an equally complete know-
ledge of the occiput and palate. For reference I shall refer to
that Munich skull which has a shoulder-girdle belonging to it
as A, the other as B.

Basvoccipital.—This bone is very well shown in my skull. It
is a small bone, remarkably thin dorso-ventrally and of consider-
able width. The dorsal surface appears to be completely covered
by the exoccipitals and forms no part of the floor of the brain-
cavity. The posterior surface forms part of the condyle, which
is wide from side to side, shallow and rounded. The ventral
surface is widely exposed as a quadrangular area bounded in
front by the suture with the basisphenoid. The posterior part
of the lateral border is in contact with the exoccipitals, in advance
of which it is cut out into a shallow notch, the lower border of
the fenestra ovalis. Anteriorly the lateral margin is produced
downward and outward to take part in the formation of the
tuber basisphenoidalis.

Basisphenoid.—The basisphenoid is completely known, so far
as concerns its inferior surface, from the Munich skulls and that
of my skeleton.

Posteriorly it has a suture with the basioccipital, the ends of
which le on the summits of the well-marked tubera. From the
tubera a pair of prominent ridges run forward on the ventral
surface, so that this face forms a smooth concavity between them.
At the level of the basipterygoid processes these ridges die out,
so that the surface becomes gently convex. The basipterygoid
processes (shown in Munich A & B) are extremely short and end |
in a flat articular surface, which does not directly support the
pterygoid but is attached to a separate small bone very clearly
shown in Munich B. The lateral surface of the basisphenoid,
between the tuber and the basipterygoid process, 1s concave and
passes indistinguishably into that of the prootic. About three
millimetres above the lower edge of the tubera a minute foramen

THE MOST PRIMITIVE KNOWN REPTILE. 269

opens out from the basisphenoid. A broken face about 5 mm. in
advance of the tubera shows that in that region the basisphenoid
is a thin plate scarcely a millimetre thick medially.

Text-figure 1.

mf ae
Ex 0c.

Seymouria bayloriensis Broili.—Restoration of the palate, x 3.

‘

General shape and palate from the types in Munich. Details of maxillary dentition
from fragments in my possession. Basisphenoid and occipital region from
my skeleton.

Reference letters :—B.Oc., basioccipital; B.Sp., basisphenoid; Ecr.Pr., ectoptery-
goid ; Ex.Oc., exoccipital; Mx., maxilla; P.Mx., premaxilla; PAt., palatine ;
Par.Oc., paroccipital ; PR.O., prootic; Pr.V., prevomer; Qu., quadrate ;
X, autogenous basipterygoid process ?

Parasphenoid.—The parasphenoid is well shown in Munich B
as a short narrow rostrum projecting for ae in the palate
between the pterygoids.

The type of Conodectes shows that it supports a large ethmoidal
ossification which surrounds the anterior part of the brain.

Exoccipital.—The right exoccipital is well shown in my skull,

270 CAPT. D. M. 8S. WATSON ON SEYMOURIA,

in which unfortunately a piece about 2°5 mm. thick is missing
from the middle of its height, this not having been collected.

The exoccipital rests on the upper and lateral surface of the
basioccipital, meeting its fellow above that bone so as to form
the floor of the brain-cavity in the occipital region. Posteriorly
it projects behind the basioccipital and is provided with an arti-
cular face looking downwards and backwards which forms about
a quadrant of the occipital condyle. Above the body the bone is
continued up so as to form the side wall of the brain-cavity until
its posterior surface is overlapped by the occipital flange of the
dermo-supraoccipital.

On the inner surface the exoccipital is excavated into a deep
pit from which a rather large hypoglossal foramen starts to pass
through the bone and open into the large vagus foramen. In

Text-figure 2.

Seymouria bayloriensis Broilii—Restoration of the occipital view, X }.
Membrane-bones from Munich A. Quadrate from a young individual collected
by the writer. Occiput from my skeleton.

Reference letters as before with :—D.S.Oc., dermo-supraoccipital ; Fen.Ov., fenestra
ovalis; Qu.J., quadrato-jugal; Se., squamosal; Tas., tabular.

front of the ridge which - bounds the hypoglossal foramen
anteriorly, the inner surtace of the exoccipital turns outward
and forms the posterior margin of the foramen for the Xth
nerve. The upper end of the exoccipital is fused with the par-
occipital, no suture being visible. It approaches its fellow of the
opposite side over the foramen magnum, but leaves a space for
the cartilaginous supraoccipital. In my skull the space where
the supraoccipital should be is entirely free from bone, but in the
type of Conodectes there is some evidence of a slight supra-
occipital ossification.

Paroccipital.—The paroccipital is fused with the exoccipital on
the back of the skull. It is separated from the lower part of
that bone by a large notch, the foramen for the vagus nerve.

THE MOST PRIMITIVE KNOWN REPTILE. 271

Laterally to this it forms a deep plate extending downwards
nearly to the level of the ventral surface of the basioccipital,
where it ends in a suture with the prootic laterally and in a free
margin, part of the border of the fenestra ovalis, at its inner end.
The upper outer part of the posterior surface is overlapped by the
occipital flange of the tabular. Between this region and the top
of the exoccipital the upper part of the posterior surface is turned
forward so as to form a groove, the lower part of the post-temporal
fossa. The cranial end of the paroccipital appears to be narrow,
so that the inner ear is widely open to the cranial cavity in the
bony skull.

Prootic.—The prootic is fused with the lateral margin of the
basisphenoid below and has a suture with the paroccipital
behind. Its lateral face, which alone is satisfactorily shown, has

Text-figure 3.

oe DSO. Tab.

Seymouria bayloriensis Broili.—Occipital view of the type-skull
Munich A, X 3.
Reference letters as before with :—F.B.PR., facet on the pterygoid for articulation
with the “ basipterygoid process.”’

a powerful ridge running horizontally along it at about the
middle of its height. Below this it is concave, its lower border
forming part of the rim of the fenestra ovalis mesially and laterally
uniting with the paroccipital, the joint bones forming the massive
paroccipital process which in section has a V-shaped lower surface.
Above the ridge the prootic is essentially flat, having however
a deep groove leading to a foramen towards the outer end. The
anterior edge of the bone is damaged, but the minute foramen for
the facial nerve is just preserved.

The endocranial end of the bone is shown to be formed by an
extremely thin shell.

Dermo-supraoceipital and Tabuiar.—The dorsal exposure of
these bones on the upper table of the skull has been accurately
described by Broili. Munich A shows their occipital aspect
perfectly. Each dermo-supraoccipital has a small descending

272 CAPT. D. M. 8S. WATSON ON SEYMOURIA,

lappet passing vertically downward, the inner border of this
forms part of the margin of the foramen magnum, the outer, that
of the post-temporal fossa. |

The tabular has a similar occipital flange forming the outer
margin of the post-temporal fossa and covering a large area of
the posterior surface of the paroccipital.

My skull suggests the presence of some sort of a flange on the
ventral surface of the tabular articulating with the end of the
paroccipital process.

Text-figure 4.

R PAL i “9 , \N

Yue

/ hoy
’ Fo
. IgE 4
; Z
f i LE
f

—S
: : Yj
VOY i

Seymouria bayloriensis Broili.—Palate, X }. Drawn from Munich A.

Reference letters as before with :—B.Pr., the basipterygoid articulations on the
basisphenoid; R.Pau.TH., right palatine tusk.

Pterygoid.—The pterygoid is completely exposed in Munich A.

The quadrate ramus forms a vertically standing plate whose
posterior surface is concave. ‘The outer margin has a long
suture with the squamosal below the otic notch, ventrally it
separates from that bone so as to leave exposed an area of the
-quadrate, with which bone it has a large powerful suture.

The inner margin of the quadrate ramus approaches the
anterior border of the prootic closely, the bone in this region
though deep not nearly reaching the skull-roof. This part of the
ramus bears towards its ventral edge a large, well-developed,
slightly cupped articular facet.

When the perfectly preserved pterygoids and basisphenoids of
Munich A and B are compared, it is seen that there is a large

‘HE MOST PRIMITIVE KNOWN REPTILE. DA ips:

triangular space between this facet on the pterygoid and that on
the basipterygoid process with which in ordinary reptiles it
should articulate. In Munich B, this space is filled up by a
small separate bone on the right side of the skull. The fact that
although in A this bone is not preserved, the facets on the
pterygoids and basipterygoids are identical in the two specimens,
_shows beyond dispute that it is a separate bone. Nothing similar
is known in any other Tetrapod.

Text-figure 5.

B Oe.

Seymouria bayloriensis Broilii—Palate, X 3. Drawn from Munich B.

Reference letters as before with:—Paxr.Sp., parasphenoid.

In advance of the quadrate ramus the pterygoid is represented
by a large flat bone in the palate which articulates with its
fellow in the middle line in advance of the parasphenoid, and
laterally has sutures with the transverse, palatine, and prevomer.
That margin of the palatal part of the pterygoid which forms the
anterior limit of the subtemporal fossa is slightly deflected
laterally, so that with the ectopterygoid it forms a small flange
against the inner surface of the lower jaw.

Ectopterygoid.—The ectopterygoid is a bone of few features.
It forms merely a plate in the palate articulating with the
pterygoid in a suture only shown incompletely in my skull, a
long suture with the palatine and presumably another with the
maxilla. About the middle of its area in Munich A is a small
depression probably intended to receive a tooth on the lower
aw.

Palatine.—The palatine is well shown in Munich A and in my

274 CAPT. D. M. S. WATSON ON SEYMOURIA,

skull. It has a long suture with the pterygoid, is bounded
posteriorly by the transverse, and in front forms certainly the
back and apparently also a large part of the lateral margin of
the internal narial opening.

Its most interesting feature is the presence of a pit surrounded
by an upstanding ridge, lying on the palatal surface just posterior
to the nostril. This pit contains two large tusks, which replace
one another so that normally only one is functional at once.
The whole arrangement is identical with the large palatine tusks
and their pits in Labyrinthodonts. It is remarkable that this
tooth has only been described by Cope in Conodectes. It is well
shown also in Munich A and my skull.

Prevomer.—Only the posterior end of the prevomer is known.
The two separate the internal nares and articulate with the
anterior ends of the pterygoids.

Quadrate.—An incomplete but isolated and extremely well-
preserved left quadrate belongs to my young West Coffee Creek
specimen. It has a well-ossified and rounded articular margin
rather conspicuously divided into two condyles. The ‘anterior
face is concave. The outer side is entirely occupied by a sutural
surface for the quadrato-jugal and squamosal, and the inner
retains the impression of the tip of the quadrate ramus of the
pterygoid. ‘The exposed posterior surface is triangular and bears
an irregular knob.

Ductus naso-lachrymalis—My skull shows quite clearly the
presence of a naso-lachrymal duct lying within the substance of
the lachrymal and extending from the orbit to the nostril.

Septomaaxilla.— Whilst removing the matrix from the sym-
physial region of the mandible of my skull, I found a single
septomaxilla which had obviously dropped down through the
posterior naris.

This element is a thin plate of bone bent round so as to clasp
Jacobson’s. organ, and has a flat face for articulation with the
dorsal surface of the prevomer.

Lower jaw.—My skeleton retains the right ramus of the lower
jaw from the symphysis to behind the anterior end of the supra-
meckelian fossa. Both inner and outer surfaces are well exposed,
and a fortunate fracture along a horizontal plane lying just above
the upper surface of the cavity of the jaw, which passes through
the dentary and the three coronoids, places the structure beyond
doubt, as it permits a definite distinction between sutures and
cracks. The sutures are usually very visible, being filled with red
iron oxide and the bone white.

Articular.—Both articulars of the young specimen from Coffee
Creek are preserved.

The bone is very short and ends anteriorly in a flat face con-
tinued in life by the remains of Meckel’s cartilage.

The articular surface is convex from back to front and is
divided into two areas, corresponding to the condyles of ‘the
quadrate, by a low ridge running obliquely across.

The outer face has a deeply depressed sutural area for the

YHE MOST PRIMITIVE KNOWN REPTILE. 275
Text-figure 6.

U,

’ a)

Cor. “De DEN.
Sl
ey

OP Fostor PRArr Ane. ar

Seymouria bayloriensis Broilii—Right ramus of the lower jaw of my
skeleton, X 1.

A. Inner surface. The area covered with parallel oblique lines is covered with
matrix; the teeth in the dentary are restored on the evidence of fragments of
that bone of a youug individual. The articular is drawn from that of a
young individual and is probably slightly too small.

B. Ventral view of a horizontal fracture passing just above the cavity of the lower
jaw of my skeleton.

C. Vertical section seen on the broken posterior end of the jaw.

D. Vertical section seen on a fracture through the splenial region.

Reference letters :—Ane., angular; Arv., articular; Cor. I, II, Til, the 1st, 2nd,
and 8rd coronoids; Den., dentary; Post.Sp., post-splenial; Pr.Arz., pre-
articular; Paw.TH., the palatine tusks, represented in dotted lines to show
their position with the mouth closed; Sp., splenial; Sur.ANG., surangular,

276 CAPT, D. M. S. WATSON ON’SEYMOURIA, ©

surangular; the inner, more faintly marked depressions for the
prearticular and angular. The posterior surface bears a minute
knob representing the postarticular process.

Dentary.—In general the sutures on the outer surface of the
jaw are not well shown. ‘The dentary, however, forms the entire
outer surface for some distance behind the symphysis, posteriorly
it ends in a point which is received in a groove in the surangular.
Fragments belonging to the young individual show that it carried
a single series of rather large round teeth. Its admesial surface
has a long perfectly straight suture with the coronoids.

Splenial.—The splenial appears to enter the symphysis. It
forms the lower border of the anterior portion of the jaw, its
suture with the dentary running along just above the lower edge
on the outer surface. The greater part of the bone, however,
forms a flat inner surface articulating above with the first and
second coronoids, and behind with the prearticular and pre-
angular.

Preangular.—The preangular is a channel-shaped bone forming
the lower part of the jaw, articulating above with the dentary
on the outer surface and the prearticular on the inner, with the
splenial in front and the angular behind.

Angular.—The angular is only very incompletely preserved,
but is a bone of the ordinary reptilian pattern.

Prearticular.—The prearticular is a very large bone running
from the articular to the splenial. Its lower edge has long
sutures with the angular and preangular, interrupted so as to
leave an anterior internal mandibular vacuity between the pre-
articular and preangular and a posterior vacuity bounded by
prearticular and angular.

Coronoids.—The fracture referred to above shows that there
are three coronoids forming together a continuous strip wedged
in between the dentary and surangular on the outside and the
prearticular and splenial within.

The lst or anterior bone is small and incompletely exposed, it
is not shown to bear any teeth.

The 2nd is of considerable size, and in the middle of its length,
which in life lies immediately outside the palatine tusks, is
thickened and carries a tightly packed mass of small granular
teeth.

The 3rd, posterior, coronoid forms the anterior border of the
supra- -meckelian vacuity, extending back along the sides of that
opening in contact with the inner surface of the surangular
outside and the upper edge of the prearticular below. This 3rd
coronoid bears one large bluntly pointed tusk at about the middle
of its length: this tooth i is oval in’ cross-section and was appa- —
rently received in a special pit in the ectopterygoid.

Vavenel Column.

Atlas.—The first vertebra preserved in my skeleton is the
atlas. ‘The parts preserved are slightly displaced,

THE MOST PRIMITIVE KNOWN REPTILE, Duan

There is a large hemicylindrical intercentrum whose width
agrees with that of the basioccipital part of the condyle. The
lower surface is smooth, short blunt-ended backwardly directed
processes forming its lateral margin. The upper surface of this
bone has laterally two large flat triangular areas for articulation
with the neural elements; between these areas the anterior end
of the bone is excavated into a conical pit, only half of which is
present, but which resembles exactly that in the end of an
ordinary centrum.

Text-figure 7.

Seymouria bayloriensis Broili.—Three anterior vertebrae, X 2.
Right lateral aspect. Left surface.
The atlas is composed of drawings of the elements in my skeleton replaced in what
appears to be their natural relations, the centrum of the axis and the inter-
centrum before it are hypothetical.
C2, C3, centra of 2nd & 3rd vertebree; In.C.1)2,3, intercentra 1, 2, & 3;
Op., odontoid ; N! & N2, neural arches of 1st & 2nd vertebree.

One pair of other elements is preserved. Each of these has
a high laterally compressed but antero-posteriorly elongated
neural spine, whose posterior margin descends directly to the
hinder end of a well-formed posterior zygapophysis. The
anterior margin of the spine descends vertically about to the
level of this articular face, and then turns forward so as to
lie horizontally and form the upper edge of a squarish area of
bone whose front edge is rounded and articular and of consider-
able width. The lower part of this region ends in a short blunt
point. It is obvious that the neural spines of the pair of elements
lay in contact with one another and that the zyg sapophyses were
supported by those of the succeeding vertebra. The anterior end
could have articulated with the exoccipital part of the condyle,
the intercentrum supporting the basioccipital part.

Proc. Zoou. Soc.—1918, No. XX. 20

278 CAPT. D. M. S. WATSON ON SEYMOURIA,

The odontoid is a single bone. Its upper surface is slightly
rounded but presents no trace of any articulation. ‘The posterior
surface is not exposed. The anterior surface is slightly convex,
rising to a small point. This surface is in general triangular
with the point at the mid-ventral surface. The lateral edges are
slightly emarginate at about the middle of their height so as to
divide the whole area into three; of these, the lowest articulates
with the anterior intercentrum, and the two dorsal with the
neural arches. The lateral surface is nearly flat, but rises some-
what towards the ends; it is triangular, so that there must be a
gap fora triangular intercentrum between the odontoid and the
axial centrum,

Text-figure 8.

bo

Seymouria bayloriensis Broili. X

A. Odontoid from in front.

B. Atlantal intercentrum from in front.

C. Atlantal intercentrum from below, anterior end upward.
D. Third intercentrum and centrum from below.

EK. Fourth centrum and fifth intercentrum.

Axis.—The second vertebra preserved differs in no important
respect from the third and fourth. It has large well-formed and
somewhat swollen zygapophyses, which are not very much pro-
duced laterally, the lateral part of the neural arch immediately
below the articular surface being extended outward into a short

THE MOST PRIMITIVE KNOWN REPTILE. 279

but well-formed transverse process bearing a rib-facet distally.
The neural spine of this vertebra is destroyed.

The restoration (text-fig. 7) is only uncertain in the shape
and size of the intercentrum between the odontoid and axial
centrum and of that element itself. The sutures between the
neural arch and centrum of the axis are obliterated.

The 8rd and 4th vertebre exactly resemble the 2nd, except that
the arches are wider and the transverse processes longer. The
neural spine is represented only by a little ridge.

The 5th vertebra exactly resembles that in front of it except
that it has a low but very massive neural spine.

The centra of the 3rd, 4th, and 5th vertebre are short antero-
posteriorly and have a very well-marked keel, their sides being
excavated into deep concavities on each side of the middle line.
The intercentrum between the 4th and 5th vertebre is well
displayed, it is nearly as long as the centrum before it, and
laterally is carried out into well-marked processes for articulation
with the capitula of the ribs.

The next vertebra completely preserved is the 9th, from which
the series is complete and naturally articulated to the 10th
caudal.

The 9th is a typical Seymouria vertebra with a small well-
rounded centrum, considerably longer than that of the 5th. The
arch is enormously wide and massive, but the ends of the com-
paratively slender transverse processes project well beyond the
extremities of the zygapophyses. The 9th vertebra has a low but
very massive spine whose summit is bifid.

From this vertebra to the sacrum the structure of the vertebre
remains very uniform, the only changes being that the transverse
process gradually shortens, the spine becomes lower and more
slender, and the ventral surface of the centrum becomes flattened.

The 10th vertebra is unique in that it is entirely devoid of the
slightest trace of aspine. I myself removed the matrix which
covered it, and the surface is still practically perfect.

My skeleton is abnormal in that the 23rd vertebra is asym-
metrical, on the left side agreeing exactly with that in front,
whilst the right side of the neural arch is much elongated and
carries a sacral rib.

The 24th vertebra, the sacral, is only exposed on the left side,
where it carries a very massive sacral rib, applied to the inner
aspect of the ilium in the usual way. This vertebra has a rela-
tively high and massive spine.

As Prof. Williston has already remarked, the first caudal differs
markedly from all that precede it in its narrow and unexpanded
neural arch. It has a fairly long transverse process. From here
to the sixth the caudals do not differ much, except that the spine
gets progressively more and more slender and leans more steeply
backward.

The seventh and succeeding caudals bear no ribs and the
transverse process has disappeared from them; their centra are

20*

280 CAPT. D. M. 8. WATSON ON SEYMOURIA,

small and hourglass-shaped and separated from one another by
an interval of more than their own length. The neural arch
rests on the anterior part of the centrum and presents a large
articular face to the space between the centra; the lower part of
this space is in part filled up by the top of the chevron, but there
can be no doubt that the condition in life was essentially embolo-
merous, the cartilaginous intercentrum not having ossified.

Text-figure 9.

HY

Seymouria bayloriensis Broili. Ribs, X 1.

The numbers are those which the bones drawn occupy in the series.
Cav. IV. & V. are caudal ribs.

Ribs.—The general distribution and structure of the ribs have
been well described by Williston, to whose account, however, I
ain able to add some details of interest.

The long slender atlantal rib is double-headed, the capitulum
and tuberculum being carried at the ends of two widely separated
branches.

The upper part of the 5th rib, which has a very expanded
distal end, is extraordinarily deep and massive. The two heads
ave widely separated, the capitulum articulating with a special
process on the intercentrum.

From here backwards the ribs are more slender, and in the
middle of the back the tuberculum is merely a facet on the upper
edge of the rib, and not carried out as a special process.

The sacral ribs cannot be described in detail. The caudal ribs
are remarkably long and massive, they are well curved, and if
found separately would have been regarded as dorsals,

THE MOST PRIMITIVE KNOWN REPTILE. 281

The ribs on the 4th and 5th caudals are very distinctly double-
headed, having a distinct emargination between the well-marked
tuberculum and capitulum. This feature is, I believe, known in
no other vertebrate.

Text-figure 10.

Seymouria bayloriensis Broili.—Outer surface of the incomplete right scapula
and precoracoid, X §.

The fragment lying on the anterior end of the coracoid is clavicle.
te)

Reference letters ;—CL., clavicle; F.Gu., glenoid foramen; F.PR. Cor. ., precoracoid
foramen; F.S.GL., supra-glenoid foramen; Pr.Cor., precoracoid; Sc., scapula.

Shoulder-girdle.-—The shoulder-girdle has already been well
described by Broili and Williston, but some features of its
structure have so far escaped observation and others are worthy

further emphasis. The scapula is exactly as Williston has
described it, with a very broad supra-glenoid fossa, high up in
which les the small supra-glenoid foramen. My specimen shows
clearly not only that it only articulates with a single coracoidal
element, but also that it differs from the somewhat similar con-
dition in Varanoops in not presenting an articular face for a
cartilaginous posterior coracoidal element. In fact the evidence
makes it quite plain that there was only a single coracoid,
corresponding to the anterior one of most Cotylosaurs, the
posterior not being represented even by a cartilage. The pre-
coracoid is as Williston has described it, with a very deep fossa
on its outer surface behind the front of the glenoid cavity. A
small glenoid and a much larger precoracoid foramen open into
the fossa.

The glenoid cavity is represented by a deep groove, V-shaped
in section. This groove during life must have been filled up
with cartilage, its margins map out a typical screw-shaped glenoid
articulation.

Sternwm.—The hard matrix lying on the inner surface of the

282 CAPY. D. M. 8S. WATSON ON SEYMOURIA,

scapula and coracoid shows quite distinctly traces of an ossification
or ossifications over an area about 2°5 cm. by 1-5 cm. This bone
is of very open texture and has no definite surface, it was exposed
so irregularly in development and its matrix is so extremely hard
that no account of its shape can be given. It is certain that it
does not represent a series of abdominal ribs or other dermal
ossifications. It does not show any tendency to form long
columns such as sternal ribs would be, and must apparently re-
present a sternum, the first evidence of an ossification in this
element in a Lower Permian reptile.

Text-figure 11.

Seymouria bayloriensis Broilii—Right side of the pelvis, X 3.
From my skeleton.

Pelwis.---The perfect pelvis of my skeleton agrees very well
with Prof. Williston’s figures and description, but the very
well exposed ilia show certain significant differences from the
young bone figured by Williston (‘American Permian Vertebrates,’
pl. xxix. fig. 7). In them that part of the bone which lies
behind the acetabulum and articulates with the ischium is quite
large, and a more interesting difference is that the upper margin
of the bone has a distinct projection towards its anterior end.
This feature is shown by both right and left ilia.

Femur.—The two well-preserved femora of my skeleton agree
generally with Williston’s figures, but appear to be considerably
less massive. ‘The lower end is very markedly divided into
condyles. ;

Tarsus.—Both hind limbs are in position with the femur
directed forward with its head in the acetabulum. The knees
are very strongly flexed. The right foot lies naturally articu-
lated, but has been slightly laterally compressed, so as to slide
the head of the third over that of the fourth metatarsal. Only
four tarsals are preserved, and it appears extremely unlikely that
any more were ossified at the death of the individual. The
tarsus and foot are exposed from the plantar surface.

There are three proximal tarsals all closely articulated with
one another and with the tibia, but separated by an interval
of about 8 mm. from the fibula.

THE MOST PRIMITIVE KNOWN REPTILE. 283

The #ibiale is a small squarish bone with rounded edges and
corners. It has a somewhat definite edge towards the inter-
medium. F

The intermedium is a well-formed bone with a cylindrically
concave ventral surface. Towards the tibia and fibula it has
well-formed edges, which are separated proximally by a smooth
well-rounded notch of considerable breadth. It articulates with
both tibiale and fibulare, the faces towards those bones being
separated distally by a small smooth notch.

Text-figure 12.

Seymouria bayloriensis Broilii—Right hind leg of my skeleton, x 3.

The bones are represented as they lie in the matrix. The dotted part of the fibula is
exposed on the lower surface of the block and is represented as seen through
it, and its distal end beyond the line which crosses it is drawn reversed from
the bone of the left side.

- The fibulare is a small rounded bone resembling the tibiale in
general characters. It is separated from the intermedium by a
definite layer of matrix.

The only other tarsal preserved is a small rounded element
above the upper end of the 3rd metatarsal.

In text-fig. 12 I have represented the right femur, tibia, fibula,
and proximal tarsals in the position in which they he in the
matrix. The part of the fibula indicated in dotted lines 1s
exposed on the other side of the block, and the distal strip of the
fibula about 1 mm. wide is restored from the bone of the left
side, as the right bone is slightly weathered here.

In text-fig. 13 I have restored the tarsus by moving up the
fibula into contact with the proximal tarsals, the resulting
accuracy of fit is good evidence for the reliability of the figure.

This leg differs from Prof. Williston’s figure somewhat in the
shape of the lower end of.the fibula, and much more importantly
in showing the clearest evidence of an intermedium. There is no

284 CAPT. D. M. S. WATSON ON SEYMOURIA,

doubt that the distal notch in that bone formed part of the fora-
men which occurs in all Permian reptiles between the ‘“tibiale”
and fibulare. |

Text-figure 13.

Seymouria bayloriensis Broili.

Right tibia, fibula, tarsus, and pes, reconstructed from my skeleton,
the cross-hatched phalanges are not preserved.

The left foot retains a complete 2nd toe and a third which has
obviously been complete but from which part of the first and
the whole of the second phalange are missing, the piece of
matrix containing them not having been collected. These toes
have respectively 3 and 4 phalanges, the terminal one being in
each case a little conical bone.

The foregoing description when read in connection with those
given by previous authors renders Seymowria nearly as well
known as any reptile recent or fossil, and should allow of a very
thorough study of its taxonomic relationships and of the bearing
of its structure on morphological problems. Broili in his original
discussion recognised the extraordinary Stegocephalian appear- —
ance of its skull. Williston, from a study of the whole skeleton,
concluded ‘That the relationships of Seymouria are not very
intimate with any other reptiles known from the Permian ; at

285

THE MOST PRIMITIVE KNOWN REPTILE.

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286 CAPT. D. M. S. WATSON ON SEYMOURIA,

least any that are tolerably well known. In the skull the
presence of the deep otic notch, the arrangement and number of
the temporal bones, the slender, elongate teeth, longest in the
maxille, and the shape, all differentiate the genus widely from
elther Diadectes or Limnoscelis, as well as the chief forms now
referred to the Pariotichide, and also from the known foreign
forms.” “In*any event, it is certainly very remarkable that
this cotylosaur reptile with all its other strange amphibian
affinities should mimic so closely the temporal structure of the
veal amphibians.” ‘ However, it is very much of a question
whether these resemblances [to Amphibia in skull and limbs]
are so much the result of heredity and relationships as of
adaptive, parallel, or convergent evolution. We have been
speculating on the assumption that the known temnospondylous
amphibians hke Cacops, Hryops, Huchirosaurus, are, if not the
actual ancestors of the reptiles, their first or second cousins.
But this presumption is, in my opinion, quite unjustified.”

In several papers during the last five years, I have upheld the
view that Seymouria is the most primitive of all known reptiles,
and that its resemblances to Temnospondyls, particularly to the
Embolomeri, are due to inheritance. I now propose in the hight
of a practically complete knowledge of the skeleton in the
Cotylosaurians Diadectes, Labidosaurus, Captorhinus, Limmnoscelis,
Pariasaurus, and Procolophon, and in the Temnospondylous
Amphibia Eryops, Cacops, Trematops, Trimerorachis, Archego-
saurus, Lydekkerina, and Rhinesuchus, and a good knowledge of
the skull and much of the skeleton in many Embolomeri, to
discuss the position of Seymowria in detail.

Labidosaurus and Captorhinus belong to the same Foals:
each of the other forms represents an independent family; Dia-
dectes, Paraisaurus, and Procotophon represent the superfamily
Diadectomorpha. The other Cotylosaurs are Captorhinomorphs
(Watson, 1917).

The skull and lower jaw of Seymouria as a whole resemble those
of Labyrinthodonts in their reticulate ornamentation. Similar
sculpture is retained by the Captorhinomorpha alone amungst
Cotylosaurs, that of other types being less markedly composed of
pits, sometimes elongated to form long twisted channels.

Basis Cranii.—The basioccipital of Seymouria agrees with those
of all other Lower Permian Cotylosaurs, rachitomous Amphibia,
and certain Embolomeri, in being excluded from the brain-cavity
by the meeting above it of special flanges from the exoccipitals.
Certain Embolomeri (Pteroplax,) have the basioccipital entering
into the border of the foramen magnum, a condition which
appears to be the primitive one for Tetrapods. Seymouria 1s
unique amongst Cotylosaurs in the very large part which the
exoccipitals play in the condyle. They form well-marked, down-
wardly-projecting areas strongly reminiscent of those of Era "YOPS,
and not exactly paralleled in any other reptile.

THE MOST PRIMITIVE KNOWN REPTILE. 287

The large ventral exposure of the basioccipital is a reptilian
character not achieved in Amphibia.

Basisphenoid.—The presence of large powerful tubera basi-

sphenoidales in the basis cranii of Seymouria is a reptilian
character. Definite tubera do occur in certain ’emnospondyls
(Eryops), but these are always small. In Embolomeri tubera are
absent.
. The short basipterygoid processes of Seymouria supporting the
pterygoid through the intervention of a special bone are unique,
nothing similar occurring in any other known adult reptile or
amp! hibian. The shape and position of the pterygoid render it
certain that these special bones cannot be the epipterygoids,
which in Dimetrodon are known to articulate with the basi-
sphenoid. Swinnerton and Howes showed that in the develop-
ment of the skull of Sphenodon special articular cartilages are
developed between the basipterygoid processes and the pterygoid,
and it is not impossible that these are the representatives of the
articular bones in Seymouria.

On the other hand, Gaupp has shown that in Lacerta the
basipterygoid processes contain independent centres of ossification,
and it is feasible and attractive to regard the Seymouria bones as
permanently separate autogenous basipterygoid processes.

The way in which the sides of the lower surface of the basi-
sphenoid of Seymowria pass on directly into the outer surface of
the prootic is reptilian, agreeing exactly with the conditions in
most Cotylosaurs. The structure of this region is, however,
equally similar to that of Pteroplaw and another Middle Coal-
Measure Embolomerous amphibian; but it is quite different
from the conditions in Rachitomi.

The parasphenoid of Seymouria is of normal reptilian type,
and also agrees with that of Embolomeri. It differs in its small
size from that of Rachitom1.

Brain-case and Otic region.—The brain-case and otic region of
Seymouria present a wonderful mixture of features occurring in
other Cotylosaurs and in Labyrinthodontia.

Seymouria 1s unique amongst reptiles in not. possessing an
ossified supraoccipital. It also differs in this respect from the
Embolomeri, but agrees exactly with the conditions in most
Rachitom1.

The whole structure of the exoccipital, its relations to the
basioccipital and paroccipital, and to the skull-roof especially in
its connection with a special descending lappet from the dermo-
supraoccipital, is that of such an amphibian as Zrimerorachis,
which resembles Seymouria even in the unusual position of the
hypoglossal foramen.

The Rachitomous amphibian in which the brain-case is most
completely known is Eryops, where the structure has been
described by v. Huene, Broom, the present writer, and Williston.

Although Prof. Williston eriticises certain points of my

288 CAPT. D. M. 8S. WATSON ON SEYMOURITA,

description, the discrepancies between his account and mine seem
to be almost entirely, and in all important points are entirely,
differences of interpretation and not of structure. The only sig-
nificant difference is that the foramen I hold to be for the Xth
nerve Williston takes to be that for the XIIth. It is undeniable
that it does greatly resemble a hypoglossal foramen, but in the
several specimens in the American Museum which show both its
outer opening and the suture between the exoccipital and par-
occipital, there is no doubt that it opens between these two bones
as a vagus foramen always does. ‘The course of the Xth nerve
demanded by Williston’s interpretation leads through the large
cavity, which is definitely shown by an American Museum skull,
which I have described and figured, to have housed the vestibule
of the inner ear, and passes out high up in the lateral walls of
the skull through a foramen which is certainly absent in certain
well-preserved New York material, and therefore cannot be for
any cranial nerve. I thus still prefer my own interpretation
of the structure, although for theoretical reasons I should wish
to see a XIIth nerve in Hryops as in so many other Labyrin-
thodonts.

The brain-case and otic region of Seymouria resemble the
corresponding regions of a Rachitomous amphibian in the follow-
ing features :—

1. The large opening from the brain-cavity to that for the
inner ear.

2. The position of the inner ear in the side-wall of the skull.

3. The way in which the inner anterior corner of the prootic
reaches up to the skull-roof.

4, ‘The extremely massive paroccipital process.

5. The upward direction of the paroccipital process.

6. The presence of an occipital flange from the tabular cover-
ing a large area of the back of the paroccipital.

The only difference of importance between the ear-region of
Seymouria and Rachitomous amphibia is that in the reptile the
fenestra ovalis lies very low down, being bounded below by the
basisphenoid and basioccipital along the posterior margin of the
tubera basisphenoidalis, whilst in the Amphibia the lower border
of the fenestra is formed by the parasphenoid, the basioccipital
at any rate never entering into its margin.

-In characters 4, 5, 6, Seymouria differs from and is obviously
more primitive than all other reptiles, for it is easy to derive
the very diverse conditions in other Cotylosaurs from those in it.

The condition of the fenestra ovalis in Seymouria is paralleled
amongst Cotylosauria only by the Captorhinide, whieh in this
region present an almost identical structure.

The conditions in such types as Diadectes and Pariasaurus
seem to depend on a reduction in size of the stapes and a corres-:
ponding diminution of the fenestra which receives its proximal

THE MOST PRIMITIVE KNOWN REPTILE. 289

end. The reduction seems to have taken place at the ventral
end of the opening. The Captorhinide retain a very large
stapes. :

Sphenethmoid.—The anterior end of the brain of Seymouria is
surrounded by a badly known bone, which agrees exactly in its
main features with the sphenethmoid of “ryops, and with the
same bone in Pariasaurus and similar bones in other Cotylosaurs.

Palate.—The paiate of Seymouria is in essentials identical with
that of the Embolomerous Labyrinthodonts.

The resemblances depend on the identity in shape of the large
pterygoid, meeting its fellow in the middle line anteriorly,
laterally articulating with the prevomer, palatine, and ectoptery-
goid, and having the quadrate ramus formed by a vertically
standing plate reaching nearly up to the skull-roof, and bending
round behind the quadrate to meet the squamosal in a long
suture.

This bone is covered, apparently all over, by a shagreen of
granular teeth in Sea ymouria and Pteroplax.

The palatine of Seymouria agrees exactly with that of 'Temno-
spondyli in structure, the most striking similarity lying in the
tusk. This tooth is unique amongst reptiles in its mode of
insertion in a pit which also encloses the replacing tooth, but
agrees exactly with all the large palatal teeth of Stegocephalia.

The ectopterygoid of Seymouria agrees exactly with that of
Eryops in contributing to a rudimentary flange which is applied
to the lower jaw, and consists essentially of a deflected corner
formed by pterygoid and transverse. The only feature of the
palate of Seymouria which cannot be matched in 'Temnospondyli
is the approximation of the posterior nares to the middle line.

The palate of Seymouria differs from those of all other
reptiles in the presence of the palatine tusks and the very weak
development of the flanges which face the lower jaw.

The Captorhinid palate more nearly resembles that of Seymouria
than those of other Cotylosaurs, but differs in not having the
quadrate ramus of the pterygoid reaching the squamosal, in the
stronger transverse processes, and the loss of the palatine tusk.
The maxillary teeth of Seymouria exactly resemble those of
Temnospondyls in being fused to their base and to a labial wall
of bone, in having fluted roots, and in being replaced alternately.
No other Cotylosaur has at all a similar dentition.

Prof. Broili has already shown that in its general build, in the
deep otic notch, and the backwardly-inclined quadrate, and in the
presence of the intertemporal, the skull of Seymouria resembles
that of the Temnospondyls. In all these features it differs from
all other reptiles.

The elongated lachrymal reaching from the orbit to the nostril
is, however, a reptilian character istic known in scarcely any
Temnospondyl.

The septomaxilla of Seymourta agrees with that of Temno-

290 CAPT. D. M. 8S. WATSON ON SEYMOURIA,

spondyli in lying within the nostril and not appearing in the
face. The condition of this bone is known in few other Coty-
losaurs.

Lower gaw.—The mandible of Seymouria is sy pies Laby-
rinthodont in structure, agreeing very closely with that of
Trimerorachis, in possessing a post-splenial and three coronoids.

The presence of a patch of small granular teeth on the second
coronoid is also a point of resemblance to most Rachitomi.

The single large tusk on the 3rd coronoid is unique amongst
Tetrapods, being unparalleled in either Amphibia or Reptilia, but
of constant occurrence in Osteolepid fish.

The non-fusion of the articular with the surangular is a point
in which Seymouria differs from the Temnospondyls and agrees
with the majority of reptiles.

Vertebral column.—The atlas of Seymouria is unique im the
lateral compression and great antero-posterior length of its
neural elements. This feature comes out most clearly in the
large spine composed of two apposed halves. The pair of
neural elements together much more resemble a normal dorsal
neural arch than in any other known Tetrapod, and are un-
doubtedly extraordinarily primitive. The odontoid is ann
reptilian.

The axis of Seymouria in its complete unspecialisation, resem-
bling as it does the vertebra next behind it, is quite different
from that of any other reptile, whilst it agrees with that of
Temnospondyls, where the 2nd and 3rd vertebrze have a similar
resemblance to one another.

Prof. Williston in a recent paper has given an account of the
mode of evolution of a Seymouria vertebra from an Embolo-
merous type, with which I am in perfect agreement. He shows
that the Seymouria vertebra is more primitive than that of any
other known reptile in having a larger intercentrum, only
slightly reduced from the disk which represents that bone in
Embolomert.

The presence of distinct processes for the head of the rib in
certain Seymouria intercentra is a condition known in many
Rachitomi and occurring in no other reptile.

The presence of a single sacral vertebra only is an Amphibian
feature very rare in reptiles. The wide separation of the centra
of the caudal vertebre suggesting the presence of a ring-shaped
intercentrum, is a very primitive feature.

On the other hand, the massive neural arches of the pre-sacrals
with horizontally placed and widely separate zygapophyses are
typically Cotylosaurian, and occur in no Labyrinthodont what-
soever. The caudal neural arches are also of typical reptilian
type.

Ribs.—In possessing double-headed ribs throughout the whole
pre-sacral part of the vertebral column, Seymouria differs from
all other known Cotylosaurs and agrees with the Embolomeri.
There can be no doubt that two- headed ribs are primitive, and

THE MOST PRIMITIVE KNOWN REPTILE. 291

that they are preserved on the atlas and axis in all early reptiles
together with a permanent primitive temnospondylous structure.

The double-headed caudal ribs of Seymouria are unique
amongst Tetrapods.

Shoulder-girdle.—I have endeavoured to show that primitively
the Tetrapods had only a single cartilage element on each side of
the shoulder-girdle, that a precoracoid was then added to this,
and subsequently a coracoid appeared. If this view be justified,
Seymouria, which has only a precoracoid and presents no trace
even of a cartilaginous coracoid, is primitive, more so indeed than
any other known lower Permian reptile.

The preservation of all three foramina of the Rachitomous
shoulder-girdle is a primitive feature known in few other
reptiles.

Pelvis.—The ilium of Seymouria differs in the antero-posterior
elongation of its dorsal end from that of any Rachitomous
Labyrinthodont, but in this character it exactly resembles the
four known Embolomerous ilia. It agrees with these also in the
process from the upper edge toward the anterior end of that
border. Several cotylosaur ilia resemble that of Seymouria in
the production of the caudal end of the upper part of the bone.

Fore limb.—Williston has already called attention to the
remarkably Temnospondyl appearance of the humerus, but he
has also pointed out that this bone différs from all amphibians,
and resembles all early reptilian humeri in the presence of an
entepicondylar foramen.

Hind limb.— Williston has shown how much the Seymowria
femur resembles that of such a Rachitomous amphibian as
Er YOPs, and. I have nothing to add to his account.

It is, however, of great interest to note that it also presents a
still more striking resemblance to the Lower Carboniferous
(Lower Mississippian) femur which I recognised as probably
reptilian and called Papposaurus traquart. This bone is very
nearly as old as the oldest known amphibian bone, and far older
than any other known reptile.

The tarsus as revealed by my skeleton is thoroughly amphibian
in retaining a separate intermedium. In all other reptiles
except Limnoscelis, where the fibulare is unossified, and the
Plesiosaurs and Ichthyosaurs, there are only two proximal
tarsals, the fibulare and a compound bone formed of the fused
intermedium and tibiale.

The phalangeal formula is apparently that common to all early
reptiles.

The foregoing series of comparisons show how wonderful a
primitive and annectant form Seymouria is. In every part of its
skeleton it shows a mixture of Temnospondyl and Reptilian
characters, each recognisable, and in general showing’ little
evidence of an intermediate condition, The whole effect of its

292 CAPT. D. M. 8. WATSON ON SEYMOURIA,

structure is that of a mosaic of separate details, some completely
amphibian, some completely reptilian, and very few, if any,
showing a passage leading from one to the other. In this fea-
ture our study of Seymouwria lends support to the belief first
upheld by the Mendelians and now accepted and used by many
paleontologists, that an individual animal consists of an aggre-
gate of separate characters which are to a large extent capable of
separate evolution, and may indeed proceed with their evolu-
tionary change at very different relative rates in different stocks,
although of course they react on one another so as to produce a
workable result.

It thus appears desirable to have a tabular statement of the
characters in which Seymouria shows an advance on Temno-
spondyl structure, using so far as possible that of the Embolomeri
as standard, and pointing out where these advances are in
different directions from those which occur in the Rachitomous
Labyrinthodonts, the cousins of the Reptilia.

List of characters in which Seymouria has advanced
above the Embolomeri :—

1. The basioecipital is depressed and no longer presents a
round concave condyle. Rachitomi parallel.

There are tubera basisphenoidales. Rachitomi parallel.

The exoccipitals exclude the basioccipital from the brain-

cavity. Some Embolomeri and Rachitomi parallel.
. A hypoglossal foramen is present. ¢If really absent in
Kmbolomeri, Rachitomi parallel. |

5. The exoccipitals reach up to be overlapped by the occi-
pital flanges of the dermo-supraoccipitals. Rachitomi
parallel.

6. The supraoccipital is not ossified. Some Rachitomi
parallel.

7. The inner ear, though widely open to the brain-cavity, is
distinctly separated by arim of bone. Some Rachitomi
parallel.

8. The fenestra ovalis is large and placed low down so that
its lower margin is formed by that of the basipterygoid
process. Rachitomi not parallel.

9. The paroccipital is partly covered behind by an occipital
flange from the tabular. Rachitomi parallel.

10. The lachrymal extends from the orbit to the nostril.
Rachitomi not parallel.

11. The ductus naso-lachrymalis lies in the substance of the
lachrymal. Rachitomi parallel (M/cropholis).

12. The choane lie near to the middle line. Rachitomi not
parallel.

13. The articulating surfaces of the zygapophyses are hori-
zontally placed and the neural arches wide and swollen.

Rachitomi not parallel.

Gaal
ee eo

THE MOST PRIMITIVE KNOWN REPTILE. 293

14. Intercentra with reduced. and unossified dorsal halves
oceur. Rachitomi parallel.

15. A distinct sacral vertebra is present. Rachitomi parallel.

16. A cleithrum is absent. Rachitomi not parallel,

17. An additional bone occurs on each side of the cartila-
ginous shoulder-girdle. Some Rachitomi parallel.

18. An entepicondylar foramen is present in the humerus.
Rachitomi not parallel.

19, The presacral part of the vertebral column is shortened.

20. An articular not fused with the surangular occurs.
Rachitomi not parallel.

21. The odontoid is a single bone. Rachitomi not parallel.

The only characters which show that Seymouria is a reptile
are: 8, 10, 12, 138, 16, 18, 20, 21, of the foregoing list.

It is specially to be noted that some of these [10, 18, 20]
are features apparently of very minor interest, and that the
really important characters, in the absence of knowledge of which
no one would be justified in recognising the reptilian nature of
Seymouria, are those of the vertebral column. Nothing like the
large swollen neural arches with horizontally placed articulations
is known in Temnospondyls, and the small notochordal centra
when taken in conjunction with the crescentic intercentra are
equally distinctive.

It thus appears that the vertebral column affords the best
evidence for the reptilian nature of any Paleozoic tetrapod.

It is of interest to compare Seymouria in more detail with
other Cotylosaurs and putative Cotylosaurs.

Sauravus.—The small, rather imperfectly-known animal from
the upper part of the Stephanian of France described by
Thevenin as Sauravus costei was regarded by its describer as a
reptile, and referred by Case to the Cotylosauria. This attribu-
tion has not yet been challenged; it appears to have been
founded on the presence in the hind foot of two proximal tarsals
and a digital formula 2, 3, 4, ? %

The structure of the vertebral column shows beyond all doubt
that this form is a Lepospondylous amphibian. There are no
intercentra in any part of the column, and in the caudal region
the hemal spines, which in all reptiles are supported by inter-
centra (which in Mososaurs may fuse with the centra), project
from the lower surface of the whole length of the “ centrum.”
They have peculiar fluted ends, exactly similar to Ceraterpeton
and other types forming Miall’s family Nectridia. The dorsal
‘“centra’”’ are slender hourglass-shaped bones shown by the
section figured by Thevenin to be continuous with the neural
arch. The presacral neural arches bear a flat lamellar expansion,
the middle of which forms the rib-carrier, the whole structure
being identical with that of a nectridian vertebra. In fact the
whole column differs so fundamentally from that of a Cotylosaur

Proc. Zoou. Soc.—1918, No. X XI, 21

294 CAPT. D. M. 8. WATSON ON SEYMOURIA,

as to be incapable of ready derivation from an Embolomerous
type, while it seems to resemble that of living Amphibia in
being ‘‘ pseudocentrous,” 7. e. with the apparent centrum derived
from a membranous ossification lying between the notochordal
sheath and the cartilaginous blocks of the embryonic vertebra.
The animal is of considerable interest as showing that a reptilian
type of foot may occur in Amphibia in no way connected with
reptilian ancestry.

Hosauravus.—The remarkable skeleton from the Coal Measures
of Linton named by Williston Hosauravus copei, and described
by Cope, Williston, Moodie, and Case, seems to be certainly a
Cotylosaur if Case is correct in stating that the “ neural arches
are low and broad with horizontal zygapophysial faces and short,
heavy spines.” The ‘“intercentral” position of the ribs, which
has troubled certain American authors, really means very little.
The skeleton (which I have not seen) is probably not well
preserved, and if it be that of a reptile as primitive as Seymouria,
no doubt had double-headed ribs of which the capitulum articu-
lated with the intercentrum. The slab may have been split so
that only this head is preserved.

The chief interest of the specimen lies in the proportions of
the animal. Twenty-three left dorsal ribs are preserved, and the
position of the left hand suggests that twenty-eight or more
presacral vertebra were present. There are preserved twenty-
three caudals, the latter half of which series shows no signs of
tapering, so that the tail may easily have been quite as long
as the presacral part of the body. These proportions at once
recall the aquatic Embolomeri. Pteroplax has more than
28 presacrals, so has Pholidogaster, which animal has a tail as
long as its body. The long slender form of Cricotus is familiar
from Cope’s two skeletons. Thus it is not improbable that
Eosauravus retains a primitive form.

In possessing only two proximal tarsals, it is undoubtedly
more advanced than Seymouria.

Papposaurus.—I have already pointed out that the Lower
Carboniferous femur which is the type and only known material
of Papposaurus traquari distinctly recalls Seymouria. This
resemblance increases the probability (admitted by Williston
and Broili) that the form is a reptile. If so, it is by far the
earliest known.

Limnoscelis.—The large Cotylosaur from New Mexico described
by Williston as Zimnoscelis is in its limb-structure one of the
most primitive known. The skull is, however, far more advanced
than in Seymouria; it retains no trace of the typical Eryopine
shape of the latter skull, in particular the long slender otic
notch of the more primitive form is lost, the posterior surface
appearing to be truncated. The occiput unfortunately has not
yet been described in detail, but from Prof. Williston’s out-
line drawing it appears to be derivable from that of Seymouria
by a migration downwards of the ends of the paroccipital

THE MOST PRIMITIVE KNOWN REPTILE. - 295

processes, the fenestra ovalis retaining its position quite at the
lower surface of the skull.

The palate of Limnoscelis has advanced far beyond that of
Seymouria in the wide interpterygoid vacuity and in the
pronounced pterygo- -transverse flanges. |

The lower jaw seems to have lost the post-splenial and _ pro-
bably the anterior coronoids.

The vertebre in retaining a quite long spine are probably more
primitive than those of Seymouria, for all the Temnospondyls
have high neural spines.

The single- headed ribs are, however, an advanced feature.

In the shoulder- -girdle Limnoscelis is advanced in having a
posterior coracoid element, but retains in its cleithrum a pr imitive
bone lost by Seymouria.

The pelvis of Limnoscelis much resembles that of Sez ymouria mn
the general form of the ventral surface, which is very reminiscent
of Rachitomi in the short pubes. The posteriorly produced ilium
is also a primitive feature retained by both reptiles.

The hind leg of Zimnoscelis resembles that of Seymouria in
having a separate intermedium, but the conditions are different
in the non-ossification of the tibiale in the former animal.

Limnoscelis is thus in most ways more advanced than Sey-
mouria, although it is still a very primitive reptile.

Diadectide.— The Diadectids, known from New Mexico, Texas,
and probably also Europe (Stephenospondylus), are obviously
much more advanced reptiles than Seymouria. They have a
remarkably specialised ‘palate and occiput—which region I have
recently discussed in detail. The brain-case is theoretically
derivable from that of Seymouria, but is so extremely modified
that no useful end will be served by a further discussion.

Pariasauride.— With the Middle and Upper Permian group of
the Pariasaurs, we pass to reptiles which haveadvanced much from
the primitive Seymouria structure in all features of skull and
skeleton. Some of these differences I have already pointed out,
others are at once obvious from any comparison. One important
feature which deserves emphasis is the high position of the
fenestra ovalis, which lies on the front of the paroccipital process
far removed from the basioccipital and the tuber basisphenoidalis.
The opening is small and the stapes is a slender rod, without the
much expanded base-plate it must have had in Seymouria.

Procolophonide.—Procolophon and its allies are the most
advanced Cotylosaurs known. They differ extremely from Sey-
mouria in nearly all features. The differences which at present
most interest me are those in the brain-case and otic region. In
essentials these regions resemble the condition in Sphenodon,
and differ from that in Seymouria in ways I have already pointed
out, perhaps most markedly in the slender paroccipital process
with a grooved lower surface in which the stapes lies, and in the
high position of the fenestra ovalis.

The Cotylosaurs so far considered all differ markedly from

Zi"

296 CAPT. D. M. S. WATSON ON SEYMOURIA,

Seymouria in the condition of the brain-case and of the ear.
Although their structure could in practically all points be derived
from that of Seymouria, they do not in any way recall that
reptile. In the Captorhinids we find features which distinctly |
hark back to the structure of Seymouria, although the group as
uw whole is one of the most advanced of the ‘l'exan Cotylosaurs.

Through the work of Cope, Broili, Case, and especially Willis-
ton, we have a nearly perfect knowledge of the skeleton of both
Captorhinus and the more advanced Labidosauurus, and I have
given a somewhat more detailed account of the occiput than is
to be found in earlier writings.

The skull of Captorhinus is advanced in the loss of the
intertemporal and tabular and ‘the very great reduction of the
supratemporal; it retains the primitive feature of a lachrymal
reaching the nostril, and of a septomaxilla not appearing on
the face.

It resembles Seymouria in the following important characters :—

1. The occipital condyle is tripartite, the exoccipitals forming
a good deal of its surface.

. The fenestra ovalis is large, surrounded by the prootic,
paroccipital, basioccipital, and basisphenoid ; it is
placed low down in the skull so that its margin below
lies on the border of the tuber basisphenoidalis.

3. The basisphenoid is remarkably similar in the two
enera
4. The palate of Captorhinus, though advanced in the loss
of the palatine tusks and in the greater development
of the pterygo-transverse flanges, is on the whole not
unlike that of Seymouria.

In the vertebral column the remarkably swollen neural arches
of the presacral vertebre and the short nearly obsolete neural
spines, are points of resemblance between the two forms.

The Captorhinids have advanced over Seymouria in the
following ways :—

1. In the downgrowth of a process of the paroccipital between
the lateral margins of the basioccipital and the inner
border of the fenestra ovalis, so as to separate that
opening more widely from its fellow.

2. In the reduction of the paroccipital processes and the

rotation of their ends downwards.

. In an enormous widening of the supraoccipital.

. In a great deepening of the whole brain-case.

. In a reduction of the prootic so that it no longer reaches

the skull-roof.

6. In the loss of the tabular and the restriction of the
dermo-supraoccipital to the occipital surface. .

7. In the loss of the intertemporal and in Labidosaurus of
the supratemporal.

Ot He OO

THE MOST PRIMITIVE KNOWN REPTILE. 297

8. In the narrowing of the face.
9. In the loss of the palatine tusks.

10. In the better development of the pterygo-transverse
flanges.

11. In the lateral compression of the quadrate so that its
horizontal section is no longer U-shaped.

12. In such reduction of the pterygoid that it no longer
reaches the squamosal.

13. Ina considerable compression of the lower part of the
articular.

14, In the loss of the two anterior coronoids and the post-
splenial.

15. In the somewhat more modified axis.

16. In the presence of two sacral vertebre.

17. In the development of single-headed ribs.

18. In the reduction of the intercentra.

19. In the development of a coracoid in addition to the pre-
coracoid.

20. In the reduction and loss of ornament of the lower parts
of the clavicles and the anterior end of the interclavicle.

21. In the loss of the long posterior extension of the dorsal
part of the ilium.

22. In the more advanced humerus.

23. In the loss of the long adductor crest in the femur.

24. In the fusion of the intermedium and the tibiale.

25. In the obliteration of the otic notch by the swinging
backward of the upper end of the quadrate, so that the
posterior end of the skull seems cleanly truncated.

Thus the Captorhinids alone amongst Cotylosaurs present
definite significant resemblances to Seymouria, which in the case
of the palate are due merely to the retention of a primitive
structure, but in the important characters of the fenestra ovalis
and the vertebre are due to the possession by both types of a
structure which is certainly not derived from amphibian ancestors.

The position and character of the fenestra ovalis common to
Seymouria and Captorhinus is preserved throughout the whole
series of mammal-like reptiles, the Pelycosaurs and all the later
S. African types. This fact is clearly illustrated in the drawings
of text-fig. 15, where the stapes is indicated by shading.

This position of the fenestra is of course connected with that
restriction of the inner ear to the lower part of the side wall of
the brain-case which I have shown to be one of the most constant
and significant characters of the Anomodontia, and one which is
entirely restricted to them and the Captorhinids, which Prof.
Williston and the writer believe to be their ancestors. Com-
parison of the occipital views of skulls in text-fig. 15, with similar
drawings of any other reptiles, will bring out the difference
between the two types of structure of this region clearly.

Apart from the Captorhinids, only Limnoscelis and Seymowa

298 CAPT. D. M. S. WATSON ON SEYMOURIA,

Text-figure 15.

Series of occipital views, reduced to the same width, of the skulls of Anomodonts
and of Cotylosaurs connected with that group. To show the low position, at
the back of the tuber basisphenoidalis, which the fenestra ovalis occupies in
these reptiles, its situation is indicated by the stapes.

A. Seymouria bayloriensis.. « B. Captorhinus sp.

C. Varanosaurus acutirostris. D. Arctops willistoni.

KE. Diademodon browni. F. Deiopeus leptocephalus.
'G. Dimetrodon sp. H. Mormosaurus seeleyi.

A, from text-fig. 2; B, from Watson, 1916. C. Restored from the type-skeleton
in Munich; D, E, & H, from Watson, 1914; KF, modified from Watson, 1916;
G, from a skull figured by Case, modified by other material.

A-E form an approximate morphological series; F' & G represent an aberrant side
line arising from C; and H another distinct line also arising from a form near C.

THE MOST PRIMITIVE KNOWN REPTILE. 299

have this peculiar structure. I have already brigaded Limnoscelis
with the Captorhinids. Seymouria is so much more primitive
that it is remarkable to find in it a character of this kind
connecting it clearly with a definite series of more advanced
reptiles.

It remains to be seen whether the other reptiles have lost this
type of fenestra ovalis, or whether, as seems more probable, their
ancestors never possessed it.

It is a remarkably interesting fact that the primitive Pelyco-
saurs such as Varanosaurus retain primitive features found in
Seymouria which have been. lost by the Captorhinids. Such
are :—

The retention of a supratemporal lost in Labidosaurus.

The retention of a quadrate which is U-shaped in horizontal
section.

The retention of a suture between the pterygoid and squamosal
behind the quadrate.

The non-ossification of a posterior coracoidal element in
Varoops.

The retention of a considerably expanded lower end of the
clavicle and a largely expanded interclavicle.

The retention of a posterior process on the iJium.

The characters in the above list represent features in which
the Captorhinids are more advanced than the early Pelycosaurs ;
they are, however, all acquired by later Anomodonts and represent
parallel evolutionary changes which have gone on in the Capto-
rhinids and Pelycosaurs independently since their separation.
The most striking of these characters is the peculiar flattening
and lateral compression of the quadrate. In principle the
quadrates of Captorhinus and Dimetrodon are identical, differing
only in the wide articular condyle of the Cotylosaur. As this
type of quadrate is restricted to the members of the Superfamily
Sphenacodontide, we have a very remarkable case of parallel
evolution, which in the original stock does not follow the main
line of the evolution of the derived group, but duplicates a
structure restricted to a single aberrant side line.

The study of the Seymouria skeleton which fills this paper
shows that that reptile is far more primitive than any other,
presenting a strange mosaic of characters derived without change
from the Embolomerous Labyrinthodonts with common reptilian
characters marking distinct advance over an amphibian structure,
and one or two which are restricted to the mammal-like reptiles
and such Cotylosaurs as can be brought into connection with
them.

These facts have an. important bearing on reptilian classifica-
tion, suggesting as they do that all reptiles may be divided into
two groups, one composed only of the Anomodontia and the
Captorhinomorpha, the other of all other reptiles.

300 CAPT. D. M. S. WATSON ON SEYMOURIA,

In the Cotylosaurs this division corresponds with that which
I have previously founded on the character of the otic notch,
and the mode in which a vertically placed quadrate is arrived at.
I have recently found that this division on the character of the
otic notch applies to all reptiles, affording a sharp distinction
between the Anomodonts and all other groups, so that we have
now two quite independent series of characters, the otic notch
and the structure of the fenestra ovalis, which allow us to split all
reptiles, Seymouwria excepted, into two groups. Seymouria itself
shows characters definitely connecting it with one of these.

The curious way in which the structure of Seymouria is built
up of perfectly well-developed amphibian characters and equally
decisive reptilian features, those of intermediate type being very
rare, affords a magnificent example of the way in which the
evolution of great groups may have taken place by the rapid
change of all the definite morphological entities of which it
may be regarded as made up, the changes occurring quite inde-
pendently and over a considerable time, the passage from the
structure of the more primitive to the advanced group being quite
gradual when viewed as a whole, but when further considered
and analysed found to depend on a rapid evolution of separate
regions apparently independently of each other.

I hope that this study of Seymowria will be regarded as placing
beyond dispute the origin of the reptiles from the Embolomerous
Labyrinthodonts.

I wish to express my thanks to the Percy Sladen Trustees, who
paid the expenses of my visit to Texas, to Prof. 5. W. Williston,
who allowed me to study freely the superb collection of Lower
Permian Tetrapods which he has built up in Chicago, to President
Osborn and the Staff of the American Museum for similar
privileges there, and to’Prof. F. Broili, who placed the treasures
at Munich at my disposal. I regret that circumstances at
present forbid my obtaining his permission to publish a descrip-
tion of the Seymouria material of which he gave so excellent a
description. Finally, I wish to thank my wife for editing this
paper and thereby removing many obscurities.

Titerature.

Broiui, F. 1904. ‘“‘ Permische Stegocephalen und Reptilien aus
Texas.” Paleontographica, Bd. li. pp. 1-120.

——, 1908. “ Ein montiertes Skelet von Labidosaurus hamatus
Cope, einen Cotylosaurier aus dem Perm von Texas.”
Zeitsch. deutsch. geo]. Gesell. Bd. lx. Heft 1, pp. 63-67.

Broom, R. 1913. “Studies on the Permian Temnospondylous
Stegocephalians of North America.” Bull. Amer. Mus.
Nat. Hist. vol. xxxu. pp. 563-595.

Casg, E.C. 1911. A Revision of the Cotylosauria of North
America.

THE MOST PRIMITIVE KNOWN REPTILE. 301

Corse, HE. D. 1896. ‘‘ Second Contribution to the History of the
Cotylosauria.” Proc. Am. Phil. Soc. vol. xxxv. pp. 123-
149.

v. Huene, F. 1913. The Skull Elements of the Permian
Tetrapoda in the American Museum of Natural History,
New York.

Tueventn, A. 1906. “Amphibiens et reptiles du terrain houiller
de France.” Ann. de Paléontologie, tome 1. pp. 145-

eAGa:

Wiuiston, 8. W. 1908. “The oldest known Reptile.” Journ.
Geol. vol. xvi. pp. 175-192.

——. 1910. “ Desmospondylus anomalus.” Bull. Geol. Soc.
AIMer XX1. p. 280.

——. 1911. American Permian Vertebrates. Chicago, 1911.

——. 1916. “Synopsis of the American Permo-Carboniferous
Tetrapoda.” Contributions from the Walker Museum,
vol. 1. No. 9, pp. 193-236.

——. 1917. “ Labidosaurus Cope, a Lower Permian Cotylosaur
reptile from Texas.” Journ. Geol. vol. xxv. pp. 309-
321.

——. 1918. “(1) The Evolution of Vertebre.” <‘ (2) The
Osteology of some American Permian Vertebrates, III.”
Contributions from the Walker Museum, vol. 11. pp. 75-

pee hi.

Watson, D. M.S. 1912. “The Larger Coal-Measure Amphibia.”
Manch. Memoirs, vol. li. pp. 1-14.

——. 1914. “Notes on Varanosaurus acutirostris Broil.” Ann.
& Mag. Nat. Hist. ser. 8, vol. xiii. pp. 297-310.

——. 1914. “The Deinocephala, an Order of Mammal-like
Reptiles.” Proc. Zool. Sec. 1914, pp. 749-786.

——. 1914. “Notes on some Carnivorous Therapsids.” Proc.
Zool, Soc. 1914, pp. 1021-1038,

——. 1914. “ Procolophon trigoniceps, a Cotylosaurian Reptile
from South Africa.” Proc. Zool. Soc. 1914, pp. 735~
747,

——. 1914. “On the Skull of a Pariasaurian reptile, and on
the Relationship of that Type.” Proc. Zool. Soc. 1914,
pp. 155-180.

——. 1916. On the Structure of the Brain-case in certain
Lower Permian Tetrapods.” Bull. Amer. Mus. Nat.
Hist. vol. xxxv. pp. 611-636.

——. 1917. “A sketch Classification of the Pre-Jurassic Tetra-
pod Vertebrates.” Proc. Zool. Soc. 1917, pp. 167-186.

——. 1917. “The Evolution of the Tetrapod Shoulder-girdle
and Fore Limb.” Journ. of Anat. vol. lii. pp. 1-63.

ON 'THE FRESHWATER FISHES OF GREAT BRITAIN. 303

EXHIBITIONS AND NOTICES.
May 7th, 1918.

Prof. E. W. MacBripg, D.Sc., F.R.S., Vice-President,
in the Chair.

The Ages of Hlephants, as inferred from the Molar Teeth.

Mr. R. I. Pocock, F.R.S., F.Z.8., Curator of Mammals, ex-
hibited a series of the molar teeth of Elephants that had died in
the Gardens, and drew particular attention to the state of wear
of the last tooth of an elephant known to be about 50 years old.
Since this tooth comes into use in about the 40th year and had
lost by wear more than one-third of its lamine in 10 years,
Mr. Pocock concluded that the animal would have been toothless
and would have come to the end of her time before she was 70.

Mr. E. Heron-Aten, F.L.S., F.R.M.S., F.Z.S., gave a lantern
exhibition of Arenaceous Foraminifera of the genus Thurammina.

May 28th, 1918.

Dr. 8. F. Harmer, F.R.S., Vice-President,
in the Chair.

The Secrerary read the following Report on the Additions
to the Society's Menagerie during the month of April 1918 :—

The registered additions to the Society’s Menagerie during the
month of April were 209 in number. Of these 129 were acquired
by presentation, 77 were received on deposit, and 3 were bred in
the Menagerie.

The following may be specially mentioned :—.
_ 1 Barbary Sheep (Ammotragus lervia), born in the Menagerie
on April 18th.

1 Blossom-headed Parrakeet (Palwornis cyanocephala) 3 , from
India, presented by Miss H. F. Dunbar on April 27th.

Mr. C. Tate Recan, M.A., F.R.S., F.Z.8., gave an account,
illustrated by lantern-slides, of the Freshwater Fishes of Great
Britain, with special reference to their value as food and to the
possibilities of increasing their economic use.

——

304 LT.-COL. 8. M. COPEMAN ON A COLONY. OF

June 11th, 1918. «
A. Ezra, Esq., Vice-President, in the Chair.

The SecrerAry read the following Report on the Additions
to the Society’s Menagerie during the month of May 1918 :—

The registered additions to the Society’s Menagerie during
the month of May were 68 in number. Of these 14 were
acquired by presentation, 46 were purchased, and 8 were bred in
the Menagerie.

The following may be specially mentioned :—

2 Grecian Ibexes (Capra egagrus), born in the Menagerie on
May 9th and 15th.

1 Ruby-throated Warbler (Calliope calliope), from India, pre-
sented by W. H. St. Quintin, F.Z.S., on May 29th.

1 Indian Chameleon (Chameleon calcaratus), from Calabar,
presented by A. M. Kinloch on May 1ith; new to the Collection.

Observations on a Colony of Burrowing Bees (Andrena fulva)
on Primrose Hill.

Lt.-Col. 8. Monckx'ron Copeman, M.D., F.R.S., F.Z.S., exhibited
examples of the Burrowing Bee, dndrena fulva, and made the
following remarks :—

The existence of a flourishing colony of a burrowing bee
(Andrena fulva) on the western slope of Primrose Hill, where,
during the present year, it has spread over a much more extended
area than previously occupied by it, appears worthy of record
for the reason that one locality only in the neighbourhood of
London—a small area on Hampstead Heath—is cited as a
habitat of this insect in the British Museum Catologue of
Hymenoptera.

For the past six years this particular colony has been kept
under observation, during which period the number of nests has
not markedly varied from season to season until the present year.
The unexampled spread of the colony this year (1918), curiously
enough, was consequent on the manceuvres of a Cadet Battalion
last autumn having converted a previously grassy slope into a
quagmire of mud, now represented by a smooth bare surface of
clayey loam. ‘This patch of ground the bees have found so much
to their liking for nesting-purposes that (as indicated by a scale-
plan drawn by my son last year) they have, in large measure,
deserted their former haunts alongside a neighbouring path to
seize upon the new territory so.conveniently provided for them.

Notwithstanding the hardness of the ground, the female bee
excavates in it a tunnel which in many instances, on probing
with a fine flexible wire, I have found to extend to a depth of

BURROWING BEES ON PRIMROSE HILL. 305

ten or twelve inches, and occasionally even more. This insect is
usually about the size of the ‘“‘ worker” honey-bee, which she
somewhat resembles in form. She is, however, handsomer, and
also appears larger than her actual size owing to the abdomen
especially being covered with a thick pubescence, which ranges in
colour from a pale orange to a tawny-red in hue. This bee is,
in consequence, a conspicuous object on a sunny morning as she
flies in mazy flight, usually a few inches only above the ground.
The male insect 1s utterly unlike her, being smaller, less pubescent,
and more sombre-toned in hue.

As the outcome of six years’ observations it has been found
that over a period which has varied, according to prevalent
meteorological conditions, from the middle of April to the first
week in May the bees make their first appearance quite suddenly,
following on one or two days of warmer weather. For the first
few days the number of males emerging greatly exceeds that of
females, the relative divergence in numbers being as great
fifty to one.

Careful examination of bare patches of ground in the neigh-
bourhood, so soon as the bees are first seen, discloses a. number of
small round holes which look as though they had been punched
out with the point of a pencil. These are the open doorways of
ae burrows in which eggs were laid in the previous spring and

n which the bees, passing through the stages of larva and pupa,
ae finally developed into the perfect insect. Herein the bee
awaits the stimulus of warmth to give it strength to break
through the slight covering of earth beneath which it has been
sheltering since the previous autumn.

Within a few days further, mating having taken place, the

male bees are no longer met with, their brief life ending when
the object of their existence has been attained. Almost 1mme-
diately the females begin to dig their burrows, as indicated by the
appearance of numerous little mounds like miniature volcanoes,
eventually three or four inches in width and somewhat less in
height. These are gradually built up of the tiny particles of
earth thrown out, a few grains.at a time, as the work progresses.
In the first stages of the excavation the bee throws out the
particles of earth between her hind legs, after the fashion of a
dog when digging for a rat or rabbit, aiding the process by
sweeping motions of her abdomen. Later on, however, when
progress has been made beneath the surface, she brings up the
grains of earth she has dug out, on the top of her head, generally
waiting a few moments on reachiag the mouth of her burrow,
before pitching them over the edge with a sudden jerk. The
resulting mounds are easily recognizable owing to the material
of which they are composed being lighter in colour than the
surrounding soil.

On completion of the main burrow small nurseries are opened
out from its sidesf in each of which is deposited a carefully
kneaded pellet of pollen and honey, which has to suffice for the

306 THE SECRETARY ON ADDITIONS TO THE MENAGERIE.

needs of the grub which will develop from the egg laid upon
its surface. This done, the little chamber is sealed up, and the
process again repeated.

During the numerous foraging expeditions necessary for obtain-
ing sufficient pollen and nectar for the sustenance of her future
broods, no care is taken to protect the burrow, a fact of which
advantage is occasionally taken by a Vomada bee, too lazy herself
to build nurseries or to gather stores of food. Waiting therefore
till the rightful owner has set out on her travels, the ‘ cuckoo”
bee slips in and deposits an egg on any pellet of food of adequate
size not yet.sealed up.

The Andrena bee not infrequently appears to experience con-.
siderable difficulty in recognizing her own burrow, as she may
often be seen to tentatively explore the entrance to several before
suddenly diving down her own. Marking the bee with a little
powdered chalk as she leaves her burrow will render it easy to
recognize the rightful owner on her return.

The burrowing bee having finished her labours underground
does not take the precaution of closing up the mouth of the
burrow, but on her death, within about a month from emergence
from her winter quarters, leaves it to be graduallv obliterated by
the levelling of the mound outside under the influence of rain
and wind.

October 22nd, 1918.

Dr. A. SurrH Woopwarp, 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 months of June,
July, August, and September, 1918 :—

JUNE.

The registered additions to the Society’s Menagerie during the
month of June were 38 in number. Of these 22 were acquired
by presentation, 14 were bred in the Menagerie, and 2 were
_ deposited.

The following may be specially mentioned :—

1 White-bearded Gnu (Connochetes albojubatus), born in the
Menagerie, June 17th.

4 Taurus Lizards (Lacerta taurica), new to the Collection, from
Calamaria, Salonika, presented by G. H. Colt, F.R.C.S.,on June
20th.

JULY.

The registered additions to the Society’s Menagerie during the
month of July were 328 in number. Of these 24 were acquired

ON WHEAT WEEVIL IN AUSTRALIA. 307

by presentation, 9 were bred in the Menagerie, and 295 were
deposited.

The following may be specially mentioned :—

1 Kiang (Zquus kiang), from Tibet, born in the Menagerie,
July 15th.

1 Eland (Taurotragus oryx), from South Africa, born in the
- Menagerie, July 16th.

AUGUST.

The registered additions to the Society’s Menagerie during the
month of August were 81 in number. Of these 59 were acquired
by presentation, 9 were bred in the Menagerie, and 13 were
deposited.

The following may be specially mentioned :—

1 Thar (Hemitragus jemlaicus), from the Himalayas, born in
the Menagerie, August 10th.

5 Coypus (Myocastor coypus), from the Argentine, born in the
Menagerie, August 16th.

2 Verticillated Geckos (Gecco verticillatus), from Rangoon, pre-
sented by Mrs. Hilda D. Sedgwick, August 10th.

SEPTEMBER.

The registered additions to the Society’s Menagerie during the
month of September were 31 in number. Of these 15 were
acquired by presentation, 2 were bred in the Menagerie, 8 were
deposited, and 6 were received in exchange.

The following may be specially mentioned :—

Two hybrid Porcupines (Hystri« cristatus x H.africe australis),
born in the Menagerie on September 20th.

On behalf of Mr. E. Gerrard, Mr. R. I. Pocock, F.R.S.,
exhibited the skin of an abnormally coloured red deer stag shot
by Major R. C. Forster in Scotland, where it had been known
for many years. It was a partial albino, the red hairs of the
body being mixed with grey, the face, throat, and legs being
white, the hoofs pale horn-colour, the muffle pink, and the eyes
blue.

ae

Professor H. M. Lerroy, F.Z.S., Hon. Curator of Insects, gave
an account, illustrated by lantern-slides, on the work he had
accomplished for the Wheat Commission on Wheat Weevil in
Australia.

y

308 THE ‘“‘ NEW” RABBIT DISEASE.

November 5th, 1918.
Prof. E. W. MacBripg, D.Sc., F.R.S., Vice-President,

in the Chair.

Mr. D. Sera-Smiru, F.Z.8., Curator of Birds, exhibited a
mounted specimen of a hybrid Cockatoo (3 Roseate Cucatua
roseccapilla x 2 Lesser Sulphur-Crested C. sulphurea), bred in
1917 at Hartwell House, Aylesbury, by Mrs. Lee.

Diagnosis of Helminth Infections.

Dr. R. T. Lerper, M.D., D.Sc., F.Z.S., gave a lantern
exhibition on Diagnosis of Helminth Infections from the cha-
racter of the eggs in the feces. He stated that, by examination
of the faeces of a living animal, the extent and specifie nature of
most helminthic infections could be accurately determined, and
the method had been applied successfully as a routine practice
in the case of man, rabbit, dog, cat, and pig, and was apparently
capable of indefinite extension.

The eggs of parasitic worms were constant in character and of
great systematic importance. The ground-plan of the egg-shell
indicated the genus or even subfamily to which the parasite
belonged, and specific differences were found in slight but con-
stant peculiarities in relative Jength and breadth, and in the
conformation of excrescences on the surface of the shell.

The * New” Rabbit Disease.

Dr. R. T. Lereer also gave a demonstration on the “new”
rabbit disease. Examination of a large number of rabbits shows
that the chief cause of mortality is a coccidial invasion of the
intestinal wall or of the lining of the bile-ducts. According to
Fantham and others the causal agent in both types of disease is
Kimeria stiedew, but Dobell holds that the intestinal lesion is due
to a distinct species. In many cases changes in the liver attri-
buted to coccidiosis were the result of infection with Cysticercus
pisiformis, the larval stage of the dog tapeworm Tenia serrata.
Large swellings in the region of the head and neck, suspected to
be cancerous, were due to Caenurus serialis, the larva of the
dog tapeworm Tenia cenurus. Of relatively small economic
importance are infections with the threadworm Oxyuris ambiguus
and the tapeworm Ctenotenia leuckarti. There is some evidence
that a bacterial infection may occasionally be the cause of death,

The coccidial infections pass from infected to healthy animals
through the feces. When freshly passed the coccidial oocysts

THE SECRETARY ON ADDITIONS TO THE MENAGERIE. 309

are not infective. They only become so after a period of delay,
in which certain developmental changes take place. These
changes proceed more rapidly in dry than in wet feces. Pre-
vention depends upon the systematic periodical removal and
destruction by burning of all pellets and contaminated bedding,
and the use of some fluid which will destroy such oocysts as
remain in the hutch.

Although several cases of coccidial infection in man have been
recorded, Dobell maintains that in none of these cases is Himeria
stiedee the causal agent. There would appear therefore to be no
risk of infection to man.

The cystic stages of the tapeworms of the dog appear to occur
chiefly in those rabbits fed with dandelions and other green
stuffs collected from the roadsides, where the vegetation is
especially liaple to contamination with feces of dogs which have
acquired their infections from eating uncooked rabbit offal.

Professor H. M. Lerroy, F.Z.S., exhibited a series of lantern-
slides, from photographs taken during the recent visit to the
Zoological Gardens, Sydney, N.S. Wales.

November 19th, 1918.

Dr. A. Smrra Woopwarp, LL.D., F.R.S., Vice-President,
in the Chair.

The Secrerary read the following Report on the Additions to
the Society’s Menagerie during the month of October, 1918 :—

The registered additions to the Society’s Menagerie during the
month of October were 22 in number. Of these 8 were acquired
by presentation, and 14 were deposited.

The following may be specially mentioned :—

1 Black-cheeked Cercopitheque (Cercepithecus ascunius), from
the Lower Congo, presented by Chevalier E. Carton de Wiart on
October 2nd.

2 Mississippi Alligators (Alligator mississippiensis), from North
America, deposited on October 28th.

1 Reticulated Python (Python reticulatus), from the East
Indies, deposited on October 11th.

Special attention may also be directed to a captured German
Carrier-Pigeon Loft, with 35 of the captured Pigeons, taken by
the Canadians at Folies, France, on August 9th, and presented
to the Society on October 30th by the British Armies in France,
at the suggestion of the War Office.

Proc. Zoot. Soc.—1918, No. XXII, 22

310 CHIMPANZEE WITH PULMONARY TUBERCULOSIS.

Miss K. Lanper, F.Z.S., described the method of preparing
skeietons by the use of trypsin, and exhibited a number of
successful examples from the Society’s Prosectorium.

Mr. E. HarscueK described his investigations into the forms
assumed by drops and vortices of gelatin in various coagulants,
exhibited a series of the formations he had obtained which
simulated animal structures, and demonstrated the method by
which he obtained his results.

——

Professor F. Woop-Jonés, F.Z.8., exhibited a cast and a set of
Rontgen-ray photographs taken from a Chimpanzee belonging
to the Society, which had recently died from pulmonary tuber-
culosis, and called attention to the possibility of diagnosing
tubercle by this method in living subjects.

Errata in Mr. L. A. Lanz’s communication supra, pp. 11-17.

Table, last line.
For 26, read 36.

Page 13. Footnote, line 3 Seni bottom.
For 24, read 28.

Page 14. Line 9 from bottom.
For 37, read 39.

Page 15. Last line.
For 121, read 171.

INDEX.

1918.—Pages 197-310.

[New names in clarendon type.

Systematic references in italics.

(z.8.L.) indicates additions to the Society’s Menagerie. |

Alligator mississippiensis (z. s. L.), 309.
Ammotragus lervia (z. s. L.), 303.
Andrena fulva, 304.
AVEs:
Exhibition of a mounted specimen of
a hybrid Cockatoo, 308.

Calliope calliope (z. s.u.), 304.

Capra egagrus (z. 8. L.), 304.

Cercopithecus ascanius (z..8. L:), 309.

Chameleon calcaratus (z. s. L.), 304. |

Connochetes albojubatus (z. s. L.), 306. |

Crossorhinus jurassicus, sp. n.,
231.

Equus kiang (z. s. L.), 307. |
|

Gecco verticillatus (z. s. u.), 307.
GEOGRAPHICAL:
Madagascar Frogs of the genus Mai-
tidactylus, 257.

ed

Hemitragus Jemlaicus (z. s. u.), 807.
Hypnomys, gen. n., 210.

— mahonensis, sp. n., 218.
—— morpheus, sp. n., 219.

Hystrix cristatus (z. s. L.), 307.

INSECTA :
Observations on a colony of burrowing
Bees, exhibited, 304; On Wheat
Weevil in Australia, 307.

Lacerta taurica (z. s. L.), 306.

Mamnanta:
On a new genus of Extinct Muscar-
dine Rodent from the Balearic
Islands : structure,
209; Notes

Leonardslee, 255;

systematic,

on the Beavers at

The ages of
Elephants, as inferred from the
Molar Teeth, exhibited, 303; An
abnormaliy coloured Red Deer,
skin exhibited, 307; The ‘“‘New”
Rabbit
308,

Mantidactylus ambohimitombi,

sp. n., 260.

Mo.uuvsca:

Disease, demonstration,

Ciliary action in Pleurobrachia pileus,
structure, 263 ; Lantern-exhibition
of Arenaceous Foraminifera, 303.
Morpnoitocy. See Srrucrure.
Myocastor coypus (z. s. L.), 3807.

Xil ' INDEX.

Paleornis eyanocephala (z, s. L.), 303.
PT HO TO GY.

The function of Pathological states
in Evolution, 237; Diagnosis of
Helminth infections, 308.

Piscus:

On two new Elasmobranch Fishes:
systematic, 231; Account of the
Freshwater Fishes of Great
Britain, 303.

Protospinacide, fam. n., 2382.
Protospinax, gen. n., 233-
—— annectans, sp. n., 233.

Python reticulatus (z. s. u.), 309.

ReEprILta :

Comparison between the lower jaws
of the Cynodont Reptiles Gompho-
gnathus and Cynognathus: struc-
ture, 197; On Hermaphroditism in
a Lizard: structure, 223; Mada-
gascar Frogs of the genus Manti-

systematic,

dactylus: structure,

eS SE ee

250 > On
primitive known reptile: structure,
267.

Seymouria, the most

STRUCTURE,

Mamuatia: On a new genus of ex-
tinct Muscardine Rodent from the
Balearic Islands, 209.

ReprintA: Comparison between the
lower jaws of the Cynodont Rep-
tiles Gomphognathus and Cyno-
gnathus, 197; On Hermaphro- .
ditism in a Lizard, 223; Mada-
gascar Frogs of the genus Manti-

257 On
Seymouria, the most primitive
known reptile: structure, 267.

Mouuvsca: Ciliary action in Prewro-
brachia pileus, 263.

dactylus, systematie,

Taurotragus oryx (Zz. 8. L.), 807.

PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET,

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-

[| PROCEEDINGS

OF THE

GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

OF THE

ZOOLOGICAL SOCIETY

OF LONDON.
1918.

PARTS I. & II.

- conraininc Paces 1 to 196, with | PuLatEe

AND 82 TExT-FIGURES.

AUGUST 1918.

PRINTED FOR THE SOCIETY,
SOLD AT ITS HOUSE IN REGENT’S PARK.
LONDON :

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

[Price Twelve Shillings. |

Ww

LIST OF CONTENTS,

1918, Parts I. & II. (pp. 1-196)

EXHIBITIONS AND NOTICES.

The Szcrurary. Report on the Additions to the Society’s Menagerie during the months
of November and December, 1917

eure ee ee eevee ® ee ers e# Fe eo wo ee ee eeee et ce oo

Mr. D. Szru Smitu, F.Z.S., Curator of Birds. Exhibition of, and remarks upon, a series
of lantern-slides made from photographs of Reptiles taken in the Gardens

Dr. P. Cuaumers Mrrcuett, F.R.S., Secretary to the Society. Communication of a letter
from Mr. T. E. Whitehead on the Wild Dingo ......

eorrteete ee tee OCF esses se 8 oe

Page

188

Dr. R. Broom, C.M.Z.S. Exhibition of, and remarks upon, a New and a Rare Species of ~

the Golden Moles. 005 ae a Pay ee ote ae tater ened en ee

The Szcreraky. Report on the Additions to the Society’s Menagerie during the month
of January, 1918....... o/e id vs ie eo wie ww ee hs ae Ohta Siasts clas) ean

Dr. Suita Woopwarp, F.RB.S., V.P.Z.8. Exhibition of a copy of an incised drawing of a
hunted deer... .. 6. cee cece e eee cee cece reece teens etre er ee eaes ig eses

Professor Poutton, F.R.S., F.Z.S. Communication of a letter from Captain G. D. Hale
Carpenter on an African Civet attacking Human Beings .........--2eee+eesesee

Professor E. W. MacBripn, M.A., D.Se., F.R.S., F.Z.S. An account, iWushiaesd by
lantern-slides, of his recent investigations into the development of the Sea- Urchin. .

189
190
191
19}

192

_ Mr. D. Sern Smiru, F.Z.S., Curator of Birds. Exhibition and description of skins of the

Hoatzin, illustrated by lantern-slides” ...-....'55 06.5% esses oes meiiem ar crane pee

Mr. C. Tarr Ruaan, M.A., F.R.S., F.Z.S. Exhibition of photographs of an Indo-Pacifie
Chetodont Fish. -(Text=figure 17) io) sieiurno eset os oes cle bere 06 tes 4

192

192

The Srcrzetary, Report on the Additions to the Society’s Menagerie during the nek

of February; 1918 5; os sed pees Pelee Pie beter ee oe ee eee Pate Pa!

Miss L. E. Cunzsman, Assistant Curator of Insects. Exhibition of specimens of an Hast-
African homopterous inséet (55.2 ve. bees ek olga, op ee aan ee plea ca een

Dr. A. Smita Woopwarp, F.R.S., V.P.Z.S. Exhibition of, and remarks upon, the fossil
rostral teeth of Hopristis and Pristis ..-+..00.ceees ee cern sn ceeeneecenes Dee

193
194

194

Contents continued on page 3 of Wrapper.

Exhibitions and Notices (continued).

Mr. G. A. Bounzncer, F.R.S. Exhibition of the head of a Congo Fish, Hydrocyon
ME aie ee ask ne sb suet SA a, gusts 2 cme pala MN leo pln Sie)e viele oe wt 195

Mr. D. Sura Saru, F.Z.8., Curator of Birds. Exhibition of, and remarks upon, a Zulu
Meer tO A Sela ee leit os we gle ese Avant @ a@leatoe vs hs sce « 195

Dr. P. Onaumers Mircnutt, F.R.S., Secretary to the Society, Remarks on an advertise-
ment announcing Fur Sales in the United States .....-.. ..-eee ee ewer ee ee ee ee 195

Professor Woop-Jonzs, F.Z.S., Honorary Acting Prosector. Exhibition of, and remarks
upon, specimens from the Prosectorium illustrating the effects of Rickets.......... 196

PAPERS.

Page

1. Skull of Rana tigrina Daud. By B. L. Buaria, M.Se., and Batnt Prasnap, M.Sc.
(Assistant Professors of Zoology, Government College, Lahore). (Text-figures 1-9.) 1

2. Description of a new Snake of the Genus Oligodon from Upper Burma. By G. A.
. Seemann rs) ob 45. -(hext-fienre 1.) 2. 5 vo ce eee te sec eee e ene e thes 9

8. Reptiles from the River Tajan (Transcaspia). By L.A. Lanrz. (Plate I.) ........ 11

_ 4. On the External Characters of the Lemurs and of Tursius. By R. I. Pocock, F.R.S.
(Text-figures 1-16.) =... ..-. cee ase Sh Oe Sear ae eM Aap eit, rh oe BS bs

5, A Olassification of the Pyrauip#, subfamily Hypsorropinm. By Sir Grorce Hampson, ~
(oy eS 2 ere See 2 Ny FeSO pe NRE RS Sica TE RE A ae Pi eae ee ee Seat OO

6. First Report on the Inheritance of Visible and Invisible Characters in Silkworms.
By Miss Marpsz L. Cunauorn, F.Z.8., FLAS, FBS... 2. ee eee een ees 133

7. Notes on Oetacea stranded on the British Coasts during 1913-1917. By Srpney F.
Harmer, Sc.D., F.R.S., F.Z.S., Keeper of Zoology in the British Museum (Natural
PPISLOEY).°-....- .- abel NUS GAL, 2 0 SIS BS ee Als AC ea 147

8. On the Variation of the Pit-Viper, Lachesis atror. By Miss Joan B. Procrmr, F.Z.S.
OWemteneures 1-5.) ee eek ce eee eens Pacts PN ichetar speed aI ta) = 163

9. Report on Deaths of Animals in the Gardens in 1917. By J. A. Murray, M.D.,
B.Sce., F.Z.8., Director, Imperial Cancer Research Fund, Pathologist to the Society.. 183

a vig, SEMI ip Rn irs et oie Ps ere Odi Dy see
| Meteo Council and Officers. css eccs.ccecdceeve coveveves: anette Apier ates Beate neessl ie ali
ER IW eS lla mes Secs ie ie Pe acne see + waeatoe ll cued ii
Alphabetical List of Contributors ........ ... Re ee ee Rt anne tle MM Mene are ek aula Vil

Index eeee eo eeee seer sce soon eesee2Faresee- es SCG ee +e et Coe oe eeeeneetiereest see se se FS Be ww XI

PLATE,

1918, Parts I. & II. (pp. 1-196).

Plate Page
I. 1. Gymnodactylus microlepis, sp.n., d, River Tajan. 2. G, fedt-) —
schenkoi Str., 6, Samarkand. 3. G. caspius Hichw., 9, 17
River Tajan: s. 0... pases seek sees 50 2 Dele se eee a
NOTICE,

The ‘ Proceedings’ for the year are issued in four parts, paged consecutively,
so that the complete reference is now P. Z. 8.1917, p.... The Distribution
is usually as follows, but on account of war conditions Parts I. & II. are
published simultaneously and Parts III. & IV. will probably be -similarly
issued :-—

Part I. issued in March,
Sa oh! Brae aes June. -

. III. ,, September.
Peter rigs ti" December.

‘ Proceedings,’ 1917, Parts III. & IV. (pp. 217-388), uae published _
together on February 19th, 1918.

The Abstracts of the ‘ Proceedings,’ Nos. 175- 183, are
contained in this Part.

PROCEEDINGS

OF THE

OF THE

| GENERAL MEETINGS FOR SCIENTIFIC BUSINESS

ZOOLOGICAL SOCIETY
OF LONDON.

1918.

PARTS III. & IV.

CONTAINING Paces 197 to 310, with 2 Puarzs

AND 80 TExtT-FIGURES.

MARCH 1919.

PRINTED FOR THE SOCIETY,
SOLD AT ITS HOUSE IN REGENT’S PARK.
LONDON :

| MESSRS. LONGMANS, GREEN, AND CoO,,
PATERNOSTER ROW.

of [Price Twelve Shillings. } Sei

LIST OF CONTENTS:

1918, Parts III. & IV. (pp. 197-310).

EXHIBITIONS AND NOTICES.

Mr. R. I. Pocock, F.R.S., F.Z.8., Curator of Mammals. Exhibition of, and remarks upon,
a series of the molar teeth of Elephants ................ Frere rr sci sutaeiae

Mr. E. Heron-Auven, F.L.S., F.R.M.S., F.Z.S. Lantern-exhibition of Arenaceous
Foraminifera... 006 sece ve oe oe he en cde a eias. arse te goalies > clase

The Szcretary. Report on the Additions to the Society’s Menagerie during the month
Of April; TOUS! o.0'S a winless § # ti ele mised a lag aa naa ne ae ECS ae. ey i

Mr. C. Tarz Ruan, M.A., F.R.S., F.Z.8. Account, illustrated by lanterns of the
Freshwater Fishes of Great Britain .... 0. 5.¢.2 02: « «= sicdiys « cea eee eee ine

The Secretary. Report on the Additions to the se ti Menagerie during the month
of May; T9018) ica oe nM eon ome sb ee ee ee eh gies ae er er

Lt.-Col. S. Monckton Copeman, M.D., F.R.S., F.Z.8. Exhibition of, and remarks upon, a
colony of Burrowing Bees on Primrose Hill ...... 5.0 ees... 5 +s 55a em eee

The Szcrutary. Report on the Additions to the Society’s Menagerie during the months
of June, July, August, and September, 1918 2.0000. 6.550 e ee eons) sie see

Mr. R. I. Pocock, F.R.8S., F.Z.S., Curator of Mammals. Exhibition, on behalf of Mr. E.
Gerrard, of the skin of an abnormally coloured Red Deer .....- «00 os ae aie

Prof. H. M. Lerroy, F.Z.S., Hon. Curator of Insects. Account, illustrated by lantern-
slides, of Wheat Weevil in Australia ......cee0ee..eeeece od ore ob 0'd oie Ae ee

Mr. D. Satu Sarru, F.Z.S., Curator of Birds. Exhibition of a mounted specimen of a

hybrid Oockatoo. 2 isc. One a's stent =< 4 + wince ovine sie ocots mele aie Oe eae atte

Dr. R. T. Lerner, M.D., D.Se., F.Z.8. Lantern-exhibition on Diagnosis of Helminth
Infections < oo ised iia ck ei Be ae hee Secs oinlla ail te ae ella o ages Gu ule eae ia

Dr. R. T. Lurene, M.D., D.Sc., F.Z.S. Demonstration on the “new” Rabbit disease

Prof. H. M. Lzrroy, F.Z.S., Hon. Curator of Insects. Exhibition of a series of lantern-
slides from photographs taken in the Zoological Gardens, Sydney, N.S. Wales

Contents continued on page 3 of Wrapper. |

Page
303

303
303
303
304

— 804

309

SE NS, ee ee ee eee eat

Exhibitions and Notices (continued).

Page

The Secrerary. Report on the Additions to the Society’s Menagerie during the month
PEOctober, 1018 so... .c ek ee es we pee ees nat SiR a shedatnne: Metre cine. s 3 sik a Ua oOo

Miss K. Lanpur, F.Z.S. Description and exhibition of the method of preparing skeletons
Mpeme lise Of Trypsin 2.2. 6. eh ees ee ice ee ee ae eee Payette a iat s)srclas a ese- 310

Mr. E. Hatscunn. Description and exhibition of investigations into the forms assumed by
drops and vortices of gelatin in various coagulants ...............00. Se cesar ete beens 3H)

Prof. F. Woov-Jonus, F.Z.S. Exhibition of a cast and set of Rontgen-ray photographs
taken from a Chimpanzee which had died from pulmonary tuberculosis .......... 310

PAPERS.

10. Comparison between the Lower Jaws of the Cynodont Reptiles Gomphognathus and
Cynognathus. By Dr. Branisuay Petrontevics. (Text-figures 1-8.) ......6...0. 197

11. On a new Genus of Extinct Muscardine Rodent from the Balearic Islands. By
Dorotnea M. A. Barz, Hon. M.B.O.U. (Plate I. and Text-figures 1 & 2.) ........ 209

12, A Case of Hermaphroditism in a Lizard, Lacerta viridis, By Nor. Tayrer, B.Sc. (Lond.).
(From the Zoological Department, University of London, University College.) (Text-
See eR TT Saale Oh cn cie isle die ah 2 wees cis wn ees Cea g ce ence ee Sele aah sieileto an 223

13. On two new Elasmobranch Fishes (Crossorhinus jurassicus, sp. nov., and Protospinax
annectans, gen. et sp. nov.) from the Upper Jurassic Lithographic Stone of Bavaria.
‘By Artnvr Suita Woopwarp, LL.D., F.R.S., V.P.Z.S. (Plate I.) .............. 231

14. The Function of Pathological States in Evolution. By Moruny Roperts .......... 237

15, Notes on the Leavers at Leonardslee, 1916-1918. By Sir Epmunp G. Lover, Bart.,
ites PESIUCNt ZG ose ns baa wees st ee ce we Ree PRR an Seer ie, URSA ORL iL arabe, oe at O's 255

16, On the Madagascar Frogs of the Genus Mantidactylus Blgr. By G. A. Boutznczr,
Se eee eyes eae tie ote we aicig Wear dole’ a Rieye eee wana Sale die, Ubi wily. Aine bee ae 257

17. Ciliary Action in the Internal Cavities of the Ctenophore Pleurobrachia pileus Fabr.
By Jamus F. Gemuiuy, M.A., M.D., DSe., F.Z.S. (Text-figures 1&2.) .......... 268

18. On Seymouria, the most primitive known reptile. By D. M. 8. Watson, M.Sc., Capt.
_ RA.F., Lecturer in Vertebrate Paizontology, University College, London. (Text-

SO LLESE sO DUS a al TOE eve ace er eee 267
Ee Paces 0 acts nace n'y alah mee apa ik oie Tee PUN Biale s malslee’y je belas ee Sane + « i
SMe t Ooo 6.) os. sae as oe acs beeing sed Debsepeeeiionsvceaycs ii
MMMM te sie cia a aloie s al SS os em tele eS ee Ck WEN we BBS ewe shee weet iii
Meee row MistiG COMPIDMEONS 6 62 eo Se ines wa eb ec guewte a dues st onesies vii

PLATES.

1918, Parts III. & IV. (pp. 197-310).

Page

Plate
«. 209

Miss Bate: I. Hypnomys, gen. nov. et Leithia ..ccccsssocsseees

Woopwarp: I. 1. Crossorhinus jurassicus. 2,3. Protospinar annec-
CONS wigs ae te web 8 oe eae Bie wl wls, paid ce te eeee aot

NOTICE.

The ‘ Proceedings’ for the year are issued in fowr parts, paged consecutively,
so that the complete reference is now P. Z. 8. 1917, p.... The Distribution

is usually as follows, but on account of war conditions Parts I. & II. were
published simultaneously and Parts III. & IV. are similarly issued :—

Part I. issued in March.

| II. a June.
Pee eure. September.
1 oa aes December,

‘ Proceedings,’ 1918, Parts I. & II. (pp. 1-196), were published
together on August 18th, 1918.

The Abstracts of the ‘ Proceedings,’ Nos. 184-189, are
contained in this Part.

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if.

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